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VOL. 107 JANUARY 2005 NO. 1 


“io, PROCEEDINGS 


EE NK 7 of the 


ENTOMOLOGICAL SOCIETY 
of WASHINGTON 


cau ARTE RLY 
20 , 


| JAN 25 


CONTENTS 


BEAULIEU, FREDERIC and TERRY A. WHEELER—Diptera diversity in a homogeneous habitat: 
Brachycera associated with sedge meadows (Cyperaceae: Carex) in Quebec, Canada........ 176 


CAMBRA, ROBERTO A., VICTOR H. GONZALEZ, and WILLIAM T. WCISLO— Description 
of the male, host associations, and new distribution records for Lophostigma 


Gincian(auibaysson))(Eymenoptera-. autillidac):meeere saat a ete aes cheapie eek ees hteee 229 
DeWALT, R. EDWARD and B. D. HEINOLD—Summer emerging Ephemeroptera, Plecoptera, 
and Trichoptera of Abrams Creek, Great Smoky Mountains National Park.................... 34 


EDMISTON, JAMES F. and WAYNE N. MATHIS—A review of two Nearctic species of the 
shore-fly genus Philygria Stenhammar: P debilis Loew and P_ nigrescens (Cresson) 
(Dipte rasa ps ly Cerda ey: Ps 3k yee iro Sits, See taey eS Ol eae dy ces meted ual Hla Ms ACY ae A Be Pe 7 


GATES, M. W., S. N. MYARTSEVA, and M. E. SCHAUFF—A new Baryscapus Forster 
(Hymenoptera: Eulophidae) parasitic on Diorhabda elongata Brullé (Coleoptera: 
Chrysomelidae) and implications for the biological control of saltcedar (Tamaricaceae: 
TOma aa spp:)anithe South Wwestennt Umited States sens 4a nance sense ae oe) en) ae eee 28 


HALL, JASON P. W.—A review of the Metacharis syloes group (Lepidoptera: Riodinidae), with 


the description of two new species from west of the Andes..................0ccceeceeeceeeeees 200 
HARRISON, T. L. and M. R. BERENBAUM—Rutaceae-feeding Agonopterix Hiibner 

(Hepidoptera Plachistidae)) Mm TUiMOlsie ae access oe la ae ee atic ers a oe Basin I nee a a 162 
HINOJOSA-DIAZ, ISMAEL A. and CHARLES D. MICHENER—A new bee of the genus 

Chilicola Spinola (Hymenoptera: Colletidae: Xeromelissinae) from central Mexico.......... ] 


HUSBAND, ROBERT W. and DAVID O. HUSBAND—A new species of Dorsipes Regenfuss 
(Acari: Podapolipidae), ectoparasite of Amara latior Kirby (Coleoptera: Carabidae) 
HE OMIR-ANTEZ OTA Me ott nt al ar satel Ree MRT ay Sis lana «aN cma R ORG I Rte rs erect ckt ae ohs san 71 
KULA, ROBERT R. and GREGORY ZOLNEROWICH—A new species of Epimicta Forster 
(Hymenoptera: Braconidae) from North America and new distribution records for Epimicta 


ST AIPILAST, WVINATLOING PCa ec ete aed ie ee aire RVG ee caralics ta Sho hid ele cist MVE es SUED Haale Oa Me smn 78 
MAWDSLEY, JONATHAN R.—Extirpation of a population of Cicindela patruela 
DeJean (Coleoptera: Carabidae: Cicindelini) in suburban Washington, D.C., USA .......... 64 


(Continued on back cover) 


THE 


ENTOMOLOGICAL SOCIETY 
OF WASHINGTON 


OFFICERS FOR 2005 


JASON P. W. HALL, President Jon A. Lewis, Custodian 
STEVEN W. LINGAFELTER, President-Elect MicHAEL G. PoGuE, Treasurer 
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Hots B. WitLiAMs, Menibership Secretary E. E. GrisseELt, Past President 


Davin R. SmitH, Editor 


Publications Committee 
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Title of Publication: Proceedings of the Entomological Society of Washington. 
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PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 1-6 


A NEW BEE OF THE GENUS CHILICOLA SPINOLA (HYMENOPTERA: 
COLLETIDAE: XEROMELISSINAE) FROM CENTRAL MEXICO 


ISMAEL A. HINOJOSA-DiAZ AND CHARLES D. MICHENER 


Entomology Division, Natural History Museum and Biodiversity Research Center and 
Entomology Program, Department of Ecology and Evolutionary Biology, University of 
Kansas, Lawrence, Kansas 66045, USA. (e-mail: hinojosa@ku.edu; michener@ku.edu) 


Abstract.—Chilicola (Hylaeosoma) yanezae, new species, the second known Mexican 
species of the group of Chilicola megalostigma Ducke, is described from the mountains 
in the State of Morelos in central Mexico. It differs from C. polita Michener in its smaller 
size, yellow-testaceous areas on the thorax (only the prothorax of the male), and characters 
of the hidden sterna of the male. 


Resumen.—Chilicola (Hylaeosoma) yanezae, especie nueva, es la segunda especie 
mexicana del grupo de Chilicola megalostigma Ducke, proveniente de las montafias del 
estado de Morelos en el centro de México. Difiere de C. polita Michener en su menor 
tamano, torax (solo prot6rax en el macho) con areas de color amarillo 0 marr6n claro y 


en caracteres de los esternos ocultos del macho. 


Key Words: 


The Mexican and Central American spe- 
cies of the principally South American ge- 
nus Chilicola Spinola were reviewed by 
Michener (1994). Five species are included 
in that paper, all of them found in Mexico. 
Of these, four are members of the subgenus 
Hylaeosoma (characterized by Michener 
1994, 1995, 2000) and one of those, C. pol- 
ita Michener, was segregated as a member 
of group B, which was called by Brooks 
and Michener (1999) the group of C. me- 
galostigma (Ducke). This group differs 
from all other Chilicola by the smooth, 
shining body with insignificant microsculp- 
turing between widely spaced punctures on 
many parts of the body; the strong, lamel- 
late preoccipital carina; the elongate thorax 
with the dorsal surface of the pronotum 
about twice as long as the flagellar diame- 
ter; and the apical processes on the first two 


bees, Colletidae, Xeromelissinae, Chilicola, taxonomy, Mexico 


front tarsal segments of the female (illus- 
trated by Brooks and Michener 1999). This 
distinctive group contains three previously 
known species, C. megalostigma (Ducke) 
and stenocephala Brooks & Michener from 
South America and C. polita which, al- 
though rare, ranges from Panama to Ta- 
maulipas, Mexico. We here describe anoth- 
er species of the C. megalostigma group; it 
is from the mountains of Morelos, Mexico, 
and is a close relative of C. polita (although 
C. polita and the Brazilian C. megalostigma 
are even more similar). 

Abbreviations used in the description are: 
S, sternum; T, tergum; E flagellar segment 
or flagellomere; OD, ocellar diameter. 

Measurements of the holotype are fol- 
lowed in parentheses by those of the two 
male paratypes (except body length, due to 
the dissection of one paratype). For the fe- 


No 


Fig. 1. 
(Hylaeosoma) yanezae. 


Lateral view of holotype male of Chilicola 


male measurements of both paratypes are 
provided in parentheses. 

The term imbricate, to describe some mi- 
crosculpturing, is applied following Harris 
(ID72), is 32), 

Details and ecological aspects of the type 
locality were described by Hinojosa-Diaz 
(2003). 


Chilicola (Hylaeosoma) yanezae, 
new species 
(Figs. 1-8) 


Chilicola (Hylaeosoma) aff. polita: Hino- 
josa-Diaz 2003: 12; Yanez-Ordonez and 
Hinojosa-Diaz 2004: 179. 


Diagnosis.—Resembles other species of 
the C. megalostigma group in elongate head 
and body and shining integument with little 
and weak microsculpture. Differs from oth- 
er Chilicola in light brown to yellow color 
of most of pronotum of male and of entire 
mesosoma except nearly black scutum, scu- 
tellum and metanotum of female. Most sim- 
ilar to C. polita Michener but smaller, with 
pale mesosomal coloration described above, 
with distal thickened apical process of S8 
of the male, and rounded rather than an- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


gular medial thickenings of S7 of the male 
(Figs. 4—7). 

Description.—Male: Body length 5.1 
(5.1) mm. Coloration: Brownish_ black, 
face black, except for brown malar area, 
distal border of clypeus, labrum, mandible, 
dorsal border of preoccipital carina; anten- 
nal scape and pedicel yellow amber, F1 to 
F4 light brown or brown, remainder of fla- 
gellum brownish black above, brown be- 
low, except last segments entirely blackish; 
mesosoma very dark brown, except central 
area of pronotal lobe and tegula transparent 
amber, dorsal area of pronotum light brown, 
paler laterally and continuing yellow brown 
to pronotal lobe and posterior third of upper 
lateral area of pronotum; fore and mid legs 
yellow amber, with brown to dusky areas 
on mid femur and mid tibia; apices of pre- 
tarsal claws and arolia brown; posterior leg 
mainly brown with yellow brown on parts 
of coxa, trochanter, apex of femur, base and 
apex of tibia; metasomal terga dark brown, 
except anterior halves of T2 and T3, ante- 
rior third of T4 yellow, posterior margins 
of Tl to T3 yellow brown; S1, S2 trans- 
parent dusky amber, S3 yellow amber, S4 
to S6 brown; wings transparent with green 
and red highlights, veins and stigma black- 
ish brown. 

Body surface: Polished, without micros- 
culpturing except as indicated below; face 
with punctures scattered, denser along mid 
surface between subantenal sulcus and 
compound eye, as well as on the frons, 
where separated by less than a puncture 
width; central part of supraclypeal area im- 
punctate; gena and hypostomal area largely 
impunctate; mesosoma polished, shiny, 
with punctures separated by | to 2 puncture 
widths on mesoscutum and by 2 to 3 punc- 
ture widths on dorsal area of pronotum, 
scutellum and metanotum; anteriormost 
mid surface of mesoscutum minutely im- 
bricate, microsculpturing otherwise absent; 
mesepisternum largely impunctate; anterior 
lateral surface of pronotum, metepisternum, 
lateral and posterior surfaces of propodeum 
minutely imbricate or lineolate; basal area 


VOLUME 107, NUMBER 1 


of propodeum with about 12 longitudinal 
striae slightly radiating posteriorly; meta- 
somal terga and sterna shiny, each anteri- 
orly transversely lineolate, becoming pol- 
ished, impunctate posteriorly, particularly 
on first three metasomal segments. 

Pubescence: Hairs minute, sparse, lon- 
gest on hind tibiae (about 2 OD) and pos- 
terior half of metasoma (up to nearly 4 
OD). Face with sparse, simple, short, erect, 
white hairs, longer on clypeus, shorter on 
frons, minutely branched hairs on edge of 
preoccipital carina, appressed, more notice- 
ably branched hairs on alveolocular area 
adjacent to antennal socket; gena with more 
abundant, longer, minutely branched hairs, 
especially on posterior half; hypostomal 
area with sparse, short hairs; pronotum with 
sparse, short hairs, except pronotal lobe 
margin and lateral-anterior and posteror 
borders of pronotum with appressed plu- 
mose hairs; mesoscutum and scutellum with 
sparse, simple, short, erect, white hairs; me- 
tanotum with long, erect hairs on sides, be- 
coming shorter and converging mesally; an- 
terior and lower surface of mesepisternum 
with appressed or erect plumose hairs; me- 
tepisternum with dense, long, branched 
hairs; lateral surface of propodeum with 
short, simple, suberect hairs, longer and 
branched on upper border, basal area hair- 
less; terga and sterna with sparse, simple, 
erect hairs, longer on ventral side and on 
discs of T4 to TO. 

Structure: Head elongate, formed about 
as in C. polita (Brooks and Michener 1999, 
figs.7,8), length 1.25 (1.26,1.26) mm, width 
0.85 (0.87, 0.87) mm, eyes convergent be- 
low, upper interorbital distance 0.55 (0.54, 
0.57) mm, lower interorbital distance 0.28 
(0.30, 0.30) mm, paraocular area with well- 
developed depression for antennal scape ex- 
tending to upper ocular tangent; inner bor- 
der of depression above antennal socket 
with ovoid protuberance slightly smaller 
than 1 OD [equivalent to elongate welt de- 
scribed for C. stenocephala (Brooks and 
Michener 1999); this protuberance scarcely 
evident in male C. polita]; Fl shorter than 


oS) 


pedicel, about twice as long as broad, F2 to 
F4 broader than long, F5 as long as broad, 
remaining segments longer than broad, F1 1 
over twice as long as broad; malar area over 
twice as broad as long (three times in one 
paratype); ocelloccipital distance about 1.5 
OD measured to apex of high preoccipital 
carina; genal area above about as wide as 
eye seen from side, tapering below. Max- 
illary palpus longer than prementum, about 
as long as head, first two segments about 
half length of third, segment 6 slender. Dor- 
sal surface of pronotum continuing curva- 
ture of mesoscutum as seen in profile but 
lower than mid mesoscutum which is nearly 
four times as long as mid-dorsal pronotum. 
Legs slender (Fig. 1). Forewing length 3.4 
(3.4, 3.6) mm; apex of marginal cell mi- 
nutely truncate. Tl longer than broad, 
length 0.78 (0.78, 0.84) mm, width 0.56 
(0.57, 0.57) mm, in profile T2 and T3 con- 
stricted basally; T7 with apex truncate; pos- 
terior half of S6 mesally with flat, shining 
triangular surface surrounded by hairy areas 
posterolaterally with small tuft of erect, 
long, curved hairs (nearly 4 OD). S7, S8 
and genitalia as in Figs. 2 to 7; S7 with 
prominent membranous medial projections, 
folding towards ventral surface; genitalia 
with gonoforceps with apex broadly round- 
ed-truncate and produced inward. 

Female: As described for male except as 
follows: Body length (4.68, 5.16) mm. All 
flagellar segments dusky brown; mesoscu- 
tum, scutellum and metanotum dark brown, 
rest of mesosoma yellow except central area 
of pronotal lobe and tegula transparent am- 
ber, striae of basal area of propodeum 
brown; mid leg mainly light brown; hind 
coxa and trochanter yellow; S1 transparent 
amber, S2 to basal half of S4 yellow amber, 
rest of sterna brown amber. Scopa well de- 
veloped on SI to S3, plumose hairs denser 
and longer on S2. Head length (1.25, 1.32) 
mm, width (0.94, 0.96) mm, upper inter- 
orbital distance (0.60, 0.60) mm, lower in- 
terorbital distance (0.28, 0.30) mm. Protu- 
berance on inner border of depression for 
absent. Forewing length 


antennal scape 


4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 2-7. Male of Chilicola (Hylaeosoma) yanezae. 2-3, Genitalia, dorsal and ventral views. 4—5, S7, dorsal 
and ventral views. 6—7, S8, dorsal and ventral views. 


VOLUME 107, NUMBER 1 


0.5 mm 


Inner view of anterior tarsus of female of Chilicola (Hylaeosoma) yanezae. 


Fig. 8. 


(3.60, 3.70) mm. Anterior tarsus bristly, tar- 
someres 1 and 2 each with apical process 
and single curved hair at tip (Fig. 8). T1 
length (0.84, 0.85) mm, Tl width (0.67, 
0.69) mm. Apex of metasoma unmodified. 

Type material (all from Mexico).—Ho- 
lotype male: Km 4.5 Autopista México- 
Cuautla, Tepoztlan, Morelos. 28/04/1996 
[28 April 1996], O. Yanez OY-602, 1,940 
m [meters above sea level] 19°00’15”[N], 
99°07'46"[W], Bosque de encino // OY-602/ 
/ Museo de Zoologia, Hymenoptera, 23868 
[Catalog number] // Chilicola sp. 3 3, Det. 
T. Griswold 1998, in the Museo de Zoolo- 
gia “‘Alfonso L. Herrera’’, Departamento de 
Biologia Evolutiva, Facultad de Ciencias, 
Universidad Nacional Autonoma de Méxi- 
co. Paratypes: 2 6, 2 2, same locality as 
holotype, all 17/05/1997 [17 May 1997], I. 
Hinojosa collector. Two paratypes were col- 
lected while in copula. One female paratype 
on flowers of Sida rhombifolia L. (Malva- 
ceae), the other three paratypes on flowers 
of Tithonia tubaeformis (Jacq.) Cass. (As- 
teraceae). One female paratype in the same 
institution as holotype. A male paratype in 
the Division of Entomology, Natural His- 
tory Museum and Biodiversity Research 
Center, University of Kansas. The two other 
paratypes (one of each sex) in the U. S. 
National Pollinating Insects Collection, 
USDA Bee Biology & Systematics Labo- 
ratory, Utah State University, Logan Utah. 

Etymology.—Named in honor of Olivia 
Yanez of the Universidad Nacional Aut6n- 


oma de México, who collected the holo- 
type, and contributed toward preparation of 
this paper. 


ACKNOWLEDGMENTS 


For the loan of the specimens used in this 
description we thank the personnel of the 
Museo de Zoologia ““Alfonso L. Herrera,” 
Departamento de Biologia Evolutiva, Fa- 
cultad de Ciencias, Universidad Nacional 
Autonoma de México. Additional speci- 
mens, including a male of C. polita, were 
provided by Terry L. Griswold of the U. S. 
National Pollinating Insects Collection, 
USDA Bee Biology & Systematics Labo- 
ratory, Utah State University, Logan Utah, 
who first identified this species as new. Mi- 
chael S. Engel assisted with the photograph 
of the holotype. 

This is contribution number 3356 of the 
Division of Entomology, Natural History 
Museum and Biodiversity Research Center, 
University of Kansas. 


LITERATURE CITED 


Brooks, R. W. and C. D. Michener. 1999. The Chili- 
cola megalostigma species group and notes on 
two lost types of Chilicola (Hymenoptera: Colle- 
tidae, Xeromelissinae). Journal of Hymenoptera 
Research 8: 132-138. 

Harris, R. A. 
Occasional Papers in Entomology, California 
State Department of Food and Agriculture no. 28: 
1-31. 

Hinojosa-Diaz, I. 


1979. A glossary of surface sculpturing. 


A. 2003. Abejas silvestres (Hyme- 
noptera: Apoidea) del declive sur de la Sierra del 


6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Chichinautzin, Morelos, México. Folia Entomo- 
l6gica Mexicana 42: 1—20. 

Michener, C. D. 1994. Mexican and Central American 
species of Chilicola (Hymenoptera: Colletidae). 
Folia Entomol6gica Mexicana 85(1992): 77-93. 

. 1995. A classification of the bees of the sub- 

family Xeromelissinae (Hymenoptera: Colletidae). 

Journal of the Kansas Entomological Society 68: 

332-345. 


. 2000. The Bees of the World. Johns Hopkins 
University Press, Baltimore, xv + 913 pp. 

Yanez-Ordonez, O. and I. Hinojosa-Diaz. 2004. La co- 
lecci6n himenopterolégica (Insecta) del Museo de 
Zoologia ‘Alfonso L. Herrera” de la Facultad de 
Ciencias, UNAM, México. Acta Zool6gica Mex- 
icana (n.s.) 20(1): 167-197. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 7-20 


A REVIEW OF TWO NEARCTIC SPECIES OF THE SHORE-FLY GENUS 
PHILYGRIA STENHAMMAR: P. DEBILIS LOEW AND P. NIGRESCENS 
(CRESSON) (DIPTERA: EPHYDRIDAE) 


JAMES E EDMISTON AND WAYNE N. MATHIS 


(JFE) Franciscan Project for Russia and Kazakhstan, Novosibirsk, Russia (email: 
edmisja@att.net); (WNM) Department of Entomology, MRC 169, PO. BOX 37012, 
Smithsonian Institution, Washington, D.C. 20013-7012, United States (email: 
mathis.wayne @nmnh.:si.edu) 


Abstract.—Two species of the shore-fly genus Philygria Stenhammar, P. debilis Loew 
and P. nigrescens (Cresson), are reviewed to reassess their taxonomic status, which was 
recently questioned by Hollmann-Schirrmacher. Both species are distinct, and evidence is 
presented to support their revised status. For perspective and to facilitate their identifi- 
cation, the appropriate subfamily, tribe, genus, and species are diagnosed and keys to the 
genera of the tribe Hyadinini and to the Nearctic species of Philygria are presented. 


Lectotypes, as appropriate, are designated. 


Key Words: 


review, Diptera, Ephydridae, [lytheinae, Hyadinini, shore flies, Philygria 


debilis, P. nigrescens, lectotype designations 


This paper is an analysis of the taxonom- 
ic status of two Nearctic shore-fly species: 
Philygria debilis Loew and P. nigrescens 
(Cresson). Philygria was recently revised 
on a world basis (Hollmann-Schirrmacher 
1998), and among the 39 species that were 
included, four (P. debilis, P. dimidiata 
(Sturtevant and Wheeler), P. longicornis 
(Sturtevant and Wheeler), and P. puncta- 
tonervosa (Fallén)) occur in the Nearctic 
Region. Hollmann-Schirrmacher (1998) 
also included Philygria picta (Fallén) in his 
revision as a fifth Nearctic species, but our 
study of this species suggests that it be in- 
cluded in Nostima Coquillett, a genus we 
consider to be separate from Philygria (Ed- 
miston and Mathis 2004). 

Most Nearctic species of Philygria are 
relatively uncommon in collections and per- 
haps in nature. For example, in the USNM 
collection, Nearctic species of Philygria are 
represented with the following numbers of 


specimens: P. dimidiata (1), P. longicornis 
(1), P. nigrescens (17), P. punctatonervosa 
(96), and P. debilis (over 1,000). The ex- 
ceptional number of P. debilis in research 
collections supports other observations that 
this species is one of the most common 
shore flies in temperate North America, 
probably occurring in most urban lawns 
(Cresson 1944, Scheiring and Foote 1973, 
Zack 1998). Although common and some- 
times abundant, the taxonomic status of this 
species and P. nigrescens is unclear and 
was recently questioned (Hollmann-Schi- 
rrmacher 1998). Hollmann-Schirrmacher 
(1998) considered both species to be con- 
specific and their names to be senior and 
junior synonyms respectively. 

Cresson (1930b) described Hydrina_ ni- 
grescens from specimens collected in Lon- 
don Hill Mine near Bear Lake, British Co- 
lumbia. He wrote (1930b: 80) that his new 


species was “*... very similar to Philygria 


8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


debilis Loew, but more blackish, not so 
brownish; frons much longer, about six- 
tenths as long as broad; cheeks broader; ab- 
domen mostly shining; at most the second 
costal section one and one-half as long as 
third.”” Wirth and Stone (1956) subsequent- 
ly resolved the generic nomenclature of this 
species when they transferred some species 
of the genus Hydrina Robineau-Desvoidy, 
including N. nigrescens, into Philygria. 

In his revision of Philygria, Hollmann- 
Schirrmacher (1998: 71) wrote that the 
characters Cresson used to describe P. ni- 
grescens (body color, genal size, radial vein 
indices) were “‘within the variability” of 
Philygria debilis, and noted further that he 
could not detect any differences in struc- 
tures of the male terminalia. In his “‘Dis- 
cussion”’ section to that species, he con- 
cluded and wrote that (1998: 71): ““The 
name Philygria nigrescens is therefore a ju- 
nior synonym to P. debilis.” Evidently, 
Hollmann-Schirrmacher did examine the 
holotype of P. nigrescens, as the label data 
from the type locality were cited, but there 
is no indication that any other specimens 
were examined. For example, the label data 
from Alaskan specimens in the USNM 
were not included in his revision. He did 
write, however, that a darker variety of 
Philygria debilis did occur in the northern 
locations of British Columbia and Alaska 
(Hollmann-Schirrmacher 1998). 

In August of 2002 and 2003, the second 
author collected Philygria in the Matanus- 
ka-Susitna Borough of Alaska, in or near 
the village of Talkeetna. Some specimens 
appeared to be distinct from the commonly 
collected P. debilis, and that observation 
prompted this review of P. debilis and P. 
nigrescens. As part of our study, the perti- 
nent primary types were examined, struc- 
tures of adults, including those of the male 
terminalia, were studied and described, new 
specimens were examined, and consider- 
able evidence was found to distinguish be- 
tween P. debilis and P. nigrescens. To pro- 
vide a broader context for this paper and 
also to facilitate the identification of Phil- 


ygria and its included Nearctic species, we 
have provided brief diagnoses for the sub- 
family Ilytheinae and tribe Hyadinini and 
keys to the genera of Ilytheinae and Nearc- 
tic species of Philygria. 

Unlike most shore flies, adults of Phily- 
gria, including the two species that are be- 
ing reviewed here, occur more commonly 
in somewhat dry, grassy habitats that are 
not immediately associated with aquatic 
habitats. Zack (1979:85) collected many P. 
debilis from the flowers of Anaphalis mar- 
garitacea (L.) Bentham and Hook (Astera- 
ceae) in a habitat he described as being lo- 
cated at “‘considerable distances from per- 
manent water sources.” The larvae of Phil- 
yeria are closely linked with habitats where 
blue-green algae are abundant, and small 
patches of blue-green algae can almost al- 
ways be found growing on the soil’s surface 
in grassy habitats. Larvae feeding on blue- 
green algae are typical for many taxa in the 
subfamily Ilytheinae (Foote 1995). 


MATERIALS AND METHODS 


The descriptive terminology, with the ex- 
ceptions noted in Mathis (1986), and Math- 
is and Zatwarnicki (1990a), and below, fol- 
lows that published in the Manual of Ne- 
arctic Diptera (McAlpine 1981). We have 
followed the terminology for most struc- 
tures of the male genitalia that other work- 
ers in Ephydridae have used (see references 
in Mathis 1986, Mathis and Zatwarnicki 
1990a, 1990b), such as surstylus. Zatwar- 
nicki (1996) has suggested that the pre- and 
postsurstylus correspond with the pre- and 
postgonostylus and that the subepandrial 
plate is the same as the medandrium. Al- 
ternative spellings for some localities are 
cited in parentheses, especially for locality 
names that were transliterated into English. 
The species descriptions are composite and 
not based solely on the holotypes. 

Label data from each specimen were re- 
corded and are listed alphabetically accord- 
ing to country, state or province, county, 
and specific locality, such as city. As avail- 
able, dates of collection, collector, sex, and 


VOLUME 107, NUMBER 1 


specimen deposition are listed. Label data 
from primary type specimens are recorded 
verbatim, and clarifying information, such 
as script style and label color, is enclosed 
within brackets. 

Although most specimens are in the Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 
(USNM), we also studied numerous speci- 
mens from the following collections: Acad- 
emy of Natural Sciences of Philadelphia 
(ANSP), Philadelphia, Pennsylvania, and 
the Museum of Comparative Zoology 
(MCZ), Harvard University, Cambridge, 
Massachusetts. 

Distribution maps were made using ESRI 
ArcView® GIS 3.2. Longitude and latitude 
coordinates were obtained for the locality 
where each specimen was collected. If 
available, the longitude and latitude were 
obtained directly from the specimen labels. 
For specimen labels that did not have lon- 
gitude and latitude, gazetteers and maps 
were used to determine the geographical 
coordinates. The geographic coordinates 
were entered into ESRI ArcView tables. 
The specimen locales were plotted on a 
world land projection, presented within 
ESRI ArcView layouts and exported as en- 
capsulated postscript (EPS) files. 

External morphological structures were 
observed and recorded using a dissecting 
microscope. Continuous characters were 
measured using a calibrated ocular micro- 
meter attached to either a compound or dis- 
secting microscope. Quantitative characters 
used commonly in the descriptions are de- 
fined as: 


1. Body length: maximum distance in lat- 
eral view from anterior margin of head 
to posterior abdominal apex. 

2. Scutal length: maximum straight-line 
distance in dorsal view from the anterior 
margin of the scutum to the scutum-scu- 
tellar suture. 

3. Scutellar length: maximum straight-line 
distance in dorsal view from the scutel- 


lar suture to the posterior margin of the 
scutellum. 

4. Costal vein ratio: straight-line distance 
between the apices of veins R,., and 
R,,;/distance between the apices of 
veins R, and R,,;. 

5. M vein ratio: straight-line distance along 
vein M between crossvein dm-cu and r- 
m/distance apicad of crossvein dm-cu. 


Dissections of male and female genitalia 
and descriptions were performed using the 
method of Clausen and Cook (1971) and 
Grimaldi (1987). Microforceps were used 
to remove abdomens, which were macer- 
ated in a potassium hydroxide solution. 
Cleared terminalia were rinsed in distilled 
water and 70% ethanol and then transferred 
to glycerin for observation. If necessary for 
proper orientation, the specimen was trans- 
ferred from glycerin to glycerine jelly. The 
glycerin jelly was heated, and the specimen 
appropriately oriented. After cooling, the 
embedded specimen in glycerin jelly be- 
came immobilized. Abdomens were placed 
in an attached plastic microvial filled with 
glycerin and attached to the pin supporting 
the remainder of the insect from which it 
was removed. 

External morphological characters were 
drawn using an ocular grid attached to a 
stereoscopic dissecting microscope. Internal 
genitalic features were drawn using a cam- 
era lucida with a Wild M-400 compound 
microscope and corroborated with views 
from a Nikon SMZ-1500 dissecting micro- 
scope. 


SYSTEMATICS 
Subfamily Ilytheinae Cresson 


Ilytheinae Cresson 1943: 2 (as the tribe II- 
ytheini). Type genus: //ythea Haliday in 
Curtis 1837.—Zatwarnicki 1992: 89 [di- 
agnosis, classification, phylogeny ].— 
Mathis and Zatwarnicki 1995: 186—214 
[world catalog].—Hollmann-Schirrmach- 
er 1998: 17—44 [classification, phyloge- 
ny]. 


10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Diagnosis.—Adult. Head: Fronto-orbital 
setae proclinate and reclinate if present, fre- 
quently one or both setae absent. Face pro- 
truding with large oral opening (Ilytheini) 
or moderately protruding, often carinate 
(Hyadinini). Spinelike seta on pedicel in- 
conspicuous. 

Thorax: Insertion of posterior notopleur- 
al seta moderately to conspicuously elevat- 
ed, especially compared to anterior seta; 
presutural (sometimes sutural) dorsocentral 
setae present or absent. 

Abdomen: Male terminalia: Surstylus 
usually fused with ventral margin of epan- 
drium, often indistinguishably; subepandri- 
al plate usually fused with hypandrium and/ 
or gonites, sometimes reduced to a remnant 
or forming a gonal arch; phallapodeme tri- 
angular in lateral view; ejaculatory apode- 
me present or secondarily absent; pre- and 
postgonite fused into a single gonite. Fe- 
male genitalia: Extending process of female 
ventral receptacle sinuous, lacking an oper- 
culum. 

Larva.—Anterior spiracle divided into 2 
basal, elongate branches. 


KEY TO GENERA OF ILYTHEINAE CRESSON 


1. Posterior notopleural seta inserted near ven- 
tral margin of notopleuron and at about same 


Well AS AiMSMOr SHA socs0cccceccocnce 2 
— Posterior notopleural seta inserted at conspic- 
uously higher level than anterior seta .... 10 


N 


Vein R,,, short, costal section II about % 
length of II, and with a stump vein. A single, 
proclinate, fronto-orbital seta 
AL Ltd ovens net Nacioe et Uy es Parydroptera Collin 
— Vein R,,, long, costal section II at least half 
section III, lacking a stump vein. Usually a 
reclinate and proclinate fronto-orbital seta .. 3 
3. Costa short, extended at most to slightly be- 
yond vein R,,;. Tergite 4 at least 3x length 
OF TSTMS S) oieolc ces 06 oo 6 Axysta Haliday 
— Costa long, extended to vein M. Tergite 4 at 


most twice length of tergite5 .......... 4 
4. Wing with vein R,,, long; costal II section 
nearly 3X length of III. Face flat or weakly 
carinate, not prominent medially. Flagello- 
mere 1 rounded at apex above .......... 5 
— Wing with vein R,,, short; costal section II 
less than twice length of III. Face with low 
conical medial prominence. Flagellomere 1 
usually angulate at apex above .......... 8 


1 


. Dorsocentral setae 


Both inner and outer vertical setae well de- 
veloped; fronto-orbital setae usually moder- 
ately well to well developed, lateroclinate 
(EWEN) ssscacccreaeso0. Pelina Haliday 
Only inner vertical seta present, outer seta 
lacking; lacking well-developed fronto-orbital 
setae (Neotropical; genus Pelinoides Cresson) 


Femora and palpus yellow to yellowish red; 
arista with dorsal rays, length of rays sube- 
qual to basal aristal width; eye height about 
twice genal height ....... the pallipes group 
Femora and palpus black or dark colored; 
arista mostly bare, if short hairs present, these 
shorter than basal aristal width; eye height 
only slightly longer than genal height, some- 
times smaller 
Tergite 4 bare, shiny, enlarged, length sube- 
qual to combined length of 2nd and 3rd ter- 
gites the cyclocerus group 
Tergite 4 microtomentose, appearing dull, at 
most subshiny, length only slightly longer 
than 3rd the sulcatus group 
Tergite 4 1.3—2 length of tergite 5, both con- 
spicuously punctate. Inner vertical seta pre- 
sent, outer vertical seta absent. Lateral margin 
of scutellum not densely microtomentose, not 
appearing velvety Lytogaster Becker 
Tergite 4 subequal in length to tergite 5, nei- 
ther conspicuously punctate. Usually both 
vertical setae present, if outer absent then lat- 
eral margin of scutellum densely microtomen- 
toseappeannomvelVctyes ma seen eee 9 
Dorsocentral seta 1. Lacking well-developed 
fronto-orbital setae Hyadina Haliday 
Dorsocentral setae 2. One well-developed 
fONtO-OLrbitalisetauen a eeneien enn een 
Parahyadina Tonnoir and Malloch 


. Fronto-orbital setae either lateroclinate and 


inconspicuous or lacking; prescutellar acros- 


tichallgsetacglackan' Cieem relia iia enna 11 
Fronto-orbital setae conspicuous, well devel- 
oped, mostly reclinate or proclinate or both; 
prescutellar acrostichal setae present, well de- 
veloped (Ilytheini Cresson) ........... 13 


Outer vertical seta lacking; fronto-orbital se- 
tae lacking Garifuna Mathis 
Both inner and outer vertical setae usually 
present; fronto-orbital seta present, sometimes 
NECUCEO! Cree. cena, © Rubee carat Nee eae 12 


. Arista bare or minutely branched; 2 rows of 


facial setae. Presutural or sutural dorsocentral 
seta present Philygria Stenhammar 
Arista with short to long branches; 1 row of 
facial setae. Presutural or sutural dorsocentral 
seta lacking Nostima Coquillett 
Donaceus Cresson 
bee te Re: 14 


(14+2) 
Dorsocentral setae 2 (1+1) 


WwW 


VOLUME 107, NUMBER 1 


14. Vein R,,; long, subparallel to C:; costal sec- 
tion II over twice section III .. Jlythea Haliday 

— Vein R,,,; short, running almost straight to C: 
costal section II subequal to section III 

Zeros Cresson 


Tribe Hyadinini Phillips et al. 


Hydrinini Cresson 1944: 175. Type genus: 
Hydrina of authors, not Robineau-Des- 
voidy 1830 ( = Philygria Stenhammar 
1844), unavailable, based on a junior 
homonym. 

Hyadinini Phillips et al. in Cresson 1949: 
251. Type genus: Hyadina Haliday in 
Curtis, 1837.—Hollmann-Schirrmacher 
1998: 29, 45-56 [discussion]. 

Philygriini [nomen nudum].—Wirth and 
Stone 1956: 469.—Wirth 1965: 745 [Ne- 
arctic catalog]; 1968: 16 [Neotropical 
catalog].—Cogan and Wirth 1977: 335 
[Oriental catalog].—Cogan 1980: 666 
[Afrotropical catalog]; 1984: 149 [Pale- 
arctic catalog]. 

Philygriini Lizarralde de Grosso 1989: 51. 
Type genus: Philygria Stenhammar 
1844.—Mathis and Zatwarnicki 1995: 
190—200 [world catalog].—Hollmann- 
Schirrmacher 1998: 50 [synonymy with 
Hyadinini]. 


Diagnosis.—This tribe is similar to Ilyth- 
eini but is distinguished by the following 
combination of characters: Aristal branches 
usually reduced; acrostichal setae reduced, 
prescutellar pair lacking; dorsocentral setae 
frequently reduced in size and/or number 
(1+2, 0+2, or sutural+1); subepandrial 
plate fused with gonite dorsally, forming a 
rounded projection, sometimes joined me- 
dially over aedeagus to form a gonal arch; 
gonites and hypandrium usually fused; and 
gonite produced posteriorly as a long tri- 
angular to almost parallel sided projection, 
with apex variously modified. 


Genus Philygria Stenhammar 


Philygria Stenhammar 1844: 154 (as “‘Sec- 
tio 4” of Notiphila). Type species: Noti- 
phila flavipes Fallén 1823, by subsequent 
designation (Coquillett 1910: 588).— 


11 


Loew 1860: 24 [generic status].—Froese 
1993: 89-96 [immature stages]. 

Cressoniella Saether 1970: 107. Type spe- 
cies: Cressoniella montana Saether 1970 
( = Philygria debilis Loew 1861), by 
monotypy [preoccupied, Mitchell 1934 
(Hymenoptera)].—Mathis and Wirth 
1977: 520 [synonymy]. 

Hydrina of authors, not Robineau-Desvoidy 
1830 [misidentification]: Cresson 1930a: 
93-100 [review]; 1944: 177 [review of 
Nearctic fauna].—Sturtevant and Wheel- 
er 1954: 237-239 [review of Nearctic 
fauna]. 

Pseudohyadina Clausen 1983: 224. Type 
species: Hyadina longicornis Sturtevant 
and Wheeler 1954, by original designa- 
tion.—Hollmann-Schirrmacher 1998: 58 
[synonymy]. 

Philhygria, error for Philygria. 


Description.—Minute to small shore 
flies, body length 0.75—2.20 mm. 

Head: Frons wider than high; ocellar se- 
tae well developed, | pair, divergent, pro- 
clinate; pseudopostocellar setae minute, di- 
vergent, proclinate; both inner and outer 
vertical setae usually well developed, outer 
seta shorter than inner seta, rarely absent: 
fronto-orbital setae small, 3—5, proclinate. 
Antenna with segments often darker on dor- 
sum; arista appearing essentially bare, bear- 
ing numerous, minute, dorsal branches, 
length of branches less than basal aristal 
width. Face generally paler than frons; fa- 
cial setae small, in 2 rows on shiny strip 
medially along parafacial, lateral row later- 
oclinate and % length of medial row, medial 
row medioclinate; dorsal facial seta longest, 
setae ventrad decreasing in length; genal 
seta minute, numerous smaller setulae. Eye 
round to oval. Palpus prominent, varying 
from yellow to dark brown. 

Thorax: Scutellum trapezoidal, with pos- 
terior apex slightly rounded; chaetotaxy as 
follows: Dorsocentral setae 3 (1+2), ante- 
rior seta smaller than posterior setae, some- 
times sutural, with 3—5 presutural setulae; 
acrostichal setae in | row; postsutural su- 


12 


pra-alar seta 1; lateral scutellar setae usually 
2, rarely lateral seta absent, lateral seta 
shorter than apical seta; notopleural setae 2, 
posterior seta longer and inserted at posi- 
tion above level of anterior seta; anepister- 
nal seta 1, small, inserted along posterior 
margin, with many dorsal setulae; katepis- 
ternal seta |. Wing usually mostly hyaline 
to faintly infuscate, sometimes with distinct 
maculation pattern; crossveins sometimes 
white or black; generally lacking spurious 
veins. Legs yellow to dark brown; femora 
and tibiae often banded; tarsi usually yel- 
lowish, apical |—2 tarsomeres darker. Halter 
whitish yellow to brownish yellow. 

Abdomen: Male and female tergites with 
many setae and setulae. Male terminalia as 
follows: epandrium reduced, often absent; 
cerci present, sometimes fused to epan- 
drium; epandrium and surstylus fused, bear- 
ing row of large setulae; lacking surstylar 
projections; phallapodeme triangular in lat- 
eral view; aedeagus variously sclerotized, 
often divided into basiphallus, mediophal- 
lus, and distiphallus with various projec- 
tions; pre- and postgonites distinct. 

Discussion.—Hollmann-Schirrmacher 
(1998) recently treated the genus Nostima 
as a junior synonym of Philygria, which he 
then divided into four species groups. Holl- 
mann-Schirrmacher (1998) recognized a 
broader concept of Philygria and diagnosed 
its species groups almost exclusively by 
structures of the male terminalia. He further 
proposed the synonymy of Nostima with 
Philygria according to a hypothetical de- 
velopment of setae and projections on the 
gonite-hypandrium complex. Emphasis 
solely on genital characters, however, is 
problematic because divergence of external 
characters in extant species does not always 
correlate well with divergence of genitalic 
characters. Furthermore, his phylogenetic 
consideration of Nostima was limited to 
two exemplars, N. picta (Fallén) and WN. 
flavitarsis Canzoneri and Meneghini, and 
he did not consider updates after 1993, such 
as description of the closely related genus 
Garifuna (Mathis 1997). 


2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Our analysis, which included a wider 
sampling of characters and species from 
Philygria, Nostima, and Garifuna, resulted 
in a classification that only partially adheres 
to Hollmann-Schirrmacher’s concept of 
Philygria (we exclude his picta and flavi- 
tarsis groups), and we recognize Nostima 
and Garifuna as separate genera (Edmiston 
and Mathis 2004). 


KEY TO NORTH AMERICAN SPECIES OF 
PHILYGRIA STENHAMMAR 


1. Wing hyaline, except for spot over cross veins 


CN LEN ne eT eS me A EOS SIN 4 
= Wineyspotted! ais cas ac cee te eee) omen: a) 
2. Wing with spurious vein(s) ............. 3 


— Wing without spurious veins 
BR a Sree P. longicornis (Sturtevant and Wheeler) 
3. Single spurious vein projected from vein CuA, 
MS St P. dimidiata (Sturtevant and Wheeler) 
— Many spurious veins projected from veins R,,, 
and R,.; P. punctatonervosa (Fallén) 
4. Abdominal tergites 1-3 with dense microto- 
mentum. Parafacial area yellowish brown 
sues 2, carats Sade merce eee P. debilis Loew 
— Abdominal tergites 1-3 shiny, dark brown with 
sparse microtomentum. Parafacial area dark 


brown P. nigrescens (Cresson) 


Philygria debilis Loew 
(Figs. 1—3) 


Philygria debilis Loew 1861: 357.—Osten 
Sacken 1878: 202 [Nearctic catalog].— 
Aldrich 1905: 627 [Nearctic catalog].— 
Jones 1906: 187 [list]—Johnson 1925: 
273 [list, Maine, New Hampshire, Ver- 
mont].—Wirth and Stone 1956: 469 [Cal- 
ifornia].—Deonier 1965: 501 [biolo- 
gy].—Wirth 1965: 745 [Nearctic cata- 
log].—Cole 1969: 400 [western United 
States].—Scheiring and Foote 1973: 160 
[habitat in Ohio].—Zack 1979: 93 [list, 
Washington, natural history]; 1983: 216 
[list, Washington]; 1998: 136 [list, Wash- 
ington, natural history].—Steinly 1984: 
753-754 [biology]; 1990: 188-193 [bi- 
ology].—Mathis and Zatwarnicki 1995: 
195 [world catalog].—Hollmann-Schi- 
rrmacher 1998: 66—71 [revision]. 

Hydrina debilis: Slosson 1902: 8 [generic 
combination].—Cresson 1944: 177 [re- 


VOLUME 107, NUMBER 1 


1-3. 
State Park and Utah. Utah: Goshen Hot Springs (39°57.8’N, 112°51.2'W)). 1, Epandrium, cerci, and surstylus, 
lateral view. 2, Phallapodeme, aedeagus, and gonites, lateral view. 3, Same, ventral view. Scale bar = 0.1 mm. 


Figs. 


view].—Sturtevant and Wheeler 1954: 
237-238 [review]. 

Philygria fuscicornis Loew 1862: 155.— 
Johnson 1925: 273 [synonymy]. 

Hydrina fuscicornis: Slosson 1902: 8 [ge- 
neric combination]. 

Cressoniella montana Saether 1970: 107.— 
Mathis and Wirth 1977: 520 [synonymy]. 


Diagnosis.—This species is distinguished 
from Nearctic congeners by the following 
combination of external characters: Parafa- 
cial area yellowish brown; wing mostly hy- 
aline, infuscate spots only over crossveins, 
lacking spurious veins; basal 1—4 tergites 
densely microtomentose, grayish brown, 5" 
tergite of male sparsely microtomentose to 
bare, subshiny to shiny. 

Description.—Small shore flies, body 
length 1.30—1.86 mm; yellowish brown to 
brown with yellowish-silver and gray mi- 
crotomentum. 

Head: Frons ventrolateral triangle brown 


Structures of the male terminalia of Philygria debilis (Virginia. Westmoreland: Westmoreland 


with yellowish-silver microtomentum, an- 
terior semicircle brown, laterally paler than 
ventrolateral triangles with yellowish-silver 
microtomentum, medially same color as 
ventrolaterial triangles. Occiput dark brown 
with yellowish-silver microtomentum. Out- 
er vertical seta %4 length of inner vertical 
seta. Scape and pedicel brown; flagellomere 
1 brown dorsally, yellowish brown ventral- 
ly; arista with minute branches. Facial 
background coloration yellowish brown to 
brown with yellowish-silver microtomen- 
tum; narrow band of yellowish-silver mi- 
crotomentum along eye margin beginning 
at antenna, extended and gradually blended 
with yellowish-silver microtomentum on 
gena. A combination of bands 
ventrally from antennae: Laterally along 


extended 


parafacial suture a narrow yellowish brown 
band gradually darkening ventrally; medi- 
ally along parafacial suture a narrow band 
and 


of yellowish-silver microtomentum, 


14. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


medially along this band a yellowish brown 
band. Gena covered with dense, silvery- 
gray microtomentum; postgena covered 
with silvery gray microtomentum, less 
dense than on gena. Palpus yellowish 
brown; prementum yellowish brown. 
Thorax: Scutal length 0.44—0.59 mm; 
scutellar length 0.17—0.24 mm. Mesonotum 
vittate, background brown; with brown 
stripe medially, along dorsocentral and in- 
tra-alar tracks, between brown stripes yel- 
lowish-silver microtomentum; scutellum 
medially brown with yellowish-silver mi- 
crotomentum, laterally with silvery gray 
microtomentum, anteriorly sparse; anepis- 
ternum brown with yellowish-silver micro- 
tomentum; katepisternum brown with yel- 
lowish-silver microtomentum; subscutellum 
dark brown with sparse, yellowish-silver 
microtomentum; anatergite shiny dark 
brown with sparse, yellowish-silver micro- 
tomentum. Chaetotaxy: Presutural dorso- 
central seta % to %4 length of anterior post- 
sutural dorsocentral seta; anterior postsutur- 
al dorsocentral seta *% to %4 length of pos- 
terior dorsocentral seta; lateral scutellar seta 
¥ length of apical seta. Wing: Length 1.62— 
2.13 mm; width 0.66—0.92 mm; costal vein 
ratio 0.39—0.50; M vein ratio 0.30—0.52; 
background, veins and crossveins brown, 
posteriorly wing slightly darker; r-m cross- 
vein dark brown dorsally, yellowish brown 
ventrally; dm-cu crossvein dark brown; 
wing cells anteriorly to r-m and posteriorly 
to dm-cu slightly lighter. Halter yellowish 
brown. Legs yellowish brown; femora yel- 
lowish brown; tibiae yellowish brown; tarsi 
yellowish brown with tarsomere 5 brown. 
Abdomen: Brown; male tergites 1—4 
brown with yellowish-silver microtomen- 
tum, tergite 5 shiny brown, tergite 5 medi- 
ally % length of tergite 4; female tergites 1— 
5 brown with yellowish-silver microtomen- 
tum, tergite 5 medially % length of tergite 
4. Male terminalia (Figs. 1—3): epandrium- 
cerci-surstyli fused; epandrium a broad, U- 
shaped band with small rounded anterov- 
entral projections; cercus completely fused 
laterally with epandrium with small y- 


shaped space dorsally separating cercus 
from epandrium, and bearing many long se- 
tulae; surstyli fused dorsally with epan- 
drium, rounded ventrally with field of a few 
long setae and many smaller setulae; phal- 
lapodeme triangular in lateral view, poster- 
oventral projection spatulate, anteroventral 
projection narrow and rounded; aedeagus 
elongate triangular in lateral view; subepan- 
drial plate-gonite-hypandrium fused; sube- 
pandrial plate reduced to region between 
gonite and hypandrium, with small rounded 
dorsal projection; gonite with broad base, 
rounded posterior projection with pointed 
bifurcate apical projections, prominent ven- 
tromedial setulae; hypandrium broadly 
fused with subepandrial plate, narrow an- 
teromedially with pointed anterolateral pro- 
jections and rounded ventral projections. 

Type material—The lectotype male of 
Philygria debilis Loew, here designated to 
preserve stability and make more universal 
the use of this name, is labeled “‘[United 
States.] Penns[ylvania; green; handwritten ]/ 
Loew Coll./Debilis m [handwritten]/Type 
11160 [red; number handwritten]/Philygria 
debilis Lw. det WWirth °61 [all except “det 
W Wirth” handwritten|/LECTOTYPE Phil- 
ygria debilis Loew 3d By Mathis/Edmiston 
liblacks border: FallzexcepmadeE Ch@ IMMER S 
and “By” handwritten].”’ The lectotype is 
double mounted (short pin in a rectangular 
block of cork), is in good condition (pin 
obscures much of mesonotum), and is de- 
posited in the MCZ (11160). There are no 
paralectotypes. 

The lectotype female of Philygria fusci- 
cornis Loew, here designated to preserve 
stability and make more universal the use 
of this name, is labeled “‘[United States.] 
mittel st [Middle States; green; handwrit- 
ten|/Loew Coll./fuscicornis [handwritten]/ 
Type 11159 [red; number handwritten] / 
LECTOTYPE Philygria fuscicornis Loew 
2 By Mathis/Edmiston [black border; all 
except “LECTOTYPE” and “By” hand- 
written].’’ The lectotype is double mounted 
(short pin in a long and narrow rectangular 
block of cork), is in good condition (pin 


VOLUME 107, NUMBER 1 


obscures some of mesonotum), and is de- 
posited in the MCZ (11159). There are no 
paralectotypes. 

The syntypes of Cressoniella montana 
Saether were collected in Colorado. Boul- 
der: North Boulder Creek (northern branch) 
and were deposited in the Zoological Mu- 
seum at the University of Oslo (ZMO). In 
an attempt to examine these syntypes, we 
wrote to the author, Ole A. Saether (Uni- 
versity of Bergen, Norway), who in turn 
corresponded with the staff from the Zoo- 
logical Museum in Oslo (ZMO). Based on 
feedback from ZMO, Dr. Saether wrote to 
us that the syntypes of C. montana could 
not be located despite a diligent search and 
that they are apparently lost. Thus, we sug- 
gest continued recognition of the synonymy 
that Mathis and Wirth (1977) published and 
as indicated in the generic and this species’ 
synonymy. 

Other specimens examined.—Because 
specimens of P. debilis are abundant in col- 
lections and the species is widespread in the 
Nearctic Region we have not recorded spe- 
cific locality data. 

Distribution.—Nearctic: Canada (Alber- 
ta, British Columbia, Manitoba, Ontario, 
Quebec, Saskatchewan), Mexico (Baja Cal- 
ifornia), United States (Alabama, Arizona, 
Arkansas, California, Connecticut, Colora- 
do, Delaware, District of Columbia, Idaho, 
illinois, Indiana, Iowa, Kansas, Maine, 
Maryland, Massachusetts, Michigan, Min- 
nesota, Mississippi, Missouri, Montana, 
Nebraska, Nevada, New Hampshire, New 
Jersey, New Mexico, New York, North Car- 
olina, North Dakota, Ohio, Oklahoma, 
Oregon, Pennsylvania, South Carolina, 
South Dakota, Tennessee, Texas, Utah, Ver- 
mont, Virginia, Washington, West Virginia, 
Wisconsin, Wyoming). 

The distribution locales are so numerous 
that a map with discrete points would be 
redundant. Locales from preserved speci- 
mens include all the contiguous states ex- 
cept Florida, Georgia, Kentucky, and Lou- 
isiana. The northernmost distribution seems 
to be near 50°N with one specimen collect- 


Nn 


ed from Churchill, Manitoba (58°N). The 
locale for the southernmost specimen ob- 
served was from northern Baja California 
Norte, Mexico. 

Remarks.—Philygria debilis is one of the 
most common and widespread Nearctic 
species, probably because of its occurrence 
on lawns associated with human habitations 
and recreation, such as golf courses (Cres- 
son 1944, Zack 1998). Also, the overall 
body and wing length often exceeds 2 mm, 
allowing this species to be easily collected 
with larger mesh nets. 

Although common and widespread, the 
immature stages of this species are un- 
known, and information on its natural his- 
tory is limited to habitat descriptions. Pre- 
sumably the larvae feed on blue-green al- 
gae. 

This species is sexually dimorphic, and 
not recognizing this dimorphism is proba- 
bly the reason why Loew described this 
species twice. The lectotype of P. debilis is 
a male with the apical tergite being shiny 
black and contrasted with the mostly mi- 
crotomentose anterior tergites. The lecto- 
type of P. fuscicornis is a female of the 
same species, and all tergites, including the 
apical one, are more or less uniformly mi- 
crotomentose and appear dull. 


Philygria nigrescens (Cresson), 
revised status 
(Figs. 4—7) 


Hydrina nigrescens Cresson 1930b: 80: 
1944: 177 [review, list, Idaho, Utah].— 
Sturtevant and Wheeler 1954: 239 [re- 
view].—Hollmann-Schirrmacher 1998: 
66 [synonymy with P. debilis}. 

Philygria nigrescens: Wirth and Stone 
1956: 469 [generic combination, Califor- 
nia]— Wirth 1965: 745 [Nearctic cata- 
log].—Cole 1969: 400 [western United 
States]|_—Mathis and Zatwarnicki 1995: 
197-198 [world catalog|.—Zack 1998: 
93 [list, Washington, natural history]; 
1983: 216 [list, Washington]; 1998: 136 
[list, Washington, natural history]. 


16 PROCEEDINGS OF THE 


y, 
/) 


I, 
VI/ |. 


% 


Figs. 4-6. 


ENTOMOLOGICAL SOCIETY OF WASHINGTON 


6 


Structures of the male terminalia of Philygria nigrescens (Alaska. Matanuska-Susitna: Talkeetna 


(62°18.9'N, 150°6.3’W)). 4, Epandrium, cerci, and surstylus, lateral view. 5, Phallapodeme, aedeagus, and gon- 
ites, lateral view. 6, Same, ventral view. Scale bar = 0.1 mm. 


Diagnosis.—This species is distinguished 
from Nearctic congeners by the following 
combination of external characters: Parafa- 
cial dark brown; wing mostly hyaline, in- 
fuscate spots only over crossveins, lacking 
spurious veins; tergites brownish black to 
black, basal 3 sparsely microtomentose, 
subshiny, apical 2—3 mostly bare, shiny. 

Description.—Small shore flies, body 
length 1.52—1.80 mm; generally gray to tan, 
abdomen black, subshiny to shiny. 

Head: Frons ventrolateral triangles dark 
brown with yellowish-silver microtomen- 
tum, anterior semicircle brown, laterally 
paler than ventrolateral triangles with yel- 
lowish-silver microtomentum, medially 
darker circle. Occiput dark brown with yel- 
lowish-silver microtomentum. Outer verti- 
cal seta %4 length of inner vertical seta. 
Scape and pedicel brown; flagellomere 1 


brown dorsally, yellowish brown ventrally; 
arista with minute branches. Facial back- 
ground coloration brown with yellowish- 
silver microtomentum; narrow band of sil- 
ver microtomentum along eye margin be- 
ginning at antenna, extended and gradually 
blended with silver microtomentum on 
gena. Laterally along parafacial suture a 
narrow brown band. Gena covered with 
dense, silvery gray microtomentum; post- 
gena covered with silvery gray microto- 
mentum, less dense than on gena. Palpus 
yellowish brown; prementun brown. 
Thorax: Scutal length 0.59 to 0.62 mm; 
scutellar length 0.26 mm. Mesonotum vit- 
tate anteriorly and mostly unicolorous pos- 
teriorly; background coloration brown, me- 
dially brown, lateral from medial line yel- 
lowish-silver microtomentum, dorsocentral 
line slightly darker, laterally along dorso- 


VOLUME 107, NUMBER 1 


central line yellowish-silver microtomen- 
tum, intra-alar line slightly darker, laterally 
along intra-alar line yellowish-silver micro- 
tomentum; scutellum medially and laterally 
brown with yellowish-silver microtomen- 
tum; anepisternum dark brown with yellow- 
ish-microtomentum, darker ventrally; kate- 
pisternum dark brown with silver microto- 
mentum, less dense ventrally; subscutellum 
dark brown with sparse, yellowish-silver 
microtomentum; anatergite shiny dark 
brown with sparse, yellowish-silver micro- 
tomentum. Chaetotaxy: presutural dorso- 
central seta “2—%4 length of anterior postsu- 
tural dorsocentral seta; anterior postsutural 
dorsocentral seta %—%4 length of posterior 
dorsocentral seta; lateral scutellar seta 4% 
length of apical seta. Wing: Length 2.11— 
2.24 mm; width 0.95—0.98 mm: costal vein 
ratio 0.37—0.43; M vein ratio 0.27—0.38; 
veins and crossveins brown; r-m crossvein 
dark brown dorsally, brown ventrally; dm- 
cu crossvein dark brown. Halter yellowish 
brown. Legs brown; femora brown; tibia 
brown, yellowish brown near tibio-femoral 
joint; tarsi yellowish brown with tarsomere 
5 brown. 

Abdomen: Dark brown; male tergites 1— 
3 dark brown with sparse yellowish-silver 
microtomentum, tergites 4—5 shiny, dark 
brown; tergite 5 medially % length of tergite 
4; female tergites 1-3 brown with yellow- 
ish-silver microtomentum, tergites 4—5 
shiny dark brown and dorsomedially % 
length of tergite 4. Male terminalia (Figs. 
4—6): Epandrium-cerci-surstyli fused; epan- 
drium broad U-shaped band with small 
rounded anteroventral projections; cerci 
completely fused laterally with epandrium, 
with small v-shaped spaces dorsally and 
ventrally separating cerci from epandrium, 
with a few setae and many setulae; surstyli 
dorsally fused with epandrium, rounded 
ventrally with field of a many long setae 
and many smaller setulae; phallapodeme 
triangular in lateral view, posteroventral 
projection spatulate, ventrally with broad v- 
shaped flange, and broad rounded anterov- 
entral projections; aedeagus elongate trian- 


17 


gular in lateral view; 10" sternite dorsad of 
aedeagus, rectangular in ventral view; su- 
bepandrial plate-gonite-hypandrium fused: 
subepandrial plate reduced to region be- 
tween gonite and hypandrium with small 
rounded dorsal projection; gonite with 
broad base, rounded posterior projection 
with pointed bifurcate apical projections, 
prominent ventromedial setula; hypandrium 
broadly fused with subepandrial plate, nar- 
row anterormedially with rounded antero- 
lateral projections and large rounded ventral 
projections. 

Type material—The holotype male of 
Nostima nigrescens Cresson is labeled 
“London Hill Mine/Bear Lake 21.7.03 [21 
Jul 1903; date handwritten] 
B[ritish]C|Columbia]/altitude 7,000 ft [nu- 
meral 7 handwritten|]/R P Currie Collector/ 
Type No. 21849 U.S.N.M. [numeral hand- 
written] [red]/TYPE No. Hydrina nigres- 
cens [name handwritten] E.T. Cresson, Jr 
[red].°’ The holotype is glued by the right 
thorax to a paper point, is in good condition 
(left tarsus missing and left wing broken but 
attached to body), and is deposited in the 
USNM (21849). 

Other specimens examined.—(7 6, 19 
2) CANADA. ALBERTA. Laggan (Lake 
Eouise) 5 Zon 19285708 Bayane Ciec- 
USNM). 

NORTHWEST TERRITORIES. Akla- 
vik, 2—22 Jun 1931, O. Bryant (1 6, 1 2; 
USNM). 

UNITED STATES. ALASKA. Matanus- 
ka-Susitna: Matanuska (rotary trap), 17 
May-—10 Jun 1944, J. C. Chamberlin (2 6, 
7 2; USNM): Matanuska Valley, 16 Jun 
1945, J. C. Chamberlin (2 ¢; USNM); Tal- 
keetna (62°18.9'N, 150°6.3’W) 7 Aug 2002, 
D. and W. N. Mathis (2 6; USNM), 4 Aug 
2003, D. and W. N. Mathis (1 6, 6 Q; 
USNM). 

ARIZONA. Coconino: Long Valley 
(34°31.3'N, 111°19:7'°W) 16 Aug 1951 (1 
2; USNM). 

CALIFORNIA. San Bernardino: Moun- 
tain Home, 12 Sep 1953, E. Schlinger (1 2; 
USNM). 


18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 7. 


Distribution map for Philygria nigrescens. 


OREGON. Lake: Quartz Mountain 
(42°19.3'N, 120°48.9'W; 1,600 m), 14 Jun 
1984, R. Danielsson (1 @; ZIL). 

UTAH. Kane: Coral Pink Sand Dunes 
(37°2.8'N, 112°40.7'W), 16 May 2001, D. 
and W. N. Mathis (1 2: USNM). 

Distribution (Fig. 7).—Nearctic: Canada 
(Alberta, British Columbia, Northwest Ter- 
ritories), United States (Alaska, Arizona, 
California, Idaho (literature record), 
Oregon, Utah, Washington (literature re- 
cord)). 

Remarks.—This species is rarely collect- 
ed, probably because we have not discoy- 
ered the requirements of its microhabitat. 
The few specimens available were usually 
collected along with much greater numbers 
of P. debilis, although at northern latitudes, 
such as sites in Alaska and the Northwest 


Territories, we only collected specimens of 
this species. 

The most reliable character for distin- 
guishing this species is the brownish black 
to black abdominal tergites with the last 
two tergites being shiny, almost completely 
devoid of microtomentum. 


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 Jon 
K. Gelhaus and Jason D. Weintraub 
(ANSP) and Philip D. Perkins (MCZ), who 
loaned specimens, we express our sincere 
thanks. We also thank Ole A. Saether (Uni- 
versity of Bergen) for attempting to find the 
syntypes of Cressoniella montana. 


VOLUME 107, NUMBER 1 


Hollis B. Williams provided technical 
support and produced the maps and Young 
T. Sohn skillfully inked the line drawings. 
We also thank Amnon Freidberg, Tadeusz 
Zatwarnicki, and Irina Brake for reviewing 
a draft of this paper. 

Field work in Alaska in 2002 was funded 
by a grant from the Bateman Fund, Nation- 
al Museum of Natural History, Smithsonian 
Institution, and we gratefully acknowledge 
this financial support. 


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Cogan, B. H., and W. W. Wirth. 1977. Family Ephy- 
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PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 21-27 


NEW GENERA AND SPECIES OF LEAFHOPPERS (HEMIPTERA: 
CICADELLIDAE) FROM KYRGYZSTAN 


D. V. Novikov, G. A. ANUFRIEV, AND C. H. DIETRICH 


(DVN and CHD) Center for Biodiversity, Illinois Natural History Survey, 607 E. Pea- 
body Dr., Champaign, IL 61820 (e-mail: dietrich@inhs.uiuc.edu); (GAA) Department of 
Zoology, Nizhny Novgorod State University, Nizhny Novgorod, Russia 


Abstract.—Two new genera and species of Paralimnini, Ctenotettix kirgysicus Novikov 
and Anufriev, n. gen. and n. sp., and Triasargus ancoratus Novikov and Anufriey, n. 
gen. and n. sp., and a new species of Ulopini, Neobufonaria milkoi Novikov and Anu- 
friev, n. sp., from the western Tien Shan of Kyrgyzstan are described and illustrated. 


Ecological notes are included for N. milkoi. 


Key Words: 


During the summers of 1998—2000, three 
month-long collecting expeditions to the 
steppe grasslands of Kyrgyzstan were un- 
dertaken by researchers from the Illinois 
Natural History Survey, Nizhny Novgorod 
State University (Russia), and the National 
Academy of Sciences of Kyrgyzstan. Sam- 
pling by sweep net and gasoline-powered 
vacuum in these grasslands yielded speci- 
mens of several hundred species of Au- 
chenorrhyncha (Hemiptera: Cicadomorpha 
and Fulgoromorpha). Among these are two 
new deltocephaline leafhopper species, 
each apparently representing a separate, un- 
described, genus of Paralimnini related to 
Jassargus Zachvatkin. Vacuum sampling in 
a high elevation boulder field in the western 
Tien Shan Mountains also yielded speci- 
mens of an undescribed species of the ul- 
opine genus Neobufonaria Emeljanov, pre- 
viously known from two species endemic 
to Central Asia (Emeljanov 1963, 1996) 
and another from Lebanon (Abdul-Nour 
2000, Szwedo 2002). The new taxa, au- 
thorship of which should be attributed to 
Novikov and Anufriev, are described and 
illustrated herein. Holotypes are deposited 


leafhopper, Central Asia, Tien Shan, grasslands, taxonomy 


in the insect collection of the Illinois Nat- 
ural History Survey, Champaign, Illinois, 
U.S.A [INHS]. Paratypes are in the Zoolog- 
ical Institute, Russian Academy of Scienc- 
es, St. Petersburg [ZIN], and the personal 
collection of G. A. Anufriev, Nizhny Novy- 
gorod State University, Russia [GAA]. 


Deltocephalinae Dallas 1870 
Paralimnini Distant 1908 


Ctenotettix Novikov and Anufriey, 
new genus 


Type species.—Crenotettix kirgysicus 
Novikov and Anufriev, new species. 

Diagnosis.—C?fenotettix resembles Jas- 
sargus, but differs in the weakly produced 
head, the presence of supranumerary cross- 
veins in the forewing clavus, the rocket- 
shaped connective, and the well-developed 
preapical lobe of the male style. Also char- 
acteristic of the new genus is the pecten of 
toothlike processes at the caudal edge of the 
pygofer lobe. 

Description.—General appearance and 
body structure, apart from genitalia, similar 
to those of Psammotettix Haupt, 1929. 
Moderately robust leafhoppers with re- 


2D, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


duced hind wings and slightly reduced fore- 
wings. Head (Fig. 1): Crown rounded, 
slightly angular apically, ratio of length to 
width 0.8; ocelli on border between face 
and crown closer to eyes than to each other. 
Thorax: Pronotum slightly convex, slightly 
wider posteriorly, narrower than head, with 
convex lateral margins. Mesoscutum basal- 
ly % of width of pronotum. Forewing ru- 
gose, slightly reduced, reaching abdominal 
tergum VII or VIII (specimen in Fig. | has 
the abdomen unusually distended), with su- 
pranumerary apical cells of variable num- 
ber. Hind wing reduced, half as long as 
forewing. Male genitalia: Subgenital plate 
(Fig. 3) triangular, rounded posteriorly, ap- 
proximately same length as valve, lateral 
margin straight, apex broadly rounded, mar- 
ginal row of about 10 long macrosetae end- 
ing subapically. Genital capsule closed in 
posterior view along caudal edges of py- 
gofer lobes. Pygofer lobe elongate (Fig. 2), 
with numerous long macrosetae in dorso- 
caudal area, caudal edge dentate forming 
dark pecten consisting of about 20 small tri- 
angular projections prominent in dorsal sec- 
tion, reduced ventrally to rugosity, inner 
surface in ventrocaudal corner with trian- 
gular leaflike fold directed dorsally. Style 
(Fig. 4) with well-developed preapical lobe; 
apophysis tapered, curved laterally, dentate 
preapically on posterior surface with 4 
small rounded projections, apex narrowly 
rounded, basomedial lobe short. Connective 
(Fig. 4) linear, rocket-shaped, stem short, 
minimum width of base equal half its 
length; anterior arms 3.3 length of base, 
with greatest spread not significantly wider 
than apex of stem, fused apically, apposing 
inner margins weakly concave. Aedeagus 
very distinctive (Figs. 5—6), base in poste- 


rior view wide with pair of dorsolateral 
arms, shaft in lateral and posterior view 
wide, gonopore dorsal subapical on poste- 
rior surface; apical portion of shaft in dorsal 
view with compressed lobe dorsally, 
tonguelike in posterior view; pair of short 
basolaterally directed spines just basad of 
gonopore. Female: Abdominal sternum VII 
(Fig. 7) with central round notch bordered 
by triangular teeth and semicircular dark 
spots. 

Notes.—Ctenotettix keys to Cleptochiton 
Emeljanov in Emeljanov’s key to leafhop- 
per genera of the European U.S.S.R. (Emel- 
janov 1967), but Cleptochiton differs mark- 
edly in having the subgenital plates strongly 
divergent with macrosetae restricted to the 
basal half, and in lacking preapical teeth on 
the style. Among known genera of Paral- 
imnini, the only other genera having su- 
pranumerary claval crossveins are Erras- 
tunus Ribaut and Triasargus, n. gen. (de- 
scribed below). 

The name of the new genus, which is 
masculine, combines Greek words ktenos 
(comb) and feftix (cicada) referring to the 
shape of the caudal edge of the pygofer 
lobe. The genus is described based on one 
new species collected in dry to mesic 
steppe grasslands in the western Tien Shan. 


Ctenotettix kirgysicus Novikov and 
Anufriev, new species 
(Figs. 1-7) 


Description._VMeasurements (mm). Male: 
length 3.3 + 0.3; pronotum width 1 + 0.05; 
head width 1.05 + 0.05. Female: length 3.2 
+ 0.2; pronotum width 1.04 + 0.05; head 
width 1.09 + 0.02. Coloration (Fig. 1): 
Yellowish gray, spotted with dark gray and 
brown to black. Crown with 4 small trian- 


Figs. 1-16. 


= 


1-7, Ctenotettix kirgysicus. 1, Dorsal habitus. 2, Pygofer, lateral view. 3, Subgenital plates and 


valve, ventral view. 4, Connective and style, dorsal view. 5, Aedeagus, posterior view. 6, Same, lateral view. 7, 
Female sternite VII. 8-14, Triasargus ancoratus. 8, Dorsal habitus. 9, Pygofer, lateral view. 10, Subgenital plates 
and valve, ventral view. 11, Connective and style, dorsal view. 12, Aedeagus apex, anterodorsal view. 13, 
Aedeagus, lateral view. 14, Female sternite VII. 15-16, Neobufonaria milkoi. 15, Aedeagus, lateral view. 16, 


Same, apex, anterodorsal view. 


isa) 
(as 


VOLUME 107, NUMBER 1 


24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


gular spots between ocelli, 2 large lighter 
triangular spots between apical corners of 
eyes, another row of four less distinct spots 
more posteriorly, and 4 spots by posterior 
margin, 2 inner of which more conspicuous. 
Pronotum with spots forming 6 indistinct 
longitudinal stripes, consisting of smaller 
and darker spots anteriorly and much larger 
but less intensely pigmented spots posteri- 
orly. Mesoscutum and scutellum of general 
background color with small dots and no 
distinct patterns. Forewing with dark dots 
merging at cell borders framing most cells 
and rendering veins and wing border light. 
Abdominal terga dark under forewing, be- 
coming lighter posteriorly with spots form- 
ing 2 transverse and 8 longitudinal groups 
on each tergum. 

Material examined.—Holotype 6, KYR- 
GYZSTAN: Dzhalal-abad, ca. jct. Kara 
Kysmak & Chatkal R., 2,240 m, 42°4’0’N, 
71°35'41"E, D.V. Novikov, 18 June 1999, 
sweeping, 99-58-06 [INHS]. Paratypes: | 
3d, same data; 2 2, same locality, C.H. Die- 
trich, 18 June 1999, sweeping, 99-58-07; 2 
?, same data, vacuum, 99-58-08; 1 5,1 @, 
same data, 19 June 1999 99-58-15; 1 d, 1 
2, same locality, G.A. Anufriev, June 18— 
19 1999 [ZIN]; 3 3, same data, [GAA]; 1 
36, KYRGYZSTAN: Chuy, Boom Ravine 3 
km N Kyz-Kujo, 1,380 m, 42°39'28’N, 
TS-S3 918; © Yulky 1OYY), (Cast, IDiginidn, Y= 
84-01 [INHS]. 

Notes.—The name is from the country of 
origin, a noun in apposition. 


Triasargus Novikov and Anufriev, 
new genus 


Type species.—Triasargus ancoratus 
Novikov and Anufriev, new species. 

Diagnosis.—Triasargus resembles Cten- 
otettix in general habitus, the presence of 
supranumerary cClaval crossveins, and in the 
shape of the subgenital plates, styles, and 
connective, but differs in the bizarrely mod- 
ified pygofer lobes, the emarginate poste- 
rior margins of which together form a cir- 
cular opening that reveals the aedeagal shaft 
in posterior view. 


Description.—General appearance re- 
sembling that of robust species of Psam- 
motettix Haupt. Head (Fig. 8): Crown 
rounded, slightly angular apically, ratio of 
length to width 0.7, ocelli on border be- 
tween face and crown considerably closer 
to eyes than to each other. Thorax: Pron- 
otum slightly convex, slightly wider poste- 
riorly, narrower than head, with convex lat- 
eral margins. Mesoscutum basally half 
width of pronotum. Forewing submacrop- 
terous, rugose, slightly shorter than abdo- 
men reaching abdominal tergum VIII or IX, 
with extra apical cells. Hind wing reduced, 
half as long as forewing. Male genitalia: 
Subgenital plate (Fig. 10) triangular with 
rounded posterior margin, slightly longer 
than valve, lateral edge concave, with about 
10 macrosetae in marginal row with 1—2 
subapical macrosetae diverged transversely. 
Genital capsule open in posterior view be- 
tween caudal edges of pygofer lobes. Py- 
gofer lobe square, truncate caudally (Fig. 
9), with long macrosetae numerous in dor- 
socaudal area, 2 prominent processes 
curved ventrad, 1 extending from dorsocau- 
dal corner and another extending from ven- 
trocaudal corner. Style (Fig. 11) with well- 
developed preapical lobe, apophysis short, 
truncate, widening apically, dentate at api- 
cal edge; basomedial lobe elongate. Con- 
nective (Fig. 4) linear, stem very short, 
transverse, half as long as wide, branches 
with greatest spread slightly greater that of 
stem, fused apically; apposing medial mar- 
gins moderately concave, strongly converg- 
ing from base to apex. Aedeagus in lateral 
view (Fig. 13) U-shaped, with narrow base, 
shaft slender, tapering, with two small lat- 
eral lobes in the apical % portion of its 
length, gonopore subapical on posterior sur- 
face, apex (Fig. 12) complex with three 
lobes, two directed ventrally and one ante- 
riorly. Female: Abdominal sternum VII 
(Fig. 14) with central U-shaped notch its 
sides diverging, flanked by characteristic 
rounded processes, processes dark apically 
bearing sharp triangular tooth laterally. 

Notes.—The new genus keys to Jassar- 


VOLUME 107, NUMBER 1 


gus in Emeljanov’s (1967) key, but is dis- 
tinguishable from that genus based on the 
larger size, weakly produced head, and the 
presence of supranumerary crossveins on 
the forewing clavus. Triasargus is very 
similar to Crenotettix externally and the two 
genera have supranumerary crossveins in 
the forewing clavus, a feature rare among 
Paralimnini. Thus it might be argued that 
the two species upon which these new gen- 
era are based belong in the same genus. We 
opted to recognize two separate genera for 
these species because the differences in the 
male genitalia, particularly the form of the 
connective base (long vs. short), subgenital 
plates (straight vs. concave laterally) and 
pygofer (narrow and closed vs. broad and 
open), are as great or greater than those 
which have been used to separate other gen- 
era of Paralimnini. 

The Greek trias in the generic name 
(gender—masculine) refers to the threefold 
longitudinal configuration of the shaft of 
the aedeagus, with the gonoduct forming a 
medial thin-walled tube between lateral tu- 
bular sclerotized shaft supports. 


Triasargus ancoratus Novikov and 
Anufriev, new species 
(Figs. 8—14) 


Description._Measurements (mm). Male: 
length 3.3 + 0.2; pronotum width 0.97 + 
0.02; head width 1.05 + 0.02. Female: 
length 3.5; pronotum width 1; head width 
1.07. Coloration: Yellowish gray with dark 
spots. Crown with 4 triangular spots ante- 
riorly, 2 large lighter triangular spots be- 
tween apical corners of eyes, 4 spots pos- 
teriorly. Pronotum with 6 small spots an- 
teriorly, and 6 longitudinal stripes posteri- 
orly. Mesoscutum and scutellum with no 
patterns. Forewing with cells bordered dark. 
First 6 abdominal terga dark brown to 
black, terga VII—IX lighter with traces of 
longitudinal stripes. 

Material examined.—Holotype 6, KYR- 
GYZSTAN: Dzhalal-abad, Chandalash R. 
GCkmis Ne yct.) Chatkal RK.) 156305 im: 
AEAAOUN RIN Ors 2228.) 20 Iine 1999) 


25 
C.H. Dietrich, vacuum, 99-60-05 [INHS]. 
Paratypes: 1 6, same data; 1 ¢, same data, 
99-60-03; 1 5d, 1 2, same locality, 21 June 
1999, sweeping, 99-60-08; 3 6, same data, 
vacuum, 99-60-09; 1 3d, same data, 20-21 
June 1999, 99-60-14 [INHS, ZIN]. 
Notes.—The Greek ankyra (L. ancora) 
in the name of the species refers to the an- 
chorlike apex of the aedeagus. The aedea- 
gus of 7. ancoratus is similar to that of 
Parunculus rostratus Emeljanov, 1964, but 
the latter has a broadly rounded subgenital 
plate, a posteriorly produced pygofer, and 
the shape of style and general habitus dif- 
fer substantially from those of the new spe- 
cies. 


Ulopinae Le Peletier and Serville 1825 
Ulopini 


Neobufonaria milkoi Novikov and 
Anufriev, new species 
(Figs. 15—16, 19) 


Description.—Measurements (mm). 
Male: length 3.9 + 0.1; pronotum width 
1.1 + 0.03, head width 1.3 = 0.03. Exter- 
nally closely resembling N. oshanini (Emel- 
janov) and N. costata (Emeljanov) (Figs. 
17-18). Genital capsule and genitalia re- 
sembling those of N. oshanini, but differing 
in having distal processes of aedeagus 
much shorter, only slightly longer than 
preapical width of shaft (Fig. 15) (N. os- 
hanini has apical aedeagal processes ap- 
proximately 3X longer than preapical shaft 
width in lateral view; cf. Emeljanov 1996: 
fig. 47). 

Material examined.—Holotype ¢6, KYR- 
GYZSTAN: Naryn, Dzhumgal-Too Ridge, 
Seok R. east ravine, 3,090 m, 42°12'04'"N, 
74°59'30"E 24 July 2000, C. H. Dietrich, 
vacuum, 00-111b [INHS]. Paratypes: 1 6, 
same data; 3 d, same locality, 25 July 2000 
[INHS, ZIN]. 

Notes.—Neobufonaria Kocak 1981 (re- 
placement for junior homonym Bufonaria 
Emeljanov 1963) previously included 3 
species: N. oshanini (Emeljanov) from Tad- 
jikistan, N. costata (Emeljanov) from Kyr- 


26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 17-19. 


Neobufonaria spp., dorsal habitus. 17, N. oshanini (Emeljanoy), male lectotype. 18, N. costata 


(Emeljanov), female holotype. 19, N. milkoi, male holotype. Scale bar = 1 mm. 


gyzstan and Kazakhstan (Emeljanov 1963, 
1996), and N. hermelensis Abdul-Nour 
from Lebanon. Neobufonaria milkoi keys to 
N. oshanini in Szwedo’s (2002) key to Pa- 
laearctic Ulopini, but the new species is 
readily distinguished by its much shorter 
aedeagal processes. The type series of N. 
milkoi was collected by vacuuming Loni- 
cera sp. (Caryophyllaceae) growing on 
boulders above 3,000 m in the Tien Shan 
Mountains, less than 100 km from the type 
locality of N. costata. Occurrence of N. mil- 
koi and N. costata in the same region of 
Kyrgyzstan suggests that some Neobufon- 
aria spp. are narrowly endemic. Thus, the 
holotype female of N. costata from Kzyl- 
Su, NW slope Terskei Alatau, Kyrgyzstan, 
may represent a different species from the 
males collected subsequently (1 from ca. 
Bosogo, Atbashi range, 3,500 m; | from 28 
km S Narynkola, Kazakhstan, Terskei Ala- 
tau, Emeljanov 1996). Additional study of 
the fauna of this peculiar leafhopper genus 
in the Tien Shan and Pamir mountains will 
likely reveal the presence of additional en- 
demic species. 

We take great pleasure in naming this 


species in honor of Dmitry A. Milko, Re- 
search Entomologist at the Institute of Bi- 
ology and Pedology, National Academy of 
Sciences of Kyrgyzstan, who organized the 
joint Kyrgyz/Russian/U.S. expeditions that 
led to the discovery of the new taxa de- 
scribed in this paper. 


ACKNOWLEDGMENTS 


We are indebted to D. Milko and G. Laz- 
kov, Institute of Biology and Pedology, Na- 
tional Academy of Sciences of Kyrgyzstan, 
for facilitating our fieldwork in Kyrgyzstan. 
L. L. Deitz, D..Dmitriev, J. N. Zahniser, and 
an anonymous referee made invaluable 
comments on an earlier draft of the manu- 
script. This work was supported in part by 
grant DEB 9870187 from the National Sci- 
ence Foundation (to C.H.D.). 


LITERATURE CITED 


Abdul-Nour, H. 2000. Un nouvel Ulopinae du Liban: 
Neobufonaria hermelensis n. sp. (Hemiptera: Ci- 
cadomorpha: Cicadellidae: Ulopinae). Mitteilun- 
gen der Schweizerischen Entomologischen Ge- 
sellschaft 73: 333-336. 

Emeljanov, A. F 1963. A new leafhopper genus from 


VOLUME 107, NUMBER 1 


the subfamily Ulopinae (Auchenorrhyncha, Cica- 
dellidae). Zoologicheskii Zhurnal 42: 1581-1582. 
1967. Suborder Cicadinea (Auchenorrhyn- 
cha), pp. 421-551. /n Bei-Bienko, G. Y., ed. Keys 
to the Insects of the European U.S.S.R. I. Apter- 
ygota, Palaeoptera, Hemimetabola. Israel Program 
for Scientific Translations, Jerusalem (first pub- 
lished in Russian by Nauka, Moscow-Leningrad, 
1964). 


in) 
~l] 


. 1996. Contribution to the knowledge of leaf- 
hoppers of the family Ulopidae (Homoptera, Ci- 
cadinae). Entomologisheskoe Obozrenie 75: 278— 
293 (English translation, Entomological Review 
76: 327-341). 

Szwedo, J. 2002. Ulopidae of the Palaearctic—The 
state of the art (Hemiptera: Clypaeorrhyncha: 
Membracoidea). Denisia 4: 249-262. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 28-33 


A NEW BARYSCAPUS FORSTER (HYMENOPTERA: EULOPHIDAE) 
PARASITIC ON DIORHABDA ELONGATA BRULLE (COLEOPTERA: 
CHRYSOMELIDAE) AND IMPLICATIONS FOR THE BIOLOGICAL 
CONTROL OF SALTCEDAR (TAMARICACEAE: TAMARIX SPP.) 
IN THE SOUTHWESTERN UNITED STATES 


M. W. Gates, S. N. MYARTSEVA, AND M. E. SCHAUFF 


(MWG, MES) Systematic Entomology Laboratory, PSI, Agricultural Research Service, 
U.S. Department of Agriculture, % National Museum of Natural History, Smithsonian 
Institution, RO. Box 37012, MRC 168, Washington, D.C. 20013-7012, U.S.A. (email: 
mgates @sel.barc.usda.gov, mschauff@sel.barc.usda.gov); (SNM) Rio Tamesi #515 Pon- 
iente, Fraccionamiento Zozaya 87070, Tamaulipas, Mexico (e-mail: smyartse@uamac. 
uat.mx) 


Abstract.—Baryscapus diorhabdivorus Gates and Myartseva, new species, (Hyme- 
noptera: Eulophidae) is described and illustrated. This species was reared from the salt- 
cedar leaf beetle, Diorhabda elongata Brullé (Coleoptera: Chrysomelidae), in Turkmen- 
istan. It is compared to closely related species, and its implications for the biological 


control of Tamarix spp. are discussed. 


Key Words: 


The genus Baryscapus Forster contains 
over 130 nominal species worldwide whose 
biologies are quite diverse in terms of host 
taxa and life stages attacked (Graham 1991, 
LaSalle 1994, Noyes 2001). This is the first 
record of Baryscapus attacking the chrys- 
omelid Diorhabda elongata Brullé (saltce- 
dar leaf beetle), but not the first record for 
species of Baryscapus attacking the family 
Chrysomelidae. Baryscapus chlamytis Ash- 
mead parasitizes the eggs of Chlamisus 
spp., Neochlamisus platani (Brown), and 
Exema spp. Baryscapus microrhopalae 
Ashmead is known from larvae/pupae of 
Microrhopala spp., and B. erynniae Do- 
menichini is both a larval/pupal hyperpar- 
asitoid of Tachinidae and a primary para- 
sitoid on Pyrrhalta luteola (Miiller). The 
saltcedar leaf beetle has been released re- 
cently in the western United States as part 
of a biological control program against in- 


Hymenoptera, Eulophidae, Baryscapus, Diorhabda, Tamarix, biocontrol 


vasive Tamarix spp. (Tamaricaceae) (De- 
Loach, personal communication). A new 
species of Baryscapus was discovered par- 
asitizing the saltcedar leaf beetle in Turk- 
menistan, the source of introduction into the 
United States. Some background informa- 
tion, presented below, is necessary to place 
the potential importance of this new species 
into context in terms of biological control 
programs of Tamarix spp. in the western 
United States. 

The economic impact of this new para- 
sitoid, were it to be inadvertently intro- 
duced into the United States, could be sig- 
nificant because its host has been released 
and established in at least five western 
states: California; Colorado; Nevada; Utah; 
and Wyoming (released but not established 
in Texas), for the control of Tamarix spp. 
(DeLoach, personal communication). Re- 
leases in Arizona and New Mexico are on 


VOLUME 107, NUMBER 1 


hold until the results of the other test re- 
leases are known, because saltcedar has 
supplanted native willow (Salix spp.) in ri- 
parian areas in these areas and has become 
a common nesting site for an endangered 
bird, the southwestern willow flycatcher. 

The Old World genus Tamarix consists 
of 54 species found from southern Europe 
to Central Asia. Approximately 10 species 
were introduced into the United States in 
the early 1800s as ornamentals, windbreaks, 
or to stabilize soil. Tamarix parviflora DC. 
and 7. ramosissima Ledeb. have displaced 
native willows and cottonwoods (Populus 
spp.) in nearly every riparian area in the 
western United States. These species oc- 
cupy >1.5 million acres, with T. parviflora 
confined to coastal and central California 
and 7. ramosissima found in arid inland re- 
gions. Saltcedar is disruptive for many rea- 
sons: 1) It uses twice as much water as na- 
tive vegetation; 2) it draws salt from deep 
in the soil and deposits it on the surface, 
eliminating understory plants; 3) it inter- 
feres with access to public and private riv- 
ers and lakes; 4) it causes flood damage by 
impeding flow during high water stages; 5) 
it burns easily, which favors saltcedar over 
native riparian vegetation due to rapid re- 
growth; and 6) and its dense roots and rhi- 
zomes spread out, slow river flow, increase 
siltation and can result in rechannelization 
(Baum 1978, Everitt 1980, Truman 1996, 
Weisenborn 1996). 

Larval and adult populations of D. elon- 
gata effectively defoliate saltcedar as a 
larva and adult with complete defoliation of 
T. raimosissima by D. elongata document- 
ed in Kazakhstan and Turkmenistan (De- 
Loach 2001). Field cage trials began in 6 
western US states in 1999 and open field 
releases began at 6—7 sites in May, 2001. 
Success has ranged from partial to complete 
defoliation upon establishment (DeLoach, 
personal communication). Adults produce 
at least two generations per year, depending 
on conditions. Adults only eat green vege- 
tation, creating holes in the leaves through 
which water escapes, causing branches to 


29 


wither and die. The beetles appear to aes- 
tivate south of the 38° parallel during mid 
summer, reducing their effectiveness. As 
Tamarix has no close relatives in North 
America, exotic biological control candi- 
dates that feed on saltcedar are unlikely to 
find suitable native American food plants or 
crops and become pests. Similarly, B. dior- 
habdivorus, n. sp., might also attack related 
chrysomelid taxa in the New World were it 
to be introduced. Unfortunately, not enough 
is known about the host range of species of 
Baryscapus that attack chrysomelids to 
speculate on this. 

Abbreviations used are USNM (National 
Museum of Natural History, Smithsonian 
Institution, Washington, DC), BMNH (The 
Natural History Museum, London), and 
ZIN (Zoological Institute, Kiev, Ukraine). 


Baryscapus diorhabdivorus Gates and 
Myartseva, new species 
(Figs. 1—6) 


Etymology.—Named for the host genus 
of Chrysomelidae, Diorhabda, and the Lat- 
in root vor, meaning “eat.” 

Diagnosis and discussion.—The follow- 
ing diagnosis of B. diorhabdivorus should 
be compared to the subsequent diagnoses of 
similar species that also attack Chrysome- 
lidae. The male of B. diorhabdivorus has 
the ventral plaque occupying only ~0.3X 
the length of the scape (Fig. 6), and gaster 
~1.8xX as long as broad. Both sexes are 
dark brown to black with bluish to greenish 
metallic reflections; have a bilobate clype- 
us; stouter stigmal veins; 3—4 adnotaular se- 
tae on the mesoscutum, and a fine median 
line (Figs. 2, 4); tibial banding on all legs; 
and the propodeal spiracle in a teardrop- 
shaped depression (Fig. 4). 

Baryscapus microrhopalae: dark with 
metallic blue reflections, male scape with 
longer ventral plaque occupying ~0.5X its 
length; clypeus not bilobate; more slender 
stigmal vein; propodeal spiracle not in tear- 
drop-shaped depression; male with a more 
elongate gaster with longer first gastral ter- 
gum; tibial banding only on hind leg. 


30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


5 


Figs. 1-6. 


Baryscapus chlamytis: brown with very 
faint metallic (blue/purple) reflections; male 
scape with ventral plaque in medial half and 
occupying 0.3—0.5x its length; clypeus 
shallowly bilobate; more gracile and shorter 
stigmal vein rounded apically with finer un- 
cus; propodeal spiracle not in teardrop- 


Baryscapus diorhabdivorus. 1, Lateral view of gaster, female. 2, Lateral view of mesosoma, 
female. 3, Anterior view of head, female. 4, Dorsal view of mesosoma. 5, Antenna, female. 6, Scape, male. 


shaped depression; only two adnotaular se- 
tae, very faint median line on mesoscutum. 

Baryscapus erynniae: extensive bright 
metallic green reflections; male scape with 
ventral plaque in basal half and occupying 
~0.5 its length; clypeus truncate; more 
slender stigmal vein; propodeal spiracle not 


VOLUME 107, NUMBER 1 


in teardrop-shaped depression; male with 
anterior third of gaster translucent brown; 
tibial banding on all legs; sometimes begin- 
nings of second row (1—2 only) of adnotau- 
lar setae anteromedially, distinct median 
line on mesoscutum. 

Herein we place B. diorhabdivorus in 
Graham’s (1991) key and compare it diag- 
nostically with other species of Baryscapus 
associated with Chrysomelidae. This spe- 
cies may mistakenly key to B. servadeii 
(Domenichini) (couplet 9) IF the gaster is 
collapsed and the hypopygium appears to 
extend to 0.7—0.73 the length of the gas- 
ter. However, unlike B. servadeii, the malar 
sulcus of B. diorhabdivorus is not strongly 
curved. Continuing beyond couplet 9’ re- 
quires that IF the hypopygium is 
situated beyond the middle of the gaster, 
then there are OFTEN >1 rows of adnotau- 
lar setae (only one row in B. diorhabdivo- 
rus), thus it is possible to proceed. Barys- 
capus diorhabdivorus ultimately keys to 
couplet 37. This result is obtained by pro- 
ceeding through the following couplets: 
10’—clava <4.2—4.3X as long as broad, 
mesoscutum with median line; 1 1’—gaster 
<2.0X as long as head + thorax; 12’—ped- 
icel+ flagellum <1.4X as long as width me- 
soscutum; 13’—gaster not strongly acumi- 
nate and <1.5*X as long as thorax; 19’—all 
funiculars not distinctly longer than broad: 
20’—gaster at least 1.5 as long as broad; 
21’—F1 not at least 1.5 as long as broad; 
34’—pedicel + flagellum usually at least as 
long as width mesoscutum, host not Quer- 
cus (Fagaceae); 35’—antennal clava lack- 
ing prominent spine; 36—gaster 1.5—2.1 xX 
as long as broad with last tergite broader 
than long), after which it would come out 
near B. obesulus Graham (couplet 38). 
However, B. diorhabdivorus does not quite 
match the characters of B. obesulus in 
which the gaster is longer and the propo- 
deum is greater than 0.25 the length of 
the scutellum. Furthermore, B. obesulus is 
only known from France and Sweden from 
unknown hosts. 

Description.—Female: 


Length 1.1—1.7 


31 


mm. Color: Body dark brown to black, 
with dark greenish-blue metallic reflections; 
lateral surfaces of mesosoma, coxae and 
gaster lustrous brown with faint metallic re- 
flections. Antenna yellow, scape and pedi- 
cel basally slightly darkened. Legs yellow, 
except femora in basal % and pretarsomeres 
brown. Fore wing hyaline, veins light yel- 
low, setae on disk short and sparse. 

Head: Lineolate to finely reticulate, oc- 
cipital region imbricate, slightly broader 
than thorax, 1.2-1.3x as broad as high, 
sparse setation ventrad toruli and laterad 
scrobal depression. Scrobal depression shal- 
low with fine intrascrobal carina in apical 
half, converging at “V” of frontal arms. 
Eye 1.6—1.8X as long as malar space. Malar 
sulcus slightly curved. Posterior ocelli 
placed on margin of vertex, separated from 
dorsal eye margin by 1.4 ocellar diameter 
and from each other by 3.0 ocellar di- 
ameter. Distance between posterior ocelli 
~2.0 greater than between lateral ocellus 
and eye margin. Scrobes connected dorsal- 
ly, converging with fine intrascrobal carina 
and frontal arms. Toruli level with lower 
margin of eyes, separated by ~2.0 torulus 
diameter. Clypeus bilobate (Fig. 3). Width 
of oral fossa equal to 0.67 width of head. 

Antenna: Scape 3.8—4.5X as long as 
broad, pedicel 1.4—1.8 as long as broad. 
Anellus transverse. Funicles roughly quad- 
rate to longer than broad, Fl 0.8—1.2X as 
long as broad; F2 0.8—1.0X as long as 
broad. Clava 3-segmented, slightly wider 
than funicle, 2.5—3.4* as long as broad 
(Bice): 

Mesosoma: Pronotum short, transverse, 
finely reticulate with row of distinct setae 
along apical margin. Mesoscutum 1|.1—1.5x 
as long as broad, dorsally lineolate and be- 
coming more broadly lineolate to reticulate 
on lateral surfaces of mesoscutum and scu- 
tellum, with median line and with a single 
row of 3—4 adnotaular setae (Figs. 2, 4). 
Scutellum 0.8 as long as broad; submedia 
and sublateral lines present, submedial lines 
distinct and closer to each other than either 
is to sublateral line; 2 pairs setae laterad of 


32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


submedial lines, anterior seta placed near 
middle of scutellum and ~1.0X setal base 
diameter from submedial line, posterior seta 
placed near posterior edge of scutellum and 
~2.0X setal base diameters from submedial 
line; circular placoid sensillum present ap- 
proximately midway to just anteriad of 
midpoint between scutellar setae. Metano- 
tum transverse, rounded medially, with 
faint reticulate sculpture, ~0.5 length of 
propodeum. Propodeum faintly reticulate 
with fine median carina; spiracles ~0.5X 
their diameter from anterior margin of pro- 
podeum, located in shallow, teardrop- 
shaped depression that tapers posteriorly, 
two long setae just anterolaterad and one 
long seta posterad of spiracle above base of 
metacoxa (Fig. 4). Prepectus finely imbri- 
cate, mesopleuron glabrate dorsally becom- 
ing finely imbricate-reticulate ventrally, 
femoral depression shallow, arcuate. Meta- 
coxa ~2.0X as long as mesocoxa, broadest 
medially, finely imbricate. 

Wing: Fore wing 2.0—2.1x as long as 
broad, marginal fringe short, setation sparse 
and fine. Submarginal vein with three dor- 
sal setae, two in basal third and one at mid- 
point, 0.9-1.1x as long as marginal vein. 
Marginal vein 7.0—10.0 as long as post- 
marginal vein and 2.5—3.0X as long as stig- 
mal vein. Stigmal vein slightly less than 
3.0X times shorter than marginal vein. Bas- 
al portion of fore wing glabrous. 

Metasoma: Gaster 1.8—2.0X as long as 
broad (uncollapsed specimens), finely retic- 
ulate-imbricate, each tergum with single, 
transverse row of setae near apex of tergite; 
hypopygium extending ~0.7X length of 
gaster. Ovipositor sheaths exserted, but 
barely visible in dorsal view (Fig. 1). 

Male: Length 0.9—1.0 mm. Similar to fe- 
male in coloration and structure except as 
follows: Scape 3.0 as long as broad with 
ventral plaque in apical third (Fig. 6). Anel- 
lus transverse. Fl 0.8X as long as broad, 
F2 1.2X as long as broad; clava 3.5X as 
long as broad. Setae of flagellomeres as 
long as width of corresponding segment. 
Gaster 1.8 as long as broad. Genitalia 


(phallobase) slightly more than 3.0 longer 
than its width. Digital sclerites slightly less 
than 6.0 shorter than phallobase, with one 
thin tooth apically, their length equal to 
length of parameres and slightly enlarged to 
apex (in dried specimens). 

Type material.—Holotype 2. Turkmen- 
istaine IDiey Sjoerd Wale, YZ NINO, &- 
Myartseva, ex adults of beetle Diorhabda 
elongata (USNM). Paratypes 14 2, 1 6, 
same data as holotype (10 2, 1 6, USNM; 
2 2 BMNH; 2 2 ZIN). The exact location 
of Dry Sport Lake is unknown, though ap- 
parently in the vicinity of Ashgabat. 

Other material—Turkmenistan: Ashga- 
bat and vicinity, 1.x.1993, S. Myartseva, ex 
larvae of beetle Diorhabda elongata Brullé 
On Marans Soo, (@ 2 il @  WSINMMl): 
vc NOE (@Q 2p MOnatin M4 (BG 2 
ix 1994" (Gi S)s 8-x1993, (4G Se ail 
USNM). 


ACKNOWLEDGMENTS 


We thank Alex Konstantinov and Dave 
Smith (USDA, ARS, Systematic Entomol- 
ogy Laboratory, Washington DC), Greg 
Evans (Florida State Collection of Arthro- 
pods, Gainesville), John LaSalle (CSIRO, 
Canberra, Australia), and an anonymous re- 
viewer for providing critical reviews of this 
manuscript and Raymond Carruthers 
(USDA, ARS, Albany, CA) and Jack 
DeLoach (USDA, ARS, Temple, TX) for 
insightful discussions regarding the biolog- 
ical control of saltcedar. 


LITERATURE CITED 


Baum, B. 1978. The genus Tamarix. Israel Academy 
of Sciences and Humanities, 209 pp. 

DeLoach, J. 2001. General news: Saltcedar biocontrol 
reconciling risks. Jn CABI Biocontrol News & In- 
formation 22(3): 1-5. 

Everitt, B. 1980. Ecology of saltcedar: A plea for re- 
search. Environmental Geology 3: 77-84. 

Graham, M. W. R. de V. 1991. Reclassification of Eu- 
ropean Tetrastichinae. Memoirs of the American 
Entomological Institute No. 49, 322 pp. 

LaSalle, J. 1994. North American genera of Tetrasti- 
chinae (Hymenoptera: Eulophidae). Journal of 
Natural History 28: 109-236. 

Noyes, J. 2001. Interactive Catalogue of World Chal- 


VOLUME 107, NUMBER 1 


cidoidea. CD-ROM, 2nd edition. Taxapad and The 
Natural History Museum, London. 

Truman, K. 1996. Collaborative efforts for tamarisk 
management and control. Problem statement: Fax. 
University of Nevada Environmental Studies Pro- 
gram, Las Vegas, Nevada. 


BS 


Weisenborn, W. 1996. Saltcedar impacts on salinity, 
water, fire frequency and flooding, pp. 9-12. In 
Proceedings of the Saltcedar Management 


Workshop. June 12, 1996, Rancho Mirage, Cal- 
ifornia. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 34—48 


SUMMER EMERGING EPHEMEROPTERA, PLECOPTERA, AND 
TRICHOPTERA OF ABRAMS CREEK, GREAT SMOKY MOUNTAINS 
NATIONAL PARK 


R. EDWARD DEWALT AND B. D HEINOLD 


Illinois Natural History Survey, 607 E. Peabody Dr., Champaign, Illinois 61820, U.S.A. 
(e-mail: edewalt @inhs.uiuc.edu) 


Abstract.—Abrams Creek drainage was surveyed for adult mayflies (Ephemeroptera), 
stoneflies (Plecoptera), and caddisflies (Trichoptera) during summer 2001 as part of the 
All Taxa Biodiversity Inventory in the Great Smoky Mountains National Park (GRSM). 
Nine reaches were collected, up to five times, using primarily ultraviolet light trapping. 
Of the 35,710 specimens examined, 164 species resulted. These consisted of 35 species 
of mayflies, 36 stoneflies, and 93 caddisflies. Eight species are new to Tennessee, while 
39 are new GRSM records. Three rarely collected mayflies, Epeorus vitreus (Walker), 
Leucrocuta thetis (Traver), and Nixe spinosa (Traver), were taken. Rare caddisfiies in- 
cluded Ceratopsyche macleodi (Flint), Cheumatopsyche helma Ross, Hydroptila chattan- 
ooga Frazer and Harris, H. talladega Harris, and Chimarra augusta Morse. No rare stone- 
flies were collected. Additional specimens of two undescribed (but known to specialists) 
species were taken in Goera (Trichoptera) and Jsoperla (Plecoptera). Specimens of Hy- 
droptila nr. amoena Ross (Trichoptera) and a Caenis nr. mccafferti Provonsha (Ephem- 
eroptera) are possibly new to science, but require more specimens and study. More effort 
concentrated in southwestern GRSM and in Ephemeroptera and Trichoptera should yield 


additional significant records. 


Key Words: 


Ephemeroptera, Plecoptera, Trichoptera, Great Smoky Mountains National 


Park, All Taxa Biodiversity Inventory 


The Great Smoky Mountains National 
Park (GRSM) straddles the border between 
Tennessee and North Carolina, USA. It re- 
sides in one of the most species-rich tem- 
perate zones in the world and has been des- 
ignated as an International Biosphere Re- 
serve (Sharkey 2001). The National Park 
Service, aided with funding and logistical 
support by the non-profit Discover Life in 
America (DLIA), has been documenting the 
presence, distribution, and biology of 
GRSM species as part of an All Taxa Bio- 
diversity Inventory (ATBI). The many ben- 
efits of the ATBI project have been docu- 
mented by Sharkey (2001). 


Abrams Creek is the westernmost drain- 
age in the GRSM and is entirely contained 
within Blount County, Tennessee. It arises 
from elevations near 1,300 m along the 
Tennessee and North Carolina border and 
empties into Lake Chilhowee, an impound- 
ment of the Litthke Tennessee River at ap- 
proximately 320 m elevation. This drainage 
is largely pristine in its headwaters but has 
been cleared of its forested riparian zone 
within its middle third, an area referred to 
as Cades Cove. One Cades Cove reach is a 
“losing stream’’ at times, where porous 
bedrock allows water to escape the channel, 
only to resurface downstream. Beyond 


VOLUME 107, NUMBER 1 


Table 1. 


85 


Streams, elevation, coordinates, and dates visited for nine repeatedly sampled sites in the Abrams 


Creek drainage of the Great Smoky Mountains National Park, summer 2001. All dates are ultraviolet trap events 


unless otherwise noted. 


Stream and Description m asl Latitude 
Anthony Cr., Anthony Cr. Trail 821 35.5868 
Abrams Cr., Cades Cove CG 584 35.6047 
Abrams Cr., Sparks Ln. 553 35.6025 
Abrams Cr., Abrams Falls Trlhd. 532 35.5921 
Mill Cr., Abrams Falls Trlhd. 540 35.5901 
Forge Cr., Gregory Rg. Tr., CG 12 733 35.5472 
Trib. Forge Cr., Gregory Rg. Tr. 715 35.5485 
Trib. Forge Cr., Parson Br. Rd. 671 35.5588 
Abrams Cr. at Abrams Cr. CG 367 35.6103 


Longitude 


83.7516 
83.7757 
83.7939 
83.8520 
83.8522 
83.8321 
83.8349 
83.8546 
83.9327 


Dates Visited 


UV 5/27; Sweep 5/26, 6/9 
5/24, 6/9, 6/19, 7/5, 7/17 
5/25, 6/7, 6/19, 7/6, 7/17 
5/25, 6/8, 6/18, 7/6, 7/18 
6/6, 6/18, 7/6, 7/18 
Sweep 5/25; UV 5/26, 6/7 
Sweep 5/25; UV 5/26, 6/7 
5/27, 6/9, 6/19, 7/7, 7/18 
5/29, 6/10, 6/20, 7/8, 7/22 


, 7/6 


5 G2, Wis WAY 
, 6/21, 7/7, 7/20 


Cades Cove the stream begins a steep de- 
scent through a narrow canyon culminating 
at Abrams Falls, a bedrock outcropping 
some 4—5 m high. The stream eventually 
flattens near its mouth, providing long plac- 
id runs and pools, interspersed by short rif- 
fles. 

Ephemeroptera (mayflies), Plecoptera 
(stoneflies), and Trichoptera (caddisflies), 
hereafter collectively referred to as EPT, are 
common inhabitants of streams throughout 
much of the world. In North America, they 
are especially diverse in the southeastern 
United States, where they have undergone 
much speciation, due in part to the relative- 
ly stable geology and climate of the area 
(Brigham et al. 1982). Their current status 
and risks to survival in the Southeast have 
been reviewed by Morse et al. (1993). 

EPT species have a wide range of toler- 
ance to organic pollution and habitat alter- 
ation, with many species being extremely 
intolerant of environmental change. As an 
ecological indicator, the number of EPT 
taxa is an efficient measure of stream 
health, provides a surrogate for more costly 
measures of ecosystem function, and has 
the ability to monitor habitat specific im- 
pacts (Barbour et al. 1992, Lenat and Pen- 
rose 1996, Wallace et. al. 1996). Ecological 
tolerance values have been established for 
many commonly encountered taxa in the 
Southeast (Lenat 1993). 

Several stream locations in GRSM are 
routinely monitored using immature life- 


stages with identification being facilitated 
using important works such as Brigham et 
al. (1982). Although many immatures of 
EPT species have been associated with 
adults, a large component remains unde- 
scribed or underdescribed; therefore, these 
data are insufficient to inventory GRSM 
EPT species. ATBI permanent plots (see 
Sharkey 2001), using Malaise traps, have 
provided some adult EPT. Small, system- 
atics-oriented projects have also helped to 
expand the number of known species, How- 
ever, basin-wide inventories for adults are 
necessary to accurately determine the EPT 
species that occur in GRSM and to associ- 
ate species with biotic and abiotic factors 
for predictive purposes. 

The objective of this study was to con- 
duct an intensive inventory of EPT in the 
Abrams Creek drainage during early and 
midsummer months. This drainage was 
chosen after the authors reviewed speci- 
mens at the Illinois Natural History Survey 
(INHS) insect collection. Plecoptera alone 
constituted 600 records, and georeferencing 
of these locations demonstrated a hole in 
coverage west and south of Cades Cove— 
the Abrams Creek drainage. Charles Parker, 
an aquatic entomologist working in GRSM, 
confirmed that a study of this drainage 
would help close a gap in information for 
streams in GRSM. 

MATERIALS AND METHODS 

Nine sites (Table 1, Fig. 1) were repeat- 

edly sampled in the Abrams Creek drainage 


36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Tennessee @ Collection Sites N 
ea Abrams Creek Watershed hese) 
Park Roads 0 05 1 2 3 Miles 
SS eS ee 
North Carolina Great Smoky Mts Nat Pk 
80 
» 70 @ Mayflies C1 Stoneflies (1 Caddisflies 
>) 
= 
= 60 
4 
Zz 50 
2 40 
5 
2. 30 
N 
E 20 
a 
= ji - 
0 
G S . S 
eS © * RS Re oo s S & 
> & < «Ss «S$ S & oN S 
> > = oS S g ey S$ & 
BIC ois ET PESO TE ce g 
= <> aS < “8 <0 oe Ne 
Sy S ENS “se s 
Re ceotio<Wr ssh eniety: Ree 
y 
Y. ~ «Ss s 
Figs. 1-2. 1, (top) EPT sampling locations in the Abrams Creek drainage of the Great Smoky Mountains 


National Park, summer 2001. 2, (bottom) EPT species richness at nine locations in the Abrams Creek drainage 
of Great Smoky Mountains National Park, summer 2001. 


VOLUME 107, NUMBER 1 


between 25 May and 22 July 2001. These 
locations were spread throughout the drain- 
age (Fig. 1) and over an elevational gradi- 
ent of 454 m. Sites ranged from seepage 
areas (tributaries of Forge Creek) to a 30 
m-wide reach of Abrams Creek at Abrams 
Creek Campground. 

Most reaches were sampled using a Bio- 
quip™, 12-v, ultraviolet light. Effort was 
standardized by time (approximately one 
hour beginning at sunset), by reflective 
sheet size, and by common weather condi- 
tions. Often, several sites were sampled in 
a single evening, necessitating the use of 
DC timers that permitted remote lighting of 
traps. Two trays with 80% EtOH were po- 
sitioned at the bottom of the sheet to cap- 
ture insects. At attended sites, mayfly sub- 
imagoes (a subadult with hair-covered 
wings that molts to an imago) were cap- 
tured and stored in a container until trans- 
formation. Additionally, males of perlid and 
perlodid stoneflies were captured and their 
intromittent organ extruded to facilitate spe- 
cies identification. Anthony Creek was 
sampled using sweep-netting and handpick- 
ing during daylight hours, with the excep- 
tion of one ultraviolet trap event. Night 
sampling was avoided due to higher than 
normal bear activity in that area. Hence, the 
results at this site are not strictly compara- 
ble to that of other sites. Geographic coor- 
dinate data were captured using a Garmin™ 
12XL at each site. 

Some Abrams Creek Campground trap 
events produced in excess of 5,000 speci- 
mens. Consequently, subsampling at 25% 
was conducted inconjuction with a search 
for large and rare taxa. An extraordinary 
event occurred on 22 July necessitating an 
overall 1/16 subsample rate for microcad- 
disflies, producing 797 specimens from an 
estimated 12,832 hydroptilids. Full sample 
abundance was estimated at 51,328 individ- 
uals. For this site, species richness values 
are reflective of specimens actually inspect- 
ed and no effort was made to estimate rich- 
ness based on total sample abundance. No 
subsampling occurred for other sites. 


Oo 
~ 


Specimens were identified to species 
when possible. Often, only the males of 
species could be identified. Females were 
determined where descriptions existed or 
where it seemed that color pattern, size, or 
wing veination was consistent with males 
of known identity. Specimens are housed in 
the INHS insect collection and are denoted 
as GRSM and ATBI related specimens. 
Specimen records are available via the In- 
ternet at http://ctap.inhs.uiuc.edu/insect/ 
search.inhs.asp. 


RESULTS 


Forty-five site visits were made in the 
Abrams Creek drainage during this project 
(Table 1), resulting in 35,710 identified 
specimens. This project recorded 164 EPT 
species (Table 2). Most species were cad- 
disflies (56.7%), with the remainder evenly 
split between mayflies and stoneflies. 

Heptageniid mayflies were especially 
species rich (17), while baetids, ephemer- 
ellids, and leptophlebiids provided only 
five, four, and two species respectively. The 
latter three families were drastically under 
represented in our samples compared to the 
regional species pool (Unzicker and Carl- 
son 1982). The dominance of subimagoes 
in our unattended ultraviolet traps has un- 
doubtedly reduced taxonomic resolution in 
these families. Morse et al. (1989), working 
in similar elevations straddling the North 
and South Carolina border collected 29 
mayfly species with low representation in 
the same families. 

Stoneflies were dominated by perlids (10 
species) and perlodids (9). The Chloroper- 
lidae (six species) and Leuctridae (four) 
were probably under represented (Unzicker 
and McCaskill 1982). Ultraviolet light col- 
lections and sweep-netting frequently pro- 
duced many more females than males in 
these families, often limiting taxonomic 
resolution beyond genus. Nelson (1996) 
conducted a similar inventory, but with im- 
matures, in the Litthe Pigeon River of 
neighboring Sevier County, Tennessee. He 
reported 29 stonefly taxa over a year-long 


38 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


effort. His use of immatures limited deter- 
minations to genus in some species rich 
groups such as Alloperla and Leuctra. 

Caddisflies were represented by 93 spe- 
cies. Surprisingly, microcaddisflies (Hy- 
droptilidae) were the most diverse family 
with 17 species. Until this study, few hy- 
droptilids were reported from GRSM (C. 
Parker, personal communication) and were 
not thought to be a rich component of the 
fauna. Leptocerids and hydropsychids pro- 
vided 15 and 14 species respectively. Morse 
et al. (1989) reported a similar, but more 
diverse, fauna from comparable elevations. 
Much of the increase in diversity in their 
study was attributable to continued sam- 
pling into the fall season when adults of 
Pycnopsyche and Neophylax were avail- 
able. Otherwise, the distribution of species 
among families, with hydroptilids, hydrop- 
sychids, and leptocerids providing the bulk 
of the species, was comparable to the pre- 
sent study. 

Species richness varied greatly across 
sites (Table 2, Fig. 2). Abrams Creek at 
Abrams Creek Campground produced the 
greatest number of EPT species. This rich- 
ness is undoubtedly due to many diverse 
habitats, including long, placid runs, under- 
cut banks, and deep pools not common in 
other sections of the watershed. Abrams 
Creek at Abrams Falls Trailhead also pro- 
duced many species. This is the only reach 
with an intact flatwoods, an area of minimal 
gradient that is heavily forested and well 
protected from foot traffic. This site shares 
many species with Abrams Creek Camp- 
ground such as several long-horn and mi- 
crocaddisflies, but also offers some distinct- 
ness such as a large population of Cheu- 
matopsyche oxa Ross found only in the flat- 
woods during this study. 


SIGNIFICANT FINDS 


Species were compared to published lit- 
erature and electronic species lists including 
Berner (1977), Brigham et al. (1982), DLIA 
list for aquatic insects (www.discoverlife. 
org /nh/cl/GSMNP/aquatic_insects-GS MNP. 


html#overview), Etnier et al. (1998), Lenat 
and Penrose (1987), Long and Kondratieff 
(1996), Mayfly Central (www.entm.purdue. 
edu/entomology/mayfly/mayfly.html), 
NatureServe (www.natureserve.org), and 
Pescador et al. (1999) to determine new 
GRSM and Tennessee state records. A dis- 
cussion of the more significant finds follows. 


EPHEMEROPTERA 
Caenidae 


Caenis anceps Traver.—Taken from 
Abrams Creek Campground. Known from 
central and eastern Tennessee (Long and 
Kondratieff 1996, Provonsha 1990). A new 
GRSM record. 

Caenis nr. mccafferti Provonsha.—Four 
females from Abrams Creek Campground 
resemble Caenis mccafferti Provonsha, but 
lack the fleshy, fingerlike posteromedial 
projection of abdominal tergite two of this 
species (A. Provonsha, personal communi- 
cation). More specimens are needed to de- 
termine its identity. 


Ephemerellidae 


Ephemerella invaria (Walker).—One 
male from Tributary to Forge Creek at 
Gregory Ridge Trail. Initially identified as 
E. fratercula McDunnough and recently 
synonymized with E. invaria (Jacobus and 
McCafferty 2003). Known locally from 
eastern Tennessee (Long and Kondratieff 
1996). A new park record. 

Serratella molita (McDunnough).-Abun- 
dant in lower Abrams Creek. Known from 
central and eastern Tennessee (Allen and 
Edmunds 1963, Long and Kondratieff 
1996) and North and South Carolina (Ja- 
cobus and McCafferty 2002). Jacobus and 
McCafferty (2002) recently synonymized 
with Serratella serratoides (McDunnough) 
with this species. A new GRSM record. 


Ephemeridae 


Ephemera varia Eaton.—Found at 
Sparks Lane and Mill Creek. Several Ten- 
nessee records reported (Long and Kondra- 
tieff 1996). Southeastern records are gen- 


VOLUME 107, NUMBER 1 39 


Table 2. Ephemeroptera, Plecoptera, and Trichoptera from nine repeatedly collected locations in the Abrams 
Creek drainage of the Great Smoky Mountains National Park, summer 2001. Sites 1 = Anthony, 2 = Abrams, 
Cades Cove CG, 3 = Abrams, Sparks, 4 = Abrams, Fall Trailhead, 5 = Mill, Falls Trailhead, 6 = Forge CG12, 
7 = Trib. Forge Gregory, 8 = Trib. Forge Parson, 9 = Abrams CG. 


Sites 
Ephemeroptera 
Baetidae spp. 60 60 
Acentrella turbida 2 628 630 
Acerpenna pygmaea ] 1 
Baetis brunneicolor ] 
Baetis flavistriga 1 28 10 39 
Baetis intercalaris 53 
Baetis sp. 7 1 1 2 ] 12 
Caenidae 
Caenis anceps 1 138 139 
Caenis n.sp. 4 4 
Ephemerellidae spp. 13 104 32 ] 150 
Ephemerella dorothea 3 3 
Ephemerella hispida 1 | 
Ephemerella invaria 1 
Ephemerella sp. 2 D, 1 17 9 31 
Serratella serrata 18 18 
Serratella molita S) 6 6 6 497 520 
Serratella sp. 1 5 6 
Ephemeridae 
Ephemera guttulata 1 | 
Ephemera varia 1 4 5 
Heptageniidae 
Cinygmula subaequalis 13 13 
Epeorus dispar 2; 2 
Epeorus vitreus 95 25 332 42 3 4 1 84 586 
Heptagenia marginalis 17 17 
Leucrocuta aphrodite 5 2 205 212 
Leucrocuta juno 59 968 15 312 3 1,358 
Leucrocuta thetis 6 5 l 1, 
Nixe spinosa 1 1 Pp; 4 
Stenacron carolina 1 2 1 4 
Stenacron interpunctatum 1 68 69 
Stenacron pallidum 3 3) 
Stenacron sp. 2 3 1 2 8 
Stenonema ithaca 1 105 106 
Stenonema meririvulanum | 
Stenonema modestum 4 3 62 69 
Stenonema pudicum 10 63 5 5 l 84 
Stenonema terminatum ] | 2 
Stenonema vicarium 3 92 95 
Stenonema sp. l 2 l 2 5 11 
Isonychiidae 
Isonychia bicolor 13 | | 15 
Isonychia serrata | l 
Isonychia sp. l 3 | | | 17 24 


Leptophlebiidae 


—— ee 


40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 2. Continued. 


Sites 

I 2 3 4 5 6 7 8 9 Sum 
Habrophlebiodes americana | 1 
Paraleptophlebia assimilis 2 2 
Paraleptophlebia sp. l | | 3 
Siphlonuridae 
Siphlonurus typicus ] 1 
Leuctridae 
Leuctra alexanderi 14 4 5) 23 
Leuctra ferruginea 3 3 | 7 
Leuctra grandis 1 1 
Leuctra carolinensis 2 6 8 
Leuctra sp. 19 2 6 13 7 47 
Nemouridae 
Amphinemura nigritta i 1 
Amphinemura wui 20 2 8 8 3 41 
Chloroperlidae 
Alloperla caudata 1 1 
Alloperla nanina 1 1 1 3 
Alloperla neglecta 1 1 
Alloperla usa 1 1 
Alloperla sp. 2 1 3 1 1 8 
Suwallia marginata 1 1 
Sweltsa mediana 5 5 
Sweltsa sp. 6 14 2 1 1 24 
Perlidae 
Acroneuria abnormis 5 2 3 1] 16 21 47 105 
Acroneuria carolinensis 2 2 
Acroneuria filicis 1 1 Z 4 
Beloneuria georgiana 2 1 3 
Eccoptura xanthenes 6 10 16 
Neoperla coosa 11 11 
Perlesta decipiens 13 13 
Perlesta frisoni 1 1 
Perlesta nelsoni 2 3 15 20 
Perlesta placida 4 38 42 
Perlodidae 
Diploperla duplicata | 1 
Isoperla dicala 8 2 5 
Isoperla frisoni 5 5 
Isoperla holochlora 7 1 17 5) 9 3 42 
Isoperla sp. M8 1 1 
Isoperla sp. 4 4 
Malirekus hastatus 1 1 
Remenus bilobatus 3 1 2 6 
Yugus arinus 1 i 
Yugus bulbosus 3 1 4 
Yugus sp. DD 2 
Peltoperlidae 
Tallaperla anna 1 1 


Tallaperla laurie 6 ] 7 
2 Ea ee ee ae a en nan ene Mira A 


VOLUME 107, NUMBER 1 4] 

Table 2. Continued. 

Sites 
1 2 3 4 5 6 7 8 9 Sum 

Tallaperla maria 2 24 26 
Tallaperla sp. 2 2 
Viehoperla ada | ] 
Pteronarcyidae 
Pteronarcys scotti 1 ] 
Brachycentridae 
Micrasema wataga ] 526 169 1 334 1,031 
Micrasema sp. 1 1 2 
Calamoceratidae 
Anisocentropus pyraloides 2 2 
Heteroplectron americanum 1 | 
Glossosomatidae 
Agapetus rossi 24 206 230 
Agapetus tomus AN 183 136 82 1,893 2,506 
Agapetus minutus 1 ] 
Agapetus sp. 2,566 1,818 1,093 5,477 
Glossosoma nigrior 1 3 2, 8 1 16 
Goeridae 
Goera calcarata ] 4+ 100 28 Sy) 185 
Goera n.sp. ] 3 1 5 
Helicopsychidae 
Helicopsyche borealis 1 71 72 
Hydropsychidae 
Arctopsyche irrorata D; 2 
Ceratopsyche macleodi 87 87 
Ceratopsyche morosa 2 =) ] 8 
Ceratopsyche slossonae Wf 209 181 9 5 16 427 
Ceratopsyche sparna 2 S)// 78 219 110 11 3 181 721 
Ceratopsyche sp. 2 1 3 
Cheumatopsyche nr. goera ] | 
Cheumatopsyche harwoodi 305 236 136 Syl 931 Ah) 
Cheumatopsyche helma 2 27 
Cheumatopsyche oxa 211 60 271 
Cheumatopsyche sp. 364 l 858 1,223 
Diplectrona modesta ] 111 See Sie OO m2S 718 
Hydropsyche betteni 4 6 6 16 
Hydropsyche depravata 11 4 15 
Hydropsyche betteni or depravata 2 16 17 35 
Hydropsyche simulans 84 16 100 
Hydropsyche sp. 13 2 15 
Parapsyche cardis if D5 10 | 43 
Hydroptilidae 

Hydroptila alabama | 3 4 
Hydroptila nr. amoena | l | 3 
Hydroptila armata 4 3 15 22 
Hydroptila callia 4 10 14 
Hydroptila chattanooga 6 6 
Hydroptila delineata 19 19 
Hydroptila fiskei | 27 | 163 192 
Hydroptila grandiosa | 113 23 PSU 7,288 


42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 2. Continued. 


Sites 

SK SO ee, ae, a ne CN, MRT 
26 ue eee ee EEE eee EEE ee ee ee ee 
Hydroptila hamata ] 40 8 748 797 
Hydroptila talladega I 3 
Hydroptila sp. 4 1 52 7 D) 197 263 
Mayatrichia ayama 1 1 
Ochrotrichia graysoni D D) 
Ochrotrichia sp. 1 i 8 
Oxyethira novasota 2 2 
Oxyethira pallida 2 2, 
Oxyethira sp. 3 4 
Palaegapetus celsus 3 3 
Stactobiella delira 1 1 
Stactobiella martynovi 4 1 1 6 
Stactobiella sp. 15 1 1 17 
Lepidostomatidae 
Lepidostoma lydia 1 1 
Lepidostoma modestum group 2 yy 
Lepidostoma ontario 4 1 5) 
Lepidostoma pictile 3 1 4 
Lepidostoma tibiale 1 17 18 
Lepidostoma togatum 4 4 
Lepidostoma (Mormomyia) sp. 1 2 1 4 5 i 20 
Leptoceridae 
Ceraclea cancellata 1 5) 6 
Ceraclea flava 119 119 
Ceraclea nepha 1 1 
Ceraclea tarsipunctata 1 1 16 18 
Ceraclea transversa 8 8 5 6 4 1 5 8 45 
Ceraclea sp. 7 1 2 10 
Leptocerus americanus 1 1 
Mystacides sepulchralis 5 3 8 
Nectopsyche exquisita 57 57 
Oecetis avara 32 1 318 351 
Oecetis inconspicua 5 2 2 1 804 814 
Oecetis persimilis 1 26 5 139 171 
Setodes stehri 92 3 1,455 1,550 
Triaenodes ignitus 4 2 12 31 2 51 
Triaenodes perna 1 9 10 
Triaenodes taenius 1 2 3 
Limnephilidae 
Pseudostenophylax uniformis 1 1 3 5) 
Molannidae 
Molanna blenda 1 1 
Molanna ulmerina 1 1 1 6 9 
Odontoceridae 
Psilotreta amera 3 3 
Psilotreta sp. 1 1 
Philopotamidae 
Chimarra aterrima 2 2 
Chimarra augusta 38 3 
Chimarra socia 193 193 


VOLUME 107, NUMBER 1 


Table 2. Continued. 


Chimarra sp. 
Dolophilodes distinctus 
Dolophilodes major 
Dolophilodes sp. 
Wormaldia moesta 
Wormaldia sp. 


Phryganeidae 


Ptilostomis ocellifera 


Polycentropodidae 
Nyctiophylax affinis 
Nyctiophylax banksi 
Nyctiophylax celta 
Nyctiophylax nephophilus 
Nyctiophylax sp. 
Plectronemia crassicornis 
Polycentropus cinereus 
Polycentropus confusus 
Polycentropus maculatus 
Polycentropus sp. 


Psychomyliidae 


Lype diversa 
Psychomyia flavida 
Rhyacophilidae 
Rhyacophila atrata 
Rhyacophila carolina 
Rhyacophila carpenteri 
Rhyacophila fuscula 
Rhyacophila glaberrima 
Rhyacophila nigrita 
Rhyacophila teddyi 
Rhyacophila torva 
Rhyacophila vibox 
Rhyacophila sp. 


Sericostomatidae 
Fattigia pele 
Site Totals 
Ephemeroptera 
Plecoptera 
Trichoptera 

EPT Total 


6 
16 
] 
4 
39 
37) 
6 
26 20 
1,446 
1 
3 2 
7 
4 
24 
2 
2 
291 2,595 
7 11 
16 7 
16 32 
39 50 


47 


43 
Sites 
4 5 6 1 8 9 Sum 

129 129 
5 10 8 4 6 58 
11 7 ] 20 
3 16 7 3 45 
2 4 7 
2 2) ] 5 
| 
21 21 
| 
1 81 82 
] 3 3 ] 12 
11 2 3 2 320 3/7 
] a 
3 2] 16 18 D; 3 105 
3 8 2 3 13 51 
| ] 
3 i 2 11 
5 11 7 2 5 1 78 
419 260 ] 18 17 2,208 
| 
1 1 3 2) ] 14 
2 2 4 
11 15 1] 1 5) 50 
| 5) 
4 28 
1 3 
2 
| 2 
i ] 7 
2 2 
5,010 3,195 487 458 276 18,874 35,710 
16 1] 8 9 4 15 35 
8 4 10 13 6 8 36 
40 37) Dil 24 23 52 93 
64 52, 45 46 33 75 164 


erally from “high mountain areas” Mc- 
Cafferty (1975). A new GRSM record. 


Heptageniidae 


Epeorus vitreus (Walker).—Abundant in 
larger streams. A rare find in Tennessee 
(Long and Kondratieff 1996). A new 


GRSM record. 


Leucrocuta aphrodite (McDunnough).— 
Abundant in lower Abrams Creek. Known 
from the Southeast (Berner 1977) and Ten- 
nessee (Long and Kondratieff 1996). First 
confirmed record from GRSM, but see Ber- 
ner’s (1977) nonspecific record of Sevier 
County, Tennessee. 


44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Leucrocuta juno (McDunnough).—Most 
commonly encountered Leucrocuta in mid- 
dle reaches. Known from the Southeast 
(Berner 1977) and central and eastern Ten- 
nessee (Long and Kondratieff 1996). First 
confirmed GRSM record (again, see Ber- 
ner). 

Leucrocuta thetis (Traver).—Relegated 
to higher elevation sites. Possibly rare in 
Tennessee (Long and Kondratieff 1996), 
but known from GRSM. 

Stenonema ithaca (Clemens and Leo- 
nard).—Abundant at Abrams Creek Camp- 
ground. Known from central and eastern 
Tennessee (Bednarik and McCafferty 1979, 
Long and Kondratieff 1996). New GRSM 
record. 

Nixe spinosa (Traver).—Four males tak- 
en from three locations. Known from north- 
western North Carolina (Unzicker and Carl- 
son 1982, Pescador et al. 1999). A new 
Tennessee and GRSM record. 


Isonychiidae 


TIsonychia bicolor (Walker).—Fifteen 
males taken in middle reaches of Abrams 
Creek. Known from Tennessee (Long and 
Kondratieff 1996). A new GRSM record. 


Leptophlebiudae 


Paraleptophlebia assimilis (Banks).— 
Two specimens taken from Tributary to 
Forge Creek at Gregory Ridge Trail. 
Known from the Southeast (Berner 1977), 
including Tennessee (Long and Kondratieff 
1996). A new GRSM record. 


Siphlonuridae 


Siphlonurus typicus Eaton.—One male 
taken from Sparks Lane. Not known from 
the Southeast (Berner 1977, Pescador et al. 
1999), but reported from Indiana and Mas- 
sachusetts (Provonsha and McCafferty 
1982), the Midwest and eastern Canada 
(Randolph and McCafferty 1998), and New 
York (Jacobus and McCafferty 2001). A 
new Tennessee and GRSM record. 


PLECOPTERA 
Perlidae 

Perlesta decipiens (Walsh).—Thirteen 
adults from Abrams Creek Campground. A 
widespread, eastern species (Stark 1989). A 
new Tennessee and GRSM record. Any ref- 
erence to species of Perlesta prior to Stark’s 
(1989) revision must be viewed with skep- 
ticism. 

Perlesta placida (Hagen).—Several 
specimens taken from Abrams Creek at the 
Falls Trailhead and at Abrams Creek Camp- 
ground. Widespread in Atlantic and Gulf 
Coastal Plains states (Stark 1989). A new 
Tennessee and GRSM record. 

Neoperla coosa Smith and Stark.—Elev- 
en adults taken at Abrams Creek Camp- 
ground. A recently described species from 
northern Alabama (Smith and Stark 1998). 
A new Tennessee and GRSM record. 

TIsoperla n.sp. M8.—One specimen taken 
from Tributary to Forge Creek at Gregory 
Ridge Trail. An undescribed, light colored 
species, superficially similar to /. holoch- 
lora (Klapalek). Stan Szczytko is currently 
reevaluating the genus in eastern North 
America. The above naming convention is 
his. A new Tennessee and GSRM record. 


TRICHOPTERA 
Hydropsychidae 


Cheumatopsyche helma Ross.—Twenty- 
seven specimens taken from Abrams Creek 
Campground. Known from four locations in 
Tennessee, Kentucky, and North Carolina 
(Etnier et al. 1998). Not a new GRSM re- 
cord, but it is rare. 

Cheumatopsyche oxa Ross.—Taken from 
Abrams and Mill creeks at the Falls Trail- 
head. Gordon (1974) reported it from Mid- 
western and eastern USA. Known from 
eastern and middle Tennessee (Etnier et al. 
1998). A new GRSM record. 

Hydropsyche simulans Ross.—Taken 
from Abrams Falls Trailhead and Abrams 
Creek Campground. Widespread in Tennes- 
see (Etnier et al. 1998). A new GRSM tre- 
cord. 


VOLUME 107, NUMBER 1 


Hydroptilidae 


Hydroptila alabama Harris and Kelley.— 
Taken from Abrams Creek Campground. 
Known from Alabama (Harris and Kelley 
1984) and Polk County, Tennessee (Etnier 
et al. 1998). A new GRSM record. 

Hydroptila nr. amoena Ross.—Three 
males from Cades Cove area. Steve Harris 
(personal communication) states that he has 
not been able to identify them and that they 
may be new. 

Hydroptila armata Ross.—Collected 
from lowest elevation sites. Known from 
middle and eastern Tennessee (Etnier et al. 
1998). A new GRSM record. 

Hydroptila chattanooga Frazer and Har- 
ris—Abrams Falls Trailhead yielded six 
males. Described from Alabama, additional 
specimens from Ohio and Pennsylvania 
(Frazer and Harris 1991). A new Tennessee 
and GRSM record. 

Hydroptila fiskei Blickle-——Taken from 
larger reaches of the drainage. A new Ten- 
nessee and GRSM record. 

Mayatrichia ayama Mosely.—One spec- 
imen taken at Abrams Creek Campground. 
Etnier et al. (1998) reported it from nearby 
Bradley and Monroe counties. A new 
GRSM record. 

Ochrotrichia graysoni Parker and Vosh- 
ell—Two males taken at Abrams Creek 
Campground. Originally described from 
Virginia (Parker and Voshell 1980). A new 
Tennessee and GRSM record. 

Oxyethira novasota Ross.—Two males 
taken from Abrams Creek Campground. 
Known regionally from northern Alabama 
(Harris 1986, Harris et al. 1991) and eastern 
Tennessee (Etnier et al. 1998). A new 
GRSM record. 

Oyxethira pallida (Banks).—Two males 
taken from Abrams Creek Campground. A 
second record for Tennessee (Etnier et al. 
1998), a first for GRSM. 


Leptoceridae 


Ceraclea flava (Banks).—Many individ- 
uals taken from Abrams Creek Camp- 


45 


ground. Reported from middle and eastern 
Tennessee (Etnier et al. 1998). A new 
GRSM record. 

Ceraclea nepha (Ross).—One male tak- 
en from Abrams Creek Campground. 
Known from Tennessee, including nearby 
Polk County (Etnier et al. 1998). A new 
GRSM record. 

Ceraclea transversa (Hagen).—Widely 
distributed in the drainage. Reported from 
middle and northeastern Tennessee (Etnier 
et al. 1998). A new GRSM record. 

Leptocerus americanus (Banks).—One 
male taken at Abrams Creek Campground. 
Known from east-central Tennessee (Etnier 
et al. 1998). A new GRSM record. 

Oecetis avara (Banks).—Abundant in 
the three lowest elevation reaches Abrams 
Creek. Widespread in Tennessee (Etnier et 
al. 1998). A new GRSM record. 

Triaenodes perna Ross.—Small numbers 
collected at Cades Cove and Abrams Creek 
campgrounds. Known from central and 
eastern Tennessee (Etnier et al. 1998). A 
new GRSM record. 


Molannidae 


Molanna ulmerina Navas.—Collected at 
low elevation. Known from a single east- 
central Tennessee location (Etnier et al. 
1998). A new GRSM record. 


Philopotamidae 


Chimarra aterrima Hagen.— The genus 
is not well represented in GRSM. It appears 
that Dolophilodes and Wormaldia replace it 
above 400 m. Two males taken at Abrams 
Creek Campground. Known from eastern 
and central Tennessee (Etnier et al. 1998). 
A new GRSM record. 

Chimarra augusta Morse.—Three males 
taken at Abrams Creek Campground. In 
Tennessee, known only from Bradley 
County (Etnier et al. 1998). A new GRSM 
record. 

Chimarra socia Hagen.—Abrams Creek 
Campground produced many specimens. 
Known from scattered locations in Tennes- 


46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


see, including adjacent Monroe County (Et- 
nier et al. 1998). A new GRSM record. 


Polycentropodidae 


Nyctiophylax affinis (Banks).—Twenty- 
one males taken at Abrams Creek Camp- 
ground. Known from middle Tennessee (Et- 
nier et al. 1998). A new GRSM record. 

Nyctiophylax celta Denning.—Eighty- 
two males taken at Tributary of Forge 
Creek at Parson Branch and at Abrams 
Creek Campground. Etnier et al. (1998) re- 
ported it from eastern Tennessee, including 
Blount County. A new GRSM record. 

Plectronemia crassicornis (Walker).— 
Two females taken from Sparks Lane and 
Mill Creek. Known from middle Tennessee 
(Etnier et al. 1998). A new GRSM record. 


Rhyacophilidae 


Rhyacophila vibox Milne.—One female 
each were taken at Sparks Lane and at Trib- 
utary to Forge Creek on Gregory Ridge 
Trail. Known from eastern Tennessee (Et- 
nier et al. 1998). A new GRSM record. 


DISCUSSION 


This large project contributed eight new 
Tennessee and 39 new GRSM records. 
Most of these were recorded from Abrams 
Creek Campground. Other large streams 
may offer more unknowns for GRSM. Sev- 
eral other significant finds originated from 
the flatwoods portion of Cades Cove at the 
confluence of Abrams and Mill creeks. 
These areas are easily accessible and pose 
a conundrum as to how they have been 
overlooked through the 70 year history of 
the park. It is probable that most systema- 
tists have avoided Cades Cove and the 
western end of GRSM because it is heavily 
traveled and constitutes some of the lowest 
elevations. 

Several EPT specimens represent unde- 
scribed species known to specialists. These 
include /soperla sp. M8 and Goera sp. Ad- 
ditionally, specimens of Caenis nr. mccaf- 
ferti and Hydroptila nr. amoena may rep- 
resent new species. More specimens and 


further study will be required to solve these 
questions. 

Our study has also documented new lo- 
cations for several rarely collected species. 
Rarity is defined here as species being 
known from a relatively few published lo- 
cations. Since designation of species as rare 
depends on available data, those with un- 
published records may disagree with our in- 
terpretations. Publication of those records 
in paper or digital format would increase 
the accuracy of determinations of rarity. 

Rare mayflies include Epeorus vitreus, 
Leucrocuta thetis, and Nixe spinosa. The 
distribution of Epeorus vitreus is not well 
known (NatureServe) and we provide sev- 
eral records. Leucrocuta thetis is given an 
S3 (vulnerable due to a restricted range) 
status in Tennessee by NatureServe. We 
have provided three GRSM locations. Nixe 
spinosa is thought of as a North Carolina 
endemic and has been listed by Nature- 
Serve as N3 (vulnerable due to a restricted 
range). The species has no imperilment sta- 
tus in North Carolina. We provide the first 
records for Tennessee. 

There were no rare stoneflies collected, 
but five rare caddisflies are discussed. Cer- 
atopsyche macleodi was originally de- 
scribed from North Carolina (Flint 1964) 
and Etnier et al. (1998) gave a nonspecific 
reference of eastern Tennessee. Anthony 
Creek supports a large population of this 
species. NatureServe lists Cheumatopsyche 
helma as S1 (Alabama and Pennsylvania- 
critically imperiled) to S3 (Tennessee-vul- 
nerable) to SH (Kentucky-extirpated). A 
small population was taken at Abrams 
Creek. Hydroptila chattanooga has only 
been reported from five locations in Ala- 
bama, Ohio, and Pennsylvania (Frazer and 
Harris 1991). We have six males from the 
a low gradient, flatwoods section of Abrams 
Creek. NatureServe rates Hydroptila tallad- 
ega as S1| in Alabama, with insufficient data 
for other states where it occurs (Georgia, 
Kentucky, North Carolina, and South Car- 
olina). It was listed by Morse et al. (1993) 
as rare and vulnerable. We report it from 


VOLUME 107, NUMBER 1 


three locations in the drainage. Chimarra 
augusta is rated at S1 in Alabama, but has 
no rating in other states in which it occurs 
(South Carolina, Tennessee, Virginia, West 
Virginia) (Armitage 1991). 

Our data suggest that the potential for 
discovery of new state and GRSM records 
is greatest in mayflies and caddisflies. Fu- 
ture work on these and other aquatic insects 
should focus on southwestern GRSM drain- 
ages and on the streams above Lake Fon- 
tana. These areas are less accessible than 
any other region in GRSM and may hold 
additional significant records. 


ACKNOWLEDGMENTS 


We thank DLIA for funding and ac- 
knowledge the advice and guidance of C. 
Parker in choosing the Abrams Creek drain- 
age for study. He has answered many ques- 
tions and confirmed the identity of several 
specimens. DLIA employee J. Hilten has 
arranged for housing in GRSM, while K. 
Langdon and the National Park Service 
have supported our efforts and requests for 
permits. Additionally, S. Johnson, A. Zim- 
merman, and N. Lowe have helped pack in 
heavy batteries and spent long evenings 
with the ultraviolet lights in the back coun- 
try. Specimens were verified by B. Kondra- 
tieff (Colorado State University), S. Szczyt- 
ko (Univerisity of Wisconsin, Stevens 
Point), M. Meyers (Purdue University), L. 
Jacobus (Purdue University), S. Harris 
(Clarion College, PA), and B. Stark (Mis- 
sissipp1 College). 


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107(1), 2005, pp. 49-54 


REVIEW OF THE SPECIES OF ELAPHROPEZA MACQUART 
(DIPTERA: EMPIDIDAE: TACHYDROMIINAE) FROM THE 
CHINESE MAINLAND 


DING YANG AND STEPHEN D. GAIMARI 


(DY) Department of Entomology, China Agricultural University, Beijing 100094, Chi- 
na; (SDG) California State Collection of Arthropods, Plant Pest Diagnostics Laboratory, 
California Department of Food and Agriculture, 3294 Meadowview Road, Sacramento, 
CA 95832-1448, USA (e-mail: sgaimari @cdfa.ca.gov) 


Abstract.—The species of the genus Elaphropeza Macquart from the Chinese mainland 
are reviewed. The following new species are described: E. liui and E. anae. A key to 
the species of the genus from the Chinese mainland is presented for the first time. New 
combinations are created for the following Chinese species which were included in Dra- 
petis (Elaphropeza): E. centristria (Yang and Yang), E. fujianensis (Yang and Yang), E. 
fuzhouensis (Yang and Yang), E. guangxiensis (Yang and Yang), E. guiensis (Yang and 
Yang), E. jianyangensis (Yang and Yang), E. jinghongensis (Yang and Yang), E. lancan- 
gensis (Yang and Yang), EF. /ii (Yang and Yang), E. longiconica (Yang and Yang), E. 
medipunctata (Yang and Yang), E. meihuashana (Yang and Yang), E. paucipunctata (Yang 
and Yang), FE. pilata (Yang and Yang), E. postnigra (Yang and Yang), E. ruiliensis (Yang 
and Yang), FE. striata (Yang and Yang), E. triangulata (Yang and Yang), E. xanthina 
(Yang and Yang), E. xizangensis (Yang and Yang), and E. yunnanensis (Yang and Yang). 


Key Words: 


The genus Elaphropeza Macquart be- 
longs to the subfamily Tachydromiinae 
(Diptera: Empididae), and was previously 
considered a subgenus of Drapetis Meigen. 
It is very similar to Drapetis in having an 
indistinct gena, the first basal cell distinctly 
shorter than the second basal cell, and the 
long Rs, but it can be separated from Dra- 
petis by the hind tibia with ad (anterodorsal 
setae) and the usually yellow body. The ge- 
nus is distributed worldwide with 42 known 
species from the Oriental Region (Smith 
1975) when last cataloged, 17 from the 
Australasian and Oceanian Regions (Smith 
1989), and 3 from the Palaearctic Region 
(Chvéla and Kovalev 1989). Twenty-five 
species have since been reported from the 
Chinese mainland by Yang and Yang 


Empidoidea, Empididae, Tachydromiinae, Elaphropeza, China, new species 


(I989arebs 19905 19924° bs 1994572005): 
Yang et al. (2002), and Saigusa and Yang 
(2002), most of which were included in 
Drapetis (Elaphropeza). Some other major 
references dealing with Elaphropeza in- 
clude Bezzi (1904), Brunetti (1920), Melan- 
der (1927), Quate (1960), Smith (1965), 
and Chvala (1975). 

In this paper, we review the species of 
the genus Elaphropeza from the Chinese 
mainland. Two species are described as new 
to science, based on specimens collected by 
Shuwen An and Xingyue Liu with sweep 
nets and a light trap in Guangxi. Addition- 
ally, the following new combinations are 
made for the Chinese species which were 
previously included in Drapetis (Elaphro- 
peza).: E. centristria (Yang and Yang 2003), 


50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


E. fujianensis (Yang and Yang 2003), E. fu- 
zhouensis (Yang and Yang 2003), E. gu- 
angxiensis (Yang and Yang 1992a), E. 
guiensis (Yang and Yang 1989a), E. jiany- 
angensis (Yang and Yang 2003), E. jingh- 
ongensis (Yang and Yang 1990), E. lancan- 
gensis (Yang and Yang 1990), E. lii (Yang 
and Yang 1990), E. longiconica (Yang and 
Yang 1992b), E. medipunctata (Yang and 
Yang 1994), E. meihuashana (Yang and 
Yang 2003), E. paucipunctata (Yang and 
Yang 1989a), E. pilata (Yang and Yang 
1994), E. postnigra (Yang and Yang 1990), 
E. ruiliensis (Yang and Yang 1990), E. stri- 
ata (Yang and Yang 1992b), E. triangulata 
(Yang and Yang 1992b), E. xanthina (Yang 
and Yang 1990), E. xizangensis (Yang and 
Yang 1989b), and E. yunnanensis (Yang 
and Yang 1992b). A key to the species of 
the genus from the Chinese mainland is pre- 
sented for the first time. Holotypes and 
most paratypes of the new species are de- 
posited in the insect collection of China Ag- 
ricultural University (CAU), Beijing, and 
one paratype is deposited in the National 
Museum of Natural History (USNM), 
Washington, DC. 

Basic terminology follows McAlpine 
(1981) and Steyskal and Knutson (1981). 
The following abbreviations for setae are 
used: acr-acrostichal, ad-anterodorsal, av- 
anteroventral, d-dorsal, dc-dorsocentral, h- 
humeral, oc-ocellar, npl-notopleural, pd- 
posterodorsal, psa-postalar, pv-posteroven- 
tral, sa-supra-alar, sc-scutellar, and vt-ver- 
tical. 


KEY TO SPECIES OF ELAPHROPEZA FROM THE 
CHINESE MAINLAND 


1. Thorax yellow, usually with black spots ... 4 
— Thorax black 
E. henanensis Saigusa and Yang (2002) 
Antenna yellow 
3. Antenna yellow with Ist flagellomere brown- 
ish yellow; 1 vt. Guangxi ........... 
E. guangxiensis (Yang and Yang) 
— Antenna entirely yellow; 2 vt. Guangxi .... 
Se ereueear? E. liui Yang and Gaimari, new species 
4. Head black 
= aleadsyellow see Rene a uaeeeeiebe ce) ces conraiee 5) 


14. 


15. 


. First flagellomere brown. Hainan 


. Mesonotum not black posteriorly 


Thorax entirely yellow. Guangxi 
VAs bee Cerner E. pilata (Yang and Yang) 
Thorax with dark spots 
Mesonotum with brownish lateral spot. Fujian 

Bir eat cae oc E. fujianensis (Yang and Yang) 
Mesonotum without spots 
Scutellum and metanotum black medially; 
pleuron lacking spots. Yunnan 
LPNS roo, OP circu cee eee E. xanthina (Yang and Yang) 
Scutellum and metanotum brown except an- 
terolateral corner of metanotum yellow; lower 
portion of meron brown. Fujian 
E. jianyangensis (Yang and Yang) 
Sno cae 12 


Mesonotum with black spots 
Mesonotum entirely yellow 

Hind tibia with 1 ad. Yunnan 
E. ruiliensis (Yang and Yang) 
ind stibiamwathi2 ad seine eee 10 


. Metanotum entirely blackish. Yunnan, Sich- 


uan E. longiconica (Yang and Yang) 
Metanotum yellow with blackish median por- 
tion 


E. bisetifera Yang, Yang and Hu (2002) 
First flagellomere yellow. Guizhou 
E. paucipunctata (Yang and Yang) 
Rs 3. Bead c 15 


Mesonotum black posteriorly 


. Wing with wide brownish median spot; hind 


tiblalwathes ad Haman. ene eee 
E. alamaculata Yang, Yang and Hu (2002) 
Wing without spot; hind tibia with 1—2 ad 14 
Hind tibia with 1 ad. Yunnan......... 
Ro cea et ccne res et E. postnigra (Yang and Yang) 
Hind tibia with 2 ad. Yunnan......... 
E. yunnanensis (Yang and Yang) 
Mesonotum with brown median vitta, but 
without lateral spot. Fujian 
E. centristria (Yang and Yang) 
Mesonotum without median spot or vitta, 


sometimes with lateral spot ........... 16 
> labial (illo Wallin @ Cl 555500000 00005c0 20 
Eling stibiraswithel aden eee ae 17/ 


. Mesonotum with one rather large black lateral 


spot. Tibet ... 
Mesonotum with one small black lateral spot 
SRSA Bik: Makes e So Ay ae eee ee ee 18 


E. xizangensis (Yang and Yang) 


. Scutellum and metanotum yellow. Fujian. . . 


hae omeyho schasy co E. fuzhouensis (Yang and Yang) 
Scutellum and metanotum black ........ 19 


. First flagellomere yellow; scutellum entirely 


black; pleuron with spot on meron only. 
Guan exahsElain ane ee 
E. medipunctata (Yang and Yang) 
First flagellomere dark yellow; scutellum with 
yellow apical margin; pleuron with spots on 
anepimeron, meron and metapleuron. Yunnan 
E. jinghongensis (Yang and Yang) 


VOLUME 107, NUMBER 1 


2) wessrentirelyayellowsaneee cae oe eee 24 
— Legs partly dark yellow, brownish yellow, or 
Garkalbro within ses apeis ces tonto eR Se 21 


21. Arista pale; tibiae and tarsi dark yellow. Hai- 
nan .. E. pallidarista Yang, Yang and Hu (2002) 
— Arista black; tibiae and tarsi variable .... 22 
22. Femora greyish apically. Sichuan, Guizhou 
Dycacehte peace eet E. guiensis (Yang and Yang) 
—  FKemora entirely yellow .............. 23 
23. Metanotum yellow; fore and mid tibiae and 
all tarsi dark brown except tarsomere 5 black. 
(GUE GN = 2, Steer ine coer Crue 1A Hier 3. Men 
E. anae Yang and Gaimari, new species 
—  Metanotum brownish; legs only with fore tar- 
sus (except fore tarsomere 1) dark brownish 
yellow. Yunnan 


E. lancangensis (Yang and Yang) 
24. Metanotum yellow ................. 26 
— Metanotum blackish 
25. Lateral spot on mesonotum triangular. Yun- 
TAR te seas eerie E. triangulata (Yang and Yang) 
— Lateral spot on mesonotum stripe-like. Yun- 
nan, Fujian....... E. striata (Yang and Yang) 
26. Scutellum yellow; abdomen entirely brown- 
ish. Yunnan, Hainan ... E. lii (Yang and Yang) 
—  Scutellum brown medially; abdomen yellow 
with tergites 4—5 dark brown. Fujian 
E. meihuashana (Yang and Yang) 


Elaphropeza liui Yang and Gaimari, 
new species 
(Figs. 1-3) 


Diagnosis.—Head with 2 incurved vt. 
Antenna yellow; Ist flagellomere short, 
conical. Thorax black. Hind tibia with 1 ad. 

Male.—Body length 2.3 mm, wing 
length 3.0 mm. 

Head: Black with grey pollen; hairs and 
setae pale; eyes contiguous on face; ocellar 
tubercle with 2 long, strong oc and 2 short 
posterior hairs; 2 pairs of vt curved inward. 
Antenna yellow; scape bare, shorter than 
pedicel; pedicel with circlet of apical hairs; 
Ist flagellomere short, conical, 1.1 times 
longer than wide, short pubescent; arista 
long (6.0 times longer than Ist flagello- 
mere), black, short pubescent. Proboscis 
yellow with pale hairs; palpus yellow with 
4 brown hairs and 2 brown setae. 

Thorax: Black with gray pollen; hairs 
and setae pale; pleuron more or less sub- 
shining black; h absent, 2 long np/, | long 
sa, 1 very short psa, acr absent, 3 dc (pos- 


51 


teriormost dc longest); scutellum with two 
pairs of sc (basal pair very short, about % 
as long as apical pair). Legs yellow, except 
tarsomere 5 brown; hairs and setae brown; 
fore coxa with 2 d at base, apically with 3 
setae; mid coxa apically with 5—6 setae; 
fore and mid femora subequal in thickness, 
1.1 times thicker than hind femur, each with 
1 preapical seta, one row of av and one row 
of pv; fore tibia apically with 2 av and 1 
pv; mid tibia apically with 2 av and 2 pv; 
hind tibia with 1 brown ad at middle, api- 
cally with 3 short setae. Wing hyaline, veins 
brown. Calypter dark brown with black 
hairs. Halter yellow. 

Abdomen: Dark brown with grey pollen; 
hairs and setae blackish; tergites 1—3 nearly 
membranous, each with one long triangular 
lateral sclerite; tergite 4 enlarged but tergite 
5 narrowed, each with two groups of short 
black spines medially. Male genitalia (Figs. 
1—3): Left tergal lobe rather narrow basally, 
with its surstylus short and nearly quadrate 
with short obtuse apical corners; right tergal 
lobe fused with its surstylus, distinctly lon- 
ger than wide, with long setae on outer mar- 
gin, and with wide apex; left cercus long, 
fingerlike, acute apically; right cercus long, 
fingerlike, nearly acute apically. 

Female.—Unknown. 

Type material—Holotype d, deposited 
in CAU: Guangxi, Maoershan National Na- 
ture Reserve, Sanjiangyuan (1,900 m), 30 
June 2003 (light trap), Xingyue Liu. 

Etymology.—tThe species is named after 
the collector Mr. Xingyue Liu. 

Remarks.—The new species is somewhat 
similar to E. guangxiensis, from Guangxi. 
But in E. guangxiensis, the antenna is yel- 
low with the Ist flagellomere brownish yel- 
low, and only | vt is present (Yang and 
Yang 1992a). 


Elaphropeza anae Yang and Gaimari, 
new species 
(Figs. 4—6) 
Diagnosis.—Head black, with | incurved 
vt. Antenna brownish; Ist flagellomere 
blackish, long, conical. Thorax yellow, with 


Nn 
i) 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-6. 1—3, Elaphropeza liui, male. 1, Right tergal lobe. 2, Genitalia, dorsal view. 3, Left surstylus. 4—6, 
Elaphropeza anae, male. 4, Right tergal lobe. 5, Genitalia, dorsal view. 6, Left surstylus. 


a brown lateral spot on mesonotum. Fore 
and mid tibiae dark brown; hind tibia with 
2 ad. 

Male.—Body length 
length 2.0 mm. 

Head: Black with grey pollen; hairs and 
setae blackish; eyes contiguous on face; 
ocellar tubercle with 2 oc and 2 short pos- 
terior hairs; 1 vt curved inward, slightly 
longer than oc. Antenna brownish with Ist 


1.6 mm, wing 


flagellomere blackish; scape bare, shorter 
than pedicel; pedicel with circlet of apical 
hairs; Ist flagellomere long, conical, 3.0 
times longer than wide, short pubescent; 
arista long (2.0 times longer than Ist fla- 
gellomere), black, short pubescent. Probos- 
cis brownish yellow with black hairs; pal- 
pus yellow with blackish hairs. 

Thorax: Yellow with pale grey pollen; 
mesonotum with one brown lateral spot just 


VOLUME 107, NUMBER 1 


before wing base; hairs blackish, setae 
black; h absent, 2 np/ (posterior np/ longer), 
1 sa, 1 short psa, biseriate acr, 1 long 
strong dc; scutellum with two pairs of sc 
(basal pair very short, about 4 as long as 
apical pair). Legs yellow, except fore and 
mid tibiae and all tarsi dark brown except 
tarsomere 5 black; hairs and setae blackish; 
fore coxa with 2 d at base, apically with 2 
setae; mid coxa apically with 3 d; hind coxa 
with | outer seta at apical margin; fore and 
mid femora equal in thickness, 1.1 times 
thicker than hind femur; fore femur with 1 
long thin pv at extreme base; mid femur 
with | preapical anterior seta; hind femur 
with 3 d at base; fore tibia apically with 1 
short strong av and | short strong pv; mid 
tibia with one row of small black spines, 
apically with 2 av and | pv; hind tibia with 
2 ad, apically with 3 short weak setae. 
Wing nearly hyaline, veins brown. Calypter 
brown with black hairs. Halter dark yellow. 
Abdomen: Dark brown with grey pollen; 
hairs and setae blackish; tergites 1—3 nearly 
membranous except tergites 2—3 each with 
one small lateral sclerite; tergites 4—5 
blackish, each with one group of short 
black lateral spines. Male genitalia (Figs. 
4—6): Left tergal lobe rather narrow, with 
its surstylus long, finger-like, and curved 
inward apically; right tergal lobe rather 
large, fused with its apically widened sur- 
stylus; left cercus long and obtuse apically, 
right cercus short and obtuse apically. 
Female.—Body length 2.1—2.2 mm, wing 
length 2.1—2.3 mm. Similar to male, but 
scape and pedicel dark brownish yellow. 
Type material.—Holotype d, deposited 
in CAU: Guangxi, Maoershan National Na- 
ture Reserve (2100 m), 30 June 2003, Shu- 
wen An. Paratypes: Same data as holotype 
(2 2, CAU; 1 2, USNM); Guangxi, Maoer- 
shan National Nature Reserve (350 m), 27 
June 2003, Shuwen An, (1 2, CAU). 
Etymology.—The species is named after 
the collector Ms Shuwen An. 
Remarks.—The new species is somewhat 
similar to E. lancangensis from Yunnan. 
But in E. lancangensis, the metanotum is 


5)3) 


brownish, and the legs are yellow with only 
the fore tarsus (except fore tarsomere 1) 
dark brownish yellow (Yang and Yang 
1990). 


DISCUSSION 


Twenty-seven of the 69 Oriental species 
of the genus Elaphropeza are now known 
from the Chinese mainland, with 17 species 
endemic to the southwestern region of Chi- 
na, one endemic to the central region, and 
seven endemic to the southern region. Only 
two species, E. lii and E. striata are rela- 
tively widely distributed, being in both the 
southern and southwestern regions of Chi- 
na. The southwestern region of China ap- 
pears to be the center of diversity for Ela- 
phropeza on the Chinese mainland. 


ACKNOWLEDGMENTS 


Our sincere thanks are due to Ms. Shu- 
wen An and Mr. Xingyue Liu (China Ag- 
ricultural University, Beijing) for collecting 
specimens and for their help during this 
study, and to one or more anonymous re- 
viewers for their helpful comments. The re- 
search was funded by the National Natural 
Science Foundation of China (No. 
30070100, No. 30225009) and the Califor- 
nia Department of Food and Agriculture. 


LITERATURE CITED 


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Signor L. Bird. Annales Historico-Naturales Mu- 
sei Nationalis Hungarici 2: 320-361. 

Brunetti, E. 1920. Diptera Brachycera. The Fauna of 
British India, including Ceylon and Burma, Vol. 
1. Taylor and Francis, London, 401 pp. 

Chvala, M. 1975. The Tachydromiinae (Dipt. Empi- 
didae) of Fennoscandia and Denmark. I. Fauna 
Entomologica Scandinavica, Vol. 3. Scandinavian 
Science Press, Copenhagen, 336 pp. 

Chvala, M. and V. G. Kovalev 1989. Family Hyboti- 
dae, pp 174-227. In Soés, A. and L. Papp, eds. 
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sterdam and Budapest, 435 pp. 

McAlpine, J. E 1981. Morphology and Terminology— 
Adults, pp. 9-63. In McAlpine, J. E, B. V. Peter- 
son, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, 
and D. M. Wood, coords. Manual of Nearctic Dip- 


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tera, Vol. 1. Research Branch, Agriculture Canada, 
Ottawa. Monograph 27, 674 pp. 

Melander, A. L. 1927. Diptera. Fam. Empididae. Gen- 
era Insectorum, Fascicle 185. 434 pp. 

Quate, L. W. 1960. Diptera: Empididae. Insects of Mi- 
cronesia 13: 55-73. 

Saigusa, T. and D. Yang 2002. Empididae (Diptera) 
from Funiu Mountains, Henan, China (1). Studia 
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Smith, K. G. V. 1965. Diptera from Nepal. Empididae. 
Bulletin of the British Museum of Natural History 
(Entomology) 17: 63-112. 

. 1975. Family Empididae, pp. 185-211. In 

Delfinado, M. D., and D. E. Hardy, eds. A Catalog 

of the Diptera of the Oriental Region, Vol. 2. Uni- 

versity Press of Hawaii, Honolulu, 459 pp. 

. 1989. Family Empididae, pp. 382-392. In Ev- 
enhuis, N. L., ed. Catalog of the Diptera of the 
Australasian and Oceanian Regions. Bishop Mu- 
seum Press and E. J. Brill, Honolulu and Leiden, 
1,155 pp. 

Steyskal, G. C. and L. V. Knutson. 1981. 47. Empi- 
didae, pp. 607—624. In McAlpine, J. FE, B. V. Pe- 
terson, G. E. Shewell, H. J. Teskey, J. R. Vock- 
eroth and D. M. Wood, coords. Manual of Nearc- 
tic Diptera, Vol. 1. Research Branch, Agriculture 
Canada, Ottawa. Monograph 27, 674 pp. 


Yang, C. and D. Yang. 1992a. Three new species of 
Empididae from Guangxi—Diptera: Brachycera. 
Journal of the Guangxi Academy of Science 8(1): 
44-48. 

Yang, D. and C. Yang. 1989a. Four new species of 
dance flies from Guizhou Province (Diptera: Em- 
pididae). Guizhou Science 7(1): 36—40. 

. 1989b. The dance flies of Xizang (II) (Dip- 

tera: Empididae). Acta Agriculturae Universitatis 

Pekinensis 15(4): 415—424. 

. 1990. Eleven new species of the subfamily 

Tachydromiinae from Yunnan (Diptera: Empidi- 

dae). Zoological Research 11(1): 63-72. 

. 1992b. Diptera: Empididae, pp. 1089-1097. 

In Chen, S., ed. Insects of the Hengduan Moun- 

tains Region, Vol. 2. Science Press, Beijing. 

. 1994. Three new species of Maoer Mountain 

in Guangxi (Diptera: Empididae). Guangxi Sci- 

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. 2003. Empididae, pp. 258-265. Jn Huang, B., 
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PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 55-63 


FOSSIL TRIGONALIDAE AND VESPIDAE (HYMENOPTERA) IN 
BALTIC AMBER 


GEORGE POINAR, JR. 


Department of Zoology, Oregon State University, Corvallis, OR 97331-2907, U.S.A. 
(email: poinarg @casco.net) 


Abstract.—A fossil trigonalid (Trigonalidae: Hymenoptera), Eotrigonalis balticus Poin- 
ar, n. gen., n. sp., and a fossil vespid, Palaeovespa socialis Poinar, n. sp., are described 
from Eocene Baltic amber. Eotrigonalis balticus is a large, robust, heavily armored spe- 
cies, which could have parasitized members of the Vespidae. It can be separated from all 
other members of the family by the presence of large scutellar horns. Palaeovespa socialis 
has the diagnostic characters of members of the subfamily Vespinae and was probably 
eusocial. It differs from extant vespines by the shape of the clypeus, the presence of 


interparapsidal furrows and venational characters. 


Key Words: 
alis, Baltic amber 


The Trigonalidae are a monophyletic 
group of enigmatic parasitic wasps com- 
prising some 16 genera worldwide. They 
have unique morphological characters as 
well as a complex life history involving 
both a carrier and a developmental host 
(Carmean 1991, Carmean and Kimsey 
1998, Weinstein and Austin 1991). Fossil 
trigonalids are rare (Carpenter 1992, Ras- 
nitsyn and Quicke 2002) and the present 
study describes a new genus and species 
from Baltic amber. 

Social wasps belonging to the subfamily 
Vespinae of the family Vespidae are not 
commonly fossilized in amber (Carpenter 
1992, Rasnitsyn and Quicke 2002) and the 
present study describes a new species in 
Baltic amber closely related to extant mem- 
bers of the genus Vespula L. The fossil ves- 
pid is a potential developmental host for the 
Baltic amber trigonalid. 


MATERIALS AND METHODS 
The pieces of Baltic amber containing 
the trigonalid and vespid originated from 


Trigonalidae, Eotrigonalis baltica, Vespidae, Vespinae, Palaeovespa soci- 


the Kalinigrad region in Russia. Both pieces 
were recut and repolished for study. The fi- 
nal piece containing the trigonalid fossil 
weighed 1.6 grams, was 20 mm long, 13 
mm wide and 5 mm deep. The piece con- 
taining the vespid fossil weighed 2.6 grams, 
was 20 mm long, 19 mm wide and 11 mm 
in depth. Baltic amber has been dated at 
~ 40 million years (Eocene) [for a discus- 
sion of the age of these deposits, see Poinar 
(1992) and Larsson (1978)]. Observations 
and photographs were made with a Nikon 
Optiphot microscope and a Nikon SMZ- 
10X stereoscopic microscope at magnifi- 
cations of 800. Terminology follows that 
presented by Huber and Sharkey (1993), 
Mason (1993), Brothers and Finnamore 
(1993) and Duncan (1939) with some tra- 
ditional venation terminology as used by 
Michener et al. (1994). All measurements 
are in millimeters unless otherwise noted. 


Trigonalidae Cresson, 1887 
The Baltic fossil possessed the following 
characters, which are diagnostic for the 


56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


family Trigonalidae (Carmean 1991, Car- 
mean and Kimsey 1998, Mason 1993): 23 
antennal segments; the presence of groups 
of white setae on the outside of the middle 
antennal segments, fore wing with a stigma 
and 10 closed cells, Cu deflected abruptly 
posteriorly at the base of 1 m-cu, vein 2 cu- 
a less than half as long as vein | cu-a, veins 
2 r-m and 2 m-cu present, veins C and R 
separate, forming a long narrow costal cell; 
hind wing with 2 closed cells; claws cleft 
with arolia; apicoventral plantar lobes pre- 
sent on tarsomeres 1—4, hind trochantellus 
divided; metasoma pedunculate and ovipos- 
itor reduced. 

This specimen is nearly complete, with 
only the apical portions of the right fore and 
hind wings, a portion of one antenna and 
the tarsi of the right hind leg missing. Por- 
tions of the head are covered with bubbles, 
which make examination of the mouthparts 
difficult. 


Eotrigonalis Poinar, new genus 


Type species: Eotrigonalis baltica Poinar. 

Description.—Body large, length, 11.3; 
antenna with 24 segments; lateral poste- 
rior borders of scutellum each with a large 
horn; crossvein | cu-a straight, meeting M 
vein. 

Diagnosis.—Only on some extant speci- 
mens indicated as belonging to the genus 
Xanthogonalos Schulz (1907) and on the 
Early Cretaceous Albiogonalys elongatus 
Nel et al. (2003) does crossvein | cu-a meet 
M vein. Other venational differences and 
the large horns on the scutellum separate E. 
balticus from all known extant and extinct 
genera. 

Etymology.—Eo is from the Greek 
eos” for dawn, early. This gender is mas- 
culine. 


oe 


Eotrigonalis balticus Poinar, 
new species 
(Figs.1—3, 6-10) 


Description.—Holotype female; with 
characters listed under generic description. 
Head: Large, 2.9 long (without mandi- 


bles), setae absent except for very short 
hairs on clypeus; eyes protruding, length 
eye, 1.8; width eye, 1.2; torulus located be- 
neath a ridge below middle point of com- 
pound eye; malar space short; mandibles 
toothed, asymmetrical, left mandible with 3 
teeth, right mandible with 4 teeth; palpal 
segments obscured by debris; antenna 8.7 
long, with 24 segments which taper toward 
apex; lengths of segments as follows: 1, 
O4o8 2, W253 3, OL50% 4 W108 SD, 0.7102 ©, 
Osa 7, O02 3, O43 2, O4B IMO, OAs Ili, 
355 2, W302 13, OL308 14. 0.253 IS), ©)3/0? 
IG; OL.B08 17, O.30s Mh, O.252 9, O25; 20, 
O73 Zl, O22 ZZ, 0203 235 WL i53 24, 0-353 
outer areas of antennal segments 9—20 with 
patches of raised white setae (or scales). 
Mesosoma: 3.3 long; pronotum short, 
not visible from above, length mesoscutum, 
1.5; width mesoscutum, 3.1; length scutel- 
lum, 1.8; width scutellum, 1.8; length pro- 
podeum 0.6, width propodeum, 1.2; two 
slightly oblique vertical rows of foveae oc- 
cur on mesoscutum, latero-posterial corners 
of mesoscutum projecting outward; a single 
transverse row of foveae along anterior 
margin and a medial vertical row of foveae 
on scutellum; latero-posterior corners of 
scutellum bearing large robust horns; me- 
sopleuron large, with medial vertical row of 
foveae; fore and hind coxae contiguous, 
mid coxae slightly separated; hind leg with 
divided trochantellus; leg measurements: 
fore leg, coxa, 0.88; trochanter, 0.74; femur, 
0.17; tibia, 1.4; middle leg, coxa, 0.97; tro- 
chanter, 0.56;. femur, 2.64; tibia, 2.3; hind 
leg, coxa, 1.32; trochanter, 0.65; femur 
(with trochantellus); 3.34; tibia, 3.14; tarsal 
Segment lensthss toreless lle O97 2. 
0.44; T3, 0.23; T4, 0.21; T5, 0.44; middle 
leg Wil, MelWs WZ, O-2lle 13, Oss5e 104), O.268 
WD; O.53 laine! les, Wily oS; 02, O.s0s WS; 
0.38; T4, 0.29; TS, 0.44; tarsomeres 1—4 on 
all legs with apicoventral plantar lobes; 
paired claws cleft, large arolia; wings hya- 
line, membrane bare; length forewing, 11.0; 
length hind wing, 7.6; venation as in 
Figs.7—8; in hind wing, Rs not reaching 
wing margin, distal part of M not reaching 


VOLUME 107, NUMBER 1 7 


Figs. 1-3. Eotrigonalis balticus in Baltic amber. 1, Lateral-dorsal view. Note attenuated antennal segments. 
Scale bar = 3.1 mm. 2, Head and mesosoma. Note horn (arrow) on scutellum. Scale bar = 1.0 mm. 3, Apico- 
ventral plantar lobes (arrows) on first 4 tarsal segments of right leg 3. Scale bar = 0.5 mm. 


58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


wing margin, distal part of Cu reaching 
wing margin, cu-a short, oblique, 10 hamuli 
present, positioned in distal half of wing, as 
shown in Fig. 8. 

Metasoma: 5.5 long; greatest width, 4.5; 
tergites with light and dark zones; dark 
zones confined to bands along posterior 
borders of tergites 2—7; small ovipositor an- 
terior of cerci. 

Material examined.—Holotype female in 
Baltic amber, deposited in the Poinar col- 
lection (accession # Hy-10—180) main- 
tained at Oregon State University. 

Etymology.—The name is derived from 
Baltic amber. 

Discussion.—This is the first described 
Tertiary trigonalid. Several Cretaceous tri- 
gonalids have been described, but accord- 
ing to Nel et al. (2003), all of these have 
uncertain family affinities except Albiogon- 
alys elongatus Nel et al. (2003), and pos- 
sibly Cretogonalys taimyricus Rasnitsyn 
(1977). Wing venation, mesosomal arma- 
ture, number of antennal segments and size 
separate E. balticus from all previously de- 
scribed extinct and extant trigonalids. 

The large size, thick cuticle and armature 
of E. balticus suggest that it was parasitic 
on an aggressive insect, possibly a member 
of the Vespidae. Its size, mesosomal arma- 
ture and wing venation is similar to that of 
extant members of Bareogonalos canaden- 
sis (Harrington 1896), a Pacific Northwest 
species that parasitizes yellow jackets (Car- 
mean 1991). Possible hosts for E. balticus 
might have been members of the genus Pa- 
laeovespa, one of which is described below. 
There is only one extant species of trigon- 
alid in Europe, Pseudogonalos hahnii (Spi- 
nola), which has been reared from Lepidop- 
tera pupae that have been parasitized by 
Ichneumonidae and Diprionidae (Carmean 
and Kimsey 1998). Wing venation and scu- 
tellar horns separate E. balticus from P. 
hahnii. 


Vespidae Leach, 1815 


This specimen is complete and clearly 
visible. The hind wings are located under 


sy =: 

Figs 4-5. Paleovespa socialis in Baltic amber. 4, 
Lateral view. Note yellow bands (arrows) on distal 
edges of metasomal tergites. Scale bar = 2.5 mm. 5, 


Face view showing yellow areas between toruli and on 
scapes and clypeus. Scale bar = 1.2 mm. 


the fore wings and it is difficult to delineate 
their veins, however no jugal lobe could be 
detected. This character, along with the ses- 
sile metasoma, abruptly declivous tergum 1, 
absence of parategula, simple, smooth tarsal 
claws, fore wing with first subdiscal cell 


VOLUME 107, NUMBER 1 


59 


Figs. 6-9. 


neither narrowed nor projecting apically, 
metacoxae with a dorsal longitudinal cari- 
na, fore wing with vein 1 cu-a (cu-v) 
straight and much shorter that vein 2/1A 
and with the apex of the marginal cell not 
separated from anterior margin of wing, 
place the specimen in the eusocial subfam- 
ily Vespinae (Brothers and Finnamore 
1993). 

Characters which place the specimen in 
the genus Palaeovespa Cockerell 1906 are 
the oblique apex of the first discoidal cell, 
the recurrent veins joining the second sub- 
marginal cell near its ends and the basal 
vein joining the costal vein near the stigma. 
The genus Palaeovespa was erected for 
several species described from the Floris- 
sant deposits (Cockerell 1906). Later Cock- 
erell (1909) described P. balticus from Bal- 
tic amber, assigning it to this genus on the 
basis of similar wing characters. Since the 
present fossil possesses similar venational 
characters, it is placed in Palaeovespa. 


Eotrigonalis balticus in Baltic amber. 6, Head. Scale bar = 0.6 mm. 7, Fore wing. Scale bar = 
1.4 mm. 8, Hind wing. Scale bar = | mm. 9, Penultimate tarsal segment of left fore tarsus with cleft claws and 
empodium. Scale bar = 63 pm. 


Palaeovespa socialis Poinar, — 
new species 
(Figs. 4-5, 11-14) 


Description.—Holotype female; total 
length, 12.8. Head: Length, 2.6 (without 
mandibles), head bearing long setae (except 
eyes), greatest width head, 3.5, clypeus 
wider (1.6) than high (1.2), light in color, 
basal margin convex with small medial pro- 
jection; ocelli positioned close to occiput; 
antenna 12 jointed, segments thick, third 
segment nearly as long as scape; compound 
eyes deeply emarginate; light (yellow) area 
between toruli; malar space short; mandi- 
bles with three sharp teeth and an inner mo- 
lar area; a light yellow genal band extends 
a short distance behind each eye; scapes 
and clypeus yellow. 

Mesosoma: 3.8 long, bearing long setae; 
black except for following light areas: an- 
terior border of mesoscutum, edges of me- 
tanotum (a pair of light areas), scutellum 


60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 10-14. Fossil Trigonalidae and Vespidae in Baltic amber. 10, Mesoscutum and scutellum of Eotrigon- 
alis balticus. Scale bar = 1 mm. 11, Dorsal view of head and mesosoma of Palaeovesoa socialis (setation not 
shown). Arrows show interparapsidal furrows. Scale bar = 1 mm. 12, Simple tarsal claws of the left mesotarsus 
of P. socialis with basal bristles and empodium. Scale bar = 63 wm. 13, Face view of P. socialis. Scale bar = 
1 mm. 14, Fore wing of P. socialis. B = basal vein, C = costal vein, 1D = discal cell, IR = first recurrent 
vein, 1SD = first subdiscal cell, 2R = second recurrent vein, 2S = second submarginal cell, 2te = second 
transverse cubital vein. Scale bar = 0.9 mm. 


(two light areas) and sides of propodeum; er furrows between parapsidal furrows 
mesepisternum with a pair of light spots on (termed interparapsidal furrows); length 
sides; mesoscutum with a mesidan notal su- fore wing, 8.5; apex of first discoidal cell 
ture, two parapsidal furrows and two short- oblique; basal vein meets costal vein near 


VOLUME 107, NUMBER 1 


stigma, distance from apex of basal vein to 
stigma, 135 jm, wing membrane covered 
with minute setae; legs with yellow and 
black markings; leg measurements: fore 
leg, coxa, 1.0; trochanter, 0.4; femur, 2.0; 
tibia, 1.9; tarsus 1, 1.2; (rest of tarsus not 
visible); middle leg, coxa, 0.5; trochanter, 
0.3; femur, 2.2; tibia, 1.8; tarsal segments, 
Til, W@s 22; O33 U3s Ose WZ, OB! US. O68 
hind leg, coxa, 0.8; trochanter, 0.4; femur, 
1.6; tibia, 2.2; tarsal segments, T1, 2.8; T2, 
0.4; T3, 0.3; T4, 0.4; T5, 0.4; tarsal claws 
simple, each with a basal seta; empodium 
large. 

Metasoma: Length, 6.4; tergites with 
proximal portions black, distal areas yel- 
low; stinger protruding. 

Material examined.—Holotype female in 
Baltic amber, deposited in the Poinar amber 
collection (accession # H-10-175) main- 
tained at Oregon State University. 

Etymology.—The name socialis refers to 
the likely social habit of the specimen. 

Diagnosis.—This specimen has many 
features of extant members of the genus 
Vespula, including the short malar space, 
ocelli located at edge of vertex, eyes deeply 
incised, longitudinal carina on the dorsal 
side of the metacoxa and coloration (Dun- 
can 1939). While the color patterns are 
mainly light and dark, yellow shades in 
many of the light areas suggest that the true 
colors were black and yellow. The diagnos- 
tic characters mentioned above, especially 
the absence of parategula and simple, 
smooth tarsal claws, place P. socialis in the 
Vespinae. 

Earlier reports of Vespidae in Baltic am- 
ber include Vespa dasypodia Menge 
(1856), the description of which is limited 
to roughly a half page, without illustrations. 
The location of this specimen is unknown. 
However several characters separate this 
species from P. socialis. In V. dasypodia, 
the ocelli are arranged in almost a straight 
line, which differs from the triangular po- 
sition in P. socialis. Also V. dasypodia has 
reddish- yellow hairs on the ventral surface 
of the first tarsal segment of the fore leg 


61 


and hirsute hairs on the first tarsal segments 
of the middle and hind legs. Such hairs are 
lacking in the present specimen. Menge 
(1856) also stated that the posterior portions 
of the 4'", 5" and 6 abdominal sternites of 
V. dasypodia are covered with short bris- 
tles, which is not the case in P. socialis. 

The second Baltic amber vespid was a 
brief description of Palaeovespa_ baltica 
Cockerell (1909). Cockerell used venational 
characteristics to align P. baltica with the 
Nearctic Palaeovespa species, however, it is 
questionable whether the former species be- 
longs to the same genus as the New World 
fossils. 

Size differences between P. baltica and 
P. socialis do occur (P. baltica is 16 mm 
in length with a wing length of 11 mm in 
contrast to 12.8 mm and 8.5 mm, respec- 
tively, for P. socialis) but they could rep- 
resent caste or individual differences. 
However coloration differs between the 
two species. In P. baltica the apical portion 
of the abdominal segments, the venter, legs 
and wings are ferruginous, which is not the 
case in P. socialis. There are also wing 
venational differences. In P. socialis, the 
end of the second transverse cubital vein 
turns abruptly and meets the marginal cell 
at a right angle. This angle is oblique in P. 
baltica. The distance from the apex of the 
basal vein to the stigma is 425 pm in P. 
baltica, but only 225 pm in P. socialis. A 
major difference between P. socialis and 
modern vespines is the shape of the clyp- 
eus and the presence of interparapsidal fur- 
rows on the mesoscutum, but these char- 
acters were not included in the description 
of P. baltica. 

The species of Palaeovespa from the 
Florissant shales (Cockerell 1906, 1917, 
1923) are described mainly on size and col- 
or variations, which could indicate caste, 
colony or individual differences. The Flor- 
issant fossils differ from P. socialis in both 
size and coloration. Lengths are 14.5 mm 
for P. gillettei Cockerell, 18 and 22 mm for 
two specimens of P. scudderi Cockerell, 25 
mm for P. florissantia Cockerell and 17.5 


62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mm for P. relecta Cockerell. Regarding 
coloration, P. gillettei has two longitudinal 
yellow stripes on the mesothorax and a dark 
area on the apical part of the costal cell, 
which is lacking in P. socialis. Palaeovespa 
scudderi has apical light areas on only the 
last 2 abdominal segments (all abdominal 
segments have apical light areas in P. so- 
cialis), P. florissantia has no distinct ab- 
dominal markings but reddish wings and P. 
relecta has the head and thorax black, the 
first two abdominal segments pallid, with 
small lateral dark markings, and abdominal 
segments 3 to 5 with broad dark bands. 
These color patterns differ from those on P. 
socialis. Unfortunately very few body char- 
acters are described in the Florissant spec- 
imens, including the shape of the clypeus 
and presence of interparapsidal furrows on 
the mesoscutum. 

Discussion.—The present fossil is con- 
sidered eusccial since it possesses the basic 
characters of present day Vespinae, all 
members of which are eusocial today 
(Brothers and Finnamore 1993). The most 
obvious difference between P. socialis and 
extant Vespinae is the shape of the clypeus 
and the presence of interparapsidal furrows 
on the mesoscutum (Duncan, 1939). In ad- 
dition, M ,,, 1s much more oblique than on 
extant Vespines and the second recurrent 
vein is perpendicular to the second sub- 
marginal cell in P. socialis but oblique in 
extant vespines (Duncan 1939). The man- 
dibles of P. socialis are clenching a morsel 
of tissue with crochets, thus indicating that 
caterpillars were used as a protein source 
for the larvae. 


ACKNOWLEDGMENT 


I thank Roberta Poinar for comments on 
an earlier draft of this manuscript. 


LITERATURE CITED 


Brothers, D. J. and A. T. Finnamore. 1993. Chapter 8. 
Superfamily Vespoidea, pp. 161—278. Jn Goulet, 
H. and J. T. Huber, eds. Hymenoptera of the 
World: An Identification Guide to Families. Pub- 
lication 1894/E, Research Branch, Agriculture 
Canada, Ottawa, 668 pp. 


Carmean, D. 1991. Biology of the Trigonalyidae (Hy- 
menoptera), with notes on the vespine parasitoid 
Bareogonalos canadensis. New Zealand Journal 
of Zoology. 18: 209-214. 

Carmean, D and L. Kimsey. 1998. Phylogenetic revi- 
sion of the parasitoid wasp family Trigonalidae 
(Hymenoptera). Systematic Entomology 23: 35— 
76. 

Carpenter, E M. 1992. Superclass Hexapoda, pp.468— 
495. In Kaesler, R. L., ed. Treatise on Invertebrate 
Paleontology, Arthropoda 4, Part R, Vol. 4, The 
Geological Society of America, Inc., Boulder. 

Cockerell, T. D. A. 1906. Fossil Hymenoptera from 
Florissant, Colorado. Bulletin of the Museum of 
Comparative Zoology 50: 33-58. 

. 1909. Descriptions of Hymenoptera from Bal- 

tic amber. Mitteilungen aus dem Geologisch-Pa- 

laeontologischen Institut und der Bernsteinsamm- 

lung der Universitét K6nigsberg 50: 1—20. 

. 1917. Descriptions of fossil insects. Proceed- 

ings of the Biological Society of Washington 30: 

79-81. 

. 1923. Two fossil Hymenoptera from Floris- 
sant (Vespidae, Megachilidae). Entomological 
News 24: 270-271. 

Duncan, C. D. 1939. A contribution to the biology of 
North American vespine wasps. Stanford Univer- 
sity Publications in the Biological Sciences 8: 1— 
272. 

Huber, J. T. and M. J. Sharkey. 1993. Chapter 3. Struc- 
ture, pp. 13-59. Jn Goulet, H. and J. T. Huber, eds. 
Hymenoptera of the World: An Identification 
Guide to Families. Publication 1894/E, Research 
Branch, Agriculture Canada, Ottawa, 668 pp. 

Larsson, S. G. 1978. Baltic Amber—A Palaeobiolog- 
ical Study. Entomonograph, Vol. 1. Klampenborg, 
Denmark, 192 pp. 

Mason, W. R. M. 1993. Chapter 11. Superfamilies 
Evanioidea, Stephanioidea, Magalyroidea, and 
Trigonalyoidea, pp. 510—520. Jn Goulet, H. and J. 
T. Huber, eds. Hymenoptera of the World: An 
Identification Guide to Families. Publication 
1894/E, Research Branch, Agriculture Canada, 
Ottawa, 668 pp. 

Menge, A. 1856. Lebenszeichen vorweltlicher, im 
bernstein eingeschlossener thiere. Programm Of- 
fentlichen Priifung der schiiler der Petrischule, 
Danzig. A. W. Kafemann, 42 pp. 

Michener, C. D., R. J. Mcginley, and B. N. Danforth. 
1994. The bee genera of North and South America 
(Hymenoptera: Apoidea). Smithsonian Institution 
Press, Washington, DC, 209 pp. 

Nel, A., V. Perrichot and D. Néraudeau. 2003. The 
oldest trigonalid wasp in the Late Albian amber 
of Charente-Maritime (SW France) (Hymenop- 


tera: Trigonalidae). Eclogae geologica Helvetica 
96: 503-508. 


VOLUME 107, NUMBER 1 63 


Poinar, Jr., G. O. 1992. Life in Amber. Stanford Uni- Fasicle 61. Verteneuil et Desmet, Brussels. 24 
versity Press, Stanford, California, 350 pp. pp. 

Rasnitsyn, A. P. and D. L. J. Quicke, eds. 2002. His- Weinstein, P. and A. D. Austin. 1991. The host rela- 
tory of Insects. Kleuver Academic Publishers. tionships of trigonalyid wasps (Hymenoptera: Tri- 
Dordrecht, 517 pp. gonalyidae), with a review of their biology and 

Schulz, W. A. 1907. Hymenoptera. Fam. Trigona- catalogue to world species. Journal of Natural His- 


loidea. In Wytsman, P. ed. Genera Insectorum, tory 25: 399-433. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 64-70 


EXTIRPATION OF A POPULATION OF CICINDELA PATRUELA DEJEAN 
(COLEOPTERA: CARABIDAE: CICINDELINDT IN SUBURBAN 
WASHINGTON, D.C., USA 


JONATHAN R. MAWDSLEY 


Department of Entomology, National Musuem of Natural History, Smithsonian Insti- 
tution, Washington, DC 20560-0187, U.S.A. (e-mail: jonathan.mawdsley @nfwf.org) 


Abstract.—A population of Cicindela patruela patruela DeJean (Coleoptera: Carabidae: 
Cicindelini) existed until at least 1950 in a small area of eastern Washington, D.C., and 
adjacent Prince George’s County, Maryland, USA. Suitable habitat for this population 
consisted of open sandy barrens with soils derived from Cretaceous sediments of the 
Potomac Group and vegetation characterized by oak and pine species, particularly Quercus 
marilandica Miinchhausen and Pinus rigida Miller. This habitat was eliminated in the 
Washington area by extensive suburban housing construction, which was driven by rapid 
growth in the human population. Recommendations are provided for the reintroduction 


of C. patruela and for the restoration of suitable habitat at remnant natural areas. 


Key Words: 
agement 


Washington, D.C., and its suburbs are 
among the most richly sampled areas for 
tiger beetles in the world. Over the past 120 
years, scientists from the Smithsonian In- 
stitution’s National Museum of Natural His- 
tory (NMNH) and the U.S. Department of 
Agriculture’s Systematic Entomology Lab- 
oratory have repeatedly collected speci- 
mens of these beetles. A recent inventory 
by the author of material preserved in the 
NMNH collections found that 1,105 tiger 
beetle specimens, representing 14 species, 
had been collected since 1885 in the Dis- 
trict of Columbia and adjacent municipali- 
ties in Maryland and Virginia. 

Of these fourteen species, only three are 
still widespread and abundant in the Wash- 
ington suburbs (C. punctulata Olivier, C. 
repanda DeJean, and C. sexguttata Fabri- 
cius), and several species have probably 
been extirpated from this area (Glaser 1984, 
Knisley and Schultz 1997, Mawdsley, un- 


Cicindela patruela, tiger beetle, extirpation, conservation, restoration, man- 


published data). Glaser (1984) suggested 
that the declines of certain tiger beetle spe- 
cies in the Washington metropolitan area 
may be due to a loss of habitat resulting 
from rapid urban and suburban develop- 
ment. 

To test this hypothesis, I analyze data 
here from museum specimens of Cicindela 
patruela patruela DeJean. From 1918 to 
1950, adults of this species were routinely 
collected in a small area of the eastern Dis- 
trict of Columbia and adjacent parts of 
Prince George’s County, Maryland. Cicin- 
dela patruela has not been found in the 
Washington area since 1950 (Glaser 1984, 
1995; Knisley and Schultz 1997), despite 
intensive collecting efforts over the past fif- 
ty years. 

Cicindela patruela is distributed widely 
throughout eastern North America (Pearson 
et al. 1999) but occurs in small localized 
populations throughout much of its range 


VOLUME 107, NUMBER 1 


(Glaser 1995, Willis 2001). Populations of 
C. patruela are monitored by state conser- 
vation agencies (e.g., New York State De- 
partment of Environmental Conservation 
2003, Pennsylvania Department of Conser- 
vation and Natural Resources 2003). In ad- 
dition, C. patruela is currently listed as a 
species of special concern by the states of 
Massachusetts (Massachusetts Division of 
Fisheries and Wildlife 2003), Minnesota 
(Minnesota Department of Natural Re- 
sources 2003), and Wisconsin (Wisconsin 
Department of Natural Resources 2003). 
Given the current conservation interest in 
C. patruela and other tiger beetles (Knisley 
and Schultz 1997, Pearson and Vogler 
2001), it was felt that a more extensive ac- 
count of this population and its demise 
would be appropriate and relevant to con- 
servation efforts. 

Data associated with museum specimens 
and other historical data also allow the re- 
construction of some details of the biology 
of this population, including its seasonal cy- 
cle and associated vegetative communities 
and soil types. Although aspects of the bi- 
ology of C. patruela have been described 
by Lawton (1970), Boyd (1978), Knisley et 
al. (1990), and Willis (2000, 2001), uncer- 
tainties still exist regarding the habitat as- 
sociations of piedmont and coastal plain 
populations (Knisley and Schultz 1997). 


MATERIALS AND METHODS 


As part of the larger tiger beetle inven- 
tory effort described above, pinned adult 
specimens of C. patruela were examined in 
the collection of the National Museum of 
Natural History, Smithsonian Institution. 
Complete specimen label data were record- 
ed from specimens collected in Virginia, 
Maryland, Pennsylvania, and the District of 
Columbia; data relevant to this paper are 
given in the Appendix. Collecting localities 
were identified on paper and digital maps 
of the Washington, D.C., region. 

For purposes of this paper, the Washing- 
ton, D.C., metropolitan area is defined to 
include the District of Columbia and adja- 


ON 
nn 


cent municipalities in Maryland (Montgom- 
ery and Prince George’s counties) and Vir- 
ginia (Arlington and Fairfax counties and 
the City of Alexandria). 

Information on historical vegetation and 
statistics on housing development in the 
town of Cheverly, Maryland, were obtained 
from Bellamy (2000). Historical census 
data for Prince George’s County, Maryland, 
were obtained from the online databases of 
the Maryland State Data Center (2003) and 
the U.S. Census Bureau (2004). 

Information regarding recent collections 
of tiger beetles in Prince George’s County, 
Maryland, was provided by Warren E. 
Steiner, Jr. Additional information was ob- 
tained by the author through visits to nat- 
ural areas in Prince George’s County and 
the District of Columbia between 2000 and 
2004. 


ANALYSIS OF EXTIRPATION 


Cicindela patruela is undoubtedly extir- 
pated from the Washington metropolitan 
area, aS specimens of this species have not 
been collected in Prince George’s County 
or the District of Columbia since 1950. The 
tiger beetle fauna of Prince George’s Coun- 
ty was extensively collected by D. G. Shap- 
pirio in the early 1950s and by G. Hevel 
and W. E. Steiner, Jr., in the late 1960s and 
early 1970s. Specimens from these collect- 
ing efforts are preserved in NMNH and do 
not include any representatives of C. pa- 
truela. More recent surveys by the author 
and others at remnant natural areas within 
the historic distribution of C. patruela have 
failed to locate any individuals of this spe- 
cies. Warren E. Steiner, Jr. has collected ti- 
ger beetles in Cheverly, Maryland (site of 
the largest historic population of C. patrue- 
la in the Washington area), since 1991 and 
has never encountered this species in Chev- 
erly. 

The period of decline and extirpation of 
C. patruela corresponds to a period of in- 
tense growth in both the human population 
and the number of houses in Prince 
George’s County (Bellamy 2000). Table 1 


66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Comparison by decade of the rate of growth of the human population in Prince George’s County, 
Maryland, the number of new houses built in the Town of Cheverly, Maryland, and the numbers of museum 
specimens of Cicindela patruela collected from Prince George’s County and the Town of Cheverly. 


Rate of Growth— Number of New Specimens— 
Human Population, Specimens— Houses Built, Total for 
Years Prince George’s Co. Total for County Town of Cheverly Cheverly 
1901-1910 20.90% 0) 0 0 
1911-1920 19.92% 8 0) 0 
1921-1930 38.64% 0) 13 0) 
1931-1940 48.91% 42 207 27 
1941-1950 117.0% 3 731 0) 
1951-1960 84.05% 0) 400 0) 
1961-1970 84.83% 0) 504 0) 
1971-1980 0.6818% 0 165 0) 
1981-1990 9.545% 0 103 0) 


compares C. patruela collection records 
with the human population growth rate in 
this county. The last collections of C. pa- 
truela were in 1948, 1949, and 1950, at the 
end of a ten-year period in which the coun- 
ty’s population grew by 117%, the highest 
growth rate for any decade in the twentieth 
century. 

Obviously these incoming residents re- 
quired new houses. Data on housing con- 
struction were available for the Town of 
Cheverly, site of the largest C. patruela 
population in the Washington area (Table 
1). As would be expected, the number of 
new houses is strongly correlated with the 
human population growth rate (from simple 
linear regression, P = 0.0006). The C. pa- 
truela population in Cheverly was first en- 
countered by collectors in 1931 and was 
last sampled in 1934. During the decade 
1931-1940, the number of houses in Chev- 
erly nearly quadrupled, from 74 to 281 
units. Further construction and associated 
landscaping of an additional 1,635 houses 
during the period 1941—1970 eliminated the 
remaining open barrens habitat in Cheverly 
(Bellamy 2000). 

Other factors that have been proposed as 
causes of tiger beetle population declines 
include trampling by pedestrians or vehi- 
cles, over-collecting, and pesticide spraying 
(Knisley and Schultz 1997). There is no ev- 
idence that trampling or pesticide spraying 
would have played a role in the decline of 


C. patruela in the Washington area, and it 
is unlikely that the small numbers of spec- 
imens that were collected (54 specimens 
collected over 32 years) would have had a 
significant impact on the larger population. 
Knisley and Schultz (1997) review the 
available scientific evidence supporting var- 
ious hypotheses which have been proposed 
to explain tiger beetle declines and con- 
clude that habitat destruction and degrada- 
tion are easily the most important factors 
driving population declines. 

In summary, the available evidence in- 
dicates that the extirpation of C. patruela 
from the Washington metropolitan area was 
caused by a loss of suitable habitat due to 
suburban housing construction, which was 
driven in turn by substantial growth in the 
human population. 


HABITAT CHARACTERIZATION 


The historical collecting sites of C. pa- 
truela in the District of Columbia and west- 
ern Prince George’s County are all clustered 
on low hills overlooking the confluence of 
several streams to form the Anacostia River. 
It is interesting to note that C. patruela was 
not collected at other localities in the Wash- 
ington area which were well-sampled for ti- 
ger beetles during the first half of the 20th 
century, such as Plummers Island in Mary- 
land (5 species of tiger beetles recorded), 
Rock Creek Park in the District of Colum- 
bia (8 species recorded), and Falls Church 


VOLUME 107, NUMBER 1 


Table 2. 


67 


Numbers of tiger beetle specimens in NMNH collected at sites with Cicindela patruela in the 


District of Columbia and Prince George’s County, Maryland. 


D.C. 


Woodridge Bladensburg 


. patruela DeJean 

. punctulata Olivier 

. purpurea Olivier 

. repanda DeJean 

. rufiventris DeJean 

. scutellaris Say 

. sexguttata Fabricius 
. splendida Hentz 

. tranquebarica Herbst 
. unipunctata Fabricius 
Total by site 


AAAAAAYA OG © 
RORODFPOFPF OF 
ROhPNOONOOUW 


— 
— 


Maryland 
Cheverly College Park Hyattsville Riverdale Totals 
27 1] 8 3)3) 
] 0) 0) 0) i 
19 10 4 0) 34 
l ) 0) 2 9 
O D, 0) O 2 
4 8 2 l 16 
14 3 0) 4 29 
0) 10) 0) 0) l 
5 2 2 18 
2 | O 0) 3 
73 40 16 9 166 


(5 species recorded) and Mount Vernon (6 
species recorded) in Virginia. The absence 
of historical specimens from these well- 
sampled localities suggests that the distri- 
bution of C. patruela was quite restricted, 
even in the early 20th century. 

Comparisons with the geological map of 
Maryland (Maryland Geological Survey 
1968) indicate that the historical collecting 
sites discussed here are all located on soils 
derived from the Cretaceous Potomac 
Group, characterized as “interbedded 
quartzose gravels; protoquartzitic to ortho- 
quartzitic argillaceous sands; and white, 
dark gray and multicolored silts and clays.” 
This formation extends in a northeastern di- 
rection into Anne Arundel and Baltimore 
counties, paralleling the historic distribution 
of C. patruela on Maryland’s coastal plain 
(as mapped by Knisley and Schultz 1997). 
The limited ecological information on mu- 
seum specimen labels indicates an associa- 
tion with sandy soils, similar to that found 
elsewhere in this species’ range (Knisley et 
al. 1990, Willis 2000). 

Bellamy (2000) characterized the vege- 
tation of Cheverly, Maryland, as “locust 
and scrub pine”’ that had grown up on soils 
depleted of nutrients by past tobacco farm- 
ing practices. Museum specimen labels sug- 
gest that the historic collecting sites were 
open, sandy barrens with scattered pines 
and deciduous trees. Based on current veg- 


etation at the collecting sites, tree species 
present in these barrens would have includ- 
ed blackjack oak (Quercus marilandica 
Miinchhausen) and pitch pine (Pinus rigida 
Miller) (W. E. Steiner, Jr, personal com- 
munication). Cicindela patruela is associ- 
ated with oak and pine barrens in other 
parts of its range (Boyd 1978, Knisley et al. 
1990, Knisley and Schultz 1997, Willis 
2000). 


ASSOCIATED TIGER BEETLE SPECIES 


Many authors have noted the existence 
of well-defined assemblages of tiger beetle 
species in particular habitats (see discussion 
in Knisley and Schultz 1997). Data from 
the author’s museum specimen inventory 
(described above) allow a tentative recon- 
struction of historic tiger beetle assemblag- 
es in the Washington area, including assem- 
blages that included C. patruela. Table 2 
lists the numbers of specimens of nine other 
species of Cicindela which were collected 
at the same sites and at approximately the 
same time of the year as the C. patruela 
specimens listed in the Appendix. Cicindela 
purpurea Olivier and C. sexguttata Fabri- 
cius appear to have been the most common 
associates of C. patruela, although C. scu- 
tellaris Say and C. tranquebarica Herbst 
were also well represented at C. patruela 
sites. 


68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


POPULATION SEASONALITY 


Seasonality of the Washington, D.C., 
population of C. patruela can be inferred 
from dates when specimens were collected. 
Adult activity was evidently on a spring-fall 
cycle, as in other populations of this species 
(Knisley and Schultz 1997). The spring ac- 
tivity period appears to have been between 
the end of March and late June, while the 
fall activity period appears to have been 
from early September to early October. 


OPPORTUNITIES FOR HABITAT RESTORATION 
AND REINTRODUCTION 


Much of the literature on tiger beetle 
conservation has focused on identifying im- 
mediate threats to individual populations 
and documenting the causes of population 
declines (see, for example, the numerous ci- 
tations in Knisley and Schultz 1997). Often 
overlooked are two key facts: simple man- 
agement practices can greatly increase the 
available habitat for these beetles (Wilson 
1970, Kritsky et al. 1999), and equally sim- 
ple techniques can be used to successfully 
reintroduce tiger beetle species to restored 
habitat (Knisley and Hill 2001, Scherer 
1999, Brust 2002). 

Given the extent of urbanization in 
Prince George’s County, Maryland, habitat 
restoration for C. patruela and other pine/ 
oak barrens species would be most feasible 
on several large, publicly-owned tracts of 
land which are managed by the United 
States Department of Agriculture (Beltsville 
Agricultural Research Center), the National 
Park Service (Greenbelt Park) and the U.S. 
Fish and Wildlife Service (Patuxent Re- 
search Refuge). Potomac Group soils un- 
derlie much of these tracts, and the barrens 
tree species listed above are found in some 
of the closed-canopy forests on ridges and 
other upland areas. 

Restoration activities for tiger beetles on 
these properties would consist primarily of 
vegetation management, with the overall 
goal of maintaining an early successional 
native plant community with open areas of 


bare sandy soil. The presence of mosses, 
lichens, and sedges is characteristic of C. 
patruela microhabitats and oviposition sites 
in Ohio (Knisley et al. 1997) and Wisconsin 
(Willis 2000), so restoration and manage- 
ment activities should be designed to pro- 
mote the growth of these species. Periodic 
mechanical thinning or fire management (as 
appropriate for the site) would be necessary 
to control tree and shrub growth and pre- 
vent full canopy closure. Native vegetation 
control will need to focus on species such 
as Virginia pine (Pinus virginiana Miailler), 
black locust (Robinia pseudoacacia L.), 
chestnut oak (Quercus castanea Née), and 
white oak (Quercus alba L.), which ag- 
gressively colonize open sandy barrens (W. 
E. Steiner, Jr., personal communication). 
Herbicide treatments may be necessary to 
control invasive, non-native vegetation or 
aggressive natives. Knisley and Schultz 
(1997) reported that the herbicide glyphos- 
ate in its Rodeo® formulation has no ad- 
verse effects on larvae of C. dorsalis Say. 

Reintroduction of C. patruela at restored 
barrens sites may be advisable if no natural 
populations survive close enough to repop- 
ulate restored areas. Although transloca- 
tions of C. patruela have not yet been at- 
tempted, simple yet effective strategies for 
reintroducing adult and larval tiger beetles 
have been described in the literature. Brust 
(2002) reported success at establishing a vi- 
able C. formosa Say population after re- 
leasing adults of this species at a restored 
sand dune in Wisconsin. However, similar 
translocation attempts with adults of the 
threatened species C. dorsalis Say and C. 
puritana Horn failed, due probably to dis- 
persal of adults immediately after release 
(Hill and Knisley 1993, Knisley and Hill 
2001). Knisley and Hill (2001) successfully 
established a population of C. dorsalis at 
Sandy Hook, New Jersey, by translocating 
larvae. Methods for translocating adult tiger 
beetles are described by Brust (2002) and 
methods for translocating larvae are de- 
scribed by Knisley and Hill (2001). Poten- 
tial source populations for C. patruela re- 


VOLUME 107, NUMBER 1 


introductions can be found in western 
Maryland, Virginia, West Virginia, and 
Pennsylvania. 


ACKNOWLEDGMENTS 


Terry L. Erwin graciously permitted the 
examination of tiger beetle specimens in 
NMNH and sponsored my continuing work 
as a Research Associate at the Smithsonian 
Institution. Warren E. Steiner, Jr. provided 
much helpful information about tiger bee- 
tles and barrens vegetation in Maryland, 
and contributed thoughtful comments on an 
earlier draft of this manuscript. C. Barry 
Knisley of Randolph Macon College, James 
M. McCann of the Maryland Department of 
Natural Resources, and Michael A. Valenti 
of the Delaware Forest Service also provid- 
ed helpful comments on earlier drafts of the 
manuscript. 


LITERATURE CITED 


Bellamy, R. W. 2000. History: The Town of Cheverly, 
Maryland: Plantation to planned Community, pp. 
23-28. In Cheverly, Maryland, Citizen’s Hand- 
book. Town of Cheverly, Maryland. 

Boyd, H. P. 1978. The tiger beetles (Coleoptera: Ci- 
cindelidae) of New Jersey with special reference 
to their ecological relationships. Transactions of 
the American Entomological Society 104: 191— 
242. 

Brust, M. L. 2002. Reintroduction study on Cicindela 
formosa generosa in Marinette County, Wiscon- 
sin. Cicindela 34(1—2): 5—7. 

Glaser, J. D. 1984. The Cicindelidae (Coleoptera) of 
Maryland. Maryland Entomologist 2(4): 65—76. 

. 1995. Notes on two rare Maryland tiger bee- 
tles (Coleoptera: Cicindelidae). The Maryland 
Naturalist 39(1—2): 8-10. 

Hill, J. M. and C. B. Knisley. 1993. Puritan Tiger Bee- 
tle (Cicindela puritana G. Horn) Recovery Plan. 
U.S. Fish and Wildlife Service, Northeast Region, 
Hadley, Massachusetts, 45 pp. 

Knisley, C. B. and J. M. Hill. 2001. Translocation and 
establishment of a threatened tiger beetle (Cicin- 
dela dorsalis) at Sandy Hook, New Jersey. Con- 
ference Abstract, Entomological Society of Amer- 
ica 2001 Annual Meeting. World Wide Web doc- 
ument at http://esa.confex.com/2G01/techprogram/ 
paper_3188.htm 

Knisley, C. B. and T. D. Schultz. 1997. The Biology 
of Tiger Beetles and a Guide to the Species of the 
South Atlantic States. Virginia Museum of Natural 
History, Martinsville, Virginia, viii + 210 pp. 


69 


Knisley, C. B., T. D. Schultz, and T. H. Hasewinkel. 
1990. Seasonal activity and thermoregulatory be- 
havior of Cicindela patruela (Coleoptera: Cicin- 
delidae). Annals of the Entomological Society of 
America 83(5): 911-915. 

Kritsky, G., A. Watkins, J. Smith, and N. Gallagher. 
1999. Mixed assemblages of tiger beetles on sand 
piles of various ages (Coleoptera: Cicindelidae). 
Cicindela 31(3—4): 73-80. 

Lawton, J. K. 1970. A new color variant of Cicindela 
patruela. Cicindela 2(2): 1-3. 

Maryland Geological Survey. 1968. Geologic Map of 
Maryland. Baltimore, Maryland. 

Maryland State Data Center. 2003. World Wide Web 
site at: http://www.mdp.state.md.us/msdc/ 

Massachusetts Division of Fisheries and Wildlife. 
2003. Massachusetts List of Endangered, Threat- 
ened, and Special Concern Species. World Wide 
Web site at: http://www.state.ma.us/dfwele/dfw/ 
nhesp/nhrare.htm 

Minnesota Department of Natural Resources. 2003. Ti- 
ger Beetles. World Wide Web document at: http:/ 
/www.dnr.state.mn.us/ets/tigerbeetles.htm] 

New York State Department of Environmental Con- 
servation. 2003. Animal Species Actively Inven- 
toried by Natural Heritage Program—Biological 
and Conservation Data System. World Wide Web 
document at: www.dec.state.ny.us/website/dfwmr/ 
heritage/animallist.pdf 

Pearson, D.L., T.G. Barraclough, and A. P. Vogler. 
1999. Distributional maps for North American 
species of tiger beetles (Coleoptera: Cicindelidae). 
Cicindela 29(3—4): 33-84. 

Pearson, D.L. and A. P. Vogler. 2001. Tiger beetles: 
The Evolution, Ecology, and Diversity of Cicin- 
delids. Cornell University Press, Ithaca, New 
York, 333 pp. 

Pennsylvania Department of Conservation and Natural 
Resources. 2003. Invertebrates. World Wide Web 
document at: http://www.dcnr.state.pa.us/forestry/ 
pndi/fullinvertibrates.asp 

Scherer, A. 1999. Restoring the Northeastern Beach 
Tiger Beetle, Reintroducing a federally listed spe- 
cies. Field Notes, U.S. FWS New Jersey Field Of- 
fice, 1999(September):2. 

U.S. Census Bureau. 2004. World Wide Web site at: 
http://www.census. gov 

Willis, H. L. 2000. Collecting notes for Cicindela pa- 
truela in Wisconsin. Cicindela 32(3—4): 49—54. 

. 2001. Zoogeography of Cicindela patruela in 


Wisconsin (Coleoptera: Cicindelidae). Cicindela 
33(1—2): 1-18. 

Wilson, D. A. 1970. Two interesting observations in- 
volving Cicindela duodecimguttata. Cicindela 
2(3): 20. 

Wisconsin Department of Natural Resources. 2003. 
The Natural Heritage Inventory Working List: 
Rare World Wide Web 


Beetles. document at: 


70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


http:// www. dnr. state. wi. us/ org/land/er/ working- 
list/taxalists/beetles.htm 


APPENDIX 


Collections of Cicindela patruela DeJean 
from the Washington, D.C., metropolitan 
area, as documented by specimens in the 
National Museum of Natural History, 
Smithsonian Institution. 

DISTRICT OF COLUMBIA: Washing- 
ton: Woodridge, 31.11.1918 (1 @). 

MARYLAND: Prince George’s County: 
Bladensburg, 20.1V.1919 (2 3), 1.VI.1919 
GG), BILWLIG20 (j@ yep Caevenhy, 


VII.1931 (1 3), VII.1931 (2 2), 6.1X.1931 
Go 6. 2 Qy lO V1Os3 G6, Il LY), 
NINES (Bo, | By BiLWIOS3 @ S, 2 
2), IOj0< 133 GZS, 4 YS) BIW MOS! (l 
2); College Park, 21.I1V.1939, in sun on 
sparse pine & deciduous wooded hillside, 
Ssandye@OmGnn on +) isamendatamexcept 
21.1X.1939 (1 3), 5.X.1940, on sandy bare 
spot on pine-deciduous hillside (1 @), 
1O1V.1949 (1 CG), 17D 1950° Cd 2): Hy- 
arom, SWIMS Cl Gs hk VQ), GIGI). 
(2 6,4 2); Riverdale,? .V.1919 (1 2); no 
locality specified, 12.VI.1948 (1 @). 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 71-77 


A NEW SPECIES OF DORSIPES REGENFUSS (ACARI: PODAPOLIPIDAE), 
ECTOPARASITE OF AMARA LATIOR KIRBY (COLEOPTERA: CARABIDAE) 
FROM ARIZONA 


ROBERT W. HUSBAND AND DAviID O. HUSBAND 


(RWH) 1035 Scottdale Drive, Adrian, MI 49221, U.S.A. (e-mail: husbandadrian @aol. 
com); (DOH) 346 Jamacha, Apt. 61, E. Cajon, CA 92019, U.S.A. (e-mail: davidhsbnd@ 


aol.com) 


Abstract.—Dorsipes amarae, new species, is a podapolipid mite that parasitizes Amara 
latior Kirby (Coleoptera: Carabidae) in Arizona, U.S.A. It belongs to the inflatus group 
of Dorsipes Regenfuss and represents the first record of the group in the western hemi- 
sphere. The new species and three additional species of Dorsipes in the group inflatus are 
compared with each other and a key to adult females is provided. 


Key Words: 


Mites in the family Podapolipidae (Acari: 
Tarsonemina) are highly specialized ecto- 
and endoparasites of insects of the orders 
Blattaria, Orthoptera, Heteroptera, Hyme- 
noptera, and especially Coleoptera. Regen- 
fuss (1968) proposed the genus Dorsipes 
and described seven species from carabid 
beetles found in Germany, two of them in 
the inflatus group. Husband and Kurosa 
(2002) described new species in the dorsi- 
pes group from Japan and discussed differ- 
ences among the dorsipes, inflatus, and pla- 
tysmae groups. Kurosa and Husband (2002) 
added Dorsipes curtonoti from Japan to the 
inflatus group and discussed changes in in- 
flatus group concepts based on new discov- 
eries. It is the purpose of this paper to de- 
scribe the first member of the inflatus group 
found on American carabid beetles and 
compare it with other species in the inflatus 


group. 


MATERIALS AND METHODS 


Sixty specimens of Amara species (Car- 
abidae) in the University of Michigan Mu- 
seum of Zoology (UMMZ) were examined 


beetle, parasitic mite, Carabidae, Podapolipidae, Arizona 


for mites by the senior author. Many rep- 
resentatives of all life stages of a new Dor- 
sipes species were found under the elytra of 
Amara latior Kirby collected from Cocon- 
ino County, Arizona, U.S.A. 

Measurements in micrometers (j4m) were 
taken with the aid of a Zeiss phase contrast 
compound microscope with an ocular mi- 
crometer. Setae that are no longer than the 
diameters of setal acetabulae are listed as 
microsetae (m). Setae with only an acetab- 
ulum and no remnant of a setal base are 
listed as vestigial setae (v). Often long setae 
are obscured, bent, broken or at an angle 
which makes measurement difficult. Setae 
are at least as long as indicated. Terminol- 
ogy follows Lindquist (1986). 


Dorsipes amarae Husband and 
Husband, new species 
(Figs. 1-3) 


Diagnosis.—With traits of the inflatus 
group: Vagina expanded distally with open- 
ing somewhat dorsal, setae f not present, 
with strong ambulacrum I claw, tarsus II 
without solenidion, coxal setae 3a not pres- 


WZ, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 


ent, setae v, conspicuous, larval female 
without femur III setae (most species), with 
setae i, widely separated, genital capsule of 
male wider at base than at apex. Setae e 
and f on plate EF occur in dorsipes, coxal 
setae 3a occur in platysmae, setae f not 
present in platysmae and inflatus, setae 3a 
not present in inflatus nor in all but one 
dorsipes. 

Female Dorsipes amarae without stig- 


Dorsipes amarae, adult female, ventral (left) and dorsal aspects. 


mata and setae h,. Stigmata present in D. 
notopus Regenfuss, 1968, and D. curtonoti 
Kurosa and Husband, 2002. Vestigial setae 
h, clear in female D. inflatus Regenfuss, 
1968, D. notopus and D. curtonoti. Female 
D. amarae with genu III setae 1", D. inflatus 
without setae /’. Genital capsule of male D. 
amarae with broad base similar to capsule 
of D. inflatus. Setae c, 5—8 in D. amarae, 
3 in D. inflatus. Setae c, anterior to setae c, 


VOLUME 107, NUMBER 1 


Fig. 2. 


in D. amarae, l\ateral to c, in male D. infla- 
tus, D. notopus and D. curtonoti. Setae v, 
longer in larval D. amarae (25—32), shorter 
in D. notopus (12-17) and D. curtonoti 
(11—13). Genu III setae 7” 8—10 in larval D. 
amarae, not present in D. inflatus or D. no- 
topus. Metrical data of dorsal and ventral 
gnathosomal setae, idiosomal setae c,, and 


mel 


100pm 


Dorsipes amarae, male, ventral (left) and dorsal aspects. 


femur seta d in adult female, cheliceral sty- 
lets, and idiosomal setae sc, in male, che- 
liceral stylets and idiosomal setae , and c, 
in larval female potentially useful in dis- 
crimination of species (Table 1). 

Adult female (Fig. 1).—Gnathosoma: 
Length 55-60, width 50-62 (n = 6). Palp 
length 17—20; cheliceral stylet length 48—55 


74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ad 
ewe ae oe 


L| 


Fig. 3. 


with 2 basal sclerites, pharynx width 12— 
13; dorsal gnathosomal setae 24—27, ventral 
setae 12—17, distance between ventral setae 
18—25. Stigmata not evident. 

Idiosoma: Length 248-550, width 186— 
380. Prodorsal plate length 95, width 160: 
setae v, 15-18, v, and sc, vestigial, sc, 36— 


100 pm 


Dorsipes amarae, larval female, ventral (left) and dorsal apsects. 


46. Distance between setae v, 46—54, v, me- 
dial to and sc, near a line connecting v, and 
SC>. Plate C length 80, width 258; setae c, 
13—15, setae c, 18—22. Setae d 11—12, setae 
e 10—13, setae f absent, setae h, 3-5, dis- 
tance between setae h, 47—54. 

Venter: Apodemes 1 moderately devel- 


VOLUME 107, NUMBER 1 


Table 1. Comparison of selected maximum mea- 
surements for Dorsipes inflatus (Din), D. notopus 
(Dno), D. curtonoti (Dcu), and D. amarae (Dam) of 
the inflatus group or Dorsipes. Measurements are in 
micrometers. 


Character Din Dno Deu Dam 


ADULT FEMALES 


Idiosomal length 370 505 600 550 
Idiosomal width 321 415 398 380 
Cheliceral stylets 38 50 53 55 
Pharynx width 14 15 20 13 
Dorsal gnath. setae 23 30 35) 27 
Ventral gnath. setae 12 3) 20 17 
Idiosomal setae 
vy’ 9 10 8 18 
Cy 10 18 16 15 
C) 20 35 40 22 
h, v v v 0) 
Femur I seta d 3 m 5 3 
Femur II seta d O 0) 5 0) 
Tarsus II seta pv’ 3 0) 4 3 
Genu III seta 7” O 6 5) 9 
MALES 
Idiosomal length 155 NPY 7s tte) 
Idiosomal width 134 151 136 157 
Cheliceral stylets 25 20 NY 24 
Dors. gnath. setae ) V 11 8 
Idiosomal setae 
ISG> 29 33 62 40 
Cy m m 6 3 
Genit. cap. length 26 Dif 28 28 
Genit. cap. width 32 32 27 38 
LARVAL FEMALES 
Idiosomal length 230 8 208 197 188 
Idiosomal width 171 160 128 128 
Cheliceral stylets 3) 4] 44 36 
Pharynx width 14 11 10 8 
Idiosomal setae 
V; 24 17 13 32 
Gi 14 15 20 16 
C) 30 40 45 32 
h, 185 100 73 160 
h, 1] 5 5 5) 
Femur II seta d 0 0) 4 O 
Genu III seta /” 0) 0 8 10 
Dist. setae h,—h, 20 13 12 18 


oped, meeting sternal apodeme medially; 
apodemes 2 not extending to sternal apo- 
deme. Coxal setae thin, la and 2a 8—9, 3b 
9-10. 


75 


Legs: Leg setation as in Table 1. Am- 
bulacra I, II, Il with moderately developed 
claws. Tarsus I solenidion w 8. Tibia I so- 
lenidion & 10, seta k 3. Tibiae I, I, II setae 
d 36, 35, 20 respectively. Genu I seta v” 6— 
8, /’ 7-8, genu III seta 1” 7-8. 

Male (Fig. 2).—Gnathosoma: Length 
28-35, width 27-37 (n = 4). Palp length 
13—15; cheliceral stylet length 22—24, phar- 
ynx width 5—9; dorsal gnathosomal setae 5— 
8, ventral setae 7—8, distance between ven- 
tral setae 12. 

Idiosoma: Length 139-180, width 123— 
157. Prodorsal plate length 45, width 88; 
setae Vv), Vo, sc; Mm; sc, 21—40. Distance be- 
tween setae v, 14, v, medial to and sc, on 
a line connecting v, and sc,. Setae c, 3, c, 
5—8, d m-3 and e m. Genital capsule dorsal, 
length 24, width 30. 

Venter: Apodemes | moderately devel- 
oped, meeting sternal apodeme medially: 
apodemes 2 not extending to sternal apo- 
deme. Coxal setae la 5—8, 2a 6-8, 3b 7. 

Legs: Legs I, Il, Il, IV setation (includ- 
ing solenidia) for femur, genu, tibia, tarsus: 
3-3-7-110;, 0-2-4-7,,0-1-4-7~ 0-0=2-4= Am- 
bulacrum I claw 5—6, ambulacra II, Ill 
claws 3—5, no ambulacral IV claws. Femur 
I setae v” 10, no femur II setae d or femur 
III setae v’. Tarsus I solenidion w 6. Tibia I 
solenidion & m, seta k m. Tibiae I, I, Ill 
setae d 19, 17, 15 respectively. Tibia IV 
setae d m, setae v” spinelike 4: tarsus IV 
pv’ m, tc’ spinelike 5, wu’ spinelike 5. 

Larval female (Fig. 3).—Gnathosoma: 
Length 36—42, width 29-38 (n = 5). Palp 
length 12—13; cheliceral stylet length 34— 
36, pharynx width 7—8; dorsal gnathosomal 
setae 25, ventral setae 11—12, distance be- 
tween ventral setae 1 1—14. 

Idiosoma: Length 149-188, width 105— 
128. Prodorsal plate length 72—75, width 
88-97; setae v,; 25-32, v, and sc, V, Sc; 
110—120. Distance between setae v, 17—29, 
v, Slightly medial to and sc, slightly lateral 
to a line connecting v, and sc. Plate C and 
D fused, length 61—70, width 108—126; se- 
tae c, 14-16, setae c, 25-32, setae d 10. 
Plate EF length 28—30, width 40—48; setae 


76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 2. 


Leg setation for femora, genua, tibiae, and tarsi for adult females of species in the genus Dorsipes, 


group inflatus. Setation for legs IV in males is similar in all Dorsipes: 0-O-2-4. 


Leg I Leg Il Leg Ill 
F G Ti Ta F G Yi Ta le G aT Ta 
D. inflatus 3 3 7 10 0) 2 + 6 0) O 4 6 
D. notopus 3 3 7 10 O 2 4 5 0) | 4 6 
D. curtonoti 3 3 7 10 | 2 4 7 ] 4 6 
D. amarae 3 3 7 10 0) 2 4 V 0) | 4 5 
e 30-38. Plate H length 20—23, width 25— DISCUSSION 


30; setae h, 120—160, setae h, 3—5, distance 
between setae h, 15-18. 

Venter: Apodemes 1 moderately devel- 
oped, meeting sternal apodeme medially; 
apodemes 2 not extending to sternal apode- 
me. Coxal setae la 7-8, 2a 7-10, 3b 7-8. 

Legs: Legs I, II, III setation for femur, 
genu, tibia, tarsus: 3-3-7-10, 0-2-4-6, O-1- 
4-5. Ambulacra II, HI with small claws, 2. 
Femur I seta v” 12-14, tarsus I solenidion 
w 7. Tibia I solenidion 6 10-12, seta k 3. 
Genu III seta /” 8-10. 

Egg.—Length 182-205, width 127-139 
(n = 7). 

Type, host, and locality data.—Holotype 
female (RWH80803-1), allotype male and 
20 paratypes: from Tusayan National For- 
est, 8,000 feet, Flagstaff, Coconino County, 
Arizona, U,S.A. from hind wings, metano- 
tum or abdominal tergites under the elytra 
of Amara latior Kirby (Carabidae), collect- 
ed by T. H. and G. G. Hubbell, 2 September 
1993'S). 

Type deposition.—Holotype, allotype, 3 
females, | male, 3 larvae, 7 eggs, paratypes, 
and | vial with mites in 70% ethanol de- 
posited in UMMZ. Paratypes: 1 female, 1 
male, | larval female and | vial of mites 
deposited in the collection of the senior au- 
thor; | female, | male and | larva deposited 
in the Zoological Museum, University of 
Hamburg (ZMH), Hamburg, Germany; 1 
female, | male and | larva deposited in the 
collection of Kazuyoshi Kurosa, Tokyo, Ja- 
pan. 

Etymology.—The specific name is de- 
rived from the generic name of the host in- 
sect. 


Dorsipes is restricted to hosts in the fam- 
ily Carabidae (Coleoptera) and is found in 
Europe, N. America, Africa and Asia. The 
genus was reviewed and the inflatus group 
of Dorsipes discussed by Kurosa and Hus- 
band (2002). We find setae v, and sc, as 
vestigial or microsetae in all Dorsipes. In 
the inflatus group, seta v, varies in position 
from on a line drawn from seta vy, to seta 
sc, to distinctly medial to this line depend- 
ing on the instar and species. Likewise, seta 
sc, varies from being in line with setae v, 
and sc, to lateral to this line. Seta sc, varies 
from close to seta sc,, 6, to distant from sc,, 
more than 12. In respect to leg setation, 
adult female D. inflatus have the fewest se- 
tae (apomorphy), D. notopus and D. amar- 
ae have an intermediate number and D. cur- 
tonoti have the most setae (Table 2). A sim- 
ilar pattern occurs for larval females and 
males except larval D. notopus lack genu 
III setae. 


KEY TO ADULT FEMALES IN THE /NFLATUS 


GROUP OF THE GENUS DORSIPES 

1. Without femora II, I] setae ............. 
— With femora II, III setae D. curtonoti 
2. With genua III seta /” 3 
— Without genua III seta/” .......... D. inflatus 
3. Stigmata and tracheal atria present, genu I seta 

l” (m-3) shorter than % length of coxal setae 

GOST es acy Ak aps ss ee D. notopus 
— Stigmata and tracheal atria not present, genu I 

seta /” (7-8) nearly as long as coxal setae (8— 

LEG) in Peete Main Sahat eran eet D. amarae, 0. sp. 


Nw 


Studies of the distribution and variation 
in Dorsipes from a variety of carabid bee- 
tles from the eastern and western hemi- 
spheres are in progress. The three groups of 


VOLUME 107, NUMBER 1 


Dorsipes currently include four inflatus spe- 
cies from Europe, Japan and the United 
States, four dorsipes species from Europe 
and Japan, and eight platysmae_ species 
from Europe, Africa and the United States. 


ACKNOWLEDGMENTS 


We are grateful to Barry O’Connor and 
Mark O’Brien, Museum of Zoology, Uni- 
versity of Michigan, Ann Arbor, Michigan, 
for the loan of specimens of Amara (Cara- 
bidae) to search for Dorsipes. 


LITERATURE CITED 


Husband, R. W. and K. Kurosa. 2002. Two new species 
of Dorsipes (Acari: Podapolipidae) ectoparasites 


U7) 


of Carabus spp. (Coleoptera: Carabidae) from Ja- 
pan. International Journal of Acarology 28: 29— 
36. 

Kurosa, K. and R. W. Husband. 2002. A new species 
of Dorsipes (Acari: Podapolipidae), ectoparasite 
of Amara gigantea (Coleoptera: Carabidae) from 
Japan. International Journal of Acarology 28: 
147-155. 

Lindquist, E. E. 1986. The world genera of Tarsonem- 
idae (Acari: Heterostigmata): A morphological, 
phylogenetic, and systematic revision with reclas- 
sification of family group taxa in Heterostigmata. 
Memoirs of the Entomological Society of Canada 
136: 1-517. 

Regenfuss, H. 1968. Untersuchungen zur Morpholo- 
gie, Systematik und Okologie der Podapolipidae 
(Acarina: Tarsonemini). Zeitschrift fiir Wissen- 
schaftliche Zoologie. Leipzig 177: 183-282. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 78-83 


A NEW SPECIES OF EPIMICTA FORSTER (HYMENOPTERA: 
BRACONIDAE) FROM NORTH AMERICA AND NEW DISTRIBUTION 
RECORDS FOR EPIMICTA GRIFFITHSI WHARTON 


ROBERT R. KULA AND GREGORY ZOLNEROWICH 


Department of Entomology, Kansas State University, Manhattan, KS 66506-4004, 


U.S.A. (e-mail: rkula@oznet.ksu.edu) 


Abstract.—Epimicta konzaensis Kula, new species, from Kansas and Tennessee is 
described. A diagnosis and key are provided for differentiation of the three species cur- 
rently included in Epimicta Forster. New distribution records for Epimicta griffithsi Whar- 
ton are reported from Kansas, Wyoming, and British Columbia. 


Key Words: 


Epimicta Forster is an uncommon genus 
in Dacnusini with two described species 
worldwide. Epimicta griffithsi Wharton is 
the only described species from the New 
World and is known from Texas (Wharton 
1994). Epimicta marginalis (Haliday) is 
currently the only described species from 
the Old World and is known from western 
Europe (Shenefelt 1974, Tobias 1986). An- 
other Palaearctic species, Synelix rossica 
(Telenga), was originally placed in Epim- 
icta but was transferred to Synelix Forster 
by Tobias (1986). 

Relative to species in larger dacnusine 
genera (e.g., Chorebus Haliday), species in 
Epimicta are infrequently collected. For ex- 
ample, determinations (by RRK) for 1,678 
dacnusine specimens in the Canadian Na- 
tional Collection of Insects (CNCI) resulted 
in 633 specimens of Chorebus and only five 
specimens of Epimicta. Thus, the paucity of 
described species in Epimicta appears to be 
a reflection of taxon rarity, not a result of 
limited collecting effort. 

It is likely that most transient collecting 
efforts fail to acquire species in Epimicta. 
A species in Epimicta may be present at a 
particular locality but will remain undetect- 


Alysiinae, Dacnusini, Nearctic, parasitoids 


ed if the collecting effort does not coincide 
with the species’ phenology. Long-term 
collecting programs are an effective strate- 
gy for collecting rare taxa (e.g., Epimicta) 
because they circumvent the problem of 
phenology. For example, Whitfield and 
Lewis (2001) sampled the braconid fauna 
of six tallgrass prairies in the midwestern 
United States. Konza Prairie Biological Sta- 
tion (KPBS), a 3,487 hectare taligrass prai- 
rie located in the Flint Hills near Manhat- 
tan, Kansas, was among the prairies sam- 
pled. For one week each month in June, 
July, and August, two Malaise traps were 
placed in KPBS watershed 4B (54.5 hect- 
ares). In that short period of time and small 
sampling area, 293 braconids representing 
20 subfamilies and 86 species were col- 
lected. KPBS had the greatest species rich- 
ness among the prairies sampled, but spe- 
cies in Epimicta were not recovered. 

The second author (GZ) initiated a long- 
term collecting program at KPBS in April 
2001. At least five watershed units were 
continuously sampled annually, spring 
through fall, using Malaise traps and yellow 
pan traps. Additionally, sweep net samples 
were periodically taken throughout the col- 


VOLUME 107, NUMBER 1 


lecting season. During the first two collect- 
ing seasons, several specimens of Epimicta 
were acquired. In this paper one new spe- 
cies in Epimicta is described, and the 
known distribution of E. griffithsi is ex- 
panded. 


MATERIALS AND METHODS 


Most specimens used in this study were 
collected with Malaise traps and a sweep 
net at KPBS. The Kansas State University 
Museum of Entomological Prairie Arthro- 
pod Research (KSU-MEPAR), the Univer- 
sity of Wyoming Insect Museum (UWIM), 
and the CNCI provided additional speci- 
mens. All specimens were compared with 
paratypes of E. griffithsi housed at Texas 
A&M University (TAMU). Wharton (1994) 
indicated the holotype for E. griffithsi was 
deposited in the National Museum of Nat- 
ural History (NMNH), but it is currently 
missing and could not be examined. Diag- 
nostic information in Tobias (1986) and 
Wharton (1994) was used to differentiate 
Epimicta konzaensis Kula, new species, and 
E. griffithsi from E. marginalis. 

Measurements are as in Wharton (1977) 
with the following additions and modifica- 
tions. Tergite 1 (tl) length is the maximum 
length of tl in lateral view, and tl width is 
the width of the posterior edge of tl in dor- 
sal view. Thorax length and thorax height 
are referred to as mesosoma length and me- 
sosoma height, respectively. Mesonotal 
width is referred to as mesoscutal width. 
The following abbreviations are used for 
measurements in the description: head 
length (HL), head width (HW), temple 
width (TW), face width (FW), face height 
(FH), eye length (EL), eye height (EH), 
mandible length (MNL), mandible apical 
width (MNAW), mandible basal width 
(MNBW), flagellomere | length (F1L), fla- 
gellomere 2 length (F2L), mesosoma length 
(ML), mesoscutum width (MW), mesosoma 
height (MH), tl length (T1L), and tl width 
(T1W). Measurements were taken using an 
ocular micrometer in a Leica MZ APO ste- 
reomicroscope with 10 oculars. 


79 


Mandibular setation and the shape of 
each tooth are described as observed in lat- 
eral view. The numbering of teeth follows 
Wharton (1977). Terminology for anatom- 
ical features, surface sculpture, and setation 
follows Sharkey and Wharton (1997). 

Data used in the description were taken 
from the holotype, four paratype females, 
and the paratype male. Two additional fe- 
males were examined but not used for the 
description because of their poor condition. 


RESULTS AND DISCUSSION 


Epimicta konzaensis Kula, 
new species 
(Figs. 1—4) 


Diagnosis.—Several characters can be 
used to differentiate E. konzaensis from E. 
griffithsi. In konzaensis the head is yellow 
except very dark brown surrounding the 
ocelli, the mandibles are whitish yellow, the 
frons is smooth, and the 3RS vein in the 
forewing is evenly curved. In griffithsi the 
head is entirely black, the mandibles are 
dark yellow to yellow brown, the frons is 
usually (90% of type specimens) rugose, 
and the 3RS vein in the forewing is weakly 
sinuate. 

The characters Wharton (1994) used to 
differentiate griffithsi from E. marginalis 
can also be used to differentiate kKonzaensis 
from marginalis. The most distinct diag- 
nostic difference between konzaensis and 
marginalis is the location of crenulations 
within the posterior mesopleural furrow. In 
konzaensis the posterior mesopleural fur- 
row is smooth dorsal to the episternal scro- 
be and minutely crenulate ventrally. In mar- 
ginalis the entire posterior mesopleural fur- 
row is conspicuously crenulate (Fig. 5). Ad- 
ditionally, in kKonzaensis the head is yellow 
except very dark brown surrounding the 
ocelli, and in marginalis the head is entirely 
black. 

Description.—Female. Body length: 
2.06—2.34 mm. Head: HL 0.58—0.59X HW, 
HW 0.94-1.00* TW, FW 1.78—-1.89X FH, 
EL 0.50-0.58X EH, MNL 1.00-1.13X 


80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1—S. 
3, Hind wing. 4, Mesopleuron with posterior mesopleural furrow crenulate ventral to episternal scrobe. 5, Me- 
sopleuron of E. marginalis with posterior mesopleural furrow entirely crenulate. 


MNAW, MNAW 1.29-1.38 MNBW, FIL 
1.00—1.25X F2L; antenna with 21-22 fla- 
gellomeres, maxillary palp 6 segmented, la- 
bial palp 4 segmented; face smooth, seti- 
ferous; frons smooth, occasionally (50% of 
type specimens) with pit mesally, setiferous 
laterally, glabrous mesally; gena and vertex 
smooth, setiferous; occiput smooth, gla- 
brous; eyes glabrous; clypeus with apical 
rim, setiferous; mandible with 4 teeth (as 
for E. griffithsi in Wharton 1994), excavat- 
ed distal portion glabrous, rest of mandible 
setiferous, tooth 1 round, with diagonal 
ridge, tooth 2 nearly an equilateral triangle, 
tooth 3 round, tooth 4 (additional tooth) lo- 
cated along ventral margin. Mesosoma: 
ML 1.64-1.81X MW, ML 1.30-1.41x 
MH, MW 0.78—0.86 MH; lateral portion 


1—4, Epimicta konzaensis. 1, Frons with pit mesally. 2, Forewing with vein 3RS evenly curved. 


of pronotum smooth to slightly rugose, se- 
tiferous along margins, glabrous mesally, 
latero-anterior furrow smooth, latero-pos- 
terior margin almost entirely unsculptured 
(a few crenulae ventrally), entirely crenu- 
late, or with transverse ridges radiating an- 
teriorly; notauli crenulate, continuous with 
lateral margin of mesoscutum, terminating 
anteriad mesoscutal midpit; mesoscutal 
midpit slitlike, crenulate; mesoscutum (ex- 
cluding lateral margin, notauli, and midpit) 
smooth, setiferous laterally, anteriorly, and 
along lines where notauli would run if com- 
plete; dorsal surface of scutellum smooth, 
setiferous, scutellar sulcus with 4—8 longi- 
tudinal ridges; metanotum with sharp dorsal 
protuberance; propodeum areolate rugose, 
setiferous; sternaulus rugose; posterior me- 


VOLUME 107, NUMBER 1 


sopleural furrow smooth dorsal to episternal 
scrobe, minutely crenulate ventrally; me- 
sopleuron (excluding sternaulus and poste- 
rior mesopleural furrow) dorso-ventrally 
crenulate along anterior margin, anterior 
portion of subalar region rugose, smooth 
mesally, setiferous along anterior and ven- 
tral margins; metapleuron with smooth me- 
sal portion surrounded by rugosities, seti- 
ferous. Forewing: Hyaline; stigma with 
well-defined proximal and distal margins; 
vein r basad middle of stigma; vein 1m-cu 
basad vein 2RS; vein 3RS evenly curved to 
leading margin; Ist subdiscal cell closed by 
tubular veins. Hind wing: Hyaline; basal 
and subbasal cells closed by tubular veins. 
Metasoma: T1L 1.07—1.27 T1W; tl lon- 
gitudinally rugose, setiferous, dorsope pre- 
sent; t2 striate, setiferous; t3 entirely 
smooth or mostly smooth with anterior 
edge striate, setiferous mesally or in pos- 
terior half of tergite; t¢—t6 smooth, seti- 
ferous mesally or in posterior half of ter- 
gite; remaining tergites smooth, setiferous; 
Ovipositor partially exserted, slightly visible 
dorsally. Color: Head yellow except very 
dark brown surrounding ocelli, mouthparts 
whitish yellow except distal margin of man- 
dible amber with tooth 2 conspicuously 
darker than other teeth, antennal flagello- 
meres brown with proximal flagellomeres 
lighter than distal flagellomeres, scape and 
pedicel brownish yellow to yellow; meso- 
soma very dark brown except propleuron 
brownish yellow to brown; tl very dark 
brown, t2 orangish brown, t3 yellowish 
brown to brownish yellow with posterior 
edge brown, remaining tergites brown; pro- 
thoracic and mesothoracic legs yellow with 
tarsi darker than other leg parts, metatho- 
racic leg yellow except distal half of tibia 
and entire tarsus brown to dark brown. 

Male. As in female except: Head: HL 
0.63X HW, FW 1.67X FH, EL 0.69 EH; 
antenna with 23 flagellomeres. Mesosoma: 
ML 1.46 MH. Color: t2 whitish yellow, 
metathoracic leg yellow with distal half of 
tibia and entire tarsus darker than other leg 
parts. 


81 


Host.—Unknown. 

Material examined.—Holotype female: 
Top label = USA, “KANSAS:Riley Co.; 
Konza Prairie Biol. Station; Kings Creek’’. 
Second label = “39°06.20'N,96°35.77'W; 
27.1V.-1.v.2001; Zolnerowich,Kula,Brown; 
Malaise trap’’ (NMNH). Paratypes: 1 2 
same data as holotype; 1 2 same data as 
holotype expect 26.iv.-30.iv.2002, 2002- 
010; 1 female same data as holotype except 
13.v.-16.v.2002, 2002-030; 1 5 same data 
as holotype except 23.iv.-26.iv.2002, yel- 
low pan traps 2002—005; | 2 same data as 
holotype except 11.v.2002 R. R. Kula, 
sweep net (KSU-MEPAR). Other deter- 
mined material: 1 2° same data as holotype 
except 4.v.-9.v.2002, 2002-020 (KSU-ME- 
PAR); 1 2 USA, TENNESSEE: Hamilton 
Co., East Ridge, 9.v.1952, G. S. Walley 
«NED: 

Discussion.—Epimicta konzaensis and E. 
griffithsi are morphologically similar to cer- 
tain species in Dacnusa Haliday and Exo- 
tela Forster. However, the additional tooth 
along the ventral margin of the mandible 
and striate t2 clearly place both species in 
Epimicta. Epimicta marginalis is larger and 
more heavily sculptured than konzaensis 
and griffithsi and in habitus more closely 
resembles certain species in Symphya For- 
ster. Epimicta and Symphya contain all dac- 
nusines with a striate t2 and an additional 
tooth along the ventral margin of the man- 
dible. However, in Symphya t2+t3 is cara- 
pacelike, while in Epimicta t2+t3 is not 
carapacelike. The relationship between 
Epimicta and Symphya is discussed in Grif- 
fiths (1964) and Wharton (1994). 

All specimens from KPBS were collected 
in a gallery forest along an intermittent 
stream. The dominant tree species in the 
forest are Quercus macrocarpa Michx. (bur 
oak), Quercus muehlenbergii Engelm. 
(chinquapin oak), Celtis occidentalis L. 
(hackberry), and Ulmus americana L. 
(American elm) (Knight et al. 1994). Host 
utilization is unknown for all species in 
Epimicta. 

Etymology.—This species is named after 


82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 6-7. 


the collection site, Konza Prairie Biological 
Station. 


Epimicta griffithsi Wharton 
Epimicta griffithsi Wharton 1994: 630 


Distribution.—New distribution records 
are indicated by asterisks. *CANADA, 
BRITISH COLUMBIA: 1 female Vaseux 
Lake, Oliver, 13.v.1959; 1 female Oliver, 
24.v.1959 (CNCI). USA, *KANSAS: Riley 
Co. 1 female Manhattan, 25.iv.1938; 1 male 
IIS,  lomes  Cresk, 3906.20 7NI 
96°35.77'W, 25.iv.-27.iv.2001; 1 female 2 
males KPBS, Kings Creek, 39°06.20'N 
96°35.77'W, 4.v.-9.v.2001; 2 males KPBS, 
Kings Creek, 39°06.20'’N 96°35.77'W, 
11.v.-15.v.2001; 1 male KPBS, Kings 
Greek. 39-06:204N 96235. 77 Ne 3 Onive- 
4.v.2002; 1 male KPBS, watershed 4B, 
39°04.65’°N 96°35.75’°W, 4.v.-9.v.2001; 1 
female KPBS, watershed 4F 39°04.37'N 
963422 OM Walleye lisaveZ 00a litemialle 
KPBS, watershed SpB, 39°04.50'N 
96°35.25'W, 1.vi.-8.vi.2001 (KSU-ME- 
PAR); TEXAS: Bosque Co. 7 females 1 
male 3 mi. W. Laguna Park, 13.iv.1984 
(CNCI, TAMU, NMNH); Brazos Co. 1 
male College Station, 13.i11.-18.i11.1982; 1 
male College Station, 8.i11.-31.111.1991 
(TAMU); *Kerr Co. 1 male Kerrville, 
30.10.1959 (CNCI); *WYOMING: 3 fe- 


Notauli. 1, Epimicta marginalis (complete). 2, E. griffithsi (incomplete). 


males 2 males Grand Teton National Park, 
University of Wyoming-National Park Ser- 
vice Research Center, 1|.viii.-9.vii1.2002; 1 
female Grand Teton National Park, Univer- 
sity of Wyoming-National Park Service Re- 
search Center, 25.vii.1990 (UWIM). 


KEY TO SPECIES IN EPIMICTA 


1. Posterior mesopleural furrow entirely crenu- 
late; notauli complete, extending to posterior 
margin of mesoscutum (Fig. 6) (Palaearctic) 
SE or ORR RARE ety a es E. marginalis (Halliday) 

— Posterior mesopleural furrow smooth, at most 
with a few crenulae ventral to episternal scro- 
be; notauli incomplete, terminating anteriad 
mesoscutal midpit (Fig. 7) (Nearctic) 

2. Head entirely black; mandibles dark yellow to 
yellow brown; frons usually rugose; forewing 
3RS vein weakly sinuate ... E. griffithsi Wharton 

— Head yellow except very dark brown surround- 
ing ocelli; mandibles whitish yellow; frons 
smooth; forewing 3RS vein evenly curved .. 

E. konzaensis Kula, new species 


ACKNOWLEDGMENTS 


We thank Henri Goulet (CNCI), Scott 
Shaw (UWIM), and Bob Wharton (TAMU) 
for supplying material used in this study. 
We are also grateful to Srinivas Kambham- 
pati (KSU) and Paul Marsh for reading the 
manuscript. Kent Hampton (Scanning Elec- 
tron Microscope Laboratory, KSU) cap- 
tured all original scanning electron micro- 
graphs. We acknowledge the cooperation of 


VOLUME 107, NUMBER 1 


KPBS and the Department of Entomology 
at KSU. This article is Contribution No. 04- 
221-A from the Kansas Agricultural Exper- 
iment Station (KAES) and was supported 
by KAES Hatch Project No. 583, Insect 
Systematics, and the NSF Long Term Eco- 
logical Research Program at Konza Prairie 
Biological Station. 


LITERATURE CITED 


Griffiths, G. C. D. 1964. The Alysiinae (Hym. Bra- 
conidae) parasites of the Agromyzidae (Diptera). 
I. General questions of taxonomy, biology and 
evolution. Beitrage zur Entomologie 14: 823-914. 

Knight, C. L., J. M. Briggs, and M. D. Nellis. 1994. 
Expansion of gallery forest on Konza Prairie Re- 
search Natural Area, Kansas, USA. Landscape 
Ecology 9: 117-125. 

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 


83 


(Hymenoptera). International Society of Hymen- 
opterists. Special Publication 1, 439 pp. 

Shenefelt, R. D. 1974. Pars 11. Braconidae 7, Alysi- 
inae, pp. 937-1113. In Vecht, J. van der and R. 
D. Shenefelt, eds. Hymenopterorum Catalogus 
(nova editio). Dr. W. Junk, The Hague. 

Tobias, V. I. 1986. Subfamily Alysiinae, pp. 100—231. 
In Medvedev, G. S., ed. Keys to the Insects of the 
European Part of the USSR, Vol. Il, Hymenop- 
tera, Part V. Akademia Nauk, Leningrad, 501 pp. 
[in Russian, English translation published in 
1995]. 

Wharton, R. A. 1977. New World Aphaereta species 
(Hymenoptera: Braconidae: Alysiinae), with a dis- 
cussion of terminology used in the tribe Alysiini. 
Annals of the Entomological Society of America 
70: 782-803. 

. 1994. New genera, species, and records of 
New World Alysiinae (Hymenoptera: Braconi- 
dae). Proceedings of the Entomological Society of 
Washington 96: 630-664. 

Whitfield, J. B. and C. N. Lewis. 2001. Analytical sur- 
vey of the braconid wasp fauna (Hymenoptera: 
Braconidae) on six Midwestern U.S. tallgrass prai- 
ries. Annals of the Entomological Society of 
America 94: 230-238. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 84-89 


TUMORIALA, A NEW NEOTROPICAL PHYCITINE GENUS 
(LEPIDOPTERA: PYRALIDAE) 


H. H. NEUNZIG AND M. A. SOLIS 


(HHN) Department of Entomology, North Carolina State University, Raleigh, NC 
27695-7613, U.S.A.; (MAS) Systematic Entomology Laboratory, PSI, Agricultural Re- 
search Service, U.S. Department of Agriculture, % National Museum of Natural History, 
Smithsonian Institution, PO. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. 
(e-mail: asolis@sel.barc.usda.gov) 


Abstract.—Tumoriala n. gen., is proposed for the Neotropical phycitine 7. subaquilella 
(Ragonot 1888), n. comb. The previously unknown male is described and the occurrence 
of this species in Costa Rica and Brazil is reported for the first time. Males in the genus 
are characterized by a unique, raised cluster of scales on the upper surface of the forewing, 
and slender genitalia with a tegumen bearing earlike lobes. Females have a strongly de- 
veloped diverticulum about midway on the ductus bursae. Photographs of the adults, and 
line drawings of male wing venation, labial palpus and antenna, and male and female 


genitalia are included. 


Key Words: 


During an ongoing study of the Phyciti- 
nae of the Neotropics, we discovered males 
of the species Hyalospila subaquilella, Ra- 
gonot 1888, previously known only from 
the female type specimen. Heinrich (1956), 
following a study of the genitalia of the 
type, moved subaquilella to his genus 
Peadus, with the caveat “‘The generic 
placement is tentative, pending discovery 
of a male.”’ Males have a slightly-protuber- 
ant, oval tuft of specialized scales on the 
upper surface of the forewing (Figs. 1—2, 4) 
not seen previously in the Phycitinae, and 
unique genitalia, particularly with regard to 
the tegumen (Fig. 7); therefore, because of 
these features, as well as others, the species 
cannot remain in Peadus. Here we place su- 
baquilella in the new genus Tumoriala. 

Abbreviations used for depositories of 
types and other specimens are as follows: 
Instituto de Biodiversidad, Santo Domingo, 
Costa Rica [INBio]; North Carolina State 
University, Raleigh, North Carolina, USA 


Phycitinae, Guatemala, Costa Rica, Brazil 


[NCSU]; Essig Museum, the University of 
California at Berkeley, California, USA 
[UCB]; National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D. C., USA [USNM]; The Natural History 
Museum, London, England [BMNH]. 


Tumoriala Neunzig and Solis, new genus 


Type species.—Hyalospila subaquilella 
Ragonot 1888. 

Diagnosis.—The male bears a unique, 
raised, oval cluster of many, small scales on 
the upper surface of the forewing (Figs. 1— 
2, 4) and its genitalia are slender with large, 
earlike, lateral lobes projecting posterolat- 
erally from the tegumen (Fig. 7). The fe- 
male has a pronounced, well-sclerotized di- 
verticulum about midway on one side of the 
ductus bursae (fig. 9; see also Heinrich, 
IQSOGE LEVY, 7/53). 

Description.—Antenna (Fig. 6): Shaft of 
male with shallow sinus basally and asso- 
ciated tuft of scales; tuft weak basally, be- 


VOLUME 107, NUMBER 1 


Figs. 1-3. 
raised, cluster of scales on male forewing. 3, Female habitus. 


coming gradually stronger distally; shaft 
beyond scale tuft serrate; sensilla trichodea 
(cilia) about %4 length of diameter of shaft 
at sinus. Frons and vertex: Rough-scaled 
in both sexes. Labial palpus (Fig. 5): Up- 
turned in both sexes, reaching above vertex. 
Maxillary palpus: Short-scaled in both sex- 
es. Haustellum: Well-developed in both 
sexes. Ocellus: Present in both sexes. Fore- 
wing: Male with raised, oval, pale cluster 
of many, small scales at about 4% distance 
from wing base on upper surface of wing 
(Figs. 1-2); wing with 11 veins (Fig. 4); 
R;,, and R; stalked for about % distance 
beyond cell; M, from anterodistal angle of 
cell; M, and M, fused at base for about % 
distance beyond cell; CuA, from posterod- 
istal angle of cell; CuA, from slightly be- 
fore posterodistal angle of cell. Hindwing 


Dorsal view of male and female Twmoriala subaquilella. 1, Male habitus. 2, Enlarged view of 


(Fig. 4): Simple, with 8 veins (1A, 2A, and 
3A together treated as one vein); Sc + R, 
and Rs contiguous or fused for about /% dis- 
tance beyond cell; CuA, from posterodistal 
angle of cell; CuA, from well before pos- 
terodistal angle of cell. Male: Abdominal 
segment 8 simple. Male genitalia (Figs. 7— 
8): Slender; uncus triangular, weakly de- 
veloped, rounded apically; gnathos distally 
with pair of diverging, seta-bearing lobes; 
transtilla with medial pair of fingerlike pro- 
jections; tegumen produced posterolaterally 
into large, earlike lobes (lobes larger than 
uncus); juxta a V-shaped plate with short, 
setiferous, lateral protuberances; valva tri- 
partite, consisting of slender, seta-bearing 
members (sacculus with cluster of slightly 
broadened setae at base); aedoeagus simple; 
vesica with microspines and_ sclerotized 


86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


4 


Figs. 4-6. Wings, antenna and labial palpus of male Tumoriala subaquilella. 4, Right forewing and hindwing 
(the oval outline on the forewing shows the location of the raised cluster of scales on the upper surface of the 
forewing). 5, Left labial palpus, lateral view. 6, Left antenna, frontal view. 


plate; vinculum small, slightly longer than 
greatest width. Female genitalia (Fig. 9): 
Ostium bursae simple; ductus bursae with 
large, rounded, heavily sclerotized, antero- 
laterally-projecting diverticulum near its 
middle; corpus bursae with microspines in 
posterior half and with signum a small, 
thornlike spine; ductus seminalis attached 
to corpus bursae near signum. 

Etymology.—The genus name is a com- 
bination of the Latin tumor (a swelling or 
elevation) and the Latin ala (wing) refer- 
ring to the unusual, elevated scale tuft on 
the male forewing. The gender of Tumori- 
ala is feminine. 


Tumoriala subaquilella (Ragonot), 
new combination 
(Figs. 1-9) 


Hyalospila subaquilella Ragonot 1888:11. 
Peadus subaquilellus (Ragonot): Heinrich 
1956:84. 


Type locality—Cerro Zunil, Guatemala. 

Note.—The type [BMNH] from Guate- 
mala was not examined, but Heinrich’s fig- 
ure of the female genitalia of the type 
(1956: fig. 753), and Ragonot and Hamp- 
son’s color habitus illustration (1901: plate 
XXXVIII: fig. 24) leave no doubt as to the 
identity of subaquilella. 


VOLUME 107, NUMBER 1 


Description.—Head: Frons and vertex 
white to ochre and brownish red; labial 
palpus outwardly white to ochre and 
brownish red to black; maxillary palpus 
mostly white to ochre, brownish red to 
black basally. Thorax: Dorsum of protho- 
rax purple to dark purple, ochre mesially; 
tegula brownish red to purple. Forewing: 
Length 7.5—10.0 mm; mostly brownish red 
to dark purple; anterior half of wing in 
some specimens very lightly dusted with 
white; raised cluster of scales on wing of 
male chiefly pale brown; posterior half of 
wing, near margin, with white or pale 
ochre and pale brownish red longitudinal 
streak in some specimens; antemedial line, 
postmedial line and discal spots absent; 
some specimens with wing mostly brown- 
ish red with dark brown to dark purple 
streaks between veins on posterior half of 
wing. Hindwing: somewhat hyaline, but 
brown to dark brown on veins and margins 
of wing. Male and female genitalia: As 
described for genus. 

Material examined.—COSTA RICA: 
CARTAGO PROVINCE: 1 2, Paraiso, P. 
N. Tapanti-Macizo de la Muerte, del Puente 
del Rio Porras, 1,660 m., Nov. 2001, R. 
Delgado, LN 186550, #65804, INBio CR 
0003401590 [INBio]; 1 @, Paraiso, P. N. 
Tapanti-Macizo de la Muerte, 0.3 km. W. 
del Mirador, 1,350 m., Jul. 2000, R. Del- 
gado, LN 191100560650, #57138, INBio 
CR 0003145304 [INBio]. GUANACASTE 
PROVINCE: 3 o, Est. Pitilla, 9 km. S. San- 
ta Cecilia, 700 m., Oct. and Nov. 1994, C. 
Moraga, LN 330200 380200, INBio CR 
1002035227, INBio 1002041482, INBio CR 
1002041480, genitalia slides 2035227 MC, 
2041482 MC [INBio]; 2 2, Est. Pitilla, 9 
km. S. Santa Cecilia, 700 m., Feb. and July, 
1991, 1995, P. Rios, C. Moraga, LN 329950 
380450, INBio CR 1002135373, INBio CR 
1000599222, genitalia slide 2135373 MC 
[INBio]. PUNTARENAS PROVINCE: | 
3, Monteverde, 15-16 May 1980, D. H. 
Janzen & W. Hallwachs, INBio CR 
1002043394, genitalia slide 107, 755 DA 
[USNM]; 1 3, 2 km. E. Monteverde, 1,500 


87 


m., VU-24—90, Meredith and Powell, gen- 
italia slide 4855 HHN [NCSU]; 2 9, 2 km. 
E. Monteverde, 1,500 m., VIJ-24—90, Mer- 
edith and Powell, genitalia slide 4856 HHN 
[NCSU]; 4 6, Estac. Biol. Las Alturas, 
1550 m., 12 air km. NE San Vito, I-22/24— 
1993, J. Powell, genitalia slides 4861, 4862 
HHN [UCB]; 4 @, Estac. Biol. Las Alturas, 
1,550 m., 12 air km. NE San Vito, I-22/24— 
1993, J. Powell, genitalia slides 4863, 4864 
HHN [UCB]. SAN JOSE PROVINCE: 1 
2, Braulio Carrillo, 1,100 m., VII 1981, V. 
O. Becker, genitalia slide 4929 HHN 
[NCSU]; 1 2, La Montura, Braulio Carrillo 
Nats) Pke OO sm= ly) Decl S Site ask 
Janzen & W. Hallwachs, INBio CR 
1002043895, genitalia slide 107, 756 DA 
[USNM]. BRAZIL: STATE OF RIO DE 
JANEIRO: 1 63, Pq. Nat. Itatiaia, 2,400 m., 
18 X 1985, V. O. Becker, genitalia slide 
4600 HHN [NCSU]; 3 6, 1 &, Itatiaia, 
I/OO wang it Oci, WIS; axon Iviliillikere 
[USNM]. 

Remarks.—A comparison of Tumoriala 
and Peadus shows that Tumoriala males 
have a conspicuous tuft of scales at the base 
of the antenna (Fig. 6), the gnathos is well 
developed and bears apically a pair of lobes 
(Fig. 7), the tegumen has large, lateral lobes 
(Fig. 7), the transtilla is present (Fig. 7), and 
females lack sclerotized plates dorsad of the 
ostium bursae and have a large anterolat- 
erally projecting diverticulum on the ductus 
bursae (Fig. 9), whereas, Peadus males lack 
an antennal scale tuft, lack or have a weakly 
developed gnathos, have a tegumen that is 
greatly reduced, lack a transtilla, and fe- 
males have well developed sclerotized 
plates near the ostium bursae, and are with- 
out a anterolaterally projecting diverticulum 
on the ductus bursae. 


ACKNOWLEDGMENTS 


We thank the following for making ma- 
terial available for study: V. O. Becker, Bra- 
silia, Brasil; E. Phillips, Instituto Nacional 
de Biodiversidad, Santo Domingo, Costa 
Rica; and J. A. Powell, University of Cali- 
fornia, Berkeley, California, USA. 


88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


tegumen 


Figs. 7-9. 
omitted. 8, Aedoeagus. 9, Female. 


Some of the genitalia slides were pre- 
pared by D. Adamski, Systematic Ento- 
mology Laboratory, Washington, D. C., 
USA, and M. Camacho, Instituto Nacional 
de Biodiversidad, Santo Domingo Costa 
Rica. R. L. Blinn, North Carolina State Uni- 
versity, Raleigh, North Carolina, USA, 
made the photographs. An initial draft of 


ductus pbursae 


diverticulum 


lobe 


= L 


Genitalia of Tumoriala subaquilella, ventral view. 7, Male, aedoeagus and setae at base of sacculus 


the paper was reviewed by L. L. Deitz and 
D. L. Stephan, both of North Carolina State 
University. Additional suggestions for im- 
provement were given by S. H. McKamey 
and D. R. Smith, both of the Systematic 
Entomology Laboratory, ARS, USDA, and 
J. C. Shaffer, George Mason University, 
Fairfax, Virginia. 


VOLUME 107, NUMBER 1 89 


LITERATURE CITED Ragonot, E. L. 1888. Nouveaux genres et espéces de 
Phycitidae & Galleriidae, 52 pp. Paris. 


Ragonot, E. L., and (completed by) G. E Hampson. 
Heinrich, C. 1956. American moths of the subfamily 1901. Monographie des Phycitinae et des Galler- 


Phycitinae. United States National Museum Bul- iinae. Jn Romanoff, N. M. Mémoires sur les Lép- 
letin 207: 1-581. idopteres 8: 1-602, pl. XXIV—LVII. Paris. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 90-98 


CHECKLIST AND NOMENCLATURAL NOTES ON THE CHINESE 
PENTATOMIDAE (HETEROPTERA). 
Hl. PHYLLOCEPHALINAE, PODOPINAE 


DaAvip A. RIDER AND LE-YI ZHENG 


(DAR) Department of Entomology, Box 5346, University Station, North Dakota State 
University, Fargo, ND 58105, U.S.A. (e-mail: david.rider@ndsu.nodak.edu); (L-YZ) 
Department of Biology, Nankai University, Tianjin 300071, P.R. China 


Abstract.—A checklist of all known Chinese phyllocephaline and podopine pentatomid 
species is presented with synonymical, distributional, and nomenclatural notes. 


Key Words: 


Previously, checklists have been pub- 
lished for the Chinese Asopinae (Rider and 
Zheng 2002) and Pentatominae (Rider et al. 
2003). This paper, treating the Phylloce- 
phalinae and Podopinae, completes the 
checklist of the Chinese Pentatomidae. A 
brief history of Chinese pentatomid taxon- 
omy was presented in the asopine paper 
listed above. Within the past 25 years, we 
have seen a renewed interest in Oriental 
pentatomidology with the publication of 
many valuable faunal surveys and descrip- 
tive papers. The fine catalogs of Hoffmann 
(1935) and Tang (1935) have now become 
outdated. Also, while preparing the penta- 
tomid manuscript for the Palearctic catalog, 
several nomenclatural and distributional 
problems were discovered. This and the 
previous two checklists are attempts to ei- 
ther correct these nomenclatural and distri- 
butional problems, or at least draw attention 
to them, and thus perhaps stimulate further 
study. 

Entries in the checklist that are in square 
brackets indicate either species whose Chi- 
nese records need verification or species not 
recorded from China, but are likely to be 
found there. 


Pentatomidae, Phyllocephalinae, Podopinae, nomenclature 


CHECKLIST OF CHINESE PHYLLOCEPHALINAE 
Chalcopis glandulosus (Wolff, 1811) 


Distribution.—Fujian, Guangdong, Gu- 
angxi, Hainan, Jiangsu, Jiangxi, Shandong, 
Taiwan, Yunnan, Zhejiang. Also recorded 
from India, Sri Lanka, Bangladesh, south- 
east Asia, and Indonesia. 

Note.—Chalcopus Kirkaldy (1909) was 
originally proposed as a subgenus of Me- 
tonymia Kirkaldy, 1909, but several recent 
workers have recognized Chalcopus as a 
valid genus. In fact, Linnavuori (1982) 
states that Chalcopus “*... differs greatly 
from Metonymia ...” The species of Chal- 
copus that he studied was glandulosa. Yet, 
most recent workers from China still refer 
to this species as Metonymia glandulosa; if 
Linnavuori (1982) is correct, the generic 
name should be Chalcopus. Furthermore, 
Metonymia is a junior objective synonym 
of Dalsira Amyot and Serville, 1843 (see 
Rider and Rolston 1995, for further expla- 
nation). 


Cressona divaricata Zheng and Zou, 1982 


Distribution.— Yunnan. 


VOLUME 107, NUMBER 1 


Cressona rufa Zhang and Lin, 1984 


Distribution.—Jiangxi, Yunnan. 


Cressona valida Dallas, 1851 


Distribution.— Yunnan. Also known 
from India and southeast Asia. 


Dalsira scabrata Distant, 1901 


Distribution.—Yunnan. Also recorded 
from Myanmar. 

Note.—Most records of this species have 
been under the name Metonymia scabrata, 
but Metonymia is a junior objective syno- 
nym of Dalsira (see Rider and Rolston, 
1995, for further explanation). The actual 
generic placement of this species, as well 
as one recorded from Sumatra, need veri- 
fication as all other known species of Dal- 
sira are African. 


Diplorhinus furcatus (Westwood, 1837) 


Synonyms.—Phyllocephala distans Her- 
rich-Schaffer, 1844; Diplorhinus sinensis 
Walker, 1868. 

Distribution.—Guangdong (Hong Kong), 
Guizhou, Hainan, Hunan, Jiangsu, Jiangxi, 
Sichuan, Taiwan, Zhejiang. Also known 
from India and Indonesia. 


Gonopsimorpha ferruginea Yang, 1934 


Distribution.—Hube1, Jiangxi. 


Gonopsimorpha lutea Yang, 1934 


Distribution.—Jiangx1. 


Gonopsimorpha nigrosignata Yang, 1934 


Distribution.—Hube1, Jiangxi. 


Gonopsis affinis (Uhler, 1860) 


Synonym.—Macrina vacillans Walker, 
1868. 

Distribution.—Fujian, Guangdong, Gu- 
angxi, Guizhou, Hubei, Hunan, Jiangsu, Ji- 
angxi, Shaanxi, Shandong, Zhejiang. Also 
reported from Korea and the Ryukyu Is- 
lands. 


Gonopsis coccinea (Walker, 1868) 


Synonym.—Bessida_ scutellaris Walker, 
1868. 


91 


Distribution.—Guangxi, Sichuan, Xiz- 
ang, Yunnan. Also known from India, 
Myanmar, and Malaysia. 


Gonopsis diversa (Walker, 1868) 


Distribution.—Fujian, Hainan, Jiangsu, 
Zhejiang. Also recorded from India. 


Gonopsis rubescens Distant, 1887 


Distribution.—Yunnan. Also recorded 
from India. 

Note.—Only recently recorded from 
Xishuangbana in southwestern Yunnan 
Province, China (Zhang and Lin, 1990). 


[Gonopsis salvazana Distant, 1921] 


Distribution.—India, southeast Asia. 

Note.—Tang (1935) listed this species 
from China, but he was probably following 
Hoffmann (1932) who also included this 
species in a list of species from China and 
neighboring countries. This species has not 
actually been officially recorded from Chi- 
na; in fact, Hoffmann (1948) indicated that 
it probably did not occur in China. The 
same applies to the following species (ton- 
kinensis). 


[Gonopsis tonkinensis Breddin, 1904] 


Distribution.—India, Vietnam. 
Note.—See note under Gonopsis salva- 
zana. 


Megarrhamphus fuscus 
(Vollenhoven, 1868) 


Distribution.—Zhejiang. 


Megarrhamphus hastatus 
(Fabricius, 1803) 


Synonyms.—Aelia rostrata Fabricius, 
1803 (rejected name, Opinion 719, 1965); 
Megarhynchus elongatus Laporte, 1833. 

Distribution.—Anhui, Fujian, Guang- 
dong (Macao), Guangxi, Hainan, Hubei, 
Hunan, Jiangsu, Jiangxi, Taiwan, Zhejiang. 
Also recorded from India, southeast Asia, 
Malaysia, Indonesia, and the Philippines. 

Note.—Fabricius (1803) also described 
this species under the name Aelia rostrata. 


92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Later, Boheman (1852) described a true 
species of Aelia, also using the specific ep- 
ithet rostrata. Because the Fabrician name 
appeared in the literature rarely (and this 
species had usually been referred to under 
the name Megarrhamphus hastatus), while 
A. rostrata Boheman is an important eco- 
nomic pest, Brown (1962) requested that A. 
rostrata Fabricius be rejected, and A. ros- 
trata Boheman be conserved. This was ac- 
cepted in Opinion 719 (1965). 


Megarrhamphus intermedius 
(Vollenhoven, 1868) 


Distribution.—Zhejiang. 


Megarrhamphus limatus 
(Herrich-Schaffer, 1851) 


Distribution.—Guangdong (Macao). 
Also known from India and Myanmar. 


Megarrhamphus tibialis Yang, 1933 


Synonym.—Megarrhamphus tibialis var. 
antetibialis Yang, 1933. 

Distribution.—Sichuan. Also known 
from Vietnam. 


Megarrhamphus truncatus 
(Westwood, 1837) 


Synonym.—Megarhynchus 
Amyot and Serville, 1843. 

Distribution.—Fujian, Guangdong, He- 
bei, Jiangxi, Taiwan, Xizang, Zhejiang. 
Also recorded from Pakistan, India, south- 
east Asia, Malaysia, and Indonesia. 


testaceus 


Salvianus lunatus (Distant, 1901) 


Distribution.—Yunnan. Also recorded 
from India and Myanmar. 


[Salvianus vitalisanus Distant, 1921] 


Distribution.—Laos, Vietnam. 

Note.—Originally described from Laos 
and Vietnam, Tang (1935) listed this spe- 
cies from China. He probably was follow- 
ing Hoffmann (1932) who also listed this 
species from China and neighboring coun- 
tries. It has not actually been officially re- 
corded from any specific locality in China. 


The same applies for the following species 
(Tetroda denticulifer). 


[Tetroda denticulifer Bergroth, 1915] 


Distribution.—Laos, Vietnam. 

Note.—See note under previous species 
(Salvianus vitalisanus). Hoffmann (1948) 
verified that this species has not formally 
been recorded from China. 


Tetroda histeroides (Fabricius, 1798) 


Synonyms.—Aelia furcata Fabricius, 
1803; Megarhynchus quadrispinosus West- 
wood, 1837; Tetroda histeroides var. su- 
matrana Ellenrieder, 1862; Tetroda bilinea- 
ta Walker, 1868. 

Distribution.—Fujian, Guangdong, Gu- 
angxi, Guizhou, Henan, Hubei, Hunan, 
Jiangsu, Jiangxi, Shandong, Sichuan, Tai- 
wan, Yunnan, Zhejiang. Also known from 
India, Japan, southeast Asia, Malaysia, and 
Indonesia. 


[Tetroda latula Distant, 1921] 


Distribution.—Laos 

Note.—Originally described from Laos; 
Tang (1935) listed this species from China, 
but he probably followed Hoffmann (1932) 
who also included this taxon in his list of 
species from China and neighboring coun- 
tries. Hoffmann (1948) indicated that 7. la- 
tula has not formally been recorded from 
any specific locality in China. 


CHECKLIST OF CHINESE PODOPINAE 
Apidestrophus lineola Vollenhoven, 1863 


Distribution.—Hainan. Originally de- 
scribed from Borneo. 


[Apidestrophus morio Stal, 1854] 


Distribution.—Taiwan. India, southeast 

Asia, Malaysia, and Indonesia. 
Note.—Having been recorded in Taiwan, 

it is possible that this species may eventu- 


ally be found on mainland China. 


Asaroticus ogloblini Kiritshenko, 1926 


Distribution.—Nei Monggol. Also 
known from Mongolia and Russia (eastern 
Siberia). 


VOLUME 107, NUMBER 1 


[Asaroticus solskyi (Jakovlev, 1874a)] 


Distribution.—Nei Monggol. Also 
known recorded from Kazakhstan and 
southwestern Russia. 

Note.—Nonnaizab (1984) recorded this 
species from Nei Monggol, but this is un- 
doubtedly based upon a misidentification of 
Asaroticus ogloblini; A. solskyi is distrib- 
uted much farther to the west, and is not 
known to occur in China. 


Brachycerocoris camelus Costa, 1863 


Synonym.—Teressa terranea Walker, 
1867. 

Distribution.—Anhui, Fujian, Guang- 
dong, Guangxi, Hubei, Jiangsu, Jiangxi, 
Zhejiang. Also known from India and Sri 
Lanka. 


Dybowskyia reticulata (Dallas, 1851) 


Synonyms.—Eurygaster = incomptus 
Walker, 1867; Dybowskyia ussurensis Ja- 
kovlev, 1876; Svarinella inexspectata Bal- 
thasar, 1937. 

Distribution.—Anhui, Fujian, Guang- 
dong, Guangxi, Heilongjiang, Henan, Hu- 
bei, Hunan, Jiangsu, Jilin, Liaoning, Nei 
Monggol, Shaanxi, Sichuan, Zhejiang. Also 
recorded from southeastern Europe, Russia 
(eastern Siberia and the Far East), Korea, 
and Japan. 


Graphosoma lineatum (Linnaeus, 1758) 


Synonyms.—Cimex italicus Miller, 
1766; Cimex nigrolineatus Fabricius, 1781; 
Tetyra shangini Gebler, 1817; Graphosoma 
lineatum var. reductum Horvath, 1901; 
Graphosoma italicum var. ochraceum Roy- 
er, 1925; Graphosoma italicum var. obscu- 
rum Stepanek, 1926; Graphosoma italicum 
var. soosi Halaszfy, 1953; Graphosoma it- 
alicum var. nigriscutellatum Halaszfy, 
1955; Graphosoma italicum var. flavolinea- 
tum Halaszty, 1955. 

Distribution.—Xinjiang. Widespread in 
the Palearctic Region. 


Graphosoma rubrolineatum 
(Westwood, 1837) 


Synonym.—Graphosoma crassa 
schulsky, 1861. 

Distribution.—Gansu, Guangdong, Gu- 
angxi, Guizhou, Hebei, Heilongjiang, Hen- 
an, Hubei, Hunan, Jiangsu, Jiangxi, Liaon- 
ing, Nei Monggol, Shaanxi, Shandong, 
Shanxi, Sichuan, Yunnan, Zhejiang. Also 
known from Russia (far eastern), Korea, 
and Japan. 

Note.—The Xinjiang record of Lindberg 
(1934) is undoubtedly a misidentification of 
Graphosoma lineatum. 


Mot- 


Leprosoma tuberculatum Jakovlev, 1874b 


Synonym.—Leprosoma turkestanicum 
Zolotov, 1928. 

Distribution.—Xinjiang. Also reported 
from Afghanistan, Armenia, Iran, Kazakhs- 
tan, Kyrgyzstan, Russia, Tajikistan, Turkey, 
Uzbekistan. 

Note.—Only recently recorded from Chi- 
na (Zheng et al. 1992). 


Melanophara dentata Haglund, 1868 


Distribution.—Fujian, Guangdong (Ma- 
cao), Guangxi, Guizhou, Hainan, Hunan, 
Jiangsu, Zhejiang. Also known from India 
and Myanmar. 


Putonia asiatica Jakovlev, 1885 


Distribution.—Xinjiang. Also recorded 
from Afghanistan, Algeria, Armenia, Azer- 
baijan, Egypt, Iran, Kazakhstan, Kyrgyzst- 
an, Pakistan, Portugal, Russia (southwest- 
ern), Spain, Syria, Tajikistan, Tunisia, Uz- 
bekistan. 

Note.—Only recently recorded from Chi- 
na (Zheng et al. 1992). 


Scotinophara bispinosa (Fabricius, 1798) 
Distribution.—Guangdong. Also known 

from India and Sri Lanka. 

Scotinophara coarctata (Fabricius, 1798) 


Synonyms.—Podops spinosus Walker, 
1867; Podops nasalis Walker, 1867; Podops 
exactus Walker, 1867. 


94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Distribution.— Yunnan. Widespread spe- 
cies occurring from Pakistan, India, and Sri 
Lanka eastward through southeast Asia, 
Malaysia, and Indonesia to the Philippines. 

Note.—Recently recorded from Xish- 
uangbana in the southwestern corner of 
Yunnan Province, China, near Laos and 
Myanmar (Zhang and Lin 1990). 


Scotinophara horvathi Distant, 1883 


Distribution.—Fujian, Guangdong, Gu- 
angxi, Guizhou, Hunan, Jiangxi, Sichuan, 
Xizang. Described from Japan, and also 
known from Korea. 


Scotinophara limosa (Walker, 1867) 


Distribution.—Yunnan. Described from 
India, and also known from Pakistan and 
Myanmar. 

Note.—Another species recently reported 
from Xishuangbana by Zhang and Lin 
(1990). 


Scotinophara lurida (Burmeister, 1834) 


Distribution.—Anhui, Fujian, Guang- 
dong (Macao), Guangxi, Guizhou, Hainan, 
Hebei, Hubei, Hunan, Jiangxi, Jiangsu, 
Shandong, Sichuan, Taiwan, Zhejiang. Also 
recorded from India, Sri Lanka, Japan, and 
Korea. 


[Scotinophara malayensis (Distant, 1903)] 


Distribution.—Taiwan. Malaysia. 

Note.—Originally described from Ma- 
laysia; Esaki (1926) recorded this species 
from Taiwan. This record needs verifica- 
tion, but if it is correct, then it is possible 
this species may eventually be found on 
mainland China. 


[Scotinophara nigra (Dallas, 1851)] 


Distribution.—Jiangsu. Described from 
India. 

Note.—Tang (1935) in a catalog listed 
this species from Soochow, China. We have 
not been able to locate the source of this 
record, nor has there been any record of this 
species from China since. Also, Hoffmann 
(1948) questioned the validity of this re- 


cord. Its presence in China needs verifica- 
tion. 


[Scotinophara obscura (Dallas, 1851)] 


Distribution.—Jiangsu. Described from 
Myanmar, and also known from India, Pak- 
istan, Sri Lanka, southeast Asia, and Indo- 
nesia. 

Note.—Once again, the only Chinese re- 
cord for this species comes from Tang’s 
(1935) catalog. Its presence in China needs 
further verification. 


Scotinophara parva Yang, 1934 


Distribution.—Jiangxi. Also recorded 
from Japan. 


Scotinophara scottii Horvath, 1879 


Synonym.—Scotinophara tarsalis Scott, 
1874 (preoccupied). 

Distribution.—Guangdong, Jiangxi, 
Sichuan, Taiwan, Xizang, Yunnan. De- 
scribed from Japan, and also known from 
Korea. 


[Scotinophara scutellata Scott, 1880] 


Distribution.—Japan, India, Pakistan, Sri 
Lanka. 

Note.—This species was originally de- 
scribed from Japan; it has been reported 
several times from both Japan and from the 
Indian subregion. This interesting disjunct 
distribution raises the question of why it has 
not been recorded from China. Its identity 
and apparent distribution needs further 
study. 


Scotinophara serrata (Vollenhoven, 1863) 


Distribution.—Yunnan. Described from 
Indonesia, and also known from India, Ma- 
laysia, and the Philippines. 

Note.—Another species only recently re- 
corded from Xishuangbana in southwestern 
Yunnan Province, China (Zhang and Lin 
1990). 


VOLUME 107, NUMBER 1 


[| Sternodontus obtusus 
Mulsant and Rey, 1856] 


Synonyms.—Ancyrosoma obtusangula 
Fieber, 1861; Sternodontus obtusus var. de- 
bilicostis Puton, 1884. 

Distribution.—Nei Monggol. Widespread 
species in southern Palearctic, west of Chi- 
na. 

Note.—The Nei Monggol record of Non- 
naizab (1984) is undoubtedly based upon a 
misidentification of Sternodontus similis; S. 
obtusus occurs to the west and north, and 
has not been officially reported from China. 


Sternodontus similis (Stal, 1854) 


Synonyms.—Sternodontus purpureus 
Reuter, 1879; Sternodontus similis var. ehn- 
bergi Reuter, 189 1a. 

Distribution.—Nei Monggol. Also 
known from Kazakhstan, Mongolia, and 
Russia (Siberia). 


Storthecoris nigriceps Horvath, 1883 


Distribution.—Guangdong, Guizhou. 
Described from Himalaya, and also known 
from India and Pakistan. 


Tarisa elevata Reuter, 1901 


Synonym.—Tarisa elevata var. luridula 
Reuter, 1901. 

Distribution.—Nei Monggol. Also re- 
ported from Afghanistan, Armenia, Azer- 
bayan, Iran, Kyrgyzstan, Kazakhstan, Mon- 
golia, Russia, Tajikistan, and Turkmenistan. 


Tarisa subspinosa (Germar, 1839) 


Synonyms.—Tarisa subspinosa var. 
fraudatrix Horvath, 1891; Tarisa subspi- 
nosa var. rosea Horvath, 1891; Tarisa con- 
similis Reuter, 1891b; Tarisa chloris Hor- 
vath, 1904. 

Distribution.—Xinjiang, Xizang. Wide- 
spread southern Palearctic species. 

Note.—Most recent records of this spe- 
cies, including the Chinese records, are un- 
der the name Tarisa fraudatrix, but Linna- 
vuori (1986) placed fraudatrix as a syno- 
nym of subspinosa (fraudatrix was origi- 
nally described as a variety of subspinosa). 


95 


Records of fraudatrix should be transferred 
to subspinosa. 


Tholagmus flavolineatus (Fabricius, 1798) 


Synonym.—Tholagmus confinis Hor- 
vath, 1898. 

Distribution.—Xinjiang. Widespread 
species in the Palearctic Region. 


Ventocoris sp. 


Distribution.—Xinjiang. Also known 
from Tajikistan, Turkmenistan, and Uzbek- 
istan. 

Note.—Zheng et al.’s (1992) report of 
Ventocoris armeniacus (Kiritshenko) from 
China (Xinjiang) is a misidentification. This 
may actually be an undescribed species; 
further study is needed to confirm its actual 
status. 


ACKNOWLEDGMENT 


We thank Dr. I. M. Kerzhner, Zoological 
Institute, St. Petersburg, Russia, for the 
many valuable comments he provided on 
all three checklists. His remarks have great- 
ly improved the quality of these checklists. 


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PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 99-107 


REVIEW OF THE GENUS ACORDULECERA SAY (HYMENOPTERA: 
PERGIDAE) OF THE WEST INDIES, AND THE FIRST RECORDS OF 
SYMPHYTA FROM MONTSERRAT AND ST. KITTS 


DAVID R. SMITH 


Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- 
ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, 
P.O. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. (e-mail: dsmith@sel. 
barc.usda.gov) 


Abstract.—Three species of the pergid genus Acordulecera Say are recorded from the 
West Indies, A. montserratensis, n. sp., from Montserrat, St. Kitts, and Dominica, A. 
longica, n. sp., from Dominica and Montserrat, and A. insularis Ashmead from St. Vin- 
cent and Dominica. Acordulecera montserratensis and A. longica are the first sawflies 
recorded from Montserrat, and A. montserratensis is the first sawfly known from St. Kitts. 
The species are keyed, described, illustrated, and separated from other species of Acor- 


dulecera. 


Key Words: 


Few sawflies are known from the West 
Indies and most are from the Lesser Antil- 
les. Smith (1969) first reviewed the fauna, 
and no additions have been made since. 
Only three species, Acordulecera insularis 
Ashmead (Pergidae) from St. Vincent and 
Dominica, an undescribed species of Acor- 
dulecera Say (Pergidae) from Dominica 
(Smith 1969, 1990), and Hemidianeura 
thoracica Ashmead (Argidae), from Gre- 
nada (Smith 1969, 1992), have been re- 
corded from the Lesser Antilles. Elsewhere, 
only Sericoceros krugii (Cresson) (Argidae) 
occurs in Puerto Rico, the U.S. Virgin Is- 
lands, and the Dominican Republic (Smith 
1992), and three species of Diprionidae and 
two species of Siricidae occur in Cuba 
(Smith 1988). Specimens of undescribed 
species of Acordulecera Say from Montser- 
rat and St. Kitts were brought to my atten- 
tion by M.A. Ivie, Montana State Univer- 
sity. These represent the first records of 
Symphyta from these islands. Here, I de- 


St. Vincent, Dominica, Lesser Antilles, Neotropical, Symphyta 


scribe two new species and give a key to 
the Acordulecera of the West Indies. 

Acordulecera Say is a large New World 
genus distributed from Canada to Argenti- 
na. Though 46 species have been described 
(Smith 1990, Smith and Janzen 2003), more 
than 200 species are possible. The genus 
was defined by Smith (1990). Food plants 
are not known for the West Indian species. 
Larvae of Nearctic Acordulecera feed on 
Quercus spp. (Fagaceae) and Hicoria spp. 
and Juglans spp. (Juglandaceae) (Smith 
1979). Three species have been reared in 
Costa Rica, A. binelli Smith from Poso- 
queria latifolia (Rudge) Roem. & Schult. 
(Rubiaceae), A. liami Smith from Erythrox- 
ylum havanense Jacq. (Erythroxylaceae), 
and A. dashielli Smith from Arrabidaea pa- 
tellifera (Schltdl.) Sandwith (Bigoniaceae) 
(Smith and Janzen 2003). Larvae of a re- 
lated genus, Tequus Smith, feed on potato, 
Solanum spp. (Solanaceae) in Peru and Bo- 
livia (Smith 1981, 1990). 


100 


iw) 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


WUE: (2, 
wy 


Figs. 1-4. Head. 1, Acordulecera montserratensis, front view. 2, A. montserratensis, dorsal view. 3, A. 
insularis, front view. 4, A. insularis, dorsal view. 


WEST INDIES 


. Third antennal segment subequal in length to 


fourth (Fig. 8); abdomen dorsally with transverse 
black bands; female lancet with laterally project- 
ing, hooklike serrulae on apical half and spinelike 
annular armature (Fig. 11); male with antenna 
yellow, mesothorax, except sternum, black, gen- 


Key TO SPECIES OF ACORDULECERA OF THE 


italiayiny io O Rs ee A. montserratensis, Ni. sp. 


Third antennal segment longer than fourth 
(Figs. 9-10); abdomen black with basal sterna 
pale orange or black with dorsal central orange 
area extending from base to about 6th segment; 
female lancet with short, rounded or truncated, 
ventrally projecting serrulae and annuli without 
spines or hairs (Figs. 12—13); male with anten- 
nal flagellum black, thorax mostly orange, gen- 
iui rim TENAS, IANS 2 oc e cao dascaasace 


. Ocelli large, hind ocelli separated by distance of 


less than diameter of one (Fig. 4); head shining; 
dorsum of abdomen with central orange area ex- 
tending from base to about sixth segment; anten- 
na short, about 0.6 head width; serrulae of fe- 


N 


male lancet symmetrical, directed ventrally (Fig. 
12); apex of male penis valve large, triangular 
(nyse tal LAN) Canim Cotten ee Nan tea cg A. insularis Ashmead 
— Ocelli small, hind ocelli separated by distance 
more than 2X greater than diameter of one; head 
dulled by fine irregular sculpturation (Figs. 5—7); 
abdomen black with basal sterna pale orange; an- 
tenna long, about 0.8 head width; serrulae of 
female lancet asymmetrical, directed anteriorly 
(Fig. 13); apex of male penis valve small, oval 
( aiteseml l's)) outer eta go eh tea e. as mes A. longica, 0. sp. 


Acordulecera montserratensis Smith, 
new species 
(Figs. 1-2, 8, 11, 14, 19) 


Female.—Length, 3.5—4.0 mm. Antenna 
white to yellowish. Head black with supra- 
clypeal area, clypeus and mouthparts white; 
apex of mandible black. Thorax orange 
with tegula, spot on extreme upper corner 
of mesepisternum, and metapleuron gray to 


VOLUME 107, NUMBER 1 


black. Legs white with apical 3 tarsal seg- 
ments dark brown, darker to black on outer 
surfaces. Abdomen, except first segment, 
dorsally with broad, black transverse bands, 
bands narrower at center, wider laterally, 
narrower alternating bands pale orange; lat- 
erally and ventrally pale orange except for 
central black area on apical sternite. Wings 
hyaline, stigma and venation black. 

Antenna short, length 0.6 head width 
and 1.3 lower interocular distance; seg- 
ments 3—6 each 2X or less longer than 
broad and subequal in length; apical seg- 
ment slightly longer and broader than oth- 
ers and without long apical seta (Fig. 8). 
Right mandible without carina on lower 
margin. Head shining, from above narrow 
behind eyes (Fig. 2); distances between eye 
and hind ocellus, between hind ocelli, and 
between hind ocellus and posterior margin 
of head as 1.0:2.0:2.0; ocelli small, distance 
between hind ocelli more than 2 diameter 
of an ocellus; postocellar area 1.8 broader 
than long. Eyes converging below (Fig. 1), 
lower interocular distance 0.9X eye length, 
upper interocular distance subequal to eye 
length. Hindbasitarsus 1.2 length of re- 
maining tarsal segments combined. Sheath 
short, rounded in lateral view, in dorsal 
view with thick, laterally projecting scopae 
(Fig. 14). Lancet short, with about 12 ser- 
rulae, basal 4 serrulae symmetrical, directed 
ventrally, apical 8 serrulae hooklike, with- 
out subbasal teeth, and directed anteriorly 
and laterally; annuli with spinelike armature 
(Fig. 11). 

Male.—Length, 2.7 mm. Color as for fe- 
male except antenna yellow; thorax black 
with propleuron, pronotum, tegula, anterior 
edge of mesepisternum, mesosternum, an- 
terior spot on mesoprescutum pale orange; 
abdomen mostly dark brown with apical 
margins of segments, terga 2—6 laterally, 
and basal 4—5 sterna light orange. Antennal 
length subequal to distance between eyes. 
Other characters as for female. Genitalia in 
Fig. 19; parapenis evenly rounded on me- 
son; penis valve oval. 


Type material——Holotype: @, labeled 


101 


Figs. 5-7. 


Head, Acordulecera longica. 5, Front 
view. 6, Dorsal view. 7, Close-up of area between and 
behind ocelli showing sculpturation. 


‘‘Montserrat, Beattie House, nr. Cassava 
Ghaut, 14—30 June 2002, Malaise trap, 632 
ft., M. A. Ivie, 16°45.908'N, 62°12.953'W.” 
In the National Museum of Natural History, 
Smithsonian Institution, Washington, DC 
(USNM). Paratypes: Same data as for ho- 
lotype (10 2); Montserrat: Beattie House, 
16°45'N, 62°12.95'W, O8-17 Apr. 2002, 
632 ft., A. Krakower, Malaise (3 2), same 
except 18 Mar — 04 April 2002 (1 2), same 


102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


except 05—15 Feb. 2002 (1 6), same except 
04—23 March 2002 (2 2); Montserrat: hill 
above Hope Ghaut, 16° 45.169'N, 62° 
IZT36IW;, 16 May 920025 EO S7 Mitek 
Markse, canopy fogging (1 2). Deposited 
in the collections of Montana State Univer- 
sity, Bozeman; N. M. Schiff, Stoneville, 
MS; and USNM. 

Other specimens examined.—ST. KITTS: 
St. Kitts, Milikin Estate, 17°19.607'N, 
62°44.850'W, 02-04 July 2003, 1,200 ft., 
K. A. Markse, P.A.D. Orchard, Malaise trap 
(1 2). DOMINICA: Parish of St. Mark, 4 
km N Soufriere, Malaise in dry wash of de- 
ciduous forest 75 m, 17—19 March 2003, M. 
E. Irwin, M. B. Shepard, E. Benson, G. 
Crmer, 15°14.3'N, 61°22’W (1 6); Clarke 
Hall, 21-31 Jan. 1965, W. W. Wirth, light 
trap, Bredin-Archbold Smithsonian Bio. 
Surv. Dominica (1 6); S. Chiltern, 1,600’, 
II-19-65, H. E. Evans, Bredin-Archbold 
Smithsonian Bio. Surv. Dominica (1 <). 
The specimens from Clarke Hall and S. 
Chiltern, Dominica, were recorded as A. in- 
sularis by Smith (1969). 

Etymology.—The species name is from 
Montserrat, the locality of the type series. 

Discussion.—Though specimens from 
Montserrat, St. Kitts, and Dominica are 
identical, I prefer to use only the specimens 
from Montserrat in the type series. The fe- 
male from St. Kitts has the dorsum of the 
abdomen more solidly black than the more 
striped appearance in most specimens from 
Montserrat. The specimens from Dominica 
are all males, but they appear identical to 
the single male associated with females 
from Montserrat; they range in length from 
2.7—3.0 mm. 

There are no described species of Acor- 
dulecera with the long, hooklike, laterally 
projecting serrulae of the lancet, although I 
have seen undescribed species with similar 
lancets from southern Brazil, northern Ar- 
gentina, and Costa Rica. The unusual lancet 
will immediately separate A. montserraten- 
sis from all other described species of the 
genus. Of the described species of Acor- 
dulecera, A. globulicornis (Konow) from 


Peru and A. cervicata (Konow) from “‘Ita1- 
tuba” are closest in coloration, but A. glob- 
ulicornis has long antennae, nearly as long 
as the head width, and with all segments 
much longer than broad, and A. cervicata 
has the abdomen orange with the apical two 
segments and sheath black. Acordulecera 
vericulata (Konow) from Peru has similar 
serrulae, but they are straighter, not hook- 
like, directed ventrally, the lancet is much 
longer with more than 20 serrulae, and the 
sheath is simple, without scopae. 

Similar species, e.g., species with similar 
lancets from various parts of the Neotrop- 
ics, raises the question that this may be an 
introduction in the West Indies. However, 
the specimens I have seen differ in other 
characteristics from the West Indian speci- 
mens, such as color, sheath shape, and an- 
tennal structure, and none compare conspe- 
cifically. Being such a large genus in the 
Neotropics, further investigations are need- 
ed to confirm A. montserratensis as an en- 
demic species. 

A single male is associated with speci- 
mens collected on Montserrat. It differs in 
color from the female by the mostly black 
mesothorax and abdomen. Males of A. in- 
sularis and A. longica have the thorax 
mostly orange and are more similar to their 
respective females. The genitalia of A. mon- 
tserratensis differs from the other two spe- 
cies by the evenly curved inner margin of 
the parapenis and more oval penis valve 
(see Figs. 17-19). 

This species and A. l/ongica are the first 
records of Symphyta form Montserrat and 
St. Kitts. No sawflies were mentioned in the 
recent treatment of the insect fauna of 
Montserrat (Stevens and Waldmann 2001). 
The Malaise trap in which most of the spec- 
imens of A. montserrratensis were collected 
was located about 30 m away from a house 
in secondary moist tropical forest on the 
west side of Montserrat, next to the forest 
reserve of the Centre Hills in the upper part 
of Woodlands (M. Ivie, personal commu- 
nication). One specimen was collected at a 
higher altitude by canopy fogging. 


VOLUME 107, NUMBER 1 


Figs. 8-10. Antennae. 8, Acordulecera montserratensis. 9, A. insularis. 10, A. longica. 


Acordulecera insularis Ashmead 
Giesas=47 912s lel) 


Acordulecera insularis Ashmead 1900: 
298.—Smith 1969: 541.—Smith 1990: 
179, fig. 465. 


Female.—Length, 3.5 mm. Antenna with 
scape and pedicel white, flagellum black. 
Head black with supraclypeal area, clypeus, 
and mouthparts white; apex of mandible 
black. Thorax orange with tegula and me- 
tapleuron black; pronotum and mesoscutel- 
lum pale orange to white; lateral lobes dark 
orange to infuscate. Legs white with ex- 
treme tip of hind tibia and tarsi infuscate. 
Abdomen black dorsally with central, lon- 
gitudinal orange spot extending from base 
to about 6th segment; apex of 8th segment 
whitish; laterally and ventrally pale orange: 
apical margin of sheath black. 


Antenna short, length 0.6 head width 
and 1.4 lower interocular distance; 3rd 
segment nearly 1.5 length of 4th segment; 
4th and 5th segments subequal in length; 
6th segment longer than Sth; segments 
more than 2 longer than broad; apical seg- 
ment without long apical seta (Fig. 9). 
Right mandible without carina on lower 
margin. Head shining, from above strongly 
narrowing behind eyes (Fig. 4); distances 
between eye and hind ocellus, between hind 
ocelli, and between hind ocellus and pos- 
terior margin of head as 1.0:0.8:1.2; ocelli 
large, distance between hind ocelli less than 
diameter of an ocellus; postocellar area 
2.3 broader than long. Eyes slightly con- 
verging below, lower interocular distance 
0.7 eye length, upper interocular distance 
0.8 eye length (Fig. 3). Hindbasitarsus 
1.4 longer than length of remaining tarsal 


ss = 


— ne — 


Figs. 11-13. 
montserratensis. 12, A. insularis. 13, A. longica. 


segments combined. Sheath rounded at 
apex in lateral view; in dorsal view, of sim- 
ilar width throughout and bluntly rounded 
at apex, without laterally projecting scopae 
(Fig. 15). Lancet long, with about 19 wide- 
ly separated serrulae, each serrula long, 
symmetrical, directed downward, nearly 
truncate at apex, and with several very fine 
anterior and posterior subbasal teeth; annuli 
without spines or hairs (Fig. 12). 
Male.—Length, 2.8—3.0 mm. Color sim- 
ilar to that of female. Genitalia as in Fig. 
17; inner margin of parapenis concave and 
rounded medially; penis valve apex large, 
triangular; harpe with long, stiff hairs. 
Type.—Ashmead described the species 


Female lancets, entire lancet above, close-up of central serrulae below.. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


— 


UL DN RG 
Pe St - 
y 2 


11, Acordulecera 


from “St. Vincent. Described from one 2 
specimen, taken at an elevation of 2000 
feet.’ The holotype (examined) is in The 
Natural History Museum, London, BM 
1.491, labeled “‘St. Vincent, W.I., H. H. 
Smith,” “‘2000 feet,” ““W. Indies 99-331.” 
The antennae are missing. In 1990, I men- 
tioned “‘Ashmead described this species 
from two females. One specimen in London 
is hereby designated lectotype, and the oth- 
er specimen, in Washington, is designated 
paralectotype. Both have the same data and 
the specimens are conspecific.”’ Because 
Ashmead stated he had one female, the lec- 
totype designation is incorrect. 

Material examined.—DOMINICA: 


VOLUME 107, NUMBER 1 


d’LeauGommier, 15 Feb. 1965, W. W. 
Wirth (1 2); DleauGommier, 1,400’, II-15— 
65, H. E. Evans (1 [abdomen missing]); 
DleauGommier, 1,700’, Central Forest Res., 
May 26, 1966, R. J. Gagné (1 6d); Pont- 
Casse, 1.7 mi e, 10-III-65, W. W. Wirth (1 
3), same except 12-III-65 (1 ¢). ST. VIN- 
CENT: Holotype data above. [Note: Do- 
minican names given as on labels.] 

Discussion.—The short antennae with 
the apical segment lacking a long apical 
seta, the lack of scopae on the sheath, large 
ocelli with the distance between the hind 
ocelli less than the diameter of an ocellus, 
lancet, male genitalia, and color separate 
this species from described Neotropical 
species. 


Acordulecera longica Smith, new species 
(Figs. 5-7, 10, 13, 16, 18) 


Acordulecera sp.: Smith 1969: 541, 542. 


Female.—Length, 3.0 mm. Antenna with 
scape and pedicel white, flagellum black. 
Head black with clypeus and mouthparts 
white. Thorax black with tegula, pronotum, 
mesoprescutum, mesoscutellum, lower half 
of mesepisternum, and all mesosternum or- 
ange. Abdomen black with basal sterna or- 
ange. Legs orange with apical 3 tarsal seg- 
ments infuscate. Wings hyaline, veins 
black. 

Antenna long, length about 0.8 head 
width and 1.7 lower interocular distance; 
3rd segment 1.2 length of 4th segment; 
segments gradually decreasing in length to 
apex; each segment more than 2 longer 
than broad; apical segment without long 
apical seta (Fig. 10). Right mandible with- 
out carina on lower margin. Head dull, sha- 
greened, with fine irregular sculpturation, 
not shining (Figs. 6—7); from above strong- 
ly narrowing behind eyes (Fig. 7); distances 
between eye and hind ocellus, between hind 
ocelli, and between hind ocellus and pos- 
terior margin of head as 1.0:1.5:1.5; ocelli 
small, distance between hind ocelli more 
than 2X diameter of an ocellus; postocellar 
area 2.0 broader than long. Eyes slightly 


105 


converging below (Fig. 5); lower interocu- 
lar distance 0.8 eye length; upper intero- 
cular distance subequal to eye length. Hind- 
basitarsus 1.3 length of remaining tarsal 
segments combined. Sheath rounded in lat- 
eral view; nearly uniformly slender in dor- 
sal view, slightly tapering at apex, with 
short, laterally projecting scopae shorter 
than central part of sheath (Fig. 16). Lancet 
with about 16 serrulae, each serrula short, 
asymmetrical, without subbasal teeth, trun- 
cated at apex, and directed anteriorly; an- 
nuli without spines or hairs (Fig. 13). 

Male.—Length, 3.0 mm. Color similar to 
that of female except mesonotum black. 
Genitalia as in Fig. 18; parapenis broadly 
rounded medially; harpe with rounded rid- 
gelike sculpturing and with short, flexuous 
hairs; apex of penis valve small, oval. 

Holotype— @, labeled ““‘Dominica WI, 
June 15, ‘41, 800’, in forest, R. G. Fen- 
nah.”” (USNM). 

Other specimens examined.—DOMINI- 
CA: 1.5 mi. N. Pont Cassé, 1,200’, I-12- 
MSHS. lal, 1a, levams (il Gye Bay. lives 
water L., II-21—1964, collected at light trap, 
Dale E Bray (1 6, USNM). MONTSER- 
RAT: hill above Hope Ghaut, 16° 45.169'N, 
627 IDWiBO NV, wGEMay 20025 ROS att ke 
Markse, canopy fogging (1 @). 

Etymology.—The name refers to the rel- 
atively long antennae compared to the 
shorter antennae of the other two West In- 
dian species. 

Discussion.—This is the species I re- 
ferred to as ““Acordulecera sp.” in 1969 
from Dominica. The Montserrat specimen 
is the only additional specimen I have seen 
since 1969. Both males from Dominica re- 
semble the female, but due to the risk of 
associating sexes, I prefer not to designate 
them as paratypes. The female from Mont- 
serrat is identical to the holotype, but I pre- 
fer not to designate it a paratype. This and 
a female of A. montserratensis were col- 
lected on the same date during canopy fog- 
ging. 

The long antennae lacking a long apical 
seta on the apical segment, slender sheath 


106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 14-19. 14—16, Female sheaths, lateral view above, dorsal view below. 14, Acordulecera montserratensis. 
15, A. insularis. 16, A. longica. 17-19, Male genitalia, ventral aspect of left half of genital capsule on left, lateral 
view of penis valve on right. 17, A. insularis. 18, A. longica. 19, A. montserratensis. 


with small, laterally projecting scopae, lan- 
cet, color of thorax, and black abdomen 
with only basal sterna pale orange separate 
this species from the described Neotropical 
species of Acordulecera. 


ACKNOWLEDGMENTS 


Thanks are due to M.A. Ivie, Montana 
State University, Bozeman, for bringing the 
Montserrat and St. Kitts specimens to my 
attention. Work on Montserrat and St. Kitts 
was funded by The Royal Society for the 
Protection of Birds, the UK Foreign and 
Commonwealth Office, Montserrat Nation- 
al Trust, Montserrat Forestry Division, and 


the Montana Agricultural Experiment Sta- 
tion. W. K. Reeves, Clemson University, 
provided a specimen from Dominica. Cathy 
Apgar, Systematic Entomology Laboratory 
(SEL), Washington, DC, took the photo- 
graphs and prepared the plates. I thank the 
following for review of the manuscript: 
N.M. Schiff, U. S. Forest Service, Stone- 
ville, MS, and R. A. Ochoa and E. E. Gris- 
sell, SEL, Beltsville, MD, and Washington, 
DC, respectively. 


LITERATURE CITED 


Ashmead, W. H. 1900. VI. Report upon the Aculeate 
Hymenoptera of the islands of St. Vincent and 


VOLUME 107, NUMBER 1 


Grenada, with additions to the parasitic Hyme- 
noptera and a list of the described Hymenoptera 
of the West Indies. Transactions of the Entomo- 
logical Society of London, pt. II, pp. 207—367. 


Smith, D. R. 1969. Symphyta of the West Indies, in- 


cluding those collected during the Bredin-Arch- 
bold-Smithsonian Biological Survey of Dominica 
(Symphyta). Proceedings of the Entomological 
Society of Washington 71: 540-543. 

. 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, Symphyta and Apocrita (Parasi- 
tica). Smithsonian Institution Press, Washington, 
DC. xvi + 1198 pp. 

. 1981. Identification of the Acordulecera “‘po- 
tato” sawflies of Peru and Bolivia, with descrip- 
tion of these and related species from South 
America (Hymenoptera: Pergidae). Journal of the 
Washington Academy of Sciences 70: 89-103. 

. 1988. A synopsis of the sawflies (Hymenop- 
tera: Symphyta) of America south of the United 


107 


States: Introduction, Xyelidae, Pamphiliidae, Cim- 
bicidae, Diprionidae, Xiphydriidae, Siricidae, 
Orussidae, Cephidae. Systematic Entomology 13: 
205-261. 

. 1990. A synopsis of the sawflies (Hymenop- 
tera, Symphyta) of America south of the United 
States: Pergidae. Revista Brasiliera Entomologia 
34: 7-200. 

. 1992. A synopsis of the sawflies (Hymenop- 
tera, Symphyta) of America south of the United 
States: Argidae. Memoirs of the American Ento- 
mological Society, Number 39, 201 pp. 


Smith, D. R. and D. H. Janzen. 2003. Food plants and 


life histories of sawflies of the families Tenthre- 
dinidae and Pergidae (Hymenoptera) in Costa 
Rica, with descriptions of four new species. Jour- 
nal of Hymenoptera Research 12: 312-332. 


Stevens, M. and G. Waldmann. 2001. Animal biodi- 


versity of the Lesser Antillean island of Montser- 
rat (British West Indies): an annotated checklist of 
terrestrial and freshwater animals. Archiv Zoolo- 
gischer Publikation 6: 1—145. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 108-114 


THE FOURTH INSTAR LARVA AND PUPA OF THE NEOTROPICAL BITING 
MIDGE FORCIPOMYIA (FORCIPOMYIA) RIOPLATENSIS MARINO AND 
SPINELLI (DIPTERA: CERATOPOGONIDAE) 


GUSTAVO R. SPINELLI, PABLO I. MARINO, AND MARIA M. RONDEROS 


Departamento Cientifico de Entomologia, Museo de La Plata, Paseo del Bosque s/n, 
1900 La Plata, Argentina (e-mail: spinelli@museo.fcnym.unlp.edu.ar) 


Abstract.—The fourth instar larva and pupa of the Neotropical biting midge, Forcipo- 
myia (Forcipomyia) rioplatensis Marino and Spinelli, are described and illustrated using 
phase-contrast and scanning electron microscopes. Larvae, pupae and adults of Forcipo- 
myia rioplatensis were collected from nests of the monk parakeet, Mylopsitta monachus 
monachus (Boddaert), in San Vicente, Buenos Aires Province, Argentina. Larvae have 
character states typical of other species of Forcipomyia with herbivorous ceratopogonid 
larvae. This species is compared with its very similar Nearctic congener, Forcipomyia 
(F.) bipunctatapropinqua Chan and LeRoux. 


Key Words: 
parakeet, Argentina 


Forcipomyia Meigen is a large and ex- 
tremely diverse genus of Ceratopogonidae 
that is worldwide in distribution and diverse 
in morphology and habitat preference. The 
adults of some species are important polli- 
nators of cacao and other plants of econom- 
ic importance in tropical and subtropical ar- 
eas (Soria et al. 1980, Chapman and Soria 
1983, Young 1986). 

There are now 1,028 extant described 
species of Forcipomyia (A. Borkent, per- 
sonal communication). In their catalog of 
the Ceratopogonidae south of the United 
States, Borkent and Spinelli (2000) listed 
203 species. Twelve species from the Neo- 
tropics were subsequently described by Ma- 
rino and Spinelli (2001a, b, c, 2002, 2003) 
and Marino et al. (2002). 

The Forcipomyia squamitibia group in 
the subgenus Forcipomyia (Forcipomyia) 
was recently reviewed for the Neotropics 
(Marino and Spinelli 2002). Shortly before 
that paper was published, we received im- 


Diptera, Ceratopogonidae, immatures, Forcipomyia rioplatensis, nest, monk 


matures and adults of one of the included 
species, Forcipomyia rioplatensis Marino 
and Spinelli, collected by Roxana Aram- 
burt’ from nests of the “‘monk parakeet,” 
Myiopsitta monachus monachus (Boddaert) 
(Aves: Psittacidae), in San Vicente, Buenos 
Aires Province, Argentina. 

The purpose of this paper is to describe 
and illustrate the fourth instar larva and 
pupa of Forcipomyia rioplatensis, and com- 
pare it to the most similar congener F. (F.) 
bipunctatapropinqua Chan and LeRoux, as 
well as to F. (F.) elegantula Malloch and 
F. (F.) pulchrithorax Edwards. 


MATERIAL AND METHODS 


The surveyed “‘monk parakeeet”’ nests 
were located in trees of Eucalyptus tereti- 
cornis Sm. (Myrtaceae). The nesting mate- 
rial was composed of fragments of this tree, 
and to a lesser degree of Casuarina cun- 
ninghamiana Mig. (Casuarinaceae). 

For observation with light phase-contrast 


VOLUME 107, NUMBER 1 


microscope, including oil inmersion, some 
larvae and pupae were slide mounted in 
Canada balsam with their ventral side up- 
ward. Others were mounted dorsally and 
laterally to examine the chaetotaxy of the 
head capsule and abdominal segments of 
the fourth instar larva and cuticular pro- 
cesses of the cephalothorax, respiratory 
horn, and abdominal segments of the pupa. 
Only one side of the body is described. For 
examination and microphotography with 
the SEM (JOEL T100), larvae and pupae 
were mounted using the technique of Ron- 
deros et al. (2000). 

Terms follow those by Debenham 
(1987), and special terms of larval mouth- 
parts follow Hribar and Mullen (1991). 
Voucher specimens are deposited in the col- 
lection of the Museo de La Plata, Argenti- 
na. 


RESULTS 


Forcipomyia rioplatensis Marino and 
Spinelli 2002 
(Figs. 1—23) 


Forcipomyia squamitibia Lutz: Spinelli 
1983: 128 (Argentina record). 

Forcipomyia rioplatensis Marino and Spi- 
nelli 2002: 314 (female, male; Argenti- 
na). 


Diagnosis of adult—A dark brown spe- 
cies of the squamitibia group, distinguished 
from other members of the species group 
by the following combination of characters: 
Third palpal segment slightly swollen at 
base, tapering toward apex; scutum uni- 
formly dark brown; legs brown except basal 
Y% of hind femur pale, knees pale; tibial has- 
tate spines slender, pointed; spermathecae 
ovoid with short neck; sternite 9 with an- 
teromesal groove; aedeagus stout, basal 
arch very low, tip blunt; parameres widely 
separated at base. 

Fourth instar larva.—Exuvia pale yel- 
lowish. Head capsule (Figs.10—11, 13) well 
developed, hypognathous, HL 0.43 (0.37— 
0.52, n = 7) mm; HW 0.35 (0.29-0.40, n 
= 7) mm; SGW not meansurable; HR 1.24 


109 


(1.14—1.30, n = 7). Head chaetotaxy (Figs. 
10-11) as follows: 10 sensory setae, two 
pits; setae “‘p’’, ““q” lanceolate, base broad 
(Fig. 12); seta “‘s”’ stiff, 0.5 as long as “‘p” 
““q’’; setae “‘t’’, “u’’ stout, 3—4 times longer 
than “‘s’; seta “‘v” thin, shorter than “‘u”’ 
Pity [NOK Suniel, Gelkvs SO, ee, Pk 
‘“‘y”’ minute, thin, ““w”’ slightly stouter; “‘z”’ 
pore simple. Labrum (Fig. 13) short, not ex- 
tending beyond hypostoma; palatum not in 
position to describe in detail, with at least 
three sensilla visible. Messors (Fig. 1) stout, 
comma-shaped, anterior, posterior margins 
blunt. Mandible (Figs. 2, 13) stout, scoop- 
like, not articulating with head capsule, 
three blunt apical teeth, ML 0.12 (0.088— 
0.130, n = 7) mm; MW 0.037 (0.028— 
0.040, n = 7); hypostoma (Fig. 13) smooth. 
Epipharynx (Fig. 3) massive, strongly scler- 
otized, toothed along posterior end of me- 
dian sclerite, which bears conspicuous 
ridge; lateral arms stout, with two small, 
apical, pointed teeth. LAW 0.28 (0.24—0.32, 
n = 6) mm; DCW 0.13 (0.11-—0.14, n = 6) 
mm. Hypopharynx (Fig. 4) V-shaped, 
strongly sclerotized, lateral arms thin pro- 
jecting anteromesally; labium short, broad, 
not in position to describe. Maxilla (Fig. 
14) with conspicuous basal fringe, palpus 
bearing stout seta, galeolacinia triangular, 
pointed. Prothoracic pseudopod (Fig. 15) 
bifid, each ramus with six dark brown, pos- 
teriorly directed hooks, one bifid, one me- 
dial hook smaller (Fig. 16). Chaetotaxy of 
second abdominal segment (Figs. 5—6): 
dorsally, one seta *‘a”’ pale, swollen-tipped, 
spear-shaped, strong base; three more sim- 
ple very stout, brown chaetae evenly 
spaced: seta ““b’’ with conical base, seta 
*“c”?, **d’’ each with rounded base, all se- 
toceous, “‘d’”’ shorter than “‘c’’, the latter 
shorter than ““b”’; ventrally, three setae, se- 
fae ei salt | SLOMEOLOWDEs \Vehy. lone. 
smooth, “f* shorter than “‘e’ 
pale, short, without protruding base; cuticle 
with minute spicules. Anal segment (Fig. 
17) with six chetae, two of them apical, 
strong, setaceous, remaining four subapical 
insertions of strong chaetae; anal pseudo- 


>» Sela aie: 


110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


acp 


Ces 
© |- 
—. 

fee) 
— 
Dr 

To 


Figs. 1-9. Structures of immatures of Forcipomyia rioplatensis. \—6, Larva. 7—9, Pupa. 1, Messors. 2, 
Mandibles. 3, Epipharynx (median sclerite in detail). 4, Hypopharynx. 5, Second abdominal segment. 6, Setae 
of second abdominal segment. 7, Respiratory horn. 8, Cuticular processes and dorsal tubercles of cephalotho- 
raxes; (acp) anterior cuticular process, (Icp) lateral cuticular process, (pcp) posterior cuticular process, (adt) 


anterior dorsal tubercle, (pdt) posterior dorsal tubercle. 9, Setae of fourth abdominal segment. 


VOLUME 107, NUMBER 1 


pod bilobed, each lobe with two rows of 
four ventrolateral posteriorly directed 
hooks; anal papillae not visible; cuticle with 
numerous minute setae; CSL 0.21 (0.20— 
0.21, n = 2) mm, CSW 0.24 (0.22-0.27, n 
= 2) mm, CSR 0.87 (0.77—0.96, n = 2). 

Pupa (Fig. 18).—Length 2.86 (2.84— 
2.88, n = 2) mm. Exuvia pale yellowish. 
Respiratory horn (Figs. 7, 20—21) small, 
stout, with 14—15 lateral spiracular pores, 
10—11 apical ones, RH 0.18 (0.14—0.22, n 
= 6) mm, P 0.022 (0.018—0.028, n = 6) 
mm, PH 0.125 (0.114—0.130, n = 6) mm. 
Cephalothorax (Fig. 19) with three dorsal 
tapering cuticular processes; anterior pro- 
cess with truncate tip, bearing long apical 
spine; lateral, posterior processes with api- 
ces finely pointed, each with apical spine 
absent; two rudimentary dorsal tubercles, 
the anterior one bearing stout seta (Fig. 8). 
Fourth abdominal segment (Fig. 9) with 
two dorsal setae: anterior one long directed 
anteriorly, very thin, base broad not tuber- 
culate; posterior seta directed posteriorly, 
short, stout with tuberculate base; four sub- 
equal, thin ventrolateral setae, each with in- 
conspicuous base, same length as posterior 
dorsal seta. Male terminal segment (Figs. 
22-23) densely covered with very small 
spicules dorsally; genital processes dorsal, 
stout, tips blunt; posterolateral processes di- 
vergent, with blunt apices; length 0.42 
(0.35—-0.47, n = 4) mm, width 0.27 (0.26— 
0.29, n = 4) mm. 

Distribution.—Argentina (Buenos Aires 
Province), Uruguay (Departments of Arti- 
gas and Tacuaremb6). 

Specimens examined in Canada bal- 
sam.—Argentina, Buenos Aires Proy., San 
Vicente, 15-XI-2001, R. Aramburu, | larva, 
6 pupae with larval exuvia, 6 males, 3 fe- 
males (from nests of ““monk parakeets”’ ). 

Specimens examined by SEM.—Same 
data, 2 larvae, 2 pupae. 


DISCUSSION 


Forcipomyia rioplatensis is very similar 
to the Nearctic species Forcipomyia_ bi- 
punctatapropinqua, described in detail by 


111 


Chan and LeRoux (1971). Immatures of F. 
bipunctatapropinqua can be distinguished 
from immatures of F. rioplatensis by the 
larval prothoracic pseudopod lacking a bifid 
hook and bearing three ventral smaller 
hooks on each ramus, by the second ab- 
dominal segment of the pupa with the an- 
terior dorsal tubercle without a seta (ante- 
rior dorsal seta directed anteriorly with 
broad base, not tuberculate in F. rioplaten- 
sis), and by the slender male dorsal genital 
processes with pointed tips (stouter, with 
blunt tips in F. rioplatensis). The adult male 
of F. bipunctatapropinqua can be distin- 
guished from F. rioplatensis by the aedea- 
gus with a higher basal arch, tapering dis- 
tally and with a bipartite apex. 

Wirth and Grogan (1978) described and 
illustrated the larval and pupal setae and re- 
spiratory horn of the Nearctic species F. 
(F.) elegantula, and the Palearctic one F. 
(F.) pulchrithorax. Larval abdominal setae 
a, b, and d of F. pulchrithorax are nearly 
identical with those of F. rioplatensis. 
Wirth and Grogan (1978) also provided a 
detailed SEM micrograph of the pupal re- 
spiratory horn of F. elegantula which has a 
differently shaped horn than that of F. rio- 
platensis but the surface texture is very 
similar to that species. In addition, the size 
of the spiracular pores in F. elegantula are 
similar to those of F. rioplatensis, but are 
of a different shape than in that Neotropical 
species. 

The larvae of F. rioplatensis exhibit the 
character states typical of “herbivorous” 
ceratopogonids: Head capsule short and 
stout, with mouthparts directed ventrally, 
well sclerotized labium, and a mandible 
with three teeth. Larvae most probably feed 
on detritus and organic matter in the nests 
where they live. Another possibility is that 
the larvae actually live under bark, like 
many other species of Forcipomyia, and 
eggs or larvae have moved to the nests dur- 
ing its construction by the monk parakeets. 

ACKNOWLEDGMENTS 

We are very grateful to Roxana Aram- 

buri, who collected the specimens de- 


112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 10-17. Fourth instar larva of Forcipomyia rioplatensis. 10, Head capsule (lateral view), chaetotaxy. 11, 
Head capsule (dorsal view), chaetotaxy; (AN) antennae, (x) parantenal setae, (t) anterodorsal setae, (r) frontal pits, 
(s) anterior perifrontal setae, (q) postfrontal setae, (w) anterolateral setae, (u) mesolateral setae, (v) posterolateral 
setae, (p) posterior perifrontal setae, (0) parahypostomal setae, (y) ventral setae. 12, Postfrontal seta. 13, Head 
capsule (anteroventral view); (LB) labrum, (MN) mandible, (HY) hypostoma. 14, Maxilla (MX), (MP) maxillary 
palpus, (GL) galeolacinia. 15, Prothoracic pseudopod. 16, Hooks. 17, Anal segment with insertion of setae. 


VOLUME 107, NUMBER 1 113 


Figs. 18-23. Pupa of Forcipomyia rioplatensis. 18, Exuvia (lateral view). 19, Cephalothorax (lateral view). 


20, Respiratory horn. 21, Respiratory horn (pores in detail), 22, Male caudal segment (dorsal view). 23, Male 
caudal segment (ventral view). 


114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


scribed in this paper and kindly provided 
information on the “‘monk parakeet”’ nests. 
We also acknowledge Art Borkent for de- 
tailed critical review of the manuscript. 


LITERATURE CITED 


Borkent, A. and G. R. Spinelli. 2000. Catalog of the 
New World biting midges south of the United 
States of America (Diptera: Ceratopogonidae). 
Contributions on Entomology, International 4(1): 
1-107. 

Chan, K. L. and E. J. LeRoux. 1971. Nine new species 
of Forcipomyia (Diptera, Ceratopogonidae) de- 
scribed in all stages. The Canadian Entomologist 
103: 729-762. 

Chapman, R. K. and S. de J. Soria 1983. Comparative 
Forcipomyia pollination of cacao in Central 
America and southern Mexico. Revista Theobro- 
ma 13(2): 129-139. 

Debenham, M. L. 1987. The biting midge genus For- 
cipomyia (Diptera: Ceratopogonidae) in the Aus- 
tralasian Region (exclusive of New Zealand) III. 
The subgenera Forcipomyia, s.s., and Lepidohe- 
lea. Invertebrate Taxonomy 1: 269-350. 

Hribar, L. J. and G. R. Mullen. 1991. Comparative 
morphology of the mouthparts and associated 
feeding structures of biting midge larvae (Diptera: 
Ceratopogonidae). Contributions of the American 
Entomological Institute 26(3): 1-71. 

Marino, P. I. and G. R. Spinelli. 2001a. El subgénero 
Forcipomyia (Euprojoannisia) en la Patagonia 
(Diptera: Ceratopogonidae). Gayana 65(1): 11-18. 

. 2001b. Las especies del subgénero Forcipo- 

myia (Forcipomyia) en la Patagonia (Diptera: Cer- 

atopogonidae). Revista de la Sociedad Entomo- 

l6gica Argentina 60(1—4): 99-124. 

. 2001c. Los subgéneros de Forcipomyia, Thyr- 

idomyia y Synthyridomyia en la Patagonia argen- 

tina, con la descripcion de F. (S.) soibelzoni sp. 


n. (Diptera: Ceratopogonidae). Neotropica 47: 13— 
16. 


. 2002. A revision of the Forcipomyia squam- 

itibia group in the Neotropics with the description 

of three new species (Diptera: Ceratopogonidae). 

Insect Science and its Application 22(4): 307-319. 

. 2003. The Patagonian species of the subgenus 
Forcipomyia (Metaforcipomyia) (Diptera: Cera- 
topogonidae), with a key to the New World spe- 
cies. Insect Systematics and Evolution 34(1): 21— 
28. 

Marino, P. I., G. R. Spinelli, and C. G. Cazorla. 2002. 
Type-specimens of Ceratopogonidae (Insecta: 
Diptera) in the collection of the Museo de La Pla- 
ta, Argentina. Publicacién Técnica y Didactica, 
Facultad de Ciencias Naturales y Museo, UNLP 
42: 1-37. 

Ronderos, M. M., G. R. Spinelli, and P. Sarmiento. 
2000. Preparation and mounting of biting midges 
of the genus Culicoides Latreille (Diptera: Cera- 
topogonidae) to be observed with a scanning elec- 
tron microscope. Transactions of the American 
Entomological Society 126(1): 125-132. 

Soria, S. de J., W. W. Wirth, and R. K. Chapman 1980. 
Insect pollination of cacao in Costa Rica. i. Pre- 
limary list of the ceratopogonid midges collected 
from flowers. Revista Theobroma 10(2): 61—68. 

Spinelli, G. R. 1983. Notas sobre Ceratopogonidae de 
la Republica Argentina (Diptera: Nematocera) II. 
Nuevos aportes al conocimiento del género For- 
cipomyia Meigen. Neotropica 29: 121-129. 

Wirth, W. W. and W. L. Grogan 1978. Notes on the 
systematics and biology of the biting midge, For- 
cipomyia elegantula Malloch (Diptera: Ceratopo- 
gonidae). Proceedings of the Entomological So- 
ciety of Washington 80(1): 94-102. 

Young, A. M. 1986. Habitat differences in cocoa tree 
flowering, fruit-set, and pollinator availability in 
Costa Rica. Journal of Tropical Ecology 2: 163— 
186. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 115-118 


AMMONOENCYRTUS CAROLINENSIS, N. COMB. (HYMENOPTERA: 
ENCYRTIDAE), A PARASITE OF LOBATE LAC SCALE 
PARATACHARDINA LOBATA (CHAMBERLIN) (HEMIPTERA: KERRIIDAE) 


MICHAEL E. SCHAUFF 


Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department 
of Agriculture, Bldg. 005, Rm. 133, Beltsville Agricultural Research Center—West, 10300 
Baltimore Avenue, Beltsville, MD 20705, U.S.A. (e-mail: mschauff@sel.barc.usda.gov) 


Abstract.—Ammonoencyrtus carolinensis (Meyer) (n. comb.) (Hymenoptera: Encyrti- 
dae) a parasite of lobate lac scale, Paratachardina lobata (Chamberlin) (Hemiptera: Ker- 
iidae), is distinguished from other species of Ammonoencyrtus, and a diagnosis and sum- 
mary of its known biology are given. Ammonoencyrtus carolinensis was previously re- 
ported attacking Mesolecanium nigrofasciatum (Coccidae). Parasitization of Keriidae is 


an unusual host shift for this group of parasitoids. 


Key Words: 
sitoid, Hymenoptera 


The lobate lac scale, Paratachardina lob- 
ata (Chamberlin) (Hemiptera, Keriidae), 
was discovered in Florida in 1999 (Hamon 
2001). A potentially devastating pest of 
trees and shrubs, this introduced scale at- 
tacks plants in 49 families (Pemberton 
2003) and has spread quickly across a large 
section of southern Florida and has become 
a great concern to growers and homeowners 
because of its broad host range. Economi- 
cally important hosts include both orna- 
mentals, such as hibiscus, and fruit trees, 
such as citrus and mango. There is also 
great concern about the effect that the scale 
may have on natural areas and the ecology 
of the Everglades because P. lobata can at- 
tack many native plants. The Agricultural 
Research Service, USDA, and the Univer- 
sity of Florida, in cooperation with the Flor- 
ida Department of Agriculture and Con- 
sumer Services, Division of Plant Industry 
(DPI), are conducting research to find out 
more about the scale and its natural ene- 
mies. 


Encyrtidae, Kerriidae, Ammonoencyrtus, Paratachardina, taxonomy, para- 


Early in 2003, I received specimens for 
identification of an encyrtid that had .been 
reared by University of Florida scientists 
from the lobate lac scale. Tentatively iden- 
tified as Ammonoencyrtus sp., these speci- 
mens were compared with the other de- 
scribed species in the genus, and I deter- 
mined that they were not conspecific with 
A. californicus (Compere) or A. bonariensis 
(Bréthes), the only two species placed in 
this genus. Further study with the aid of Dr. 
John Noyes of The Natural History Muse- 
um, London (BMNH), indicated that the lo- 
bate lac scale parasite agreed with the de- 
scription of Anicetus carolinensis Meyer 
(Meyer et al. 2001). However, there were 
no records of any encyrtids reared from 
Kerriidae in North America. Lobate lac 
scale is native to India and Sri Lanka (Ha- 
mon 2001, Miller and Ben-Doyv 2002). 
Upon study of paratypes of A. carolinensis, 
it was determined that specimens from the 
lac scale were conspecific. Therefore, Ani- 
cetus carolinensis is hereby transferred to 
Ammonoencyrtus. 


116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Genus Ammonoencyrtus De Santis 


Ammonoencyrtus 1s placed in the Cer- 
apterocerini, a loosely defined tribe that in- 
cludes Anicetus Howard, Paraceraptroce- 
rus Girault, Homosemion Annecke, Anase- 
mion Annecke, and others. The genera in 
this tribe are generally characterized by the 
possession of: antenna with scape flattened 
and triangular to subrectangular; funicle 
flattened, with 6 segments which are gen- 
erally much wider than long (Figs. 1—2) and 
with a apically truncate, 3-segmented clava 
(some Anicetus may have the intersegmen- 
tal sulci reduced so that the clava may ap- 
pear 1-segmented, e.g., A. chinensis Girault 
as figured by Annecke 1967); frontovertex 
separated from the face by an inverted U- 
shaped carina or depression (Figs. 3—4) 
with the face below excavated; and fore- 
wing largely infuscate and usually with one 
or more hyaline areas (often with a hyaline 
band paralleling the tip of the wing, (Figs. 
5—6) and the postmarginal vein present 
(barely produced in Homosemion and Ani- 
cetus). These genera are generally parasites 
or hyperparasites of soft scales (Coccidae). 

Ammonoencyrtus is close to Homose- 
mion and Anicetus but can be distinguished 
by: carina on frontovertex interrupted me- 
dially by the interscrobal prominence (Fig. 
3) (carina separating vertex from face entire 
in other genera, (Fig. 4); and forewing ba- 
sally with an infuscate area enclosing a hy- 
aline triangular area below the parastigma 
and basal to the linea calva (Fig. 5). Ani- 
cetus annulatus Timberlake has a condition 
very similar to that of A. carolinensis and 
this character may prove of dubious value. 
Both A. carolinensis and A. californicus 
have distinct brown stripes that parallel the 
dorsal margin of the oral cavity contrasting 
with the lighter face below the toruli. De 
Santis (1963), in his redescription of A. 
bonarensis, does not mention this coloring. 
However, a series of three specimens in the 
National Museum of Natural History, 
Washington, D.C. (USNM) collected in 
Uruguay from Saissetia oleae that may rep- 


resent an additional new species, do have a 
darkened area (stripe) under the toruli. 

Little has been written about the males 
of A. californicus and A. carolinenis. They 
share the following characters: generally 
dark brown or black with metallic reflec- 
tions on the head and thorax; antennae with 
6 funiculars, all longer than wide and each 
with numerous setae which are longer than 
the width of the segment; mandibles triden- 
tate; and forewings hyaline. The antennae 
and legs vary from yellow (carolinensis) to 
brown (californicus). These characters 
agree with the illustrations and description 
of males of bonarensis given by De Santis 
(1964). 

Ammonoencyrtus was described by De 
Santis (1964) with Cerapterocerus bonar- 
iensis (Brethes) as type species. Eusemion 
californicum Compere was transferred to 
Ammonoencyrtus by Annecke (1967). 


Ammonoencyrtus carolinensis (Meyer), 
new combination 
Ces, Il, 35 5) 


Anicetus carolinensis Meyer 2001: 687. 


Diagnosis.—Anicetus carolinensis, was 
described from North Carolina, as a parasite 
of terrapin scale (Mesolecanium nigrofacia- 
tum (Pergande) (Coccidae)), and is found 
on fruit trees and shrubs in southeastern 
U.S. It can be separated from the other 
known North American species (A. califor- 
nicus) by the following: antennae, face, ver- 
tex, and dorsal thorax mostly honey yellow, 
mesoscutum sometimes darker and with 
metallic purplish reflection (dark brown to 
black in A. californicus); and femora and 
tibiae honey yellow (at least partly dark 
brown in californicus and bonarensis) 

Distribution.—North Carolina, Florida, 
Virginia. The Virginia record is heretofore 
unreported, but a female specimen was col- 
lected by D. R. Smith in a Malaise trap in 
Fairfax Co., VA, in 1989 (USNM). 

Discussion.—Use of the lobate lac scale 
as a host by A. carolinensis is unusual. The 
other hosts known for this genus are soft 


VOLUME 107, NUMBER 1 


117 


Figs. 1-6. 
dodonia Ferriére, female. 2, Antenna. 4, Face. 6, Fore wing. 


scales (Coccidae), and I must assume that 
A. carolinensis has crossed over onto the 
invasive P. lobata from its native hosts. As 
noted by Meyer et al. (2001), A. carolinen- 
sis was only reared from Mesolecanium ni- 


grofasciatum, but it would oviposit in other 


soft scales with no further development. 
That this species could complete develop- 


1, 3, 5, Ammonoencyrtus carolinensis, female. 1, Antenna. 3, Face. 5, Fore wing. 2, 4, 6, Anicetus 


ment in an introduced species from another 
family of scales is contrary to what might 
be expected. While only a few A. caroli- 
nensis have been reared from thousands of 
lobate lac scale examined (KR W. Howard, 
personal communication) it has been reared 
from more than one scale specimen and 
seems to be a primary parasite. At least one 


118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


individual was dissected directly from in- 
side the body of a P. lobata (E W. Howard, 
personal communication) so the possibility 
of an error in the rearing seems unlikely. 


ACKNOWLEDGMENTS 


I thank Dr. Michael Gates (SEL) who 
was instrumental in preparing the illustra- 
tions and specimens and who made several 
useful comments on the manuscript. Dr. 
John Noyes (The Natural History Museum) 
reviewed the manuscript and provided 
much useful information and help. Dr. EF W. 
Howard (University of Florida) supplied bi- 
ological information and specimens,and Dr. 
J. R. Meyer (North Carolina State Univer- 
sity) provided paratype specimens for the 
USNM collection. The paper was reviewed 
by J. S. Noyes, EF W. Howard, and an anon- 
ymous reviewer, and I am grateful for their 
effort. 


LITERATURE CITED 


Annecke, D. P. 1967. The genera Anicetus Howard, 
1896, Paraceraptrocerus Girault, 1920 and allies, 
with descriptions of new genera and species (Hy- 
menoptera: Encyrtidae). Transactions of the Royal 
Entomological Society of London 119: 99-169. 

De Santis, L. 1964. Encirtidos de la republica Argen- 
tina (Hymenoptera: Chalcidoidea). Anales de la 
Comision de Investigacion Cientifica Provincia de 
Buenos Aires Gobernacion 4: 9—422. 

Hamon, A. 2001. Lobate lac scale, Paratachardina lob- 
ata lobata (Chamberlin) (Hemiptera: Kerriidae). 
Pest Alert. http://www.doacs.state.fl.us/~pi/enpp/ 
ento/paratachardina.html (30 October 2002). 

Meyer, J. R., C. A. Nalepa, and C. Devorshak. 2001. 
A new species of Anicetus (Hymenoptera: Encyr- 
tidae) parasitizing terrapin scale, Mesolecanium 
nigrofasciatum (Hemiptera: Coccidae). Florida 
Entomologist 84: 686-690. 

Miller, D.R. and Y. Ben-Dov. 2002. ScaleNet http:// 
www.sel.barc.usda.gov/scalenet/scalenet.html (30 
October 2002). 

Pemberton, R. W. 2003. Potential for biological control 
of the lobate lac scale, Paratachardina lobata lok- 
ata (Hemiptera: Kerriidae). Florida Entomologist 
86: 353-360. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 119-122 


A NEW SPECIES OF THE GENUS AGETOCERA HOPE (COLEOPTERA: 
CHRYSOMELIDAE: GALERUCINAE) FROM CHINA 


LI-JIE ZHANG AND XING-KE YANG 


Institute of Zoology, Chinese Academy of Sciences, Beijing, 100080, P. R. China (e- 


mail: yangxk @io0z.ac.cn) 


Abstract.—Agetocera biclava, n. sp., is described from Guangxi Autonomous Region, 
P. R. China. The adult male, antennae and abdominal sternites of the male and female, 
male aedeagus, and female spermatheca are illustrated. 


Key Words: 


The genus Agetocera Hope (1840) was 
established as a monotypic genus contain- 
ing A. mirabilis Hope from India. The ge- 
nus currently includes 23 world species and 
is mainly distributed in the Oriental Region. 
Sixteen species are known from China, ten 
of which are endemic. 

With the studies on Agetocera by Chen 
(1942, 1964), Chujo (1962), Gressitt and 
Kimoto (1963), Kimoto (1989), Jiang 
(1992), and Yang et al. (1997, 2001), the 
taxonomy of this genus is well known. Us- 
ing the paper by Yang et al. (2001), we dis- 
covered another new species of Agetocera 
while identifying specimens from Guangxi 
Autonomous Region. This paper is a sup- 
plement to Yang et al. (2001). 


Agetocera biclava Zhang and Yang, 
new species 
(Fig. 1-8) 


Description.—Head, pronotum, and scu- 
tellum reddish brown; antenna covered with 
long hairs, yellowish brown to brown, last 
two segments dark brown; ventral surface 
of thorax yellowish brown or lighter; ab- 
domen and femora yellowish brown, apices 
of femora, tibiae, and tarsi black; elytron 


Coleoptera, Chrysomelidae, Galerucinae, A getocera, taxonomy, new species 


darkish blue. Length 10.5—12.0 mm; width 
4.8-5.0 mm. 

Male: Head exerted, narrower than pro- 
thorax, with vertex convex, smooth and im- 
punctate; postantennal tubercles strong; 
clypeus raised, with concavity in middle of 
frontal margin. Maxillary palpus large, 
third segment longer than fourth, fourth mi- 
nute and pointed at apex. Eyes strongly 
convex; interspace between eyes nearly 2 
times as wide as diameter of eye. Antenna 
(Fig. 2) extending behind to middle of el- 
ytra; first segment robust; second shortest; 
third 2 times as long as second and slightly 
longer than fourth; fourth to fifth equal in 
length; sixth longer than fourth and sube- 
qual to seventh, each broadened at apices; 
eighth to tenth longer than preceding seg- 
ments and subequal in length; eighth and 
ninth very strongly swollen, eighth with an 
oval concavity on dorsum near apex; elev- 
enth 1.6 times as long as tenth. Pronotum 
1.3 times as broad as long, narrow poste- 
riorly, broadened anteriorly; lateral margins 
and basal margin bordered, front margin not 
bordered; disc sparsely punctuate and with 
one transverse concavity on each side, with 
punctures on each of four corners. Scutel- 
lum cuneiform and impunctate, concave at 


120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


5 


Figs. 1-8. 


S 
: . | ‘ 
6 val 
7 \ 


Agetocera biclava. 1, Dorsal habitus, male. 2, Male antenna. 3, Female antenna. 4, Last visible 


sternite of female. 5, Last visible sternite of male. 6, Lateral view of male aedeagus. 7, Dorsal view of male 


aedeagus. 8, Spermatheca. 


base. Elytron 3.6 times longer than broad; 
disc convex, more broadly rounded apically 
than basally; humerus raised; a ridge behind 
humerus, parallel with lateral margin; disc 
covered with dense punctures, interspaces 
larger than diameter of a puncture; epipleu- 
ron reaching apex of elytra, slightly bend 


toward outer side, sparsely and finely punc- 
tate. Meson of last visible sternite deeply 
concave; pygidium rounded apically (Fig. 
5). Aedeagus in Figs. 6—7; in dorsal view 
lateral margins sinuate, broadened in mid- 
dle, then narrowed basally and apically, 
broadened at apex and base; in lateral view 


VOLUME 107, NUMBER 1 


apex of aedeagus bent ventrally at nearly a 
right angle. 

Female: Antennal segment three 2 times 
as long as second, third to seventh subequal 
in length, eighth to tenth subequal to each 
other in length, eleventh 1.5 times as long 
as tenth (Fig. 3). Pronotum 1.5 times as 
broad as long. Last visible sternite with 
long apical projection in middle (Fig. 4). 
Spermatheca (Fig. 8) with apex invaginated 
toward base of capsule, C-shaped, walls ob- 
viously thicker than that of ductus and sur- 
face apparently smooth at lower magnifi- 
cation. 

Types.—Holotype ¢, P. R. China: Gu- 
angxi Auto. Reg.: Tianlin, 1,600 m, 5-VI- 
2002, Coll. Jiang Guo-fang. Paratypes: | d, 
1 @, same locality as holotype, 1,300— 
1,400 m, 28-VI-2002, Coll. Liu Jian-wen; 
2 2, 1,600 m, 5-VI-2002, Coll. Jiang Guo- 
fang. All deposited in the Zoological Mu- 
seum, Institute of Zoology, Chinese Acad- 
emy of Sciences, Beijing, P. R. China. 

Etymology.—The specie epithet is based 
on the enlarged eighth and ninth antennal 
segments of the male. 

Remarks.—All species of Agetocera 
have the eighth and ninth segments of the 
male antennae modified except Agetocera 
filicornis Laboissiere. This is an important 
and easy character to identify different spe- 
cies. Also, the shape of last female sternite, 
aedeagus, and spermatheca are very differ- 
ent among species. This new species can be 
separated from other known species by the 
enlarged eighth and ninth segments of the 
male antennae, the long apical projection of 
the last sternite of the female, the apex of 
aedeagus bent ventrally in lateral view, and 
the C-shaped spermatheca. 

Agetocera biclava is similar to Agetocera 
filicornis, but differs in the shape of the 
male antennae and the last visible sternite 
of the female, the former with the eighth 
and ninth segments of the male antenna 
swollen and the last visible sternite with a 
long apical projection in the middle, with 
the projecting portion thin. In A. filicornis, 
the male antennae are normal, not swollen, 


121 


and the apical projecting portion of the last 
visible sternite of the female is broad. This 
new species is also similar to A. taiwana 
Chujo, as in the following key: 

This new species may be placed in the 
previous key to species (Yang et al. 2001) 
as follows: 


14. Male with antennal segment 8 shorter than 5 


toh7 (Combinedemiie aes eee uae 15 
— Male with antennal segment 8 longer than 5 
tO "COMbINEC ye acre cihe ech eke wey sts cee 16 


15. In male, third antennal segment 2 times lon- 
ger than second; eighth with an oval concavy- 
ity on dorsum near apex; ninth elongate, sub- 
equal to eighth in length; tibiae and tarsi black 
A. biclava, n. sp. 
— In male, third antennal segment 1.5 times lon- 
ger than second; eighth with a rectangular 
convexity apically; ninth deplanate and broad, 
almost triangular, shorter than eighth; tibiae 
and tarsi yellowish brown, last two tarsal seg- 
PROMS [NOW scogoccssogee A. taiwana Chujo 
16. In male, ninth antennal segment U-shaped, 
half as long as tenth; female with last visible 
sternite concave mesally A. similes Chen 
— In male, ninth antennal segment not U-shaped 
and more than half as long as tenth; female 
with last visible sternites convex mesally .... 17 


ACKNOWLEDGMENTS 


We thank Wen-zhu Li and Si-qin Ge for 
drawing part of the figures. This research 
was supported by the Chinese Academy of 
Sciences Innovation Program. 


LITERATURE CITED 


Chen, S. H. 1942. Galerucinae nouveaux de la Fauna 
Chinoise. Notes d’Entomologie Chinoise IX (3): 
1-67. 

. 1964. New genera and species of Galerucinae 

13(2): 


from China. Acta Entomologica Sinica 
201-211. 

Chujo, M. 
melidae (Insecta: Coleoptera) from Formosa. Part 


1962. A taxonomic study on the Chryso- 


XI. Galerucinae. Philippine Journal of Science 
91(1—2): 1-239. 

Gressitt, J. L. and S. Kimoto. 1963. The Chrysomeli- 
dae (Coleoptera) of China and Korea. Pacific In- 
sects Monograph 1B: 301—1026. 

Hope, EK W. 1840. The Coleopterist Manual. Part III. 
London, 191 pp. 

Jiang, S. Q. 1992. Coleptera: Chrysomelidae: Galeru- 
cinae, pp. 646-674. /n Chen, S. H., ed. Insects of 


122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Hengduanshan Mountains Region I. Science ince (Coleoptera: Chrysomelidae). Acta Zootax- 
Press, Beijing. onomica Sinica 22(4): 384-391. 

Kimoto, S. 1989. Chrysomelidae (Coleoptera) of Thai- | Yang, X. K., S. Q. Ge, and W. Z. Li. 2001. Revision 
land. Cambodia. Laos and Vietnam. IV. Galeru- of the Genus Agetocera Hope (Coleoptera: Chry- 
cinae. Esakia No. 27, 241 pp. somelidae: Galerucinae). Oriental Insects 35:105— 

Yang, X. K., W. Z. Li, and J. Yao. 1997. The Galer- 154. 


ucinae beetles of Xishuangbanna. Yunnan Prov- 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 123-158 


INTRODUCED SCALE INSECTS (HEMIPTERA: COCCOIDEA) OF THE 
UNITED STATES AND THEIR IMPACT ON U.S. AGRICULTURE 


DOUGLASS R. MILLER, GARY L. MILLER, GREG S. HODGES, AND JOHN A. DAVIDSON 


(DRM, GLM) Systematic Entomology Laboratory, PSI, Agricultural Research Service 
U.S. Department of Agriculture, Builg. 005, BARC-West, Beltsville, MD 20705, U.S.A. 
(e-mail: dmiller@sel.barc.usda.gov; gmiller@sel.barc.usda.gov); (GSH) Florida Depart- 
ment of Agriculture and Consumer Services, Division of Plant Industry, Gainesville, FL 
32614-7100, U.S.A. (e-mail: hodgesa@doacs.state.fl.us); (JAD) Department of Entomol- 
ogy, University of Maryland, College Park, MD 20742, U.S.A. (e-mail: jd12@umail. 
umd.edu) 


Abstract.—We provide information on 255 species of scale insects that are considered 
adventive or introduced in the United States. Included for each species, where applicable, 
is reference to: the state and earliest collection date in which the invader was first dis- 
covered; whether it is currently established in the United States; its pest status in the 
United States along with a validation citation; its principal hosts; and its zoogeographical 
region of origin. Information is provided about trends of pest introductions and on native 


scale-insect pests in the United States. 


Key Words: 
graphic origin 


Scale insects are major agricultural pests 
and pose serious problems when introduced 
into new areas of the world without natural 
enemies. Historically, introduced scale in- 
sects have fostered the development of 
strategies for dealing with exotic pests ei- 
ther after they were detected in the United 
States or for prevention of new introduc- 
tions. Devastation caused by the cottony 
cushion scale, [cerya purchasi Maskell, in 
California was one of the reasons for the 
beginning of a new subdiscipline in ento- 
mology called biological control (Van 
Driesche and Bellows 1996). The introduc- 
tion of the San José scale, Diaspidiotus per- 
niciosus (Comstock), a serious pest of stone 
fruit crops, also had major impact and was 
one of the primary reasons that Congress 
passed the U.S. Plant Quarantine Act in 
1912 (Marlatt 1920). 


scale insects, adventive species, invasive species, exotic species, pests, geo- 


In the past 20 years, 21 new exotic spe- 
cies of scale insects have been detected in 
the continental United States. Of these, 11 
currently are considered pests and several 
others have pest potential. Some of the 
most serious pest introductions are: white 
mango scale, Aulacaspis tubercularis New- 
stead, a pest of mango first detected in 
2002; cycad aulacaspis scale, Aulacaspis 
yasmumatsul Takagi, a pest of cycads de- 
tected in 1996; hibiscus mealybug, Maco- 
nellicoccus hirsutus (Green), a polyphagous 
pest that is common on hibiscus detected in 
1999; miscanthus mealybug, Miscanthicoc- 
cus miscanthi (Takahashi), a pest of mis- 
canthus grass detected in 1989; papaya 
mealybug, Paracoccus marginatus Wil- 
liams and Granara de Willink, a pest of pa- 
paya and hibiscus detected in 1999; lobate 
lac scale, Paratachardina lobata (Cham- 


124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


berlin), a pest of numerous ornamental and 
native species detected in 1999; black par- 
latoria scale, Parlatoria ziziphi (Lucas), a 
pest of citrus detected in 1985; and vine 
mealybug, Planococcus ficus (Signoret), a 
pest of fig and grapes detected in 1994. 

Although one of us (DRM) has partici- 
pated in several programs aimed at gaining 
definitive data on introduced insects in the 
United States, e.g., McGregor (1973), Sail- 
er (1978, 1983), no comprehensive analysis 
is available on the adventive scale insects 
of the United States. Therefore, the objec- 
tives of this study are to: 1) list the adven- 
tive scale insects of the United States and 
record data on: first detection date in the 
United States (with validation source), de- 
gree of certainty that the species is adven- 
tive, pest status in United States, reference 
to pest status when available, principal 
hosts, zoogeographic area of origin, vali- 
dation source of origin when available, and 
eradication status; 2) analyze these data and 
determine if patterns exist; and 3) compare 
these data with those of Sailer (1978) for 
all insects. 


MATERIALS AND METHODS 


We have treated species as introduced 
only when they are established in an exte- 
rior environment. We have excluded species 
that were discovered in nurseries or green- 
houses and ultimately were eradicated, or 
species that were taken in quarantine. As far 
aS we are aware, no species have been pur- 
posefully introduced for biological control, 
although at least one, Trabutina mannipara 
(Hemprich and Ehrenberg), has been con- 
sidered as a biological control agent of salt- 
cedar, Tamarix spp. (DeLoach et al. 1996). 

Some of the data that we have compiled 
are subjective. For example, in trying to de- 
termine the area of origin of a species, we 
have relied on statements in the literature, 
distribution of less widespread congeneric 
(and hopefully related), and, in a few in- 
stances, our best guess. Determining if a 
species is adventive or indigenous has not 
always been clear, particularly when it may 
have its origin from Nearctic Mexico or 


Central America. In some cases, we have 
been unable to find evidence or area of or- 
igin, and in other situations we have been 
unable to decide which of two zoogeo- 
graphic regions is the area of origin. Thus, 
the statistics given in the tables and charts 
do not always add up to the exact number 
of invasive species (255). 

For recording the earliest record of an ad- 
ventive in the United States, we have relied 
on published records and specimens ex- 
amined in the collections of The National 
Entomological Collection of the National 
Museum of Natural History, Beltsville, 
Maryland (USNM); The Florida State Col- 
lection of Arthropods, Gainesville, Florida 
(FSCA); and The Bohart Museum, Davis, 
California (UCD). It is possible that older 
records exist for some species, but we an- 
ticipate that the dates reported here are 
close approximations of the first U.S. de- 
tections. Although some species may have 
been introduced in 1700’s or earlier, little or 
no documentation to verify the time or lo- 
cation of these introductions is available. 
For most of these, the earliest detection re- 
cord is from the 1870’s or 1880’s when 
specimens were first deposited in collec- 
tions, even though these species might have 
been present decades earlier. 

For determining the pest status of a spe- 
cies, we have relied on the literature. If 
statements are made that a species is a pest, 
is causing harm, requires control measures, 
is economically important, or some com- 
parable comment, we are considering these 
species to be pests. Our decisions about 
pest-status intensity, i.e., minor pest, pest, 
or serious pest, are based on our experience 
or statements in the literature. We provided 
references on pest status when available. 

The principal adventive scale hosts were 
grouped in categories that associated large 
numbers of scale-insect species. These host- 
plant groupings are not always comparable, 
e.g., polyphagous vs. tree species, but they 
provide useful information on high-risk 
host categories. Our definition of polypha- 
gous is any known from 10 or more fami- 
lies of host plants. 


VOLUME 107, NUMBER 1 


In our previous papers (Miller et al. 
2002, Miller and Miller 2003), we deviated 
from the strict definition of an invasive spe- 
cies and used it synonymously with adven- 
tive, alien, exotic, and immigrant (see 
Wheeler and Hobeke 2001 for discussion of 
terminology). Here, we restrict the defini- 
tion of an invasive species to conform to 
prevailing usage, i.e., a species that has in- 
vaded a HABITAT (not necessarily a polit- 
ical unit) and is causing harm or has the 
potential of causing harm. 

In compiling statistics about the scale in- 
sect fauna, we have grouped the scales into 
five categories, i.e., all scales, diaspidids, 
pseudococcids, coccids, and all other scale 
families. 


RESULTS 


Based on information in ScaleNet (Ben- 
Dov et al. 2003), there are 7,355 species of 
scale insects worldwide, 2,369 species of 
diaspidids (32% of all scales in the world), 
2,048 pseudococcids (28%), 1,129 coccids 
(15%), and 1,179 species in all other fam- 
ilies (16%). In the continental United 
States, there are 1,019 species of scales, 320 
species of diaspidids (31% of all scales in 
the United States), 356 species of pseudo- 
coccids (35%), 105 coccids (10%), and 238 
species in all other families (23%). Assum- 
ing that these groups have been equally col- 
lected and studied (which is unlikely), it ap- 
pears that diaspidid diversity is about the 
same proportion in the United States as in 
the rest of the world, the diversity of pseu- 
dococcids and other scale families is high, 
and the diversity of coccids is low. 

Table 1 provides information on 255 
scale-insect species that are hypothesized as 
adventives in the United States (1 aclerdid, 
10 asterolecaniids, | cerococcid, 43 coc- 
cids, 1 conchaspidid, 132 diaspidids, 5 er- 
iococcids, | kerriid, 3 margarodids, 1 or- 
theziid, | phoenicococcid, and 56 pseudo- 
coccids). When comparing the number of 
introduced species in the United States with 
the total number of world species in the five 
categories, the percentages for each are 


125 
about the same. That is, 255 species of in- 
troduced scale insects comprise about 3% 
of the world’s scale fauna; 132 introduced 
diaspidids constitute about 6% of the 
world’s armored scale fauna; 56 pseudococ- 
cids is about 3% of the world’s mealybug 
fauna; 43 coccids is about 4% of the soft 
scale fauna; and 24 is about 1% of the 
world’s “‘all other families”” fauna. The suc- 
cess rate of introduction appears relatively 
similar for the five categories of scale in- 
sects. The possible exception is the taxa 
placed in the “‘all other families” category, 
which may have a slightly lower rate of in- 
troduction success. 

Examination of the relative size of the 
adventive scale diversity in the United 
States with that of the native fauna reveals 
that more than 25% of the species of all 
scale insects in the United States are intro- 
duced species. Even more surprising is that 
40% of all U.S. diaspidids and coccids 
found in the United States are introduced. 
Although the adventive loads in the pseu- 
dococcids (15%) and “‘all other families” 
(10%) is considerably lower, they still are 
high compared with other insect groups. 
About 2,000 species of introduced insects 
occur in the continental United States, com- 
prising only about 1—2% of the total insect 
fauna (Wheeler and Hoebeke 2001). The 
255 species of introduced scale insects in 
the United States represent about 13% of 
all adventive insects in the U.S. If we com- 
pare the number of United States scales 
with the total insect fauna (assuming that 
there are 90,000 species of insects in the 
United States (Kosztarab and Schaefer 
1990)), then the 1,019 scale insects in the 
United States comprise only about 1% of 
the insect fauna. These numbers reflect an 
abnormally large number of introduced 
scale insects in the United States compared 
with other insect groups. 

It is interesting to note that there was a 
peak of introductions between 1890 and 
1920 (Fig. 1), and that there has been a 
steady decline to the present. This trend is 
reasonably consistent for all five categories 


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VOLUME 107, NUMBER 1 


snosevydAjod 


snoseydAjod 


snoseyddjod 


snoseyddjod 


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‘ponunuoy “| qe. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


8e6l YIZUSSSIN 


8961 [9tueqoW 


(91ZUd JON) 


iPs(@) snoseyddjod isad B jou UuTeyID0UN 29061 ‘XL {poustqeiso DIDUAOUL DIJAIpIUOY 
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iTaYO snoseyddjod sod AJOAT €S6l “TH poystqrisa DAA{I]JAIS DIUOSU1A 
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jedioung doudIajoy “S' SOV uoneoipelg peonponuy 


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‘ponunuoy “| eqeL 


131 


VOLUME 107, NUMBER 1 


L661 ID 


L661 ILD 


6861 [PB jo NT 


Ipreuo0a] 


Wd XIMDUID sad v jou AOXT LI61 ‘ZV poysifqriso posnaja sidspuoiyy 

L661 ID ssaidAo ‘siadiu L661 ID 9S61 IIZUD MON (21031) 
Wd -nl Ajpetoadsa saayruos sod AJ ZEHI “VO poysi[qriso punuilu sidspjnivy 

ps6] Aysmoyoreg ssodru L661 I'D WNSn (pyonog) 
Wd = -nl Ajyeroadsa saayruos sod AJOX1 0061 ‘VIN poysijqrisa diadiunt sidspjnavy 

6661 [eB 19 premMoyH 666 ‘[B 19 premMoH 6661 ‘[@ 19 premoy (SPY L 
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pSol Aysmoyouyeg SNQnY Lo6l ID WNS) (pyonog) 
Wd ‘psoy snoseydostyjo ysod AJOXI] O88I ‘AN “TH ‘WO poystyqriso apsod sidspovjny 

LOOT PIOSAM WNSN YOOISWIOD 
ureyiaoun snoseydAdjod sod 10uTU AJOXT 6161 “TH poystyqeiso snsouids snjoipidsy 

8roel 
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Ivd S19JIUOS ysod AJM LE6Ll ‘AN poysyqeiso = aviuaiuojdxio snjoipidsy 

suevore 9L61 9FI9d WNSf (1]919490)D) 
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(BABSSBD) 9L6l 2PISd VOSA (1[919490)) 
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UIsLIO, s}soH smeig 1sog 01 poonpo.nuy prosody “Sn snes sa1sadg 
jediourig QUudIZJaYy “SY SOlpIV_ uoneoipelg poonponuy 
ul snqeig sad 
jpenunuoD Tf °192L 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


2 EEE 


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2 ul smvjg 1soq 
— 
Q ‘ponunuoy “TT 2x9eL 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


134 


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VOLUME 107, NUMBER 1 


snoseydAdjod 


spubuy 


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snoseydAjod 
DIADINDAY 
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snoseyddjod 


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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


136 


OHUN JO VAN 


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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


138 


8061 Hee 


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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


140 


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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


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144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


S of scale insects (Fig. 2), although a few 
os) o o& 
a3 a a more pseudococcids have been introduced 
SS is o over the past 60 years. We do not see the 
‘ eI Be increase that might have been anticipated 
BS o we) = . . . 5 > 
© 5) ES with rapid international transit and expand- 
c Ss} . 
38 s $288 2 & ed world trade. As suggested by Sailer 
= = 
Ane Om Azteca Gece ed (1978), the enactment of the Plant Quar- 
antine Act in 1912, and its intensive imple- 
mentation in 1920, might explain some ap- 
parent success in preventing more invaders 
from becoming established (Fig. 1). How- 
Blo 5 ene ever, complacency should not be the modus 
Se é o Ss He ARS 3 
es & Zi a = 'S operandi! Examination of the accumulation 
= 2 a = =| & curve (Fig. 3) shows a substantial increase 
in the adventive scale-insect load with a 
a =, rate of about one species each year. The 
vo ot : 0 é F o 
. oO a — economic impact of this load is consider- 
So) ae ° . 5 . ous 
ale a s Ee a s & | able; the impact will intensify as additional 
20 = = An Seas : 
32 2 2 > 288% » % 2] problematic invaders become established. 
me ie) i ; : 
Ba oi) BR a Es a Sg The majority of U. S. adventive scale in- 
eo i os i x 6 Rcnate 5 ° 
23 a Saas < 5 2 “ ==] sects (Fig. 4) originated in the Palearctic Re- 
aGO Fm ae £22201 gion (105 species), followed by the Oriental 
[= (67), Neotropical (43), Afrotropical regions 
uo] ° . 
oa _ = Se WS ese (22), Australasian (16), and Nearctic (16). 
Zz oD Wr) ¢ wr) oD oD © 5 O50 
z| 2 2 = I) Gate Some scales with uncertain origins (e.g., 
s- Bambusaspis bambusae (Boisduval)) are in- 
cluded within two regions. These data are 
ms 2 eee ee similar to Sailer (1978). Although he divided 
a s s See is his geographic information more finely than 
331 _ o nO ik OMe Ones ours, by combining his subunits into zoo- 
ex1NnN FY MN ARMAND hi : Pai 1 sas ed 
deaf) 2 5 A2s25 252 525] geographic regions his results were similar: 
aliens Ge “S$ 2% 2% 24] most species were from the Palearctic, fol- 
02 HS2U2 020201 Jowed by the Neotropical, Oriental, Afro- 
tropical, and Australasia. He considered 
Mexico as strictly Neotropical Region. 
In analyzing host data (Fig. 5), the over- 
whelming majority of invasive scale species 
Uv No) uo] ue} i 
Pe s a ae fe are polyphagous (112 species); 47 species 
ics || ‘a ma ey oa era are primarily found on trees; 32 are on 
es iss) isc} iS) iss} ° 3 . 
BA 3 3 Oe Sees, es grasses, including bamboo; 13 on orchids, 
6) 0 oO =) 5) 0 2) : i : 5 
bromeliads, or cacti; 12 on tropical fruit; 9 
7 v : ; 
E s 2) ge BS primarily on shrubs; and 30 on other hosts. 
3 2 Ss Sf To evaluate the pest status of invasive 
= = = SS x S 5 O 
= ales See AS Ce aS scale insects, we analyzed the data in two 
OT7,) = 3 —~ & S >= 3 D = 
Ss) Bel s ses = & 2 =6| ways: 1) Of the 255 introduced species of 
o (b) = aR SX SS ® 5 © 
“s Sul 3 8 8 os 8 S S 5S 8 scale insects in the United States, 191 
fe GO x $ i $0 So 0 © 
2 S SBSse SBSODE SG ©) are considered pests, and of these, 
2 Ba eeecae ce (75% dered pests, and of th 
N a a) 9S : : 
eS aM S LY = =) a ) acs SUAS (25%) are considered to be serious pests 
& yw a > > > 


(Fig. 6); 2) We also compared native (Table 
2) and adventive pests. There are 65 species 


VOLUME 107, NUMBER 1 145 


(1) Number of Adventive Scales at 30-year Intervals 


100 
80 


60 


Nurnber of Scales 


20 


Before 1889 1890-1919 1920-1949 1950-1979 1980-present 
Dates 


(2) Introduction of Scale Families (Overall and Selected) in the U.S. 


100 


& Before 1889 
80 @ 1890-1919 
1920-1949 


60 Hi 1950-1979 


© 1980-present 


Number of Scales 


20 


All Scale Families Diaspididae Pseudococcidae Coccidae All other Scale 
Families 
Family 


Figs. 1-2. Adventive scales. 1, Number of adventive scales in the United States recorded at 30-year intervals. 
2. Number of adventive scales (overall and selected families) in the United States reported at 30-year intervals. 


146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Accumulated Number of Adventive Scales at 30-year Intervals 


300 


250 


200 
7) 
2 
5 
172) 
S 150 
® 
Q 
E 
Ss 
z 
100 
50 
0 
Before 1889 1890-1919 1920-1949 1950-1979 1980-present 
Dates 
Origin of Adventive Scales in the U.S. 
7) 
2 
© 
15) 
7) 
x) 
~ 
o 
2 
E 
J 
2 


Australasian Nearctic Afrotropical Neotropical Oriental Palearctic 
Region 


Figs. 3-4. Adventive scales. 3, Accumulated number of adventive scales in the United States at 30-year 
intervals. 4, Origin of adventive scale species in the United States. 


VOLUME 107, NUMBER 1 147 


Host Plants of Adventive Scales to the U.S. 


120 


100 


80 


2 60 
Ss 
2 
40 
20 
0 
shrubs tropical fruit orchids &cacti other hosts grasses trees polyphagous 
Host 
Percent of Adventive Scale Pests 
- 
5 
) 
- 
VC) 
o 


All Scale Families Diaspididae Pseudococcidae Coccidae All Other Scale Families 
Family 


Figs. 5—6. Adventive scales. 5, Number of adventive scale species in the United States on specified hosts. 
6, Percent of adventive scale species that are pests in the United States. 


148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 2. List of native U.S. species of scale insects considered pests. 


Name of Native Scale Reference to Pest Status Pest Status 
Cerococcidae 
Cerococcus parrotti (Hunter) Lambdin and Kosztarab 1977 minor pest 
Cerococcus quercus Comstock Lambdin and Kosztarab 1977 minor pest 
Cerococcus kalmiae Ferris Lambdin and Kosztarab 1977 minor pest 
Coccidae 
Mesolecanium nigrofasciatum (Pergande) Kosztarab 1996 pest 
Neolecanium cornuparvum (Thro) Williams and Kosztarab 1972 pest 
Pseudophilippia quaintancii Cockerell Hamon and Williams 1984 minor pest 
Pulvinaria acericola (Walsh and Riley) Kosztarab 1996 pest 
Pulvinaria hydrangeae Steinweden Williams and Kosztarab 1972 minor pest 
Pulvinaria innumerabilis (Rathvon) Williams and Kosztarab 1972 pest 
Toumeyella liriodendri (Gmelin) Kosztarab 1996 pest 
Toumeyella parvicornis (Cockerell) MacAloney 1961 minor pest 
Toumeyella pini (King) Kosztarab 1996 minor pest 
Diaspididae 
Abgrallaspis ithacae (Ferris) Pirone 1970 pest 
Chionaspis americana Johnson Dodge and Rickett 1943 minor pest 
Chionaspis corni Cooley Baker 1972 minor pest 
Chionaspis furfura (Fitch) Kosztarab 1996 pest 


Chionaspis heterophyllae Cooley 

Chionaspis pinifoliae (Fitch) 

Clavaspis ulmi (Johnson) 

Comstockiella sabalis (Comstock) 
Cupressaspis shastae (Coleman) 
Diaspidiotus ancylus (Putnam) 

Diaspidiotus forbesi (Johnson) 

Diaspidiotus juglansregiae (Comstock) 
Diaspidiotus liquidambaris (Kotinsky) 
Diaspidiotus osborni (Newell and Cockerell) 
Diaspidiotus uvae (Comstock) 

Hemiberlesia neodiffinis Miller and Davidson 
Melanaspis obscura (Comstock) 

Melanaspis lilacina (Cockerell) 

Melanaspis tenebricosa (Comstock) 
Nuculaspis californica (Coleman) 
Quernaspis quercus (Comstock) 
Rhizaspidiotus dearnessi (Cockerell) 


Eriococcidae 

Eriococcus azaleae Comstock 
Eriococcus coccineus Cockerell 
Eriococcus quercus (Comstock) 
Kermesidae 

Allokermes kingii (Cockerell) 
Eriokermes gillettei (Tinsley) 
Nanokermes folium Bullington and Kosztarab 
Lecanodiaspididae 

Lecanodiaspis prosopidis (Maskell) 
Margarodidae 


Eumargarodes laingi Jakubski 
Margarodes meridionalis Morrison 
Matsucoccus acalyptus Herbert 


Negron and Clarke 1995 
Kosztarab 1996 

Miller and Davidson 1990 
Miller and Davidson 1990 
Michener et al. 1957 
Polavarapu et al. 2000 
Beardsley and Gonzalez 1975 
Gill 1997 

Baker 1972 

Miller and Davidson 1990 
Johnson et al. 1999 

Miller and Davidson 1998 
Stoetzel and Davidson 1971 
Miller and Davidson 1990 
Baker 1972 

Johnson and Lyon 1988 
Herbert 1936 

Lacroix 1926 


Miller and Miller 1992 
Miller and Miller 1992 
Kosztarab 1996 


Kosztarab 1996 
Miller 1991 
McConnell and Davidson 1959 


Kosztarab 1996 


Spink and Dogger 1961 
Kouskolekas and Self 1973 
McCambridge 1974 


serious pest 
serious pest 
minor pest 
pest 

minor pest 
pest 

pest 

pest 

minor pest 
minor pest 
serious pest 
minor pest 
serious pest 
minor pest 
serious pest 
pest 

minor pest 
minor pest 


pest 
serious pest 
pest 


minor pest 
minor pest 
minor pest 


minor pest 


minor pest 
pest 
pest 


VOLUME 107, NUMBER 1 


Table 2. Continued. 


Name of Native Scale 


Matsucoccus bisetosus Morrison 
Matsucoccus gallicolus Morrison 
Matsucoccus vexillorum Morrison 
Stomacoccus platani Ferris 
Xylococculus betulae (Pergande) 
Xylococculus macrocarpae (Coleman) 


Pseudococcidae 


Ehrhornia cupressi (Ehrhorn) 

Oracella acuta (Lobdell) 

Paracoccus juniperi (Ehrhorn) 

Phenacoccus acericola King 

Phenacoccus gossypii Townsend and Cockerell 
Phenacoccus madeirensis Green 


Phenacoccus minimus Tinsley 
Phenacocus solani Ferris 
Pseudococcus maritimus (Ehrhorn) 
Pseudococcus viburni (Signoret) 
Rhizoecus floridanus Hambleton 
Rhizoecus simplex (Hambleton) 
Trionymus haancheni McKenzie 


Reference to Pest Status 


McKenzie 1942 


Pest Status 


serious pest 


149 


Putoidae 


Puto cupressi (Coleman) 
Puto sandini Washburn 


Parr 1939 minor pest 
McKenzie 1943b pest 

Gill 1993 serious pest 
Gill 1993 minor pest 
Gill 1993 pest 
Herbert 1920 minor pest 
Negron and Clarke 1995 pest 
Calkins 1946 minor pest 
Britton 1924 minor pest 
Myers 1932 minor pest 
Williams and Granara de pest 

Willink 1992 

Miller 1991 minor pest 
Miller 1991 minor pest 
Geiger and Daane 2001 serious pest 
Whiting and Hoy 1998 pest 
Hamlen 1974 minor pest 
Miller 1991 minor pest 
Osborn 1952 pest 
Miller 1991 minor pest 
Washburn 1965 minor pest 


of native scale insects that are considered 
pests (Cerococcidae 3, Coccidae 9, Diaspi- 
didae 22, Eriococcidae 3, Kermesidae 3, 
Lecanodiaspididae 1, Margarodidae 9, 
Pseudococcidae 13, Putoidae 2). Of the na- 
tive pests, only 9 (14%) are rated as serious, 
considerably less than the 25% figure for 
introduced pests. A surprisingly large num- 
ber in the category “‘all other families” are 
native pest species (Fig. 7). The large per- 
centages of these are margarodids, espe- 
cially species of Matsucoccus. 

The number of native pest species (65), 
in combination with the introduced pests 
(191), totals 256 species of scale insects 
that are considered pests in the United 
States or 25% of the all the scale insects in 
the U.S. 


DISCUSSION 


Many of our findings are similar to those 
reported by Sailer (1978, 1983) for all in- 
sects. The similaries are: the majority of in- 


troduced species are indigenous to the Pa- 
learctic Region; introductions occurred at 
an increasing rate until about 1920 and de- 
clined since; the adventive species load is 
increasing at an alarming rate when the ac- 
cumulation curve data are analyzed; within 
the adventive species that occur in the Unit- 
ed States, slightly more than half are pests 
or minor pests, slightly less that one quarter 
are major pests, and slightly more than one 
quarter are non- pests. 

There are some interesting differences 
between the statistics for all insects and 
scales. Sailer (1978) indicated that about 
1% of the insect and mite fauna in the Unit- 
ed States is introduced, but for scale insects, 
it is about 25%. Although scales make up 
only slightly more than 1% of the U.S. in- 
sect fauna, they represent about 12% of the 
adventive insect fauna. Scale insects may 
represent a substantial percentage of the ad- 
ventive insect fauna in other countries as 
well. For example, Charles (1998) indicated 


150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Native and Adventive Scale Pests in the U.S. 


Percent 


All Scale Families Diaspididae 


Fig. 7. 


that 24 species of Coccoidea represented 
24% of the total “exotic” arthropod pests 
of fruit crops in New Zealand. 

Gill (1997) summarized information on 
the rather impressive list of diaspidid spe- 
cies that have been eradicated or have not 
been detected after a period of establish- 
ment in California. Species such as Hall 
scale, Mercetaspis halli (Green), and date 
palm scale, Parlatoria blanchardi (Targioni 
Tozzetti), required major, long-term eradi- 
cation effort (Boyden 1941, Fosen et al. 
1953), but the successful elimination of 
these pests may have saved of millions of 
dollars annually. Gill (1997) mentioned 25 
species of armored scales as having been 
eradicated or disappeared, an impressive re- 
cord when compared with eradication suc- 
cess in many other invasive insects. Unfor- 
tunately, no comparable information on 
eradication efforts in other states is avail- 
able. 


Pseudococcidae 


Coccidae All other Scale Families 


Adventive scales in relation to native scales as pests in the United States. 


SUMMARY 


1) There are 255 introduced scale-insect 
species in the United States and a total 
scale-insect fauna of 1,019 species. 

2) The U.S. fauna includes an unusually 
large number of introduced scale insects 
compared with most other insect groups. 

3) The largest number of introduced scale 
insects originated in the Palearctic Re- 
gion and they usually are polyphagous. 

4) Of the 255 introduced species, about 
75% are pests. 

5) Of the 766 indigenous U.S. scale spe- 
cies, only about 8% are pests. 

6) The largest number of introduced scale 
species was detected between 1890 and 
1920, and the number of introductions 
has decreased over time. 

7) Recent rates of detection are about one 
species each year, which is dramatically 
increasing the pest load, placing an ever- 


VOLUME 107, NUMBER 1 


increasing pest burden on U.S. agricul- 
ture. 


ACKNOWLEDGMENTS 


We are especially grateful to Al Wheeler 
(Department of Entomology, Clemson Uni- 
versity, Clemson, SC), Avas Hamon (re- 
tired, Florida Department of Agriculture 
and Consumer Services, Division of Plant 
Industry, Gainesville, FL), Tom Henry 
(Systematic Entomology Laboratory), Ron- 
ald Ochoa (Systematic Entomology Labo- 
ratory) and Douglas Williams (Department 
of Entomology, The Natural History Mu- 
seum, London, UK) for their careful ex- 
amination of a draft of this manuscript and 
for their thoughtful criticisms which added 
significantly to the final publication. We 
also thank Penny Gullan (Department of 
Entomology, University of California, Da- 
vis) for checking information on slides in 
the Bohart Museum. We also are indebted 
to Jonathan Eibl (Systematic Entomology 
Laboratory) for his help with manuscript 
format. 


LITERATURE CITED 


Armitage, H. M. and H. L. McKenzie. 1952. Present 
status of the olive pollinia scale Pollinia pollini 
(Costa) in California (Homoptera; Coccoidea; As- 
terolecanidae). Bulletin of the California Depart- 
ment of Agriculture 41: 115-121. 

Avidoy, Z. and I. Harpaz. 1969. Plant Pests of Israel. 
Israel Universities Press, Jerusalem, 549 pp. 
Baker, W. L. 1972. Eastern forest insects. United States 
Department of Agriculture, Forest Service, Mis- 

cellaneous Publications 1175, 642 pp. 

Balachowsky, A. S. 1948. Les cochenilles de France, 
d’Europe, du nord de |’ Afrique et du bassin Méd- 
iterranéen. IV. Monographie des Coccoidea, clas- 
sification—Diaspidinae (Premiere partie). Actuali- 
tés Scientifiques et Industrielles 1054: 243-394. 

. 1950. Les cochenilles de France, d’Europe, 

du Nord de |’ Afrique et du Bassin Méditerranéen. 

V.—monographie des Coccoidea; Diaspidinae 

(deuxiéme partie) Aspidiotini. Entomologique Ap- 

plicata Actualités Sciences et Industrielles 1087: 

397-557. 

. 1953. Les cochenilles de France d’ Europe, du 

Nord de l’Afrique, et du bassin Méditerranéen. 

VII.—Monographie des Coccoidea; Diaspidinae- 

IV, Odonaspidini-Parlatorini. Entomologique Ap- 


15] 


plicata Actualités Sciences et Industrielles 1202: 

725-929. 

. 1954. Les cochenilles Paléarctiques de la tribu 

des Diaspidini. Mémoires Scientifiques de |’nstitut 

Pasteur, Paris. 450 pp. 

1958. Les cochenilles du Sahara Francais. 
lere partie. Travaux de |’Institut de Recherches 
Sahariennes 3: 31-53. 

Bartlett, B. R. 1978. Coccidae, pp. 57—74. Jn Clausen, 
C. P., ed. Introduced Parasites and Predators of 
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PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 159-161 


DESCRIPTION OF A NEW SPECIES OF ERIOTREMEX BENSON 
(HYMENOPTERA: SIRICIDAE) FROM JAPAN 


IcHII TOGASHI 


1-chome, Honmachi, Tsurugi-machi, Ishikawa Prefecture 920-2121, Japan 


Abstract.—A woodwasp, Eriotremex makiharai, n. sp., from Japan is described and 
illustrated. A key is provided for the Japanese species of Eriotremex. 


Key Words: 


Recently, I received one specimen of a 
siricid woodwasp from Mr. H. Makihara, 
Forestry and Forest Products Research In- 
stitute, Tsukuba City, Ibaraki Prefecture, 
Honshu, Japan. This specimen is entirely 
black. According to Benson’s (1943) and 
Maa’s (1956) keys, this specimen runs to 
Eriotremex smithi (Cameron), but it is eas- 
ily separated from the latter by the form of 
the precornal basin, by the black abdomen, 
and by the smoky apical third of the hind- 
wing. Therefore, I concluded that this spec- 
imen represents a new species, which is de- 
scribed and illustrated in this paper. 


Genus Eriotremex Benson 


Eriotremex Benson, 1943, p. 42; Maa, 
1949, p. 127; Maa, 1956, p. 91. Type spe- 
cies: Tremex smithi Cameron, by original 
designation. 


Distribution.—E. India, Indo-China, 
Philippines, Borneo, New Guinea, Formo- 
sa, Japan, southeastern United States. 

Generic characters.—Head without post 
genal carina; labial palps 2-segmented; an- 
tenna swollen in middle and short; forewing 
with two cubital cross veins and radial cross 
vein situated near the base of the radial cell 
(Fig. 4); precornal basin convex in the mid- 
dle and strongly and coarsely punctured. 


Symphyta, Siricidae, Eriotremex, new species, Japan 


KEY TO THE JAPANESE SPECIES OF 
ERIOTREMEX (FEMALE) 


1. Abdominal tergites entirely black; precornal 

basin rather elongate (Fig. 8) .......... 
maktharai, 0. sp. 

— First, 2nd, 4th, 5th, and 8th tergites with yellow 
bands; precornal basin nearly circular ...... 2 

. A small spot on gena yellow; 6th abdominal 
tergite with yellow band .... yamasakii Togashi 

— Gena entirely black; 6th abdominal tergite 
black sents. 4 ier: 


Nw 


formosanus (Matsumura) 


Eriotremex makiharai Togashi, 
new species 
(Figs. 1—10) 


Female.—Length including cornus 22 
mm. Body including antenna and legs en- 
tirely black. Wings yellowish tinged, apical 
half and basal ¥; of forewing and apical % 
of hindwing smoky; stigma and veins 
black; body covered by black hairs. 

Head: Rather transverse (Fig. 1); inter- 
ocellar, postocellar, and lateral furrows in- 


. distinct; OOL:POL:OCL = 0.8:1.0:2.6; dis- 


tance between antennal sockets longer than 
scape (ratio as 1.0:0.6). Antenna 18-seg- 
mented (Fig. 2); nearly as long as costa of 
forewing: scape slightly longer than third 
segment (ratio as 1.0:0.8); third segment 
nearly as long as fourth. 

Thorax: Pronotum long, midlength about 
as long as OCL (ratio as 1.0:1.0) (Fig. 3); 
cenchrus small, distance between them 2.4 


160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


eee 


Figs. 1-7. 


Sea erm 


a= ae 


Eriotremex makiharai, holotype. 1, Head, dorsal view. 2, Antenna, lateral view. 3, Pronotum, 


dorsal view. 4, Apical half of forewing. 5, Hind tibia and basitarsus, lateral view. 6, Fore inner tibial spur, lateral 


view. 7, Tarsal claw, lateral view. 


times breadth of one. Wings: first radial cell 
(1R) narrow, rectangular (Fig. 4); first cu- 
bital cell (1C) of forewing longer than sec- 
ond radial cell (2R) (ratio as 1.0:0.46) (Fig. 
4). Legs: hind tibia shorter than hind basi- 
tarsus (ratio as 1.0:1.3) (Fig. 5); fore inner 
tibial spur as in Fig. 6; tarsal claw with a 
small inner tooth and basal lobe (Fig. 7). 

Abdomen: Sheath shorter than basal 
plate (ratio as 1.0:2.3); eighth tergite nearly 
as long as three preceding tergites com- 
bined; precornal basin rather elongate (Fig. 
8), inner surface distinctly depressed (Fig. 
8), central portion distinctly convex; cornus 
as in Figs. 8 and 9; apical portion of lancet 
as in Fig. 10. 

Punctation: Head and thorax distinctly, 
closely, and reticulately punctured, inter- 
spaces between punctures nearly impunc- 
tate, shining; mesopleuron covered with 
small but distinct punctures, interspaces be- 
tween punctures nearly impunctate, shining; 
mesoscutellum very distinctly, closely, and 
reticulately sculptured; first to third tergites 
shagreened; fourth to eighth tergites cov- 
ered with small but distinct punctures, in- 


terspaces between punctures nearly im- 
punctate, shining; precornal basin distinctly, 
closely, and reticulately sculptured; all ster- 
nites nearly impunctate, shining. 

Male.—Unknown. 

Habitat.—Japan (Ishigaki Is., Okinawa 
Prefecture). 

Food plant.—Unknown. 

Holotype.—Female, 16. V. 2003, bred 
from wood; Mt. Omoto, Ishigaki Is., Oki- 
nawa Pref., H. Makihara leg. Deposited in 
the collection of the National Science Mu- 
seum (Nat. Hist.), Tokyo. 

Remarks.—This new species keys to E. 
smithi (Cameron) from eastern India in 
Maa’s (1956) key, but it is separated from 
E. smithi by the smoky apical third of the 
hind wing (mostly hyaline in E. smithi), by 
the partially hyaline forewing (entirely 
smoky in E. smithi), and by the elongate 
precornal basin (rather rounded in E. smi- 
thi). From E. purpureipennis (Westwood), 
E. insignis (EK Smith), and E. foveopygus 
Maa, all of which have a black abdomen, 
E. makiharai is separated by the partially 


VOLUME 107, NUMBER 1 


i6] 


Figs. 8-10. Eriotremex makiharai, holotype. 8, Eighth tergite, precornal basin and cornus, dorsal view. 9, 
Posterior portion of abdomen, lateral view. 10, Apical portion of lancet. 


hyaline forewing (entirely smoky in these 
three species). 


ACKNOWLEDGMENTS 


I thank Dr. David R. Smith, USDA, 
Washington, D.C., for reading through this 
manuscript and for his kind advice. I am 
indebted to Mr. H. Makihara, Forestry and 
Forest Products Research Institute, Tsukuba 
City, Ibaraki Prefecture, for his kindness in 
giving me the opportunity to examine the 
specimen. I also thank Dr. A. Shinohara, 
National Science Museum (Natural Histo- 


ry), Tokyo, for his kindness in providing 
copies of literature. 


LITERATURE CITED 


Benson, R. B. 1943. Studies in Siricidae, especially of 
Europe and southern Asia (Hymenoptera, Sym- 
phyta). Bulletin of Entomological Research 34: 
27—S0. 

Maa, T. 1949. A synopsis of Asiatic Siricoidea with 
notes on certain exotic and fossil forms (Hyme- 
noptera, Symphyta). Musée Heude, Notes 
d’Entomologie Chinoise 8: 11—189. 


. 1956. Notes on the genus Eriotremex Benson 
(Hymenoptera: Siricidae). Proceedings of the Ha- 
waiian Entomological Society 14: 91—94. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 162-175 


RUTACEAE-FEEDING AGONOPTERIX HUBNER 
(LEPIDOPTERA: ELACHISTIDAE) IN ILLINOIS 


T. L. HARRISON AND M. R. BERENBAUM 


Department of Entomology, University of [linois at Urbana-Champaign, 320 Morrill 
Hall, 505 South Goodwin Avenue, Urbana, IL 61801, U.S.A. (e-mail: tharriso@ux1. 
cso.uiuc.edu) 


Abstract.—The most recent taxonomic treatment of North American moths of the genus 
Agonopterix Hiibner (Lepidoptera: Gelechioidea: Elachistidae) recognized two Rutaceae- 
feeding species, Agonopterix nigrinotella (Busck) and Agonopterix pteleae Barnes and 
Busck. Ecological, morphological, and molecular data presented in this paper indicate that 
four species of Agonopterix feed as larvae on Rutaceae in Illinois. Agonopterix costima- 
cula Clarke (revised status) and A. pteleae feed on hoptree, Ptelea trifoliata (Linnaeus). 
Both of these species are multivoltine. Agonopterix nigrinotella and a previously unrec- 
ognized species, described here as Agonopterix paulae Harrison, n. sp., feed on prick- 
lyash, Zanthoxylum americanum Miller. Agonopterix nigrinotella is univoltine, whereas 
A. paulae is bivoltine. Larval feeding tests determined that each of the four species is 
strictly monophagous and that literature reports of A. nigrinotella on Carya sp. and Agon- 
opterix argillacea (Walsingham) on Ptelea trifoliata are apparently erroneous. Diagnoses 
and dichotomous keys are given for identifying mature larvae and adults of the four 


Rutaceae-feeding species. 


Key Words: 


Microlepidoptera, Gelechioidea, new species, voltinism, food plant speci- 


ficity, Rutaceae, Zanthoxylum, Ptelea, mitochondrial DNA, Cytochrome 
Oxidase IT, North America, Illinois 


Although the North American component 
of the genus Agonopterix Hiibner has been 
studied more extensively than have most 
other ‘“microlepidoptera’” groups on_ this 
continent (Clarke 1941, Hodges 1974, Ber- 
enbaum and Passoa 1999), and larval food 
plants are known for almost all species, 
complete life histories have been evaluated 
for only a few (Hodges 1974). Because of 
such inadequate life history information, the 
complex of Nearctic Rutaceae-feeding spe- 
cies centered around Agonopterix nigrino- 
tella (Busck) represents an element of the 
genus that has remained unresolved. 

The earliest description of a Nearctic Ru- 
taceae-feeding Agonopterix species was that 


of A. nigrinotella (Busck 1908), which was 
described as Depressaria nigrinotella. Sub- 
sequently, Barnes and Busck (1920) de- 
scribed Agonopteryx [sic] pteleae, and 
Clarke (1941) described Agonopterix cos- 
timacula. Clarke proposed that A. pteleae 
and A. costimacula feed as larvae only on 
Ptelea trifoliata (L.), and that A. nigrino- 
tella feeds only on Zanthoxylum american- 
um Miller. 

Hodges (1974) recognized A. pteleae as 
a distinct species but relegated all other Ru- 
taceae-feeding Agonopterix to A. nigrino- 
tella (with A. costimacula sunk as a junior 
synonym of A. nigrinotella). Hodges ac- 
knowledged that more than one genital type 


VOLUME 107, NUMBER 1 


was present in the moths ascribed by him 
to A. nigrinotella. His justification for re- 
taining all of these entities under one spe- 
cies name was that a single genital type 
could be found in moths representing more 
than one of the color forms that he recog- 
nized, and that males of a single genital 
type had been reared from both Z. ameri- 
canum and P. trifoliata. 

Field investigations conducted by S. Pas- 
soa in the 1980s revealed a diverse fauna 
of Rutaceae-feeding A gonopterix in Illinois, 
with larvae utilizing both of the two native 
rutaceous plants found in the state, Prelea 
trifoliata (Linnaeus) and Zanthoxylum 
americanum Miller. The apparent presence 
of all of the relevant moth and plant species 
in this same geographic area suggested that 
a study of Rutaceae-feeding Agonopterix in 
Illinois could provide a clear resolution to 
species identities within this problematic 
group; presented herein are the results of 
such a study. 


MATERIALS AND METHODS 


For rearing and for food plant specificity 
trials, larvae were placed individually in 30 
ml plastic snap-lid cups containing a small 
amount of leaf material. In food plant spec- 
ificity trials, first-stadium larvae were sub- 
jected to no-choice feeding tests on plants 
other than the known field hosts: A. pteleae 
and A. costimacula on Z. americanum, A. 
paulae and A. argillacea on P. trifoliata, 
and A. nigrinotella on both P. trifoliata and 
on two species of Carya Nuttall, the latter 
genus having been recorded as a food plant 
by Hodges (1974). After 24 h, each larva 
was placed in a new cup containing the 
field host and was reared to adult, to con- 
firm its identity and its normal state of 
health. 

Larvae to be preserved for morphologi- 
cal study were placed for 24 h in XAA so- 
lution (10% xylene, 80% absolute ethanol, 
10% glacial acetic acid, saturated with dis- 
tilled water) and then transferred to 70% 
ethanol, 30% distilled water for permanent 
storage. Drawings of larvae, as well as all 


163 


other drawings, were done through a Wild 
M5 microscope equipped with a Wild M5 
Zeichentubus drawing tube, at magnifica- 
tions of 25x to 40X; larvae were sub- 
merged in 70% ethanol in a clear glass de- 
pression slide. Names of primary setae fol- 
low those given by Stehr (1987). 

For study of genitalia, the entire abdo- 
men of each dried specimen was pried off 
with forceps. The abdomen was macerated 
in 20% potassium hydroxide in distilled 
water for eight to 24 h, and then dissected 
in 70% ethanol. Iridectomy scissors were 
used to make a longitudinal slit in the lat- 
eral membrane, beginning at the anterior 
end of the abdomen and running posterior 
to the hind margin of the 7th (females) or 
8th (males) abdominal segment. Genitalia 
were stained overnight in 63% ethanol, 
27% distilled water, and 10% mercuro- 
chrome. While in the stain, genitalia were 
positioned for viewing in ventral aspect and 
were kept in this position with glass chips 
laid over them. After staining, they were 
cleaned with a 00000 gauge sable brush in 
70% ethanol; from here, they were either 
placed directly into a vial of 70% ethanol 
for storage or were immersed briefly in 
clove oil and then mounted in Canada bal- 
sam on microscope slides. 

For genetic analysis, adult moths were 
frozen alive, and genomic DNA was ex- 
tracted following the protocol of Sperling 
et al. (1994). A 522-bp fragment of mito- 
chondrial Cytochrome Oxidase II (COI, 
homologous with mitochondrial genome 
positions 1691 through 2212 of Yponomeu- 
ta malinellus Zeller (Sperling et al. 1995), 
was amplified by polymerase chain reaction 
(PCR) (Saiki et al. 1985, 1988) using prim- 
ersu€22J-3135 (S62 AGCGEECTEIECERE 
TAATAGAACA 3’) and C2-N-3661 (5’ 
CCACAAATTTCTGAACATTGACCA 3’) 
(Simon et al. 1994). Each PCR sample was 
sequenced for its entire length in both 5’— 
3’ and 3’—5’ directions, by using the op- 
posing terminal primers; sequencing was 
done in the University of Illinois Automat- 
ed Sequencing Laboratory. One individual 


164 


of each species was sequenced, with the ex- 
ception that partial sequence was obtained 
from a second individual of A. costimacula. 

Abbreviations of Institution and Collec- 
tion Names.—USNM: National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C., USA; INHS: Illinois 
Natural History Survey, Champaign, IIli- 
nois, USA; JRW: Collection of James R. 
Wiker, Athens, Illinois, USA; SCP: Collec- 
tion of Steven C. Passoa, Columbus, Ohio 
USA. 


RESULTS AND DISCUSSION 


One of the insects involved in the present 
study is a described species, A. costimacula. 
This species is currently placed as a junior 
synonym of A. nigrinotella. 


Agonopterix costimacula Clarke, 1941, 
revised status 


The original description and type mate- 
rial are as designated by Clarke (1941). In 
this paper, we provide ecological, morpho- 
logical, and molecular evidence to support 
the conclusion that A. costimacula is a valid 
species. 

One of the insects in this study proved to 
be a previously undescribed species. This 
species did not correspond to type material 
of the three Nearctic Rutaceae-feeding 
Agonopterix species mentioned above, nor 
with Agonopterix argillacea (Walsingham) 
or Apachea barberella (Busck), two addi- 
tional Nearctic Depressariinae species that 
have been recorded from Rutaceae (Hodges 
1974). 

Likewise, we determined that the novel 
species does not represent an introduced 
population of one of the three European 
species of Rutaceae-feeding Agonopterix, 
A. rutana (Fabricius), A. furvella (Treitsch- 
ke), and A. pupillana (Wocke), nor of either 
of two additional Rutaceae-feeding Depres- 
sariinae of Europe, Depressaria ruticola 
(Christoph) and Horridopalpus dictamnella 
(Treitschke). Larvae, adult moths, and gen- 
ital morphology of the European species 
were illustrated variously by Stainton 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(1861, 1870), Rebel (1899), and Hanne- 
mann (1953). The following is a description 
of the novel insect. 


Agonopterix paulae Harrison, 
new species 
(Figs. 1A, 2A, 3A, 4A, 5A) 


Adult (Fig. 1A).—Forewing length, 
10.0-10.5 mm. Antenna: Pedicel dark 
brown, ringed with ochreous in apical one- 
fourth; flagellum shining gray brown, 
slightly paler ventrally than dorsally. Labial 
palpus: Basal segment light brown with 
blackish-brown lateral patch; second seg- 
ment 6X as long as first, clear light brown 
medially, light brown flecked with dark 
brown laterally, numerous dark-brown 
scales ventrally where scales assume a fan- 
like arrangement, giving segment a dilated 
appearance; third segment smoothly scaled, 
acuminate, nearly as long as second, ochre- 
ous brown with dark-brown bands at one- 
sixth and two-thirds length, and _ finely 
tipped with dark brown at extreme apex. 
Maxillary palpus: Minute, folded over base 
of haustellum, whitish brown. Haustellum 
clothed in whitish-brown scales. Face: 
Smoothly scaled, shining whitish brown, 
with a small number of blackish-brown 
scales near compound eye. Head: Roughly 
scaled, ochreous with medial brown line; 
occipital scale band pale ochreous medially, 
brown laterally. Thorax: Terga and tegula 
ochreous overlaid with brown; a pair of 
whitish-ochreous tufts on metathoracic ter- 
gum. Proleg: Coxa whitish ochreous, dust- 
ed with brown except at apex; lateral sur- 
face pearl white with a few brown scales; 
femur whitish ochreous, heavily dusted 
with brown medially, with a concentrated 
brown blotch laterally; tibia slightly shorter 
than femur, with somewhat dilated appear- 
ance due to loose scaling, clear pale ochre- 
ous laterally, pale ochreous dusted heavily 
with brown medially; tarsus equal in length 
to tibia, with basal tarsomere 3X as long as 
each of remaining four, brown, apices of 
basal two tarsomeres and entire apical tar- 
somere ochreous. Mesoleg: Coxa_ pale 


VOLUME 107, NUMBER 1 


Fig. 1. 
A. nigrinotella. C, A. pteleae. D, A. costimacula. Scale = 5.0 mm. 


ochreous with central brown patch, scales 
at apex overlying region of trochanter; fe- 
mur pale ochreous dusted with brown; tibia 
subequal to femur in length, scales arranged 
to give a dilated appearance, pale ochreous 
with a brown lateral blotch near base and a 
brown band at three-fourths length; a pair 
of apical spurs, both brown, medial one 3 X 
as long as lateral; tarsus equal to tibia in 
length, basal tarsomere as long as remain- 
ing four combined, brown, apices of all tar- 
someres and entire apical tarsomere ochre- 
ous. Metaleg: Coxa and femur as for me- 
soleg; tibia slightly longer than femur, with 
loose scaling giving dilated appearance, al- 
most entirely pale ochreous, with a few 
scattered brown scales; two pairs of spurs, 
at approximately one-half tibia length and 
at apex, shining ochreous, medial one 3X 
as long as lateral in both pairs; tarsus almost 
as long as tibia, basal tarsomere slightly 
longer than remaining four combined, 


ochreous medially, brown with apex of 


Right mesothoracic wings of Nearctic Rutaceae-feeding Agonopterix; dorsal aspect. A, A. paulae. B, 


each tarsomere ochreous laterally. Mesow- 
ing: Length, 9.8 mm. Upper surface ground 
color medium ochreous brown; a small, 
blackish-brown patch at base of costa, base 
of wing otherwise ochreous to one-tenth 
length, then met by a narrow band of black- 
ish brown, this band broken above fold by 
an outward projection of basal ochreous 
color; only a slight diffusion of dark-brown 
scales extends apically from blackish- 
brown band; a small, blackish-brown spot 
on fold, a second, similar spot almost di- 
rectly anterior to it, and a third such spot 
anterior and basal to second; six or seven 
blackish-brown strigils on costa; each of 
two most prominent of these, at four-fifths 
wing length, with a similar blackish-brown 


mark immediately below it; a grayish- 
brown blotch sits over end of cell, this 


blotch not reaching costa; at its posteroap- 
ical margin containing a small but distinct 
whitish spot; veins R,, Rs, and M, delin- 
eated apically by blackish-brown streaks; 


166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


scaling becoming slightly paler than ground 
in apical one-eighth of wing; outer edge of 
wing membrane marked by four or five 
small, blackish-brown spots; fringe conco- 
lorous with ground, with no discernible 
lines; underside shining grayish ochreous, 
anterior one-sixth ochreous with occasional 
dark strigils, extreme apex ochreous with 
fine points of brown. Metawing: Slightly 
shorter than mesowing; upper surface shin- 
ing pale ochreous, outer margin of wing 
membrane finely scaled with brown; fringe 
pale ochreous basally, darker ochreous api- 
cally, with faint, incomplete lines near outer 
margin of wing membrane and near fringe 
apex; underside pale ochreous, mottled ir- 
regularly with brown in anterior one-third 
and around margin of wing membrane. Ab- 
domen: Upper surface shining pale ochre- 
ous dusted with brown; underside ochreous, 
with large, lateral black blotches on seg- 
ments III through VII forming nearly a con- 
tinuous line; small, medial, paired black 
spots on segments IV through VII; a pair 
of lateral black dashes on terminal seg- 
ments. 

Larva (Fig. 2A).—Neonate yellow with 
dark-brown head capsule; larvae of sub- 
sequent stadia pale green with roughly con- 
colorous head capsule; area of stemmata 
black in all stadia; in second and subse- 
quent stadia, a black blotch appears on lat- 
eral surface of prothorax, and in third and 
subsequent stadia a black lateral stripe ap- 
pears on anal shield. Length of mature lar- 
va, 18.5 mm (n = 5, range 18.1—19.2 mm). 

Biology.—The larval food plant is Z. 
americanum. Eggs are laid either singly or 
in small masses. In the latter case, larvae 
initially feed communally between two leaf- 
lets that they have silked together, and their 
feeding at this stage creates a distinctive 
pattern of damage in which the affected leaf- 
lets assume a singed appearance. While still 
in the first stadium, the larvae leave the 
communal leaflets and establish themselves 
individually; from this point on, they live 
in the rolled margins of leaflets. There are 
evidently two generations per year, with 


mature larvae in central [linois occurring 
in late June/early July, and again in late 
July. 

Geographic range.—Agonopterix paulae 
has been recorded from Illinois counties 
Piatt, Jo Daviess, Vermilion, Coles, Men- 
ard, and Marshall (the known populations 
of Z. americanum and A. paulae on the for- 
mer M. O. Glenn farm, historically record- 
ed as ““Putnam Co., Ill.”’, actually occur in 
the northern part of Marshall County). The 
insect may occur over the entire eastern 
USA, wherever the food plant is found, as 
was suggested for A. nigrinotella by Clarke 
(1941). The status of this species on native 
Rutaceae species that occur in areas other 
than the Midwest is not presently known, 
as is likewise true for the other three Ne- 
arctic Rutaceae-feeding Agonopterix spe- 
cies. 

Diagnosis (for quantitative measurements 
of some of the character states listed below, 
consult the keys to species)—The mature 
larva of A. paulae is unique among the Ru- 
taceae-feeding species in having black 
markings on the thorax and anal shield. 
Agonopterix nigrinotella lacks these mark- 
ings but has prominent pinacula at the bases 
of the D setae on most of the body seg- 
ments, whereas A. pteleae and A. costima- 
cula are pale green with no black markings 
or pinacula on the body (a blackish marking 
on the genal area of the head may in all 
four species display individual variation 
from absent to prominent). The adult of A. 
paulae can be distinguished from A. nigri- 
notella by the much less uniform coloration 
of the mesowing, and from A. pteleae by 
the absence of upturned scales on the dorsal 
surfaces of the mesowings and thorax. It 
can be separated from A. costimacula, 
which it most closely resembles, by the rel- 
atively dark, ochreous-brown ground color, 
presence of a diffuse, blackish-brown area 
surrounding the pale spot on the cell in the 
mesowing, and presence of several black- 
ish, longitudinal vein-streaks in the outer 
part of the forewing of A. paulae (A. cos- 
timacula with relatively pale, clay-brown 


VOLUME 107, NUMBER 1 


D2 


XDI 


xD2 \zeel 
\ 


SDI 


Fig. 2. 


167 


Mature larvae of Nearctic Rutaceae-feeding Agonopterix; left lateral aspect. A, A. paulae. B, A. 


nigrinotella. C, A. pteleae. D, A. costimacula. Segments illustrated are, from left to right, head, first and second 
thoracic, first abdominal, and eighth through tenth abdominal. Scale = 5.0 mm. 


ground color, with no dark area other than 
a narrow ring surrounding the spot on the 
cell, and no vein-streaks). In the male gen- 
italia of A. paulae (Fig. 3A), the clasper is 
much straighter (less curved), and the valve 
is relatively longer and narrower, and more 
sharply curved dorsad at about half its 
length, than in the other three species. The 


juxta is quadrate and similar to that of A. 
nigrinotella and A. costimacula. In the fe- 
male genitalia (Figs. 4A, 5A), the anterior 
margin of A8 is broadly squared (narrowly 
squared in A. nigrinotella, Figs. 4B, 5B, 
and broadly rounded in A. pfeleae, Figs. 
4C, 5C). Overall, A. paulae is similar to A. 
costimacula (Figs. 4D, 5D), except that in 


168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. 


A. paulae the “‘plate’ in the center of the 
anterior margin is thickened only at its lat- 
eral margins, whereas in A. costimacula, the 
entire anterior and lateral margins of the 
plate are thickened. Also, in A. paulae, the 
ostium bursae extends posterad almost to 
the posterior margin of A8, whereas in A. 
costimacula the posterior extremity of the 
ostium is relatively far removed from the 
posterior margin of the segment. Finally, 
the eighth abdominal sternum in A. paulae 
bears setae only in the posterolateral areas, 
whereas in A. costimacula, setae are distrib- 
uted over much of the sternum. 
Etymology.—This species is named in 
honor of Paula D. Harrison, wife of the se- 
nior author, in recognition of the immense 
contribution that she has made to his life 
and career, including valuable field assis- 
tance on some of the earliest excursions 
during which this species was collected. 
Type series.—Holotype male: Collected 


Genitalia of male adults of Nearctic Rutaceae-feeding Agonopterix; posteroventral aspect, valvae 
reflected laterally, aedeagus removed. A, A. paulae. B, A. nigrinotella. C, A. pteleae. D, A. costimacula. Scale 
= 0.5 mm. 


as larva on Zanthoxylum americanum, 
USA: Illinois, Piatt County, University of 
Illinois-Robert Allerton Park, Lost Garden 
Trail, 28-VII-1994; iss. 28-VIII-1994 
(USNM). Allotype female: same locality 
and dates as for holotype (USNM). Para- 
types: ten specimens, with same collection 
data as for holotype, emergence dates from 
26-VIII-1994 to 4-IX-1994 (USNM, INHS, 
JRW, SCP); one specimen, collected as lar- 
va on Zanthoxylum americanum, USA: Il- 
linois, Menard County, 5 mi. NW of Green- 
view, R. & C. Baugher property, T19N, 
R7W, Sec. 12, 13-VII-2001, T. Harrison/J. 
Wiker, iss. 5-VIII-2001 (JRW); five speci- 
mens, same data as for preceding except 
collected by J. Wiker, 29-VII-2002, iss. 26- 
VIII-2002 (JRW); one specimen, collected 
as larva on Zanthoxylum americanum, 
USA: Illinois, Coles County, Charleston, 
Lake View Park, T12N, ROE, NW 4% Sec. 


VOLUME 107, NUMBER 1 


24, 13-VU-2000, T. Harrison, iss. 3-VIII- 
2000 (INHS). 


BIOLOGIES OF RUTACEAE-FEEDING 
AGONOPTERIX SPECIES IN ILLINOIS 


Larval food plant specificities —Hodges 
(1974) reported that male genitalia of a sin- 
gle type were seen in specimens of “A. ni- 
grinotella” reared from both Z. american- 
um and P. trifoliata, implying that at least 
one of the entities identified as A. nigrino- 
tella is capable of utilizing both plants. In 
the present study, field-collected larvae of 
all four Rutaceae-feeding Agonopterix spe- 
cies displayed strict monophagy; further- 
more, in no-choice food plant-switching 
tests done with neonates of three of the spe- 
cies (and with fed first-stadium larvae of 
the fourth species, A. nigrinotella), each 
species accepted only the food plant that it 
was observed to utilize in the field. 

We conclude that each of the four Ne- 
arctic Rutaceae-feeding Agonopterix spe- 
cies is strictly monophagous, with A. pau- 
lae and A. nigrinotella feeding as larvae 
only on Z. americanum, and A. pteleae and 
A. costimacula utilizing only P. trifoliata. 
It is noteworthy that this is the same con- 
clusion drawn by Clarke (1941), except that 
he apparently did not examine any speci- 
mens of A. paulae, as he did not recognize 
it aS a separate species. 

Hodges (1974) reported that A. nigrino- 
tella has been reared from Carya. In the 
USNM collection are specimens of A. ni- 
grinotella (identity confirmed by our own 
examination) that are labeled as having 
been reared in Ohio, by the late Annette 
Braun, from an unidentified species of Car- 
ya. Excepting this record, Carya has never 
been listed as food plant for A. nigrinotella 
nor for any other Nearctic Agonopterix spe- 
cies. During the present study, we attempt- 
ed to rear A. nigrinotella on leaves of two 
common Carya species of the eastern USA, 
C. ovata (Miller) Britton and C. glabra 
(Miller) Britton. Larvae refused both spe- 
cies, and they fed and developed normally 
when transferred to Z. americanum. This 


169 


Lateral 
Margin = 
A Anterior 


Pa Fee /-—— Margin 


a 


tae 


Fig. 4. Medial plates of anterior margins of eighth 
abdominal sterna in female adults of Nearctic Ruta- 
ceae-feeding Agonopterix (plate absent in A. pteleae): 
ventral aspect. A, A. paulae. B, A. nigrinotella. C, A. 
pteleae. D, A. costimacula. Scale = 0.5 mm. 


test is not definitive, because only two Car- 
ya species were provided, and because fed 
first-stadium larvae (the earliest stage that 
we could obtain), rather than neonates, 
were used, and they might have been irre- 
versibly conditioned to feed only on Z. 
americanum (Zalucki et al. 2002). Given, 
however, the uniqueness of the Carya re- 
cord, the strict monophagy seen elsewhere 
in Nearctic Rutaceae-feeding Agonopterix 
(in laboratory tests involving unfed first- 
stadium larvae of three of the four species, 
and in all field observations), and the dis- 
junct taxonomic relationship of Juglanda- 
ceae and Rutaceae, we conclude that it is 
doubtful that A. nigrinotella utilizes Carya 
as a food plant. The record of A. nigrino- 
tella on Carya is tentatively assessed as a 
misidentification of the food plant, awaiting 
a definitive test in which neonates of A. ni- 
grinotella are offered all Carya species that 
occur in Ohio. 

Clarke (1941) named only “Salix spp.” 
as food plant for A. argillacea, but Hodges 
(1974) also listed P. trifoliata. In feeding 
trials conducted in the present study, unfed 
neonates of A. argillacea refused P. trifol- 
iata; they fed and developed normally when 


170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Che hat 


ik eee earn eT 
Fig. 5. 


Eighth abdominal sterna of female adults of Nearctic Rutaceae-feeding Agonopterix;, ventral aspect. 


A, A. paulae. B, A. nigrinotella. C, A. pteleae. D, A. costimacula. Scale = 0.5 mm. 


transferred to the field host, an undeter- 
mined species of Salix Linnaeus. We con- 
clude that the record of A. argillacea on 
Ptelea is an error. 

Phenology and voltinism.—L vovskiy 
(1975), commenting on the European Oec- 
ophoridae (traditional sense, including De- 
pressariinae, Amphisbatinae, and Oeco- 
phorinae), noted that almost all species are 
univoltine. The four exceptions include 
Pseudatemelia josephinae Toll, which re- 
quires two years per generation, and three 
species that go through two or more gen- 
erations per year: Endrosis sarcitella (Ste- 
phens), a synanthropic stored grain pest; 
and one species each of Depressaria 
Haworth and Amphisbatis Zeller. L vovskiy 
also outlined four different patterns of uni- 
voltinism seen in European Oecophoridae 
(Fig. 6A). Type 1, with wintering as adults, 
is seen in Agonopterix, Depressaria, and 
Martyrhilda Clarke. Type 2, in which partly 
grown larvae are the wintering stage, is 


characteristic of Oecophorinae, Pleurotinae, 
and Deuterogoniinae. Type 3, in which pu- 
pae over-winter, is seen in Cheimophila 
Hiibner, Semioscopis Hiibner, Epigraphia 
Stephens, and most Diurnea Haworth. Only 
Diurnea phryganella (Hitibner) exhibits 
Type 4, in which winter is passed in the egg 
stage. Hodges (1974) reported that North 
American species of Agonopterix for which 
life histories are known are generally uni- 
voltine and follow the Type | program, but 
that a few species lay eggs in late summer, 
and in these the egg is the over-wintering 
stage (apparently similar to Type 4). 

Our findings with Rutaceae-feeding 
Agonopterix in Illinois indicate that, al- 
though all four species over-winter as 
adults, Type 1 univoltinism is seen in only 
one of the four species, A. nigrinotella (Fig. 
6B). In this species, mature larvae are pres- 
ent in central Illinois only from late May 
into very early June. 

Although the time required to complete 


VOLUME 107, NUMBER 1 


a single generation (Fig. 6C) is similar in 
all four Rutaceae-feeding species, the sea- 
sonal cycles in the three species other than 
A. nigrinotella differ from Type I univoltin- 
ism. In A. paulae, young larvae were col- 
lected at Allerton Park, Piatt County on 3 
June, 1999, at the time when A. nigrinotella 
was completing larval development. These 
A. paulae larvae matured, pupated, and 
emerged as adults in early July 1999. These 
adults did not mate in the laboratory; how- 
ever, mature larvae appeared once again on 
Z. americanum at Allerton Park during late 
July 1999. No Agonopterix larvae have 
been found on Z. americanum later in the 
year than the late-July generation of A. pau- 
lae. Given the timing of these observations, 
it appears likely that A. paulae is bivoltine. 

The two Ptelea-feeding species display a 
pattern that is even less readily interpreted 
as strict univoltinism than that seen in A. 
paulae. In central Illinois, mature larvae of 
A. costimacula appear in a fairly synchro- 
nized initial cohort in late May. Shortly af- 
terward, larvae of all different stages can 
be found, and this situation remains contin- 
uously throughout the remainder of spring 
and summer, ending only with leaf senes- 
cence (which usually occurs in early Sep- 
tember). 

Although this pattern strongly suggests 
multivoltinism for A. costimacula, we could 
not summarily rule out the possibility that 
the different temporal cohorts of larvae that 
appear throughout the late spring and sum- 
mer are the result of remarkably staggered 
Oviposition by a single generation of over- 
wintered adults. This scenario, however, 
was cast into doubt (at least as the sole ex- 
planation for the phenology of this species) 
by our observation in the laboratory of mat- 
ing in a pair of adult A. costimacula that 
we had reared from early-season larvae, be- 
havior highly suggestive of a second gen- 
eration. 

The situation in A. pteleae is probably 
the same as in A. costimacula, although the 
evidence of multivoltinism in this species is 
not as strong. A major difficulty in studying 


171 


the biology of A. pteleae is that in central 
Illinois it occurs in markedly lower num- 
bers than does A. costimacula, to the degree 
that field collections of Agonopterix larvae 
from P. trifoliata usually yield only the lat- 
ter species. There is, however, circumstan- 
tial evidence that A. pteleae is multivoltine. 
First, a single A. pteleae adult was reared 
from larvae collected in late May at Forest 
Glen Nature Preserve, Vermilion County, 
Illinois; this corresponds to the initial co- 
hort of A. costimacula, several individuals 
of which were reared from the same collec- 
tion that produced the individual of A. pfe- 
leae. Second, adult female A. pteleae were 
collected at UV light in mid-June at Sand 
Ridge State Forest, Mason County, Illinois. 
The scale vestiture of these insects at the 
time of collection was nearly pristine (the 
vestiture of Overwintered Agonopterix 
adults is usually almost completely absent 
from months of wear). This strongly sug- 
gests that the moths at Sand Ridge State 
Forest were first-generation offspring of 
adults that had over-wintered from the pre- 
vious year, as does the timing (they were 
collected at the same time that moths would 
have appeared from the known late-May 
larval cohort at Forest Glen). Furthermore, 
when the Sand Ridge moths were caged in 
the laboratory, they laid fertile eggs, which 
the univoltine offspring of wintered adults 
would not have done. Third, A. pteleae 
adults emerged in late August from two pu- 
pae collected from sleeved P. trifoliata at 
Charleston, Coles County, Illinois. It there- 
fore appears that, as in A. costimacula, A. 
pteleae is multivoltine with several gener- 
ations produced from late spring through 
summer, although this supposition should 
be confirmed by observation of A. pteleae 
at a single site. 


GENETIC DIFFERENCES IN THE FOUR 
NEARCTIC RUTACEAE-FEEDING SPECIES OF 
AGONOPTERIX 


Sequences of the mitochondrial CO// 
fragment from Agonopterix costimacula, A. 
pulvipennella 


nigrinotella, Agonopterix 


V2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
J | M A M J J A 5 O N D 
1) adult miefll pa 
2) larva completes development p a/m/e/l partly grown larva 
3) pupa a/m/e/I p 
4) egg | p a/m/e 
A 
J ie M A M J J A S O N D 
migrinotella adult Tue ff) o 
paulae adult m/e/l p a/m/e/l p a 
pleleae adult m/e/l p a/nve/l p a/m/e/l p a 
costimacula adult n/e/ om ta/m/e/ I spmra/mi/e/ leaned 
B 
JUNE JULY 
20 21 22 23 24 25 26 27 28 29 30123456789 1011 12 13 1415 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 
e ye aes L4 ES ec a 
C 
Fig. 6. A, Four different seasonal cycles in univoltine ““Oecophoridae”’ of Europe, as outlined by L’ vovskiy 


(1975). B, Seasonal cycles of Rutaceae-feeding Agonopterix from Illinois. C, Life cycle of Agonopterix pteleae 
reared from egg to adult (four individuals, intervals averaged); similar intervals were noted in the other three 
species; e, egg; 1, larva, L1—L5, instars 1 through 5; p, pupa; a, adult; m, mating. 


(Clemens), A. pteleae, Depressaria pastin- 
acella (Duponchel), and “‘Agonopterix n. 
sp. #1” (= A. paulae) have been deposited 
in Genbank under accession num- 
bers AY527038, AY527039, AY527040, 


AY527041, AY527042, and AY527043, re- 
spectively. PCR product from one species 
(A. pteleae) yielded only 418 bp of se- 
quence readable in both directions, and 
comparisons of all four Rutaceae-feeding 


VOLUME 107, NUMBER 1 


Table 1. Pairwise comparisons of a 418-bp region 
of mitochondrial gene Cytochrome Oxidase II in Ne- 
arctic Rutaceae-feeding Agonopterix. For each species 
pair, numbers indicate total number of bases differing, 
followed by percent sequence difference (in parenthe- 
ses). 


nigrinotella costimacula pteleae 
paulae 23 (5.50) 15 G:59) 28 (6.70) 
pteleae 26 (6.22) 22 (5.26) 
costimacula 15(G259)) 


species are therefore based on this subset 
(positions 1691 through 2108) of the 522- 
bp total. Numbers of base differences and 
percent sequence difference for each spe- 
cies pair are presented in Table 1. The 
3.59—6.70% differences seen here are much 
larger than the 0.00—0.96% differences seen 
in this same region of CO/I in three species 
of Yponomeuta Latreille (Sperling et al. 
1995). Our molecular results therefore align 
with the morphological and biological evi- 
dence in supporting the conclusion that Ne- 
arctic Rutaceae-feeding Agonopterix com- 
prise four valid species. 

Generally, one individual of each species 
was sequenced; however, a partial sequence 
obtained from a second individual (repre- 
senting a second population) of A. costi- 
macula shared 100% identity with the cor- 
responding area of the complete, 522-bp se- 
quence from that species. Because COI// ap- 
pears to display considerable polymorphism 
among species of Agonopterix (but appar- 
ently a high degree of intraspecific conser- 
vation, according to our limited observa- 
tion), sequence data from this gene might 
be particularly useful in a phylogenetic 
analysis of the genus. 


CONCLUSIONS 


Four Nearctic species of Agonopterix uti- 
lize Rutaceae as larval food plants in IIli- 
nois. These species are clearly separable by 
diagnostic differences in ecological and 
morphological characters, and they display 
substantial interspecific divergence in se- 
quence of the mitochondrial gene, Cyro- 
chrome Oxidase II. Each of the four Nearc- 


173 


tic Rutaceae-feeding species of A gonopterix 
in Illinois feeds on only one of the two lo- 
cally occurring Rutaceae species, as seen in 
feeding tests with neonates and first instars 
(feeding tests conducted during this study 
also contradict previous records of A. ni- 
grinotella on Carya species, and of A. ar- 
gillacea on P. trifoliata). Only one of the 
Rutaceae-feeding species, A. nigrinotella, 
displays Type 1 univoltinism (Lvovskiy 
1975) as seen in most Agonopterix, whereas 
A. paulae is bivoltine, and both A. pteleae 
and A. costimacula are multivoltine. 

That substantially greater diversity than 
was previously recognized has been found 
to exist in associations between A gonopte- 
rix and Rutaceae in central Illinois suggests 
that much remains to be learned about even 
well-studied microlepidoptera. The depres- 
sariine Elachistidae may well serve as a 
useful group for study of such evolutionary 
phenomena as host-associated speciation 
(Howard and Berlocher 2001). 


KEY TO MATURE LARVAE OF NEARCTIC 
RUTACEAE-FEEDING AGONOPTERIX 


1. Body either with dark lateral markings on seg- 
ments Tl and A1O (but with no dark pinacula) 
(Fig. 2A), or with dark pinacula at base of D 
setae on most segments (but with no dark 
markings elsewhere on thorax or abdomen) 
(Fig. 2B); food plant, Zanthoxylum american- 
TATTUY 7) oye ON di eee ese a een eR oe oa 

— Body uniformly green, with no dark pinacula 

or markings (Figs. 2C, D); food plant, Prelea 
trifoliata 

. Body with dark lateral markings on segments 

Tl and A1O (Fig. 2A) but no dark pinacula; 
bivoltine, mature larva (central Illinois) occur- 


bo 


ee) 


i) 


ring in late June, again in late July... 
— Body with dark pinacula at base of D setae on 
most segments (Fig. 2B), but otherwise with 
no dark markings on thorax or abdomen; uni- 


A. paulae 


voltine, mature larva (central Illinois) occur- 
ring in late May/early June . A. nigrinotella 


3. Spiracular peritremes brown ....... A. pteleae 


— Spiracular peritremes black ..... J A. costimacula 
KEY TO ADULTS OF NEARCTIC RUTACEAE- 
FEEDING AGONOPTERIX 
(EXTERNAL CHARACTERS) 


1. Dorsal surfaces of thorax and mesowing cov- 


ered with upturned scales, giving these struc- 


174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


tures a “shaggy” appearance (Fig. 1C); a large, 

blackish blotch slightly beyond mid-length on 

anterior part of mesowing A. pteleae 
— Dorsal surfaces of thorax and mesowing 

smoothly scaled (Figs. 1A, B,D) ......... 2 
2. Mesowing relatively uniform brown, without 


prominent blackish spots along costa, and 
without a distinct blackish ring around the 
whitish discal spot (Fig. 1B) .... A. nigrinotella 
— Mesowing with prominent blackish spots along 
costa, and with a distinct blackish ring around 
the whitish discal spot (Figs. 1A, D) 
3. Mesowing ochreous brown, with a diffuse dark- 
gray blotch anteriorly, beyond mid length, and 
with blackish streaks in apical third, these co- 
inciding with wing veins (Fig. 1A) A. paulae 
— Mesowing grayish brown, without a dark-gray 
blotch or vein streaks (Fig. 1D) .. A. costimacula 


KEY TO ADULTS OF NEARCTIC RUTACEAE- 
FEEDING AGONOPTERIX 
(GENITAL MORPHOLOGY ) 


Female 
2. Clasper extending to or slightly exceeding dor- 
sal margin of valve; ventrobasal fold of valve 
wide, exceeding 0.4 width of valve at base; 
gnathos subglobose, length-to-width ratio 1.5 
Or less (ig, BOQ) 220660666 000000c A. pteleae 
— Clasper not extending to dorsal margin of 
valve; ventrobasal fold of valve narrow, not ex- 
ceeding 0.35 width of valve at base; gnathos 
elongate, length-to-width ratio 2.5 or greater 
GEN si SVAN IBY, ID)mg od Gus eno 6 6.4 ols oe Go celeb DD, 
3. Valve relatively long and narrow, length at 
least 2.4 width at base; clasper straight or 
meeidh7 G© (Ene, BWA) oe cose secccco. A. paulae 
— Valve relatively short and wide, length 2.1 
width at base, or less; clasper strongly curved 
(ENS, G15}5 1D) Bua g obe iano hie iuonouaseestg o-0 lo © 3 
4. Valve narrow (length greater than 2.0 width 
at base), distal half of valve tapered to a nar- 
row, acuminate apex (Fig. 3B) ... A. nigrinotella 
— Valve broad (length approximately 1.9 width 
at base), distal half not tapered to a narrowly 
acuminate apex (Fig. 3D) ...... A. costimacula 
5. Eighth abdominal sternum (hereafter referred 
to as A8) with anterior margin forming an even 
curve, without a medial plate that is differen- 
tiated from the rest of the margin (Figs. 4C, 
5S) nateee ae ee aed: aide crane eee Sie A. pteleae 
— A8 with anterior margin bearing a medial plate 
that is differentiated from the rest of the margin 
(igs! LAS BA Ds SAL B.D) is eee cee 2 
6. Lateral margins of medial plate of A8 strongly 
and evenly curved; medial plate relatively nar- 
row, occupying approximately 45 per cent of 
total width of A8 at anterior margin; anterior 


margin of medial plate relatively wide, occu- 

pying about 65 per cent of total width of plate 

(Bigst AB YoIB)) Psyereorets cle ree A. nigrinotella 
— Lateral margins of medial plate of A8 straight 

or nearly so, not strongly curved; medial plate 

relatively wide, occupying approximately 55 

per cent of total width of A8 at anterior margin; 

anterior margin of medial plate relatively nar- 

row, occupying about 45 per cent of total width 

of plate (Figs. 4A, D, 5A, D) 
7. Lateral and anterior margins of medial plate 

uniformly thickened; ostium relatively small, 

extending posterad, midventrally, to less than 

70 per cent of the distance from anterior to 

posterior margins of A8; A8 with at least 20 

setae, which are not restricted to posterolateral 

area of sternum (Figs. 4D, 5D) .. A. costimacula 
— Lateral and anterior margins of medial plate 

not thickened, except in a small area of each 

lateral margin; ostium relatively large, extend- 

ing posterad, midventrally, to more than 85 per 

cent of the distance from anterior to posterior 

margins of A8; A8 with fewer than ten setae, 

all restricted to the posterolateral areas of the 


sternum (Figs. 4A, 5A) ........... A. paulae 


ACKNOWLEDGMENTS 


We thank all of the following people for 
aiding us in completing this project. Ronald 
Hodges and Steven Passoa_ unstintingly 
shared their extensive knowledge of De- 
pressariinae. James Wiker collected Agon- 
opterix larvae and adults and directed us to 
a number of previously unknown sites in 
which these insects occur. George Balogh, 
Duane McKenna, and John Tooker allotted 
time and effort during some of their field 
excursions to collect Agonopterix larvae for 
this study. Ellen Green, Mark Carroll, and 
Claire Rutledge assisted on collecting trips. 
William McClain, the family of Murray O. 
Glenn, and the staffs of Fox Ridge State 
Park, Forest Glen County Nature Preserve, 
Robert Allerton Park, and Sand Ridge State 
Forest granted permission to collect Agon- 
opterix on lands under their care. We thank 
Ronald Hodges and two anonymous re- 
viewers for helpful comments on the man- 
uscript. T. Harrison thanks his graduate 
committee members, James Sternburg and 
Arthur Zangerl; the latter also provided 
technical assistance in photography and in 
various other aspects of the study. T. Har- 


VOLUME 107, NUMBER 1 


rison acknowledges a special debt of grat- 
itude to his wife, Paula, for her continuous 
support of his academic endeavors. This 
study was funded in part by a grant from 
the H. H. Ross Award Committee, Illinois 
Natural History Survey, and was submitted 
as part of the requirements for the degree 
of Master of Science in Entomology, Uni- 
versity of Illinois at Urbana-Champaign. 


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PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 176-189 


DIPTERA DIVERSITY IN A HOMOGENEOUS HABITAT: BRACHYCERA 
ASSOCIATED WITH SEDGE MEADOWS (CYPERACEAE: CAREX) IN 
QUEBEC, CANADA 


FREDERIC BEAULIEU AND TERRY A. WHEELER 


Department of Natural Resource Sciences, McGill University, Macdonald Campus, Ste- 
Anne-de-Bellevue, Quebec, H9X 3V9 Canada (e-mail: wheeler@nrs.mcgill.ca) 


Abstract.—Species richness and community composition of Brachycera (Diptera) were 
compared in three sedge meadows dominated by two species of Carex (Cyperaceae) in 
the Lac St. Francois National Wildlife Area in southern Quebec, Canada. More than 
10,000 specimens, representing 34 families and more than 338 species, were collected 
weekly from May to October 1999 by sweeping and yellow pan traps. Species richness 
was highest in Dolichopodidae, Chloropidae, Sciomyzidae, Syrphidae, Sphaeroceridae, 
and Ephydridae. The same families, along with Phoridae, Chamaemyiidae, and Muscidae, 
were also the most abundant. Overall species composition differed between sites and was 
apparently influenced by the type of vegetation and substrate. Correspondence analysis 
showed that habitat associations of the 39 most dominant species differed between sites. 
The assemblages of Brachycera in the two Carex lacustris meadows were similar and 
richer in saprophagous species than in the Carex aquatilis meadow. This is partly due to 


greater exposure of mud substrate in the C. lacustris meadows. 


Key Words: 
ogy 


Emergent plant communities in wetlands 
are areas of high biological productivity 
and conservation significance (Scott 1995, 
Keiper et al. 2002). The sedge genus Carex 
L. (Cyperaceae) is widespread and diverse 
in such habitats and is especially dominant 
in littoral areas, peatlands and wet meadows 
of temperate regions (Kukkonen and Toi- 
vonen 1988). 

The aquatic Diptera fauna of temperate 
wetlands is also species-rich and abundant, 
mostly due to a few nematoceran families, 
especially Chironomidae (Wrubleski 1987, 
Batzer and Wissinger 1996). Fewer studies 
have examined semi-aquatic or terrestrial 
Diptera associated with emergent plants or 
the substrate, although some (e.g., Davis 
and Gray 1966, Cameron 1972, Pollet 


Diptera, Brachycera, Nearctic, wetlands, Carex, biodiversity, habitat, ecol- 


1992, Blades and Marshall 1994, Keiper et 
al. 2002, Foote 2004) have found high Dip- 
tera diversity in marshes and peatlands. 

Although their ecological roles are not 
well-studied, it is increasingly apparent that 
Diptera are a critical component in food 
webs and nutrient cycling and are an under- 
appreciated study group in conservation bi- 
ology (Haslett 2001, Keiper et al. 2002). 
Unfortunately, the use of Diptera as model 
organisms in community ecology and con- 
servation biology is hampered by a lack of 
baseline knowledge on their diversity and 
ecology in many habitats, including fresh- 
water wetlands. 

Despite the dominance of Carex and 
Diptera in wetlands, there has been rela- 
tively little study of their association. Most 


VOLUME 107, NUMBER 1 


ecological studies have focused on selected 
families such as Chloropidae (Todd and 
Foote 1987a, Rogers et al. 1991), Ephydri- 
dae (Todd and Foote 1987b) or Scatho- 
phagidae (Wallace and Neff 1971) or were 
in sites where Carex was not dominant 
(e.g., Pollet 1992, 2001). However, Foote 
(2004) collected 93 species of acalyptrate 
Diptera associated with Carex stands in 
Ohio and provided notes on abundance, 
phenology and trophic guilds of selected 
species. Beaulieu and Wheeler (2001) iden- 
tified more than 338 species of Brachycera 
in wetland Carex meadows in Quebec, Can- 
ada, despite the apparent homogeneity of 
vegetation and physical structure in that 
habitat. In this paper, we compare the local 
species richness and composition of Bra- 
chycera in sedge meadows having different 
physical characteristics and dominated by 
different Carex species. 


MATERIALS AND METHODS 


Study sites—The study was conducted 
in the Lac Saint-Francois National Wildlife 
Area (NWA) on the south shore of Lac 
Saint-Frangois, an enlargement of the St. 
Lawrence River in southwestern Quebec. 
The NWA comprises mostly marshes dom- 
inated by Carex spp. (Cyperaceae) and Ty- 
pha spp. (Typhaceae) and swamps domi- 
nated by Acer rubrum L. (Aceraceae), Salix 
spp. (Salicaceae), or Alnus incana (L.) 
Moench. (Betulaceae) (de Repentigny and 
Fragnier 1986, Jean and Bouchard 1991). 
Three sample sites were chosen on the basis 
of dominance and homogeneity of Carex. 
All sites were open habitats with a herba- 
ceous layer between 1.05—1.10 m in height. 
Trees were present only on the periphery. 
Sites LAC1 (45°00.39'N, 74°30.99'W) and 
LAC2 (45°00.17'N, 74°30.63'W) were 
about 1 km apart; site AQUI (45°02.40'N, 
74°28.03'W) was about |1km from the oth- 
ers. Beaulieu and Wheeler (2601) listed the 
plant species recorded at each site. 

Site AQUI (Fl in Beaulieu and Wheeler 
2001) was about 0.16 ha in size and was 
dominated by Carex aquatilis Wahlenb. and 


177 


Calamagrostis canadensis (Michx.) P. 
Beauv. (Poaceae). Carex lacustris Willd. 
was abundant, but less so than at the other 
sites. Plant diversity and overall heteroge- 
neity were higher at AQUI than at the other 
sites. The substrate consisted of hollows of 
firm wet soil interspersed with tussocks of 
sedges and other vegetation. From late 
April until late May the water depth was 0— 
15 cm; for most of the summer the water 
table was near the surface and the soil was 
moist to saturated. The water depth in- 
creased to Scm in mid October and the site 
remained covered by water for the rest of 
the season. The surrounding area was dom- 
inated by C. lacustris, C. aquatilis and A. 
incana. 

Site LAC1 (T1 in Beaulieu and Wheeler 
2001) was about 0.12 ha in size with an 
almost pure stand of C. lacustris, at a den- 
sity of 277 + 14.8 stems/m? (mean + SD 
based on multiple | m? quadrats) in mid 
July. The substrate was homogeneous, flat 
and muddy. The water level was similar to 
that at AQU1 before and during the sam- 
pling period, with a gradual increase in wa- 
ter depth from O—5 cm to 15—25 cm from 
mid September until late October. The veg- 
etation of the surrounding area was more 
diverse. 

Site LAC2 (T2 in Beaulieu and Wheeler 
2001) was about 0.17 ha in size with an 
almost pure stand of C. lacustris, at a den- 
sity of 166 + 14.8 stems/m?*. The substrate 
was similar to LAC1, although the exposed 
mud was more extensive because of the 
lower density of Carex. There were more 
puddles of standing water than at LAC and 
water depth increased from about Ocm dur- 
ing most of the sampling period to 0-5 cm 
in mid-September and 10—20cm in mid-Oc- 
tober. The surrounding area contained a va- 
riety of herbaceous plants, although C. /a- 
custris was still dominant. 

Specimen collection and preparation.— 
Flies were collected every 6—8 days from 
14 May to 27 October 1999. Four yellow 
pan traps (15 cm diameter, 4 cm deep) were 
placed in a 5 m square in the center of each 


178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


On 500 


Individuals 


Fig. 1. 
pooled (B). 


site. Traps were set with their upper rim 
flush with the ground surface in sites where 
the water had receded; traps were filled 
with propylene glycol, and a drop of liquid 
detergent was used as a wetting agent. Veg- 
etation was also swept at mid-day using a 
37.5 cm aerial net for 20 sweeps along each 
of three random 20 m transects at each site. 
The location of transects within each site 
differed from week to week. 

Insects were preserved in 70% ethanol. 
Small flies were dried using hexamethyldi- 
silazane; large flies were pinned and then 
transferred to ethyl acetate for 24h before 
air-drying. All specimens were deposited in 
the Lyman Entomological Museum, McGill 
University, Ste-Anne-de-Bellevue, QC. 

Most specimens were identified to named 
species (where possible) or morphospecies; 
Anthomyiidae, Fanniidae, and most Phori- 
dae were not sorted to species. In all anal- 
yses, morphospecies were treated as equiv- 
alent to named species; unsorted genera 
were not included except for the calculation 
of the number of individuals per trophic 
group, where feeding behavior in the genus 
could be inferred. 

Data analysis.—Species richness was 
calculated based on all species collected at 
each site over the season. Data from pan 
traps and sweeping were combined in all 
analyses except in some correspondence 


1000 1500 2000 2500 3000 3500 


0 2000 


4000 
Individuals 


6000 8000 


Rarefaction (collector’s) curves for Brachycera species collected in each study site (A) and all sites 


analyses. To examine sampling efficiency in 
measuring species richness, rarefaction or 
collector’s curves were plotted using the 
program EstimateS version 6 (Colwell 
2000), with 40 randomizations for each 
curve. Non-parametric estimators of species 
richness (ACE, Chaol, Jackl) were calcu- 
lated using Estimates. 

Species composition among sites was 
compared using the Jaccard index, which is 
based on presence-absence of species, and 
the Bray-Curtis coefficient, which incorpo- 
rates species abundance (Legendre and Le- 
gendre 1998). To minimize the influence of 
accidental visitors, species represented by a 
single specimen at a site were omitted from 
that site in calculating the indices and the 
number of species shared between sites (Ta- 
ble 3), reducing the number of species in- 
cluded in the analyses from 338 to 234. 
Both indices were calculated for all species 
pooled and for the six dominant families 
individually. 

Species composition within and among 
sites was also compared using correspon- 
dence analysis (CA). Replicates of pan 
traps were used in the analysis but data 
were pooled for all three sweep samples in 
each site (within-site replicates were not 
considered because sweep samples were not 
taken along the same transects each week). 
The CA was conducted using the program 


VOLUME 107, NUMBER 1 


179 
Table 1. Number of species and specimens of dominant (ten species) families of Brachycera. 
Species Specimens 
Family AQUI LACI LAC2 Total AQUI LACI LAC2 Total 
Dolichopodidae' 26 23 30 49 184 192 282 658 
Chloropidae 34 26 20 45 398 138) 273 1,804 
Sciomyzidae 16 25 22 31 106 279 PDT) 662 
Ephydridae 8 19 25 31 19 110 186 315 
Syrphidae 12 17 12 25 110 116 107 333 
Sphaeroceridae! 12 il) 18 22 187 571 627 1,385 
Muscidae 8 9 14 21 144 150 135 429 
Empididae 12 1] 10 18 34 38 66 138 
Agromyzidae 7 6 I 1] 2] 13 9 43 
Tabanidae 5 9 8 10 18 36 3] 85 
Scathophagidae 5 6 4 10 15 8 1] 34 
All Brachycera 184 203 196 338 2,825 4,045 3,519 10,384 


' Chrysotus Meigen (Dolichopodidae) and Leptocera Olivier (Sphaeroceridae) are included in specimen totals 


but not species totals. 


SAS version 6.12 (SAS Institute 1996). Pan 
trap samples were ordinated according to 
their similarity in relative abundance of 
species and were then projected on the two 
axes that explained the largest amount of 
variation in species abundance among sam- 
ples. One CA was conducted for pan traps 
and a second for sweep samples. Only spe- 
cies represented by more than ten speci- 
mens (51 species in total, for both CA) 
were included in the analysis to minimize 
the influence of rare species. Prior to CA, 
numbers of specimens were transformed 
into percentages of the total abundance of 
the species over all sites. A third CA was 
conducted using only the 39 most dominant 
species (those represented by at least 30 
specimens). Both sampling methods were 
pooled in this analysis and the three study 
sites were plotted on the same axes. The 
dominant species were then plotted on the 
same graph to show their distribution 
among the sites. 


RESULTS 


Species richness and abundance.—Al- 
though 338 species were identified, most 
were collected in low numbers; 31.5% of 
species were singletons and 11.6% were 
represented by only two specimens (see 
Beaulieu and Wheeler (2001) for a com- 


plete species list with number of specimens 
collected, comparison of pan traps versus 
sweeping, and trophic groups). 

Species richness was similar in all sites, 
with 184 to 203 species each (Table 1). 
ACE, Chaol and Jack! all estimated a min- 
imum of 260 species at each site (Fig. 1a), 
suggesting that about 25% of species actu- 
ally present were not collected. For all sites 
pooled, the rarefaction curve (Fig. 1b) did 
not reach an asymptote, and ACE, Chao] 
and Jack! estimated that at least 434 spe- 
cies may be present. 

Species richness was highest in early to 
mid June, especially in LAC] and LAC2 
where 64—80 species were collected per 
week (Fig. 2c) but at least 15 species per 
week were collected until late October. 
Sciomyzidae, Sphaeroceridae and Ephydri- 
dae were largely responsible for the higher 
species richness in LAC sites in May and 
June (Fig. 2). 

Dolichopodidae, Chloropidae, Sciomyzi- 
dae, Ephydridae, Syrphidae, and Sphaero- 
ceridae were the most species-rich families 
in all sites (except that Ephydridae were 
less speciose in AQUI and Muscidae ex- 
ceeded Syrphidae in LAC2) (Table 1). 
These six families contributed 62.3 = 2.8% 
(mean + SD) of all species at each site. The 
same families were also dominant in num- 


180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 
Creel Pen, When ey (hm) oS 
“@....@ \ ~ wer LW, 
' 
er. * 
° ". . 
ys A Ne 


. 
. 
. 
.* 


— 


—s 
Oo N FF DO DO OD NOON fF 


Species 
Loon ©O CO 
se Oo fC Oo © & 


ES} 
Sa a RTS 
N © 
= S 


20 May 
28 May 
05 Jun 
11 Jun 
19 Jun 
26 Jun 

05 Jul 


Fig. 2. 


A 


23 Jul 
02 Aug 


een I a reo ce ose 
* . 
Op OSE Sr 


L@)) .@)) Le)) Le) Q on Q Qa on a om om 
5 3 3 S} (oO) (0) (0) oO {S) {S) [S) {S) 
qIqtqf_gf f€HNn7 nN HWW OOO O 
N foe) - ODO DK 
SRA oe SES ona mma 
Date 


Seasonal pattern of species richness in Sciomyzidae (A), Sphaeroceridae (B), all Brachycera (C) 


(excluding Phoridae, Anthomyiidae and Chrysotus (Dolichopodidae)). No pan traps were installed in site LAC2 


during the first sampling week. 


bers of individuals (except Ephydridae in 
AQU1) (Table 1). Chamaemyiidae, Phori- 
dae, and Muscidae were also collected in 
high numbers at all sites (1990, 1654, 427 
specimens, respectively). The dominance of 
Chamaemyiidae was primarily due to Plu- 
nomia elegans, which accounted for 18.5% 
of all flies collected. 

Comparison of species composition be- 
tween sites.—Only 73 species (21.6%) 
were collected at all three sites; 37 of those 
were represented by at least ten specimens. 


Chloropidae, Dolichopodidae, and Scio- 
myzidae were a major component of this 
widespread set of species. 

LAC1 and LAC2 shared 28 species rep- 
resented by at least ten individuals, and 57 
species overall. Sciomyzidae, Ephydridae 
and Sphaeroceridae were a major compo- 
nent of the assemblage collected only in 
LAC sites; all three were more speciose and 
abundant in LAC sites than in AQU 1 (Table 
1). These families also contributed to the 
higher Jaccard and Bray-Curtis coefficients 


VOLUME 107, NUMBER 1 


between LAC sites (all families, Table 3), 
than between either LAC site and AQUI. 
AQUI shared only eight species with more 
than 10 individuals (23 species overall) 
with LACI, and four species with more 
than 10 individuals and 17 species overall 
with LAC2. 

Few of the dominant species were unique 
to a single site; most species collected at a 
single site were represented by very few 
specimens. Eleven species represented by at 
least ten individuals, and 70 species in total, 
were unique to AQUI, compared to three 
dominant species (50 in total) unique to 
LAC1 and two dominant species (49 in to- 
tal) in LAC2. 

The four pan traps in each site clustered 
together in the CA of pan trap samples (Fig. 
3, inset), indicating that variation in species 
abundance within each site was lower than 
that among sites. The first and second axes 
explained 45.1% and 13.1% of the total 
variance, respectively. Two axes explained 
100% of the variance for sweeping because 
only three samples were included. In both 
analyses, the species assemblages in LAC 
sites were more similar to each other than 
to AQUI. 

Habitat associations of dominant spe- 
cies.—In the CA of dominant species (Ta- 
ble 2), two axes explained 100% of the var- 
iation in species abundance among sites be- 
cause only three sites were compared (Fig. 
3). Dominant species were divided into 
three groups. Group I species (triangles, 
Fig. 3) were collected exclusively or pri- 
marily in LAC1 and/or LAC2. This was the 
largest group, with 22 (56.4%) species. 
Twelve species were assigned to Group II 
(circles, Fig. 3), abundant in all three sites. 
Five species were assigned to Group III 
(squares, Fig. 3), collected exclusively or 
primarily in AQU1. Plunomia elegans was 
by far the most abundant species at all sites; 
comments on selected families represented 
by multiple dominant species follow. 

Twelve of the dominant species (30.8%) 
were Chloropidae; larvae of most of these 
are saprophagous secondary invaders. 


181 


Three species, Rhopalopterum soror, R. n. 
sp. nr soror and the phytophagous species 
Pseudopachychaeta approximatonervis 
were entirely or mostly associated with 
AQU 1. Rhopalopterum atriceps and Incer- 
tella bispina were more abundant in AQU1, 
whereas /ncertella n. sp. 1 was more abun- 
dant in LAC]. Eribolus longulus, Elachip- 
tera nigriceps, E. pechumani, and E. penita 
were primarily associated with LAC sites. 
Chlorops seminiger (phytophagous) and 
Dasyopa n. sp. 1 Carval habits unknown) 
were more abundant in LAC sites. 

Five of the seven dominant Sciomyzidae 
(Elgiva solicita, Pherbellia vitalis, P. anu- 
bis, Sepedon spinipes americana and Tetan- 
ocera ferruginea) were associated with 
LAC sites. The two other dominant species, 
Dictya steyskali and Poecilographa decora, 
were among the few mainly associated with 
AQUI. 

Three dominant species of Sphaeroceri- 
dae (Phthitia quadricercus, Rachispoda n. 
sp. 1, Rudolfina n. sp. 1) were almost en- 
tirely confined to LAC sites. Pullimosina 
pullula was most abundant in LAC sites, 
but was also collected in AQUI. 

Dolichopodidae, Syrphidae, and Musci- 
dae had three dominant species each. In all 
cases those species were either abundant in 
all sites (six species) or entirely or mostly 
restricted to LAC sites. 

Diversity and abundance of trophic 
groups.—The trophic group or subgroup 
(Table 4) was known for some species and 
was inferred for other species from pub- 
lished records on congeners. The trophic 
group was tentative for ten species and un- 
known for 24 species. Many species, es- 
pecially saprophages, were not assigned to 
a trophic subgroup, either because they 
have been reared from a range of decaying 
media or because their exact food source is 
unknown. As a result, total species numbers 
for trophic groups do not always corre- 
spond to the sum of species assigned to tro- 
phic subgroups (Table 4). 

Predators and saprophages were the most 
species-rich (145 and 118 species, respec- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


182 


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nusSad ae 6¢ + 9 dINa Jopueloyy Ssnoliasojoy sndjodiyv] gq [oud 
ound 44 Or IOS (M]) Dlozap vydpssoj19a0g 29Pd 
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183 
1 

R 

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ple cae 
ME 

LU 

VO 


in- 
411 
al 5: 

an 

862 

dant Ge 

bu 

d a 

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ively) a 

tive 


ing over 
ng = 
esentl s (Ta 
T imens = 
rep 1m le 
ach ec id e 
eac d sp mic S 
Ips; © ; an luno eous 
group "1es 12, dac 
g spec us re re 
i uals) all sp hago the p h the 
I divid ird of hidop If of houg were 
I < ne thir he ap for ha Alt they | or 
n Se o 4). T nted imens). cles, sitoids S 
= a ble accou specin us spe paras hagous 
3: 5 1S 20 < agous few op b- 
Oa. Y < is sg al 92 hag re apr su 
E < Pa o Ss Fak (1 hytop re we wer S in the h- 
- Ew < 3 3 fli 33 p The aaa oD 
= Bo Be us| 3 were nadine Ul ha se liv of eae: 
a = u hos eG 
i ll not a res. a A ; numb er nee 
Fs O i nivo rticu eee e few inv ‘i 
Ho ! of om les, pa he hig re Wer ndary 1, mai 
: if Z spec and t Ther 5 Seco PAGE lan. 
e za aE rate, cies. LAC2. nt in GAVE od 
i se st S spec s in nda ids, In strop 
SO as) ; a US. 
4 Sieuirmna oe 6 were pail two ch longu nt and 
N ray > Ss a : 
ey © ly beca d Eri ore a in A 
Le eg y an m han 
3) x eA eral were C2t ceTa 
a3 as) sp rs LA hyc 
| 224% dato d ON c t 
+ + yeas 5 pre Cl an CUSSI Bra insec 
7 eae gs in LA Dis nee Nera like 
cats in chyc t of Un 
aM Bra en S. m- 
5 glee te cee eens ce) ly te 
Oo = os Dive inan dge wer stan = 
om | = dominz se ites no is sea 
E apie aieOl in rs ost ; this 
2 ¢ ak are nity ou alm er; ‘be an 
iS Es 5 mmu tlands, with poe may era 
salar aq 5 co we ed, the e hye 
oF <r | © al & shinee flood ost of ater ay Brac eu 
lei 1 2 : 
a ie: ob p er d enc ula chia ding s d- 
on “ik ne wat iffer r reg Ss su bree inva 
= 2 2 || aoe al d facto lant itable ay a ed 
o ca Sr cs) son nt nt p uita nd lat 
e “22 Eas 3 porta bie esse oe us 
n or 7 é im m Oo ies, § i ous 
N = G 1 E te cie ors ene 
. a og ull ere na. stra spe dat 10g sed 
o) D Neg fau sub us re hon Os 
S d O d p e mp d 
: 281383 mu hag an The con ; an 
=Oz pls < A hytop! ores soul.” SILES; io 25 
aise Oh SS 5 p tiv et “Ls spe Srey 
= Z| aana = 55 for detri dw Uustrl ant sp spec 
2 an UN v a3) Bi Dy rs, nts an lact ne pla WSS: flora 
Z| 0 ce ae the C. a ana S. 
31% a sata sively a eneou: 
= a a) 1 us Pre in g an 
ew ae sites Aen 
> a o Z Es caine oe a juatilis “more ee = 
3 E¢ Raye! ia nl ag re - Bra rele lop 
ry oO a a) oO | cI au G: we yf bly SVeLC ~ 
N © Sort a = the -ate sity ¢ roba de yf 
= = Vas BI than bstr iversi S pre arvae rate ¢ 
= ‘a ec) su div 3 Was j la stré 
fs give | 8 mS) » 3 and eater sites any fly e sub ; could 
= S S s Ss = = iB The gr LAC : te. Me and th C sites S Or 
S 7S $ n ' strate. é A pct 
= ae Cee e oo S in bstr 8), L “itive “T= 
2 Seine lla S : su 97 e rit ael 
osecieS: SE ae 2) erento : in | ; like th f eeu: 5 Sph a- 
2 = al ere oe he 1 Irw S like so ‘ids. -eda 
s1 § & Opes 5 Ea S 2 t d ( arshes ations ydr ic pre = 
2 S gc a] nu a la = hy Cc : 
ai, & 755 1 || ES l & in n m u ep atl am 
| & = S 8 ¢ op AS in nt op a i-aque se fe 
Ais SG S Sigs By S| le: = erge rge p 3 (e.g -IM1-<¢ hese and 
sss a § ae em rt larg zers se >it ites a 
g S t e d ) 
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peebes Soeaes supp ial gt ids) anc lary Cs 
< = 2 5 3 = 5; e) 3 " i Ss icrobia rphids odid in LA 
= = 2 OES 3 plies 45 To rids, _dolichopodi 
R= 38 S 5 SE= Ss oce g., C re a 
| S288 S&| 35 ETS baie aie 
Ras Q 3 = a r 
3 Pega egil4 INEST 
g ag fe aa 2 
5 g| eZee a 
O a aale = 
= = ‘=| 
6 ) Be ~ 
ol rans page 
rm vy} 32 m 
ra) a 
CS 


drids, 
ephy 
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lichopodi 
do 

(e.g: 

some 


184 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Pan traps 


Axis 2 


Fig. 3. 


Sweeping 


AT2 


0.5 1 1.6 2 


Axis 1 


Correspondence analysis ordination of study sites (insets) and dominant Brachycera species. Shaded 


symbols = study sites; open triangles = Group I species; open circles = Group II species; open squares = 


Group III species. 


sphaerocerids) were most abundant in 
LAC2 where there was more moisture and 
exposed mud. 

According to the estimates of species 
richness, many species presumably present 
in the sites during the sampling period were 
not collected. Comparison of the species 


collected in LAC] and LAC2 supports this; 
the two sites were within lkm of one an- 
other and represent basically the same hab- 
itat but many species were collected in only 
one of the sites. Many of these species may 
have been ‘tourists’? whose presence was 
due to conditions in the surrounding habi- 


Table 3. Similarity in species composition (excluding singletons) between pairs of sites for six most dominant 
families and all Brachycera. JAC = Jaccard index, B-C = Bray-Curtis coefficient, shared = number of species 


shared. 
LAC1/LAC2 AQUI/LACI AQUI/LAC2 
Family Jac B-C Shared B-C Shared Jac B-C Shared 

Dolichopodidae 0.48 0.51 10 0.35 0.40 7 0.25 0.31 6 
Chloropidae 0.52 0.30 13 0.41 0.28 13 0.44 0.30 12, 
Sciomyzidae 0.67 0.68 16 0.32 0.18 8 0.29 0.12 6 
Ephydridae 0.44 0.63 8 0.40 0.25 4 0.18 0.16 3 
Syrphidae 0.40 0.67 4 0.60 0.49 6 0.56 0.68 5 
Sphaeroceridae 0.56 0.66 10 0.20 0.23 3 0.13 0.15 2 
All Brachycera 0.51 0.64 92 0.30 0.41 Sy7/ 0.27 0.43 51 


VOLUME 107, NUMBER 1 


tats, such as phytophagous species in fam- 
ilies like Agromyzidae or Tephritidae 
whose host plants were present near the 
sample sites. Such accidental visitors 
should not be considered equivalent to truly 
rare inhabitants of the site, but given the 
lack of information on ecology of many 
species, it is often difficult to discriminate 
between true inhabitants and tourists that 
were simply carried by wind (often sub- 
stantial in open shoreline habitats). An ad- 
ditional confounding factor is that some 
species may not inhabit sedge meadows 
throughout their larval stages but may use 
them solely for adult resting, feeding or 
mating sites (Delettre et al. 1998). Addi- 
tional inventories carried out in a range of 
habitats (e.g., marshes, swamps, meadows) 
as well as ecological studies of larval habits 
and habitats may help to resolve the status 
of resident species, transient species who 
use the habitat during part of their life cycle 
and true “‘tourists.”’ 

Habitat associations of dominant spe- 
cies.—The habits and habitats of many Bra- 
chycera species are unknown; this is an ob- 
stacle to ecological research on the group. 
Some taxa have been studied in detail, eco- 
logical data are also scattered in taxonomic 
publications and inferences about some spe- 
cies can be made by comparison with ob- 
servations on related species. Although it is 
difficult to establish habitat associations 
based on only three sites representing two 
similar habitats, some general patterns were 
noted in some dominant families and spe- 
cies. 

Dolichopodidae are diverse and abundant 
predators in wet habitats such as marshes 
and peatlands (Pollet 1992, 2001; Blades 
and Marshall 1994). Although most of our 
49 species were collected in low numbers, 
Carex meadows are clearly a suitable hab- 
itat. The fact that 18 species were repre- 
sented by singletons suggests either that ad- 
ditional rare species have been missed or 
that adults of many species range widely in 
search of prey. 

Some Syrphidae may be closely associ- 


185 


ated with Carex and mud substrates. Mai- 
bach and de Tiefenau (1993) studied larvae 
of four European species of Neoascia Wil- 
liston and found one exclusively between 
leaves of Typha, one between leaves of 
Carex, one in soil or among mosses, and 
the fourth in a variety of habitats including 
Carex and organic soils. Thus, N. metallica 
may develop in Carex stems and/or wet 
mud. It is not known whether all species of 
the genus are predaceous or saprophagous. 
Larvae of Lejops Rondani and Parhelophi- 
lus Girschner breed in mud, probably feed- 
ing on detritus and microorganisms like 
many other Eristalini (Ferrar 1987). Blades 
and Marshall (1994) collected Lejops and 
Neoascia species in Ontario fens and bogs. 
Five species of Platycheirus Lepeletier and 
Serville identified in this study have been 
previously collected in Carex marshes 
(Vockeroth 1992), although there is no oth- 
er habitat information on those species. 
Sciomyzidae thrive in wetlands and their 
abundance in all study sites suggests that 
suitable host snails are abundant. The five 
dominant species (Elgiva solicita, Pherbel- 
lia vitalis, P. anubis, Sepedon  spinipes 
americana and Tetanocera ferruginea) in 
LAC sites feed on a variety of aquatic pul- 
monate snails. Knutson and Berg (1964) re- 
corded adults of E. solicita in emergent her- 
baceous vegetation of marshes, ponds, lake 
margins, and stream banks. Bratt et al. 
(1969) found P. vitalis and P. anubis dom- 
inant in vernal marshes where the water re- 
cedes during summer, exposing snails on 
the mud or vegetation. Pherbellia anubis 
apparently prefers partially shaded Carex 
marshes but P. vitalis has been collected in 
a range of habitats from swamps to tundra. 
Adults of S. spinipes americana are found 
on low vegetation in marshes, wet meadows 
and fens (Neff and Berg 1966, Blades and 
Marshall 1994). Larvae of 7. ferruginea 
have been collected in shallow water 
among aquatic macrophytes (Foote 1999). 
The habits of Dictya steyskali, dominant in 
AQUI, are unknown but other Dictya spp. 
are found in a variety of habitats such as 


186 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 4. Brachycera species and species and specimens (in parentheses) by larval trophic group or subgroup. 
Subgroup totals do not always add up to group total (see text). 


i 


Trophic Group 
{or subgroup] AQUI 
Phytophagous 23 (99) 
[Stem-borer] 11 (46) 
[Leaf-miner] 7 (19) 
[Flowerhead feeder] 5 (34) 
Saprophagous 55 (1,491) 
[Secondary invader] 17 (334) 
[In substrate] 16 (50) 
Predaceous 78 (1,004) 
[Aphidoid predator] 13 (552) 
[Mollusc predator] 16 (106) 
[In substrate] 43 (230) 
Parasitoid 6 (40) 
Omnivore 2 (5) 
Unknown 13 (143) 


Site 

LACI LAC2 Total 
17 (60) 9 (36) 33 (195) 

5) () 4 (20) 12 (91) 

7 (14) 2 (10) 13 (43) 

3 (8) 2 (5) 6 (47) 
80 (2,164) VON@ET SO) 118 (5,411) 
19 (1,070) 14 (280) 24 (1,684) 
32 (240) S293) 49 (583) 
79 (1,427) 87 (1,431) 145 (3,862) 

9 (790) 8 (706) 22 (2,048) 
Pus) (27/2) 22 (277) 31 (662) 
40 (256) 48 (376) 76 (862) 

8 (113) 7 (100) 13 (253) 

3 (5) 1 (2) 3 (12) 
14 (234) 11 (163) 24 (540) 


ee 


marshes, wet woodland, wet meadows and 
bogs (Valley and Berg 1977). The larval 
habits of Poecilographa decora are un- 
known, but adults are associated with wet- 
lands and puparia have been collected in a 
bog and a wooded swamp (Barnes 1998). 
Chloropidae are dominant in wetlands 
and many species feed on monocots (Ferrar 
1987, Beaulieu and Wheeler 2002). They 
were one of the most species-rich families 
in this study, particularly in AQUI, where 
monocot diversity was higher. Some phy- 
tophagous species, such as Meromyza fla- 
vipalpis Malloch (stem-borer in grasses) 
and Dicraeus fennicus Duda (flowerhead 
feeder in grasses), were collected only in 
AQUI. Chlorops seminiger was dominant 
in LAC sites and has been reared from C. 
lacustris (Beaulieu and Wheeler 2002). 
Pseudopachychaeta approximatonervis has 
been reared from flowerheads of Eleocharis 
R. Br. (Cyperaceae) (Valley et al. 1969, 
Todd and Foote 1987a) and Scirpus (B.A. 
Foote, personal communication), but these 
hosts were absent in and near AQUI and 
this species may also feed in Carex. 
Secondary invaders comprised about half 
of the chloropid species (including most of 
the dominant species) and over two-thirds 
of the specimens collected. Eribolus lon- 


gulus is associated with sedges and grasses 
in wet habitats (Valley et al. 1969, Beaulieu 
and Wheeler 2002). Elachiptera pechumani 
has been reared from /ris spp. (Valley et al. 
1969) and E. penita from Phalaris arundi- 
nacea L. (Poaceae) (Beaulieu and Wheeler 
2002), but their abundance in LAC sites 
and the rarity of known host plants in the 
area suggest that C. lacustris is also a host. 
Rhopalopterum atriceps has been reared 
from Carex comosa FE. Boott. and C. lacus- 
tris (Valley et al. 1969, Beaulieu and 
Wheeler 2002) and its abundance in AQUI 
suggests that C. aquatilis is also a host. The 
dominance of Jncertella n. sp. 1 in all sites 
suggests that it is a secondary invader of 
Carex. 

Several species of Sphaeroceridae col- 
lected in our study are also found in peat- 
lands, and Marshall (1994) considered three 
of those (Phthitia quadricercus, P. ovicer- 
cus Marshall, Spelobia pappi Rohacek) 
characteristic of that habitat. Our sites share 
some characteristics with peatlands (wet or- 
ganic soil, presence of Carex). Phthitia spp. 
are usually found in wet plant material 
(Marshall 1994). Few Sphaeroceridae are 
likely to be confined to sedge meadows; 
many are associated with a range of decay- 
ing organic matter. Examples of such gen- 


VOLUME 107, NUMBER 1 


eralists collected in our study include Lo- 
tophila atra (Meigen), Pullimosina pullula, 
Rachispoda n. sp. 1, Spelobia_ clunipes 
(Meigen) and S. luteilabris (Rondani). 

Many Ephydridae are grazers of micro- 
organisms and detritus in a range of wet 
habitats including mud shores, sedge mead- 
ows and beaches (Foote 1995). Our study 
sites were a combination of these three 
types of habitats. Nine of the species col- 
lected in our study were also recorded from 
peatlands by Blades and Marshall (1994) 
and eight from a freshwater marsh by Todd 
and Foote (1987b). Notiphila caudata, 
dominant in LAC sites, is abundant in a 
range of habitats including Carex meadows 
with exposed mud, marshes, beach wrack, 
mud shores and peatlands (Eastin and Foote 
1971, Blades and Marshall 1994). 

Species of Plunomia are associated with 
wetlands and P. elegans was the dominant 
species in all sites. Adults were reared from 
larvae or puparia found between the leaves 
of C. lacustris where the larvae probably 
feed on aphids such as Thripsaphis sp. 
(Drepanosiphidae) (Beaulieu and Wheeler 
2002). 

Diversity and abundance of trophic 
groups.—Over half of all specimens and 
over one third of all species identified were 
saprophagous. Predators were more spe- 
cies-rich but not as abundant. All other tro- 
phic groups ranked far behind in both spe- 
cies and specimens. This relative ranking 
was expected, given the nature of the study 
sites. Wetlands provide ample food sources 
for saprophagous larvae and such families 
are usually dominant (Blades and Marshall 
1994 [reanalyzed for Brachycera only], 
Keiper et al. 2002). The low plant diversity 
at our sites may explain the less diverse 
phytophagous guild. 

The lower diversity of saprophagous spe- 
cies in AQUI, particularly those in the sub- 
strate, is probably related to differences in 
substrate structure. The extensive wet mud 
in LAC sites may be a better habitat for 
detritivores and microbial grazers 
ephydrids, sphaerocerids, some syrphids) 


(e.g., 


187 


than the hummocky firm substrate in 
AQUI. The lower diversity of phytopha- 
gous species in LAC2 may be partly ex- 
plained by the more heterogeneous vege- 
tation in AQUI and partly by the lower 
density of C. lacustris in LAC2. Despite the 
above differences, the similarity in trophic 
structure of Brachycera emphasizes the 
similarity of our study sites; they each con- 
sist basically of wet mud dominated by a 
single species of Carex. Given that we col- 
lected over 300 species of Brachycera in a 
single season, in three such ““homoge- 
neous” sites, Diptera diversity may be 
higher than expected even in apparently 
uniform habitats. 


ACKNOWLEDGMENTS 


We thank the Canadian Wildlife Service 
for permission to sample at Lac St. Francois 
NWA and S. Bourdon and D. Gervais for 
information about the sites. S. Boucher, S. 
E. Brooks, B. E. Cooper, J. M. Cumming, 
J. E. O’Hara and J. R. Vockeroth identified 
Diptera; Y. Bérubé identified plants; P. Le- 
gendre, P. Paquin and M. Pollet advised on 
data analysis. Financial support was provid- 
ed by an NSERC Postgraduate Scholarship 
to FB and an NSERC Discovery Grant to 
TAW. 


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PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 190-199 


THE MAYFLIES (EPHEMEROPTERA) OF ALASKA, INCLUDING A NEW 
SPECIES OF HEPTAGENITDAE 


R. P RANDOLPH AND W. P. MCCAFFERTY 


(RPR) Division of Evolution and Ecology, University of California, Davis, CA 95616 
(e-mail: rprandolph@ucdavis.edu); (WPM) Department of Entomology, Purdue Univer- 
sity, West Lafayette, IN 47907 (e-mail: pat-mccafferty @entm.purdue.edu) 


Abstract.—Forty-six species of mayflies (Ephemeroptera) are reported from Alaska, and 
27 of those are reported from the state for the first time. Nominal species in the genera 
Callibaetis Eaton, Caudatella Edmunds, Ecdyonurus Eaton, Ironodes Traver, Paralepto- 
phlebia Lestage, Plauditus Lugo-Ortiz and McCafferty, Procloeon Bengtsson, and Siph- 
lonurus Eaton are also reported from Alaska for the first time. The fauna consists mainly 
of species with widespread general or western distributions in North America, but also 
includes 11 confirmed Holarctic species. Rhithrogena ingalik, new species, is described 
from male adults; it differs from other congeners in genitalia morphology and is most 


closely related to certain Siberian species. 


Key Words: 
records 


Because of concerns of global warming 
of Arctic and alpine habitats and the poten- 
tial impacts on those ecosystems (Chapin 
and Korner 1994), it is important that doc- 
umentation of the organisms within those 
regions be completed. Studies of ecosys- 
tems of low diversity, such as those found 
in the Arctic, may provide a ready means 
of understanding these systems and any 
changes they may undergo (Danks 1981, 
Chapin and Korner 1994, Poff et al. 2001). 
In addition, such data may be valuable for 
understanding historical biogeography 1in- 
volving circumpolar species. 

Considerable recent work has contributed 
to the documentation of the Ephemeroptera 
fauna of far northern North America (e.g., 
Harper and Harper 1981, 1997; McCafferty 
1985, 1994, McCafferty and Randolph 
1998, Alba-Tercedor and McCafferty 2000, 
Randolph and McCafferty 2000). Species 
distributional data for certain subregions of 


Alaska, Ephemeroptera, mayflies, Rhithrogena ingalik, new species, new 


the North had not been readily available 
previously and as a result faunal data have 
remained fragmentary, making any ecolog- 
ical or biogeographic assessments difficult 
(Harper and Harper 1997). For example, of 
the 63 species reported from the combined 
subregions of Alaska, Yukon, Northwest 
Territories, and Nunavut, relatively few 
have been known from Alaska (McCafferty 
1985, 1994; Zloty 1996; Lee and Hershey 
2000). A more complete documentation of 
Alaska is critical for hypothesizing Berin- 
gia’s role in mayfly dispersal during the 
Pleistocene (see H6fle et al. 1994, Elias et 
al. 2000), and the possible role of Alaska 
in providing historical refugia in its ungla- 
ciated areas (McCafferty 1985, Elias et al. 
2000). 

Herein we provide new state records for 
27 species and 71 new county records for 
39 of the 46 species we document from 
Alaska. This includes a new species of 


VOLUME 107, NUMBER 1 


191 


Hoy North Slope 
ag periie 
Auuttest Si aaa 3 
i aCe retic oH 
ae PROG 
Nome ~~-~4 Yukon- 
s the Koyukuk 
I oot -7 
H ' Fairbanks; .-! 
i. - =p « North Star's 
Wad ae -” Denajj | Southeast 
e i My pase na ° 
ampton | mcd Fairbanks 


~ 
~~ 
“ws ue 


Susitna 


Fig. 1. Map of Alaska. 


Rhithrogena Eaton and the first North 
American records of Acentrella lapponica 
[nec A. feropagus| and Ephemerella nuda. 
The first records of nominal species of the 
genera Callibaetis Eaton, Caudatella Ed- 
munds, Ecdyonurus Eaton, [ronodes Traver, 
Paraleptophlebia Lestage, Plauditus Lugo- 
Ortiz and McCafferty, Procloeon Bengts- 
son, and Siphlonurus Eaton are given. This 
study also rectifies previous records based 
on misidentifications. Several species are 
shown to occur above the Arctic Circle, and 
in all, mayflies are now known from 16 of 
the 25 counties in Alaska (Fig. 1). 

Many Arctic species have widespread 
distribution patterns (Downes 1962, Chapin 
and Korner 1994). This was also noted by 


~~" Matanuska-: 


=-<4L 


1 

if 
re 
. 

1 

1 
‘ 


Ketchikan 
Gateway 


McCafferty (1985) for the Alaskan mayfly 
fauna and is further documented here. 
There are 11 Alaskan species that are wide- 
spread North American species, including 
Acentrella turbida, Baetis flavistriga, B. tri- 
caudatus, Callibaetis ferrugineus, C. fluc- 
tuans, Diphetor hageni, Ephemerella do- 
rothea, Paraleptophlebia debilis, Plauditus 
dubius, Heptagenia pulla, and Siphlonurus 
alternatus. Cinygmula subaequalis is found 
outside of Alaska only in eastern and south- 
eastern North America, and this disjunct 
pattern is suggestive of a faunal remnant 
from the Arcto-Tertiary Forest regime 
(McCafferty 1985). The predominant dis- 
tributional pattern expressed by Alaskan 
mayflies involves 17 species that are wide- 


192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


spread in, and limited to, western North 
America. These include Acentrella insigni- 
ficans, Ameletus validus, Cinygmula mimus, 
C. par, C. tarda, Drunella coloradensis, D. 
doddsii, D. grandis, D. spinifera, Epeorus 
albertae, E. deceptivus, E. grandis, E. lon- 
gimanus, Tronodes nitidus, Rhithrogena fu- 
tilis, Serratella tibialis, and Siphlonurus oc- 
cidentalis. This is also the most common 
pattern for Yukon mayflies (Harper and 
Harper 1997). Such far-reaching western 
species are often limited to mountain rang- 
es. Caudatella jacobi and Paraleptophlebia 
vaciva occur in Alaska and also occur 
southward into regions of western Canada 
and USA, but they have a more fragmented 
pattern and do not range into southwestern 
USA. 

Many Arctic arthropod species are Hol- 
arctic (Danks 1981), and the 11 Holarctic 
Alaskan species account for about one- 
fourth of the known mayfly fauna of Alas- 
ka. These include Acentrella lapponica, 
Ameletus inopinatus, Baetis bicaudatus, B. 
bundyae, Brachycercus harrisella, Ephem- 
erella aurivillii, Ephmerella nuda, Metre- 
topus alter, Ecdyonurus simplicioides, Par- 
ameletus chelifer, and Procloeon pennula- 
tum. The only occurrences in North Amer- 
ica of A. lapponica, E. nuda, and M. alter 
are confined to Alaska, and A. inopinatus 
and B. bundyae are strictly northern boreal 
species in North America. Other Holarctic 
species tend to be more widespread in 
North America. The commonness of the 
Holarctic species B. bicaudatus in western 
North America and its only sparse occur- 
rence in far East Asia may suggest east to 
west dispersal via Beringia as recently as 
the Pleistocene, whereas the opposite sce- 
nario is suggested for Ephemerella nuda. 
Acentrella lapponica and M. alter are also 
known from Scandanavia, and their restrict- 
ed North American distribution is appar- 
ently relictual. 

Acentrella feropagus is limited to Alas- 
ka, Canada, and northern conterminous 
USA. Baetis foemina is limited to Alaska, 
Northwest Territories, and Nunavut. Rhith- 


rogena ingalik, n. sp., is more closely re- 
lated to some Siberian congeners than to 
any known North American species, sug- 
gesting allopatric speciation in Alaska. 

Alaskan records below are alphabetical 
by family, genus, and species. County 
names appear in upper case. Material on 
which new records are based are larvae un- 
less stated otherwise and are held in the 
Purdue Entomological Research Collection, 
West Lafayette, Indiana. 


AMELETIDAE 
Ameletus inopinatus Eaton 


Previous records.—Zloty (1996): FAIR- 
BANKS NORTH STAR. 


Ameletus validus McDunnough 


Previous records.—Edmunds et al. 
(1976): No data provided. Zloty (1996): No 
data provided. Wipfli et al. (1998): KET- 
CHIKAN GATEWAY. 

New records.—NORTH SLOPE: Can- 
ning R 5 mi from mouth on main channel, 
70/04/30N 145/33/30W, VI-20-1972. 


BAETIDAE 


Acentrella feropagus Alba-Tercedor and 
McCafferty 


Previous records.—Harper and Harper 
(1981) as Baetis lapponica: NORTH 
SLOPE. Waltz and McCafferty (1987) as A. 
lapponica: YUKON-KOYUKUK. 


Acentrella insignificans (McDunnough) 


New records.—YUKON-KOYUKUK: 
South Slope, Sheenjek R 1 mi above pipe- 
line crossing, 67/37/45N 143/17/00W, VII- 
2921972: 


Acentrella lapponica (Bengtsson) 


New records—NORTH SLOPE: Can- 
ning R, mid-Canning gill net station, 69/27/ 
15N 146/13/15W, VIII-28-1972; Canning R 
5 mi from mouth on main channel, 70/04/ 
30N_ 145/33/30W, VI-27-1972; Middle 
Canning R % mi below Shublik Falls, 69/ 
2T/I5N 146/13/15W, VIII-8-1972; Cache 


VOLUME 107, NUMBER 1 


Cr % mi above mouth, 69/23/45N 146/05/ 
OOW, VII-27-1972; Kavik R #1, VIII-17- 
1973; Canning R at mouth unnamed trib, 
69/35/30N 146/19/30W, VII-27-1972; Can- 
ning R at unnamed Canning R spring, 69/ 
OS5/45N 145/59/30W, LX-24-1973; Kuparuk 
R, VIII-27-1971; YUKON-KOYUKUK: 
South Slope, Sheenjek R 1 mi above pipe- 
line crossing, 67/37/45N 143/17/00W, VII- 
29-1972; South Slope, Old Woman Cr | mi 
from confl Sheenjek R, 68/21/00N 144/00/ 
OOW, VII-28-1972; South Slope, Monument 
Cr, trib Sheenjek R, 67/57/45N_ 143/13/ 
OON, VIII-17-1972; South Slope, Pass Cr 5 
mi below origin, trib Coleen R, 67/53/15N 
142/50/O0W, VII-29-1972. 


Acentrella turbida (McDunnough) 


New records.—NORTH SLOPE: Anti- 
gun R, VIII-15-1971; Kuparuk R, VI-24- 
1971; Canning R 5 mi from mouth on main 
channel, 70/04/30N_ 145/33/30W, VII-27- 
1972; YUKON-KOYUKUK: South Slope, 
Coleen R 1 mi above pipeline route, 67/57/ 
30N 142/09/00W, VII-29-1972; South 
Slope, Monument Cr 1 mi above conf 
Sheenjek R, 67/57/45N_ 143/13/00W, VII- 
29-1972; South Slope, Pass Cr 5 mi below 
origin, 67/53/15N 142/50/00W, VII-29- 
1972; South Slope, Strangle Woman Cr | 
mi above mouth, 67/50/00N 141/41/30W, 
VII-29-1972. 


Baetis bicaudatus Dodds 


Previous records.—Duncan and Brusven 
(1985): PRINCE OF WALES-OUTER 
KETCHIKAN. Milner (1987): SKAG- 
WAY-HOONAH-ANGOON. McCafferty 
(1994): MATANUSKA-SUSITNA. Wipfli 
et al. (1998): KETCHIKAN GATEWAY. 

New records.—ANCHORAGE: Alyeska, 
VI-21-1997; Chugach St Prk, 18 mi from 
Old Seward, VI-22-1997; Chugach St Prk, 
6 mi from Old Seward, VI-22-1996; FAIR- 
BANKS NORTH STAR: Monument Cr, 
Chena R, VII-27-1979; JUNEAU: uniden- 
tified str, prob from water supply, N Juneau, 
VI-21-1958: KENAI PENINSULA: Dia- 
mond Cr nr Sterling Hwy, V-11-1968; main 


193 


trib Lower Russian R, VI-24-1968; Seward, 
tidewater, Northland Glacier, VI-6-2001; 
NORTH SLOPE: Arctic Foothills Province, 
Echooka R, riffle above Echooka Spr 
mouth, VI-6-1971; Arctic Foothills Prov- 
ince, Lupine Spr, VI-26-1971; Arctic Foot- 
hills Province, Nanook Cr, 69/34/30N 146/ 
16/00W, VII-27-1972; Canning R, Marsh 
Frk, lower reg, 69/06/00N 145/59/00W, VI- 
17-1972; Canning R nr Shublick Spr, 69/ 
2T/1S5N 146/13/15W, VII-9-1972; Canning 
R, unnamed spr, 69/05/45N_ 145/59/30W, 
VI-26-1973; Marsh Frk, 69/52/15N 146/00/ 
OOW, VI-17-1972; Middle Canning R %4 mi 
below Shublick Falls, 69/27/15N 146/13/ 
I5W, VIII-8-1972; South Slope, Cane Cr ca 
5 mi above confl Chandalar R, 68/39/30N 
144/54/00W, VIUI-17-1972; unnamed Can- 
ning R trib 400 m above mouth, 69/23/45N 
146/09/00W, VII-27-1972; unnamed Can- 
ning Spr 20 yds from mouth, 69/08/30N 
145/55/00W, VII-26-1972; SITKA: Nak- 
vassin Cr, Baranof Island, Port Herbert, 
VII-1-1958; SKAGWAY-HOONAH-AN- 
GOON: North Arm Hood Bay, VI-21-1969. 


Baetis bundyae Lehmkuhl 


Previous records.—Harper and Harper 
(1981): NORTH SLOPE. 

New records.—YUKON-KOYUKUK: 
South Slope, Monument Cr 10 mi below 
origin, 67/57/45N 143/13/00W, VII-28- 
1972; South Slope, unnamed trib to E Frk 
Chandalar R, 68/25/00N 145/12/00W, VII- 
28-1972; South Slope, Strangle Woman Cr 
10 mi above mouth, 67/50/00N_ 141/40/ 
30W, VII-29-1972. 


Baetis flavistriga McDunnough 


New records.—YUKON-KOYUKUK: 
South Slope, Monument Cr 10 mi below 
origin, 67/57/45N 143/13/00W, VII-28- 
1972: South Slope, Old Woman Cr | mi 
from confl Sheenjek R, 68/21/00N_ 144/00/ 
OOW, VII-28-1972; South Slope, Pass Cr 5 
mi below origin, 67/53/15N_ 142/50/00W, 
VII-29-1972; South Slope, Sheenjek R | mi 
above pipeline crossing, 67/37/45N 143/17/ 
OOW, VII-29-1972: South Slope, Strangle 


194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Woman Cr 10 mi above mouth, 67/50/00N 
141/40/30W, VII-29-1972; South Slope, un- 
named trib E Frk Chandalar R, 68/25/00N 
145/12/00W, VII-28-1972. 


Baetis foemina McDunnough 


New records.—NORTH SLOPE: Arctic 
Foothills Province, Echooka Spr, VII-1- 
1971; Cache Cr % mi above mouth, 69/23/ 
45N 146/05/O0W, VII-27-1972; Canning R 
Marsh Frk, next to pool, 69/05/45N 145/59/ 
30W, VII-23-1973; Canning R spring (CS- 
10) 275 m above weir, 69/05/45N_ 145/59/ 
30W, V-24-1973; Canning Spr 200 yds 
from spring source, 69/52/15N 146/04/ 
30W, VI-16-1972; Canning R at mouth un- 
named trib, 69/35/30N 146/19/30W, VII- 
27-1972; Echooka R, V-27-1971; Kavik R, 
VII-17-1973; Kuparuk R, VIII-29-1971; 
unnamed trib Canning R 400 m above 
mouth, 69/23/45N 146/09/00W, VIII-27- 
IQ. 


Baetis tricaudatus Dodds 


Previous records.—Duncan and Brusven 
(1985): PRINCE OF WALES-OUTER 
KETCHIKAN. Milner (1987): SKAG- 
WAY-HOONAH-ANGOON. Wipfli et al. 
(1998): KETCHIKAN GATEWAY. 

New records.—KENAI PENINSULA: 
Crooked Cr, VIII-20-1965, E Charnov; Hid- 
den Lake Cr nr Sterling Hwy, V-11-1968; 
Six Mile Cr, VII-22-1965; Slicock Cr at Ka- 
lafonsky Rd brdg, V-26-1968; KODIAK 
ISLAND: Lower Olga L, Kodiak Island, 
VI-13-1979; LAKE AND PENINSULA: 
Aniakchak National For Mon., Alas Pen- 
insula, 56/56N 158/06W, VII-1988 (adults); 
MATANUSKA-SUSITNA: 1 mi SE Was- 
illa, Cottonwood Cr at Matanusak Rd, VI- 
19-1958; Knik, Fish Cr, VI-4-1958; 
NORTH SLOPE: Arctic Foothills Province, 
Echooka Spr, V-29-1971; Arctic Foothills 
Province, Ribdon Spr, VI-2-1971; Canning 
R 5 mi from mouth on main channel, 70/ 
04/30N 145/33/30W, VI-20-1972; Canning 
R, Shublick Spr, 69/27/15N_ 146/12/00W, 
VII-9-1972; unnamed trib Canning R, riffle 
above mouth, 69/32/45N 146/15/45W, VI- 


27-1972; WRANGLE-PETERSBURG, 
Anan Cr, [V-26-1958; YUKON-KOYU- 
KUK: Birch Cr between Big Cr & Preach- 
er’s Cr, 66/00N 144/50W, VIII-17-1962; 
Rat Cr 1.6 mi upstr from Dorena Dam, VI- 
14-1958; South Slope, Pass Cr 5 mi below 
origin, 67/53/15N 142/50/00W, VII-29- 
1972; South Slope, Sheenjek R 1 mi above 
pipeline crossing, 67/37/45N 143/17/00W, 
VIT-29-1972. 


Callibaetis ferrugineus (Walsh) 

New records.—FAIRBANKS NORTH 
STAR: Mile 27, Steece Hwy, N U Alaska, 
X-2,8-1962. 

Callibaetis fluctuans (Walsh) 

New records.—FAIRBANKS NORTH 
STAR: Mile 27, Steece Hwy, N U Alaska, 
X-8-1962. 

Diphetor hageni (Eaton) 

Previous records.—Duncan and Brusven 
(1985): PRINCE OF WALES-OUTER 
KETCHIKAN. 

Plauditus dubius (Walsh) 

New records.—YUKON-KOYUKUK: 
Beaver Cr, 66/07N 146/15W, VII-21-1962. 
Procloeon pennulatum (Eaton) 

New records.—YUKON-KOYUKUK: 
Birch Cr between Big Cr & Preacher’s Cr, 
66/00N 144/S50W, VIII-17-1962. 

CAENIDAE 
Brachycercus harrisella Curtis 

New records.—YUKON-KOYUKUK: 
Birch Cr between Big Cr & Preacher’s Cr, 
VIII-17-1963. 

EPHEMERELLIDAE 
Caudatella jacobi (McDunnough) 

New records.—JUNEAU: Juneau, Fish 
Cr, VI-24-29-2000. 

Drunella coloradensis (Dodds) 


Previous records.—Allen and Edmunds 
(1962): JUNEAU; SKAGWAY-HOONAH- 


VOLUME 107, NUMBER 1 


ANGOON; WRANGLE-PETERSBURG; 
YUKON-KOYUKUK. 

New records.—HAINES: Klukshu R 
117.6 mi Haines Cutoff Hwy nr Alaska & 
BC border, VI-13-1932; SKAGWAY- 
HOONAH-ANGOON: North Arm Hood 
Bay, VI-19-1932; unidentified stream 0.5 
mi N Cape Fanshaw, VII-9-1958. 


Drunella doddsti (Needham) 


Previous records.—Allen and Edmunds 
(1962): ANCHORAGE; Wipfli et al. 
(1998): KETCHIKAN GATEWAY. 

New records.—JUNEAU: unidentified 
str, prob overflow water system, N Juneau, 
VI-21-1958: FAIRBANKS NORTH STAR: 
Mile 27, Steece Hwy, N U Alaska, X-8- 
1962; W Fk Chena R, VII-21-1979, AL 
Howe; Chatanika R Power Plant, mile 32- 
33, Steece Hwy, X-8-1962; KENAI PEN- 
INSULA: Diamond Cr nr Sterling, V-11- 
1968, E Charnov; Moose Cr at Tustamena 
L, VI-18-1968; Russian R (Lower), VII-1- 
1968; Russian R (Lower) above falls, VI- 
24-1968; Six Mile Cr, VII-22-1965; MA- 
TANUSKA-SUSITNA: Cottonwood Cr at 
Matanuska Rd 1 mi SE Wasilla, VI-19- 
1958; PRINCE OF WALES-OUTER KET- 
CHIKAN: unidentified str 1 mi N Game 
Cove in Hawk Inlet behind PE Cannery, 
Admirality Island, VII-13-1958; SITKA: 
Nakvisson Cr, Baranof Island, Port Herbert, 
VII-1-1958; WRANGLE-PETERSBURG: 
Anan Cr, [V-26-1958. 


Drunella grandis (Eaton) 


Previous records.—Ulmer (1932): 
ALEUTIANS EAST. Allen and Edmunds 
(1962): WRANGLE-PETERSBURG. 


Drunella spinifera (Needham) 
New records.—SKAGWAY-HOONAH- 


ANGOON: North Arm Hood Bay, VI-18- 
1970. 


Ephemerella aurivillii (Bengtsson) 


Previous records.—McDunnough (1924) 
as E. norda McDunnough: ALEUTIANS 
EAST. Allen and Edmunds (1965): WRAN- 


195 


GLE-PETERSBURG. Lee and Hershey 
(2000): NORTH SLOPE. 

New records.—FAIRBANKS NORTH 
STAR: North Frk Chena R, brdg 83, VI- 
11-1980 (adults); Mile 27 Steece Hwy, N 
U Alaska, X-8-1962; W Fork Chena R, VII- 
21-1979; KENAI PENINSULA: Kasilof R 
at Sterling Hwy brdg, V-21-1968, E Char- 
nov; Russian R (Lower), above falls, VI- 
24-1968; Russian R (Lower) near lake, VI- 
2-1968; Slicock Cr at Kalafonsky Rd brdg, 
V-30-1968. 


Ephemerella dorothea Needham 


Previous records.—Allen and Edmunds 
(1965), misidentified as £. inermis |Johnson 
(1978) as E. infrequens McDunnough]: 
FAIRBANKS-NORTH STAR; misidenti- 
fied as E. inermis [McCafferty (1994) as E. 
infrequens|: WRANGLE-PETERSBURG; 
YUKON-KOYUKUK. McCafferty (1985), 
misidentified as E. lacustris Allen and Ed- 
munds: NORTH SLOPE. 


Ephemerella nuda Tshernova 


New records.—YUKON-KOYUKUK: 
South Slope, Monument Cr | mi above 
confl Sheenjek R, 67/57/45N_ 143/13/00W, 
VII-29-1972; South Slope, Strangle Woman 
Cr 10 mi above mouth, 67/50/00N 141/40/ 
30W, VII-29-1972. 


Serratella tibialis (McDunnough) 

Previous records.—McCafferty (1985): 
NORTH SLOPE; Milner (1987): SKAG- 
WAY-HOONAH-ANGOON. 

New records.—PRINCE OF WALES- 
OUTER KETCHIKAN: Prince of Wales Is- 
land, Skowl Arm Inlet, Cabin Cr, VIII-14- 
1962; YUKON-KOYUKUK: South Slope, 
Sheenjek R 1 mi above pipeline crossing, 
67/37/45N 143/17/00W, VII-29-1972. 


HEPTAGENIIDAE 
Cinygma lyriforme (McDunnough) 


Previous records.—Lehmkuhl (1979): 
FAIRBANKS NORTH STAR; McCafferty 
(1985): NORTH SLOPE. 

New records.—MATANUSKA-SUSIT- 


196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


NA: Bonanza Cr, 12 mi N Old Man, VIII- 
4-1979 (adults). 


Cinygmula mimus (Eaton) 


New records.—KENAI PENINSULA: 
East Rd, Homer, VIII-26-1980 (adults); 
NORTH SLOPE: Prudhoe Bay to Can 
Pipeline, 1972 (adults); FAIRBANKS 
NORTH STAR: W Frk Chena R, VI-11- 
1980 (adults). 


Cinygmula par (Eaton) 


Previous records.—Harper and Harper 
(1981): NORTH SLOPE. 


Cinygmula subaequalis (Banks) 


Previous records.—McCafferty (1985): 
NORTH SLOPE. 

New records.—MATANUSKA-SUSIT- 
NA: Bonanza Cr, 12 mi N Old Man Cr, 
VIII-4-1979 (adults). 


Cinygmula tarda (McDunnough) 


New records.—YUKON-KOYUKUK: 
South Slope, unnamed E Frk trib Chandalar 
R, 68/25/00N 145/12/00W, VIII-13-1972 
(adults). 


Ecdyonurus simplicioides (McDunnough) 


New records.—YUKON-KOYUKUK: 
Birch Cr between Preacher Cr and Birch Cr 
Village, 66°30'’N 145°00'W, VIII-18-1962. 


Epeorus albertae (McDunnough) 


New records.—PRINCE OF WALES- 
OUTER KETCHIKAN: Prince of Wales Is- 
land, Skowl Arm Inlet, Virginia Cr, VII-29- 
1962. 


Epeorus deceptivus (McDunnough) 


New records.—FAIRBANKS NORTH 
STAR: Monument Cr, Chena R, [X-8-1972; 
PRINCE OF WALES-OUTER KETCHI- 
KAN: Prince of Wales Island, Skowl Arm 
Inlet, Virginia Cr, VII-29-1962; SKAG- 
WAY-HOONAH-ANGOON: unidentified 
stream 0.5 mi N Cape Fanshaw, mainland 
VII-9-1958; YUKON-KOYUKUK: Birch 


Cr at confl with Crooked Cr & Acme Cr, 
VIII-8-1973 (adults). 


Epeorus grandis (McDunnough) 


Previous records.—Wipfli et al. (1998): 
KETCHIKAN GATEWAY. 

New records.—ANCHORAGE: Aleyska, 
VI-21-1997; Anchorage, Campbell Cr, IV- 
21-1948; Chugach St Prk 18 mi from Old 
Seward, VI-22-1997; KENAI PENINSU- 
LA: Diamond Cr nr Sterling Hwy, V-11- 
1968; main trib lower Russian R, VI-24- 
1968; Moose Cr at Tustamena L, VI-18- 
1968; Russian R (Lower), VII-1-1968; tor- 
rential trib Lower Russian Lake, VI-23— 
1968. 


Epeorus longimanus (Eaton) 


Previous records.—Edmunds et al. 
(1976): no data provided. Wipfli et al. 
(1998): KETCHIKAN GATEWAY. 

New records.—KENAI PENINSULA: N 
Arm Hood Bay, VI-21-1969; PRINCE OF 
WALES-OUTER KETCHIKAN: Prince of 
Wales Island, Skowl Arm Inlet, Cabin Cr, 
VIII-14-1962; SITKA: Nakvissen Cr, Ba- 
ranof Island, Port Herbert VII-1-1958. 


Heptagenia pulla (Clemens) 


New records.—KENAI PENINSULA: 
Slicock Cr at Kalafonsky Rd brdg, V-30- 
1968; MATANUSKA-SUSITNA: Cotton- 
wood Cr at Matanusak Rd 1 mi SE Wasilla, 
VI-19-1958. 


Ironodes nitidus (Eaton) 


New records.—PRINCE OF WALES- 
OUTER KETHICKAN: Virginia Cr, VI-21- 
1963. 


Rhithrogena futilis McDunnough 


Previous records.—Harper and Harper 
(1981): NORTH SLOPE. 

New records.—FAIRBANKS NORTH 
STAR: Mile 69, Steese Hwy, Chatanika R, 
NE Fairbanks, VIII-7-1967 (adults). 


VOLUME 107, NUMBER 1 


5 


197 


Bee: 


4 


Nn 


Figs. 2-6. Rhithrogena male genitalia. 2, R. ingalik (ventral). 3, R. ingalik (dorsal). 4, R. ingalik titillators. 
5, R. baikovae (ventral) [after Sowa (1973)]. 6, R. lepnevae titillators [after Sowa (1973)]. 


Rhithrogena ingalik Randolph and 
McCafferty, new species 
(Figs. 2—4) 


Male adult.—Body Length: 7.6 mm; 
forewing length: 7.0—7.3 mm; hindwing 
length: 2.3-2.6 mm. Head: Color gray 
brown. Compound eyes purple, separated 
dorsally by less than diameter of median 
ocellus. Thorax: Prothorax light brown. 
Meso- and metathorax olive brown. Fore- 


wing with stigma clouded with white and 
with no anastomosed veins. Legs missing. 
Abdomen: Color generally light gray, with 
segments 8—10 gray brown, otherwise lack- 
ing distinct color pattern. Subgenital plate 
(Fig. 2) with relatively broad, shallow 
emargination; posterolateral lobes extend- 
ing posteriorly to level subequal to level 
distal margin of basal forceps segment. Pe- 
nes (Figs. 2—3) broadly rounded distally, 
extending posteriorly beyond base of me- 


198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


dially directed spine; titillators (Fig. 4) mi- 
nute, strongly narrowing to single pointed 
apex. Caudal filaments missing. 

Material examined.—Holotype: Male 
adult, Alaska, YUKON-KOYUKUK: Birch 
Creek 10 mi upstream from mile 147, Stee- 
ce Hwy, VII-11—1973 (deposited in the 
Purdue Entomological Research Collec- 
tion). Paratype: Male adult, same data and 
deposition as holotype [genitalia on slide 
(medium: euparol)]. 

Etymology.—The specific epithet is a 
noun in apposition in honor of a group of 
Native Americans known as the Ingalik 
who once inhabited the interior of Alaska. 

Remarks.—Rhithrogena ingalik is easily 
distinguished from all known North Amer- 
ican Rhithrogena by the unique shape of 
the male genitalia (Figs. 2—3), which are 
most similar to those of the far eastern Rus- 
sian species R. baikovae Sowa and R. lep- 
nevae Brodsky. The new species is differ- 
entiated from the latter two by the follow- 
ing: The titillators (best seen slide-mount- 
ed) of R. lepnevae are spatulate and have 
two or more teeth apically (Fig. 6). The ti- 
tillators of R. ingalik (Fig. 4) and R. bai- 
kovae [fig. 20 (Sowa 1973)] instead narrow 
to a single point apically. The posterior 
margin of the penes lobes of R. ingalik is 
broadly rounded and extends beyond the 
basal level of the medially directed spine 
(Fig. 3), whereas the posterior margin of the 
penes lobes of R. baikovae 1s relatively 
straight and does not extend beyond the 
base of the medially directed spine (Fig. 5). 
The subgenital plate emargination appears 
broad and shallow in R. ingalik (Fig. 2), and 
by comparison somewhat narrower and 
deeper in R. baikovae (Fig. 5). Also, R. bai- 
kovae adults have distinct abdominal pat- 
terning [figs. 5d,e (Sinitshenkova 1982 as 
R. quadrinotata Sinitshenkova)] that is not 
apparent in our specimens of R. ingalik. 


LEPTOPHLEBIIDAE 


Paraleptophlebia debilis (Walker) 


New records.—PRINCE OF WALES- 
OUTER KETCHIKAN: Old Tom Cr, 


Skowl Arm Inlet, Prince of Wales Island, 
VIII-3-1962: Cabin Cr, Prince of Wales Is- 
land, Skowl Arm Inlet, VIII-14-1962. 


Paraleptophlebia vaciva (Eaton) 


New records.—PRINCE OF WALES- 
OUTER KETCHIKAN: Cabin Cr, Prince of 
Wales Island, Skowl Arm, VIII-14-1962; 
SKAGWAY-HOONAH-ANGOON: N Arm 
Hood Bay, VII-15-1970. 


METRETOPODIDAE 
Metretopus alter Bengtsson 


Previous records.—Berner (1978) mis- 
identified as M. borealis (McCafferty 
1994): YUKON-KOYUKUK. 

New records.—NORTH SLOPE: Kavik 
R, Weir Cr, VIII-4-1973. 


SIPHLONURIDAE 
Parameletus chelifer Bengtsson 


Previous records.—Harper and Harper 
(1981): NORTH SLOPE. 


Siphlonurus alternatus (Say) 


New records.—NORTH SLOPE: un- 
named Canning R Spr above Aufies, 69/22/ 
45N 146/07/30W, VII-8-1972. 


Siphlonurus occidentalis (Eaton) 


New records.—FAIRBANKS NORTH 
STAR: Mile 27, Steece Hwy, N U Alaska, 
IX-20, X-8-1962. 


ACKNOWLEDGMENTS 


We thank George Edmunds (Salt Lake 
City, UT) and Luke Jacobus (West Lafay- 
ette, IN) for their contributions to this 
study. The research was supported by NSF 
Grant DEB-9901577 to WPM. 


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PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 200-208 


A REVIEW OF THE METACHARIS SYLOES GROUP (LEPIDOPTERA: 
RIODINIDAE), WITH THE DESCRIPTION OF TWO NEW SPECIES FROM 
WEST OF THE ANDES 


JASON P. W. HALL 


Department of Entomology, National Museum of Natural History, Smithsonian Insti- 
tution, Washington, DC 20560-0127, USA 


Abstract.—On the basis of male genitalia and wing pattern, the Neotropical riodinid 
genus Metacharis Butler is tentatively divided into four species groups, whose phyloge- 
netic inter-relationships are briefly discussed. With this taxonomic framework established, 
the derived syloes group is here characterized and taxonomically elaborated. It contains 
one described species, M. syloes Hewitson, and two that are described here, M. fergusi, 
n. sp., and M. smalli, n. sp. These are the only Metacharis species to exclusively inhabit 
montane forest, and they are distributed allopatrically in the eastern Andes, western Andes, 
and mountains of central and eastern Panama, respectively. The elevation of M. umbrata 


Stichel to species status (n. stat.) is also discussed. 


Key Words: 


The Neotropical riodinid genus Meta- 
charis Butler (Riodininae: Riodinini) con- 
tains a minimum of eight described species 
that are distributed from Nicaragua to 
southeastern Brazil. As conceived by early 
authors (Bates 1868; Stichel 1910-11, 
1930-31; Rebillard 1958; Bridges 1994; 
d’Abrera 1994), the genus was polyphylet- 
ic, and the misplaced exigua Bates, elinas 
Rebillard and chia Hiibner were only re- 
cently transferred to Symmachia Hiibner 
(Riodininae: Symmachiini) and Cariomoth- 
is Stichel (Riodininae: Riodinini) (Calla- 
ghan 1995, Hall and Harvey 2002, Calla- 
ghan and Lamas, in press). As treated by 
Callaghan and Lamas (in press), Metachar- 
is is a monophyletic group. Its species are 
characterized by having elongate wings that 
are shades of brown or iridescent blue with 
silver vein endings dorsally and four spots 
in the discal cells, male genitalia with a 
posteriorly elongate, simple (G.e. without a 
scobinate patch at the tip), “‘rod’’-like ped- 


allopatry, Andes, Ecuador, Metacharis, montane forest, Panama 


icel and a spine-tipped outer upper valve 
process, and female genitalia with a hollow, 
posteriorly projecting, trapezoid-shaped os- 
tium bursae and a ductus bursae that makes 
an asymmetrically displaced exit to the 
right (Hall 2001). Dachetola Hall has been 
suggested as the closest relative of Meta- 
charis (Hall 2001). 

Having examined the male genitalia of 
all Metacharis species, some preliminary 
phylogenetic conclusions can be drawn, 
which indicate that four main species 
groups can be recognized. Only M. ptolo- 
maeus (Fabricius) (agrius Dalman is prob- 
ably conspecific) and M. lucius (Fabricius) 
lack a long, narrow, lower valve process 
that parallels the “‘rod’’-like pedicel, and, as 
their males have more wing pattern ele- 
ments visible than those of their congeners, 
I hypothesize that these two species may be 
the most basal in the genus. Of the remain- 
ing species, M. victrix (Hewitson), M. um- 
brata Stichel (n. stat., see Note on Status 


VOLUME 107, NUMBER 1 


of Metacharis umbrata at the end of the 
paper), and M. xanthocraspedum Stichel 
have setae at the tip of the inner upper valve 
process and a curved pedicel, as in the two 
putative basal species, and M. regalis But- 
ler, M. nigrella Bates (cuparina Bates is 
probably conspecific), M. syloes Hewitson, 
and two undescribed species have stout 
spines at the tip of the inner upper valve 
process (like those on the outer upper valve 
process of all Metacharis species) and a 
horizontal pedicel that abruptly curves up- 
ward at the tip only. Of the last five species, 
the lowland Amazonian M. regalis and M. 
nigrella have sparsely distributed spines 
along the distal half of the outer upper 
valve process, whereas M. syloes and the 
two undescribed species, referred to here as 
the syloes group, have one or two, small, 
dense patches of spines on this valve pro- 
cess. Metacharis syloes group species are 
also unique within the genus in having 
males with plain brown wings dorsally, and 
in exclusively occurring in montane forest 
habitats. The widespread, east Andean M. 
syloes is the only described species of the 
group, and the two other group members, 
which occupy smaller allopatric ranges to 
the west of the Andes, are described here, 
as part of a review of the syloes group. 

It is biogeographically and evolutionarily 
interesting that most Metacharis species ap- 
pear to be allo- or parapatrically distributed 
with respect to the other members of their 
species group. In fact, all are if the distinc- 
tive M. xanthocraspedum is considered to 
be its own group. Perhaps noteworthy is the 
fact that all species in the possible sister 
genus, Dachetola, are also allo- or para- 
patrically distributed (Hall 2001). 

The following collection acronyms are 
used throughout the text: BMNH—The 
Natural History Museum, London, UK; 
JHKW—Collection of Jason P. W. Hall & 
Keith R. Willmott, Washington, DC, USA; 
USNM—National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
DESUSE: 


201 


REVIEW OF METACHARIS SYLOES GROUP 


Metacharis syloes Hewitson 1877 
(Figs. 1A, B; 2A; 3A; 4) 


Metacharis syloes Hewitson 1877: 96. Type 
locality: ““Gima”’, E. Ecuador. Syntype 
female BMNH [examined]. 


Identification and taxonomy.—Typical 
forewing length: male 24 mm; female 23 
mm. Although the male of M. syloes is vir- 
tually indistinguishable from that of M. fer- 
gusi on the basis of wing pattern, M. syloes 
actually appears to be sister to the allopatric 
west Andean clade of M. fergusi + M. 
smalli (both described below). Male M. sy- 
loes differs externally from male M. fergusi 
only by having a prominently distally dis- 
placed upper element to the black postdis- 
cal spot in cell Cu, on both wings (even 
more apparent in females), and generally 
more prominent paler submarginal blue on 
both ventral wings. However, on the basis 
of examining two dissections of each spe- 
cies, the male genitalia of M. syloes differ 
obviously from those of M. fergusi (and M. 
smalli) by having a lower valve process that 
is just shorter instead of considerably lon- 
ger than the pedicel, and an outer upper 
valve process that is just longer instead of 
very considerably longer than the pedicel 
(the inner upper valve process is also pro- 
portionately shorter), only weakly instead 
of very prominently bent downwards me- 
dially, and lacks a small spine-tipped pro- 
trusion at its inner middle. 

Female M. syloes differs externally from 
females of M. fergusi and M. smalli by hav- 
ing a more uniform, darker dorsal surface, 
and a ventral surface that is entirely brown, 
except for some submarginal yellow spots, 
instead of various shades of yellow 
throughout. The female genitalia of M. sy- 
loes differ from those of M. fergusi and M. 
smalli by having an ostium bursae that is 
broader, and has a square instead of trian- 
gular posterior tip and more anteriorly ex- 
tensive ventral sclerotization, and a poste- 
rior exit to the ductus bursae that is only 


202 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


E 


Fig. 1. 


Metacharis adults (dorsal surface on left, ventral surface on right). A, d M. syloes, La Bonita, E. 


Ecuador (JHKW). B, 2 M. syloes, Rio Negro, E. Colombia (USNM). C, Paratype d M. fergusi, Rio Chuchuvi, 
W. Ecuador (JHKW). D, Holotype 2 M. fergusi, Alluriquin, W. Ecuador (USNM). E, Holotype ¢ M. smalli, 
Cerro Campana, C. Panama (USNM). E Paratype 2 M. smalli, Cerro Campana, C. Panama (USNM),. 


slightly instead of very prominently dis- 
placed to the right. 

Note that the type locality for M. syloes 
is inaccurate. Gima is a small, remote vil- 
lage in southeastern Ecuador, lying at an el- 
evation of about 3,000 m, that Clarence 
Buckley apparently used as a base of op- 
erations. However, the material that he 
amassed in this region, and sent to Hewit- 
son for description (1877), was clearly col- 
lected between 1,000 and 2,000 m, presum- 
ably on the trail between Gima and the low- 
land town of Gualaquiza. 


Biology.—This species is uncommon in 
montane forest habitats from 950 to 1,550 
m. In Ecuador, males were encountered 
perching as solitary individuals or in small 
groups along streams and sun-dappled for- 
est paths across the hillsides above them, in 
the early morning and then again in the af- 
ternoon from 1330 to 1530 h. They made 
rapid circling sorties before returning to 
perch on the same clump of bushes from 2 
to 4 m above the ground, landing on top of 
and beneath leaves with their wings out- 
spread. During the late afternoon, their dark 


VOLUME 107, NUMBER 1 


coloration and erratic flight-path make them 
difficult to track in the air. The rarer females 
were found flying along forest trails and 
edges throughout the middle of the day. 
Distribution.—Metacharis syloes ranges 
from northern Venezuela along the eastern 
slope of the Andes to Bolivia (see Fig. 4). 


Metacharis fergusi Hall, new species 
disse CD] 2323824) 


Description.—Male: Forewing length 22 
mm. Forewing elongate, costal and distal 
margins approximately straight; hindwing 
rounded, with slightly pointed apex and tor- 
nus. Dorsal surface: Ground color of both 
wings dark brown; four black marks in dis- 
cal cell of both wings, one at base of cell 
Cu,, and three toward base of cell Cu, (dis- 
tal-most mark on forewing medially divid- 
ed), two additional black marks toward base 
of cell Sc+R, on hindwing; a disjointed 
black postdiscal band on both wings ex- 
tends from vein 2A to costa, and is distally 
displaced at vein Cu, and again at vein M,, 
especially on forewing; black submarginal 
spots on both wings surrounded by broad 
and indistinct area of russet brown scaling, 
with semicircles of dark brown immediately 
proximally, and 3 mm long intervening sil- 
ver stripes along veins between tornus and 
apex; forewing fringe brown except for 
white scaling in cell R,,;, and hindwing 
fringe brown except for white scaling in 
cell M,. Ventral surface: Ground color of 
both wings iridescent dark purple, with a 
narrow area of paler, non-iridescent purple 
at distal margin of forewing and distal and 
anal margins of hindwing, and gray scaling 
along anal margin of forewing; black mark- 
ings from dorsal surface largely obscured 
and prominently visible only at base of both 
wings and in distal half of hindwing, distal 
vein-ending silver stripes absent. 

Head: Labial palpi dark brown; eyes 
brown, surrounding scaling dark brown, 
frons dark brown, becoming paler ventrally; 
antennal segments black with white scaling 
at base, clubs black. 

Body: Dorsal and ventral surface of tho- 


203 


rax and abdomen dark brown; tegula dark 
brown; all legs dark brown. 

Genitalia (Fig. 2B): Uncus short and 
rectangular, with a diagonal margin along 
ventral posterior corner; falces of average 
size and shape for family; small, deep, 
semicircular notch in anterior margin of 
tegumen; narrow and ribbon-like vinculum 
extends dorsally over anterior portion of 
tegumen, saccus absent; valvae consist of a 
narrow and curved basal ribbon, connected 
ventrally to a long, straight and narrow low- 
er process, and dorsally to a medial, pos- 
teriorly elongate and triangular transtilla 
flanked laterally by two more processes, the 
inner process long, straight, narrow and 
tipped with short stout spines, and the outer 
process slightly longer and considerably 
broader than the inner process, prominently 
bent downwards medially and slightly in- 
ward overall, and with inwardly pointing 
spines at its tip and on a small medial pro- 
cess (which variably extends dorsally to 
nearly inwardly horizontal); aedeagus long, 
narrow and convex, with a tapering tip that 
opens to right, and a base that has soft tis- 
sue exiting directly anteriorly, cornuti ab- 
sent; pedicel extends from near base of ae- 
deagus to form a posteriorly elongate, 
smooth ‘‘rod”’ that is encircled by sclero- 
tized tissue around distal fifth only, and is 
slightly upturned at tip, which is weakly bi- 
furcate in ventral view. 

Female: Differs from male as follows: 
Forewing length 21 mm. Forewing distal 
margin convex; hindwing more rounded. 
Dorsal surface: Ground color of both 
wings brown, with a darker brown submar- 
ginal area, and a continuous, bright russet 
brown marginal band. Ventral surface: 
Ground color of both wings dark yellow, 
with an uneven, darker, tan colored sub- 
marginal area; all black spotting on both 
wings prominent, submarginal spots sur- 
rounded by large yellow rectangles. 

Head: Labial palpi dirty white; frons 
brown in dorsal half, dirty white in ventral 
half. 

Body: Dorsal surface of thorax and ab- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. 


domen pale brown, ventral surface dirty 
white; tegula pale brown; all legs dirty 
white. 

Genitalia (Fig. 3B): Corpus bursae ap- 
proximately round, signa short spine-like 
invaginations with rounded tips; membra- 
nous ductus bursae exits to right side of os- 
tium bursae, membranous ductus seminalis 
exits ductus bursae ventrally, immediately 
before ostium bursae; ostium bursae con- 
sists of a narrow, detached, sclerotized band 
along anterior margin of last sternite, and a 
hollow, posteriorly projecting, trapezoid- 
shaped sclerotized structure whose base 
only is attached to body. 

Type material.—Holotype 2, ECUA- 
DOR: Pichincha, nr. Alluriquin, 750 m, 28 
Aug (S. S. Nicolay) (USNM). 

Paratypes: ECUADOR: 1 6: Esmeral- 
das, Rio Piguambi, km. 7.5 Lita-San Lor- 


Metacharis male genitalia in lateral view. A, M. syloes. B, M. fergusi. C, M. smalli. 


enzo rd., 0°52.42'N, 78°29.55’W, 800 m, 19 
June (J. BR W. Hall) (JHKW). 1 3: Esmer- 
aldas, Rio Chuchuvi, km. 12.5 Lita-San 
Lorenzo rd., 0°53.01'N, 78°30.90’W, 900 m 
(I. Aldas) (SHKW). 1 2: Carchi, nr. Lita, 
ridge to east of Rio Baboso, 950 m, 26 Aug 
(K. R. Willmott) (JHKW). 
Etymology.—This species is named for 
my brother Fergus Monahan Hall. 
Diagnosis.—The male of M. fergusi is 
very similar to that of M. syloes, and the 
female is very similar to that of M. smalli, 
from which species it is distinguished in 
those species accounts. By the shared pos- 
session of male genitalia with a small, spine 
covered protrusion at the inner middle of 
the outer upper valve process, a unique 
character within Metacharis, M. fergusi can 
be placed as the sister species to M. smalli. 
Biology.—This uncommon species is 


VOLUME 107, NUMBER 1 


confined to premontane forest habitats from 
750 m to at least 950 m. In Ecuador, soli- 
tary males were encountered perching both 
on low secondary growth vegetation along 
streamsides during the mid to late after- 
noon, and 5 m above the ground in a hilltop 
forest lightgap from 1145 to 1530 h. They 
perched on top of and beneath leaves with 
their wings spread open. A female was col- 
lected flying at the forest edge along a ridg- 
etop during mid-day. 

Distribution.—Metacharis fergusi is cur- 
rently known only from the west Andean 
slope of Ecuador, but is presumably endem- 
ic to the Choc6 of western Ecuador and 
western Colombia (see Fig. 4). It is re- 
placed in the mountains of central and east- 
ern Panama by M. smalli, and along the 
eastern Andes by M. syloes. 


Metacharis smalli Hall, new species 
(igs EE 2 C374) 


Description.—Male: Forewing length 21 
mm. Forewing elongate, costal and distal 
Margins approximately straight; hindwing 
rounded, with slightly pointed apex and tor- 
nus. Dorsal surface: Same as M. fergusi, 
except ground color slightly paler brown. 
Ventral surface: Differs from dorsal sur- 
face as follows: Ground color of both wings 
rich russet brown, with gray scaling along 
anal margins; black markings on forewing 
slightly fainter, contrasted scaling surround- 
ing all submarginal spots except that in cell 
R,.; on forewing, and intervening silver 
stripes, absent. 

Head: Labial palpi brown; eyes brown, 
surrounding scaling brown; frons brown, 
becoming paler ventrally; antennal seg- 
ments black with white scaling at base, 
clubs black, tips orange-brown. 

Body: Dorsal surface of thorax and ab- 
domen dark brown, ventral surface paler 
brown; tegula brown; all legs brown. 

Genitalia (Fig. 2C): Same as M. fergusi, 
except valve processes slightly shorter. 

Female: Differs from male as follows: 
Forewing length 20 mm. Forewing distal 


205 
7) 
N A f 
B \\ LA\ /) 
\ \ / / 
\ fas \ \ / 
\ \ Gl SX fff 
Cn lie Ay 
ASE a / 
\ SSS 
Asque | 
iar! 1 mm 
/ | 
| VA \ 
| i | 
| / A 
| 4a \ SS 
Wy Ww 
aN | 
| y 
| Ye 
( IN 
}} \ 
\ x : 
\ \) 
\ 
\ 
/ 
\ 
\ 
\ VL 
\ ae 


Fig. 3. Metacharis female genitalia in dorsal view, 
either in their entirety (B), or with signum and ostium 
bursae only (A and C). A, M. syloes. B, M. fergusi. C, 
M. smalli. 


206 


margin convex; hindwing more rounded. 
Dorsal surface: Ground color of both 
wings a pale, slightly rufous brown, with 
an uneven, darker brown submarginal area; 
russet brown along distal margin of both 
wings slightly paler and more prominent. 
Ventral surface: Ground color of both 
wings dark yellow, with an uneven, darker, 
tan colored submarginal area; black sub- 
marginal spots prominent on both wings 
and surrounded by an elongate yellow area. 

Head: Labial palpi dirty white; frons 
brown in dorsal half, dirty white in ventral 
half. 

Body: Dorsal surface of thorax and ab- 
domen pale brown, ventral surface dirty 
white; tegula pale brown; all legs dirty 
white. 

Genitalia (Fig. 3C): Same as M. fergusi. 

Type material—Holotype ¢6, PANAMA: 
Panamd, Cerro Campana, 2,500 ft, 3 Sept 
(G. B. Small) (USNM). 

Paratypes: PANAMA: Panama, Cerro 
Campa, 2SC0 me I os WS Iwi Wl Ge 29 
July (G. B. Small) (USNM). Panama, Altos 
de Pacora, 2,000 ft, 1 d: 29 Mar; 1 d: 18 
Mar (G. B. Small) (USNM). 1 6d: Darién, 
Canay 750 aman Zon une (Gee Be smalll)) 
(USNM). 

Etymology.—As Gordon Small collected 
the entire type series, it seems only fitting 
that this species should be named after him. 

Diagnosis.—The male of M. smalli is 
readily distinguished from that of the sister 
species M. fergusi by having a rich russet 
brown instead of iridescent purple ventral 
surface. Ventral wing coloration in Meta- 
charis is clearly evolutionarily labile, as ev- 
idenced by the multiple gain or loss (the 
ancestral state is not known with any degree 
of certainty) of ventral iridescence in the 
genus, a character that recurs in every spe- 
cies group. The apparently parapatrically 
distributed, Amazonian sister species pair 
of M. regalis and M. nigrella also primarily 
differ from each other by having russet 
brown and iridescent purple ventral surfac- 
es, respectively. Having examined the gen- 
italia of two males of both M. smalli and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


100 200km 


4000m N 


3000m 
2000m 
1000m 


sea level 


PERU 


— @ Metacharis smalli 
| HE Meracharis fergusi 


Fig. 4. Topographic map of northwestern South 
America indicating the geographic distributions of Me- 
tacharis syloes group species. 


M. fergusi, the only consistent difference 
seems to be slightly shorter valve processes 
in M. smalli. 

The females of M. smalli and M. fergusi 
are very similar, but M. smalli has a more 
uniformly paler brown dorsal surface, less 
contrasting shades of yellow on the ventral 
surface, and submarginal yellow occeli on 
both ventral wings that are clearly separated 
instead of formed into a continuous band. 
This last difference is also present on the 
dorsal surface, but is less pronounced. 
There are no significant differences in the 
female genitalia of the two species. 

Biology.—Nothing is known about the 
biology of this species, except that it 1s con- 
fined to premontane forest habitats from 
about 650 m to at least 800 m. 

Distribution.—Metacharis smalli appears 
to be endemic to the mountains of central 
and eastern Panama (see Fig. 4). As the re- 


VOLUME 107, NUMBER 1 


mote Serrania de Darién and Cordillera de 
Quia straddle the Panama-Colombia border, 
it should also eventually be found in the 
extreme north of Choco province, north- 
western Colombia. It is replaced by M. fer- 
gusi along the western slope of the northern 
Andes. 


NOTE ON THE STATUS OF METACHARIS 
UMBRATA 


Stichel proposed (1929) the name um- 
brata as a form of Metacharis cuparina, 
based on a single female from Rio Micay, 
western Colombia (holotype examined in 
the Zoologische Museum fiir Naturkunde, 
Berlin, Germany), and the name was re- 
cently synonymized with M. victrix by Cal- 
laghan and Lamas (in press). This is essen- 
tially an identical nomenclatural case to 
Sarota lasciva Stichel (Riodininae: Helico- 
pini) (resolved in Hall 1998), with the con- 
tent of Stichel’s 1929 description not “‘un- 
ambiguously reveal[ing] that the name was 
proposed for an infrasubspecific entity” 
(Article 45.6.4, ICZN 1999). I therefore re- 
gard umbrata as an available name and 
raise it to the rank of species. It is the sister 
species of M. victrix, and is macrosympatric 
with it from Costa Rica to western Ecuador. 
However, the two species appear not to fly 
in the same place, with M. umbrata restrict- 
ed to wet forest and M. victrix confined to 
drier forest habitats (Hall and Willmott, un- 
published data). Their male genitalia, at 
least, do not differ significantly, but the 
male of M. umbrata is readily distinguished 
from that of M. victrix by its iridescent pur- 
ple, instead of rufous brown, dorsal surface, 
and the female is best separated by the 
broad blackish area in its forewing apex. 
DeVries (1997) figured a male of M. um- 
brata and gave it that name in his text based 
on information provided by myself, but I 
was not credited for this novel identification 
and the name on his addenda plate was un- 
fortunately misspelled “‘onorata.”’ 


ACKNOWLEDGMENTS 


I thank Phillip Ackery for access to, and 
Keith Willmott for subsequently relaying 


207 


certain specimen data from, the BMNH 
riodinid collections; Keith Willmott for al- 
lowing me to use the base map in Fig. 4; 
The National Geographic Society (Research 
and Exploration Grant # 5751-96) and The 
National Science Foundation (Biodiversity 
Surveys & Inventories Grant # 0103746) 
for financial support; Gerardo Lamas for 
helpful comments on the manuscript; and 
the Museo Nacional de Ciencias Naturales 
and the Ministerio del Ambiente (formerly 
INEFAN), in Quito, for arranging the nec- 
essary permits for research in Ecuador. 


LITERATURE CITED 


Bates, H. W. 1868. A catalogue of Erycinidae, a family 
of diurnal Lepidoptera. Journal of the Linnean So- 
ciety (London) (Zoology) 9: 373-459. 

Bridges, C. A. 1994. Catalogue of the Family-Group, 
Genus-Group and Species-Group Names of the 
Riodinidae and Lycaenidae (Lepidoptera) of the 
World. C. Bridges, Urbana, Illinois, 1,113 pp. 

Callaghan, C. J. 1995. Les types des Riodinidae du 
Muséum national d’Histoire naturelle de Paris 
(Lepidoptera, Rhopalocera). Bulletin de la Société 
Entomologique de France 100(2): 153-155. 

Callaghan, C. J. and G. Lamas. In press. Riodinidae. 
In Lamas, G., ed. Checklist: Part 4A. Hesperioi- 
dea—Papilionoidea. /n Heppner, J. B., ed. Atlas 
of Neotropical Lepidoptera. Scientific Publishers, 
Gainesville. 

d’Abrera, B. 1994. Butterflies of the Neotropical Re- 
gion, Part VI. Riodinidae. Hill House, Victoria, 
Australia. Pp. 880—1096. 

DeVries, P. J. 1997. The Butterflies of Costa Rica and 
Their Natural History, Vol. II. Riodinidae. Prince- 
ton University Press, Princeton, 288 pp. 

Hall, J. P. W. 1998. A review of the genus Sarota (Lep- 
idoptera: Riodinidae), pp. 1-21. /n Hall, J. P. W., 
ed. A Contribution to Riodinid Systematics. Trop- 
ical Lepidoptera, 9(2), Supplement 1. 

. 2001. A revision of the new riodinid butterfly 
genus Dachetola (Lepidoptera: Riodinidae: Riod- 
inini). Journal of the New York Entomological So- 
ciety 109(2): 183-195. 

Hall, J. RB W. and D. J. Harvey. 2002. A survey of 
androconial organs in the Riodinidae (Lepidop- 


tera). Zoological Journal of the Linnean Society 
136: 171-197. 

Hewitson, W. C. 1877. Equatorial Lepidoptera Col- 
lected by Mr. Buckley. 5: 81—96. J. Van Voorst, 
London. 

International Commission on Zoological Nomencla- 
ture. 1999. International Code of Zoological No- 
menclature. Fourth Edition. International Trust for 
Zoological Nomenclature, London, 306 pp. 


208 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 

. 1929. Beitraége zur kenntnis der Riodiniden- 

fauna Siidamerikas X. (Lep. Rhop.). Deutsche En- 

tomologische Zeitschrift 1929(3): 199-208. 

. 1930-31. Riodinidae, pp. 1-795. In Strand, 
ed. Lepidopterorum Catalogus 38-41. W. 


Rebillard, P 1958. Contribution a la connaissance des 
Riodinidae Sud-Américains. Mémoires du Musé- 
um d’Histoire Naturelle (A) 15: 135-216. 

Stichel, H. F E. J. 1910—11. Lepidoptera Rhopalocera. 
Fam. Riodinidae, pp. 1-452. In Wytsman, J., ed. Ee 
Genera Insectorum 112. J. Wytsman, Brussels. Junk, Berlin. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 209-213 


DESCRIPTION OF THE ADULT AND FIFTH INSTAR OF A 
MYRMECOMORPHIC PLANT BUG, BICUSPIDATIELLA CONICA 
MALDONADO (HEMIPTERA: MIRIDAE: DERAEOCORINAE), WITH NOTES 
ON ITS HABITS 


A.G. WHEELER, JR. AND THOMAS J. HENRY 


(AGW) Department of Entomology, Soils, and Plant Sciences, Clemson University, 
Clemson, SC 29634, U.S.A. (e-mail: awhlr@clemson.edu); (TJH) Systematic Entomology 
Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, % Na- 
tional Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0168, 
U.S.A. (e-mail: thenry @sel.barc.usda.gov) 


Abstract.—The deraeocorine plant bug Bicuspidatiella conica Maldonado, belonging to 
a monotypic genus in the tribe Hyaliodini, has been known only from five adults taken 
at three localities in Puerto Rico. Previous biological information consists of a record 
from guaba (nga vera Willd.). The fifth instar and adult of this myrmecomorphic mirid, 
collected at Cayey, Puerto Rico, are described and figured. The species was found on 
laurel amarillo (Nectandra turbacensis (Kunth) Nees) in association with the formicine 
ant Myrmelachista ramulorum Wheeler, the ant-attended mealybug Nipaecoccus nipae 
(Maskell) (Pseudococcidae), the soft scale Coccus moestus De Lotto (Coccidae), and an 
unidentified whitefly (Aleyrodidae). Bicuspidatiella conica, a member of an almost ex- 
clusively predacious group, probably feeds on ant-attended sternorrhynchans. 


Key Words: Heteroptera, myrmecomorphy, Puerto Rico, Sternorrhyncha, Nectandra tur- 


bacensis 


Myrmecomorphy, a morphological and 
behavioral resemblance to ants, has been 
described for more than 2,000 species of 
insects representing some 200 genera in 54 
families. Among the Arthropoda, the het- 
eropteran family Miridae contains the larg- 
est number of myrmecomorphic species 
(McIver and Stonedahl 1993). Ant-like mir- 
ids are most diverse in the Mirinae, espe- 
cially the tribe Herdoniini; the Orthotylinae, 
especially Ceratocapsini and Nichomachini; 
and the Phylinae, especially the Auricill- 
ocorini, Hallodapini, Leucophoropterini, 
and Pilophorini (Schuh 1974, 1991; McIver 
and Stonedahl 1993; Henry 1994). Myr- 
mecomorphy has arisen at least ten times in 
five mirid subfamilies, including the Der- 


aeocorinae (Schuh 1986, McIver and Sio- 
nedahl 1993). 

Maldonado (1969) described from Puerto 
Rico the new genus Bicuspidatiella in the 
deraeocorine tribe Hyaliodini and B. conica 
as the only included species. Type material, 
collected from 1914 to 1964, consisted of 
five specimens from three localities (Aibon- 
ito, Maricao, and the type locality between 
Afasco and Las Marias). Adults from the 
type locality were taken on guaba [as ““gua- 
va], Inga vera Willd. [Fabaceae]. Marto- 
rell (1976) pointed out that the common 
name Maldonado (1969) used for /. vera — 
‘“ouava’’— refers to the tropical fruit Psi- 
dium guajava L. and that guaba is the com- 
mon name for /. vera. Bicuspidatiella re- 


210 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mains a monotypic genus (Schuh 1995), 
and no additional records of B. conica have 
been published since Maldonado’s (1969) 
original description. 

Maldonado (1969) did not refer to B. 
conica aS a myrmecomorph, probably be- 
cause he had not observed nymphs. Here, 
we describe and figure the adult and fifth 
instar, and provide notes on the association 
of nymphs and adults with a formicine ant 
and ant-attended sternorrhynchans that 
might serve as prey of this presumed pred- 
atory plant bug. All specimens are depos- 
ited in National Museum of Natural Histo- 
ry, Smithsonian Institution, Washington, 
D.C. (USNM). 


Bicuspidatiella conica Maldonado 
(Figs. 1—4) 


Description of adult (Figs. 1—2).—Male 
(n=10): Elongate, subparallel, length 3.10— 
3.65 mm, width 0.85—0.86 mm. Head: 
Width 0.59—-0.62 mm, vertex 0.18—0.20 
mm; yellowish to reddish brown, wider 
than long; dorsal width of an eye subequal 
to width of vertex between eyes. Antenna: 
Segment I, length 0.29—0.33 mm; HI, 0.87— 
0.98 mm; III, 0.39—0.46 mm; IV, 0.29—0.31 
mm; pale yellowish brown. Labium: 
Length 0.78—0.82 mm, extending past pro- 
coxae to middle of mesosternum. Prono- 
tum: Length 0.65—0.72 mm, anterior width 
0.35—0.38 mm, posterior width 0.85—0.86 
mm; trapeziform; anterior lobe narrow, be- 
coming necklike, yellowish to reddish 
brown, impunctate; posterior lobe fuscous, 
more than twice as broad as anterior lobe, 
evenly punctate. Scutellum: Equilateral, 
yellowish to reddish brown; apex with a 
large, shiny conical tubercle (Fig. 2), uni- 
formly yellow except for fuscous anterior 
area. Hemelytra: Subparallel, largely hya- 
line, except for the yellowish brown clavus, 
and a narrow dark brown streak through 
middle and a larger dark brown area (pos- 
sessing a large conical tubercle [Fig. 2]) at 
apex of corium; apex of cuneus red to red- 
dish brown; membrane dark brown on basal 
two thirds, pale on apical third. Ventral sur- 


face: 


Thorax yellowish brown; abdomen 
yellowish brown basally, becoming dark 
brown to fuscous distally. Legs: Pale yel- 
lowish brown. Male genitalia (illustrated by 
Maldonado 1969): Left paramere long, 
slender, L-shaped; right paramere short, 
stout, apically blunt with a shallow notch; 
genital capsule with a blunt, dorsally di- 
rected tubercle; anal tube short but distinct. 

Female (n = 10): Length 2.94—3.17 mm, 
width 0.88—0.91 mm. Head: Width 0.59— 
0.60 mm, vertex 0.25—0.26 mm. Labium: 
Length 0.81—0.83 mm. Antenna: Segment 
I, length 0.26—0.29 mm; II, 0.72—0.83 mm; 
Ill, 0.39-0.44 mm; IV, 0.26—-0.33 mm. 
Pronotum: Length 0.68—0.69 mm, anterior 
width 0.36—0.39 mm, posterior width 0.86— 
0.87 mm. Similar to male in overall shape 
and markings. 

Description of fifth instar (Figs. 3—4) 
(n=3).—Length 2.30—2.50 mm. Colora- 
tion: Head, prothorax, abdominal segments 
II-VI, and a few fine lines and small marks 
on the meso- and metathorax deep reddish 
brown; antennae, legs, wing pads, most of 
meso- and metathorax, and abdominal seg- 
ment I-I and VI-IX pale brownish yellow. 
Structure: Head: Width 0.46—0.50 mm, 
vertex 0.27-0.32 mm, smooth, round, with 
a transverse suture between eyes and a lon- 
gitudinal suture along meson of vertex and 
frons, together forming a cross-shaped pat- 
tern; eyes relatively small, narrow, com- 
bined dorsal width about one half width of 
vertex between eyes. Labium relatively 
stout, extending to middle of mesosternum. 
Antenna: Segment I, 0.22—0.24 mm; I, 
0.64—0.68 mm; II, 0.36—-0.39 mm; IV, 
0.35—0.36 mm. Thorax: Prothorax rectan- 
gular, slightly wider than long; basal third 
of disc with two large, stout, basally con- 
nected spines (Fig. 4) strongly curved for- 
ward, each spine with a much shorter, stout, 
dorsal spinelike process, appearing 
branched or bifid; mesothorax giving rise to 
large wing pads extending to base of nar- 
rowed abdomen, with an erect, stout spine 
(Fig. 4) on inner basal margin of each wing 
pad; metathorax slender, with an erect, stout 


VOLUME 107, NUMBER | a1 


— —— 


3 i 


Figs. 1-5. 1-4, Bicuspidatiella conica. 1, Adult female, dorsal aspect. 2, Adult female, lateral aspect (arrows: 
a, conical scutellar tubercle; b, tubercle on corium). 3, Fifth instar, dorsal aspect. 4, Fifth instar, lateral aspect 


(arrows: a, spines on prothorax; b, spines on mesothorax; c, spines on metathorax; d, spines on abdominal 
segment I; e, anal tube). 5, Ant, Myrmelachista ramulorum, dorsal aspect. 


spine (Fig. 4) on each side. Abdomen: Bas- segments VII—VIII becoming more strongly 
al two segments slender, forming a narrow narrowed; segment IX or anal tube uni- 
waist, segment I with an erect spine (Fig. formly slender. Legs: Slender, unarmed. 

4) on either side of posterior margin; seg- Discussion.—Bicuspidatiella conica is 
ments III-VI broadly rounded or bulbous; morphologically most similar to members 


212 


of the hyaliodine genus Paracarnus in the 
overall slender body form, the hyaline hem- 
elytra with various brown markings, and 
short, distinct anal tube, but it is distin- 
guished from this genus and all other Der- 
aeocorinae in possessing a large conical tu- 
bercle on the scutellum and more stout tu- 
bercle on the apical third of each corium. 

Nymphs of B. conica are remarkably ant- 
like. The rounded head, quadrate thorax, 
and bulbous, basally narrowed abdomen, in 
combination with the reddish-brown head, 
prothorax, and abdomen contrasting with 
the overall pale coloration, contribute to 
their strong resemblance to the ant Myr- 
melachista ramulorum Wheeler (Fig. 5), 
with which the nymphs were associated. 
Unlike the nymphs of most other mirids, 
including other mimetic taxa, those of B. 
conica are unique in having eight stout 
spines on the thorax and abdomen. Only 
one other mirid is known to possess similar 
structures. China (1931) described the fifth 
instar of another hyaliodine, Paracarnus 
myersi China, from Cuba as having a broad 
multispined plate on the head, a large pro- 
cess on the prothorax diverging into four 
spines, and several other erect processes on 
the mesothorax and basal abdominal seg- 
ment, characteristics further suggesting the 
close relationship of these two genera. 

Field observations.—From 22 July to 2 
August 1996, we observed B. conica while 
in Puerto Rico to prepare the Hemiptera 
collection of the late Jenaro Maldonado Ca- 
priles for shipment to the USNM (Santiago- 
Blay et al. 1997). Nymphs and adults were 
found on two laurel amarillo trees, Nectan- 
dra turbacensis (Kunth) Nees [Lauraceae], 
planted as ornamentals in the front yard of 
Maldonado’s house in Cayey (Calle 6-I-1, 
Urbanizacion Aponte). This Neotropical 
tree grows in moist forests of Puerto Rico 
(Little et al. 1974). 

Nymphs and adults initially were beaten 
from branches of laurel amarillo. Exami- 
nation of branches about 4 m of the ground 
showed that the bugs were syntopic on low- 
er-leaf surfaces with a tiny (2.0—2.3 mm) 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ant, Myrmelachista ramulorum, whose 
workers were streaming up and down the 
trunks. This arboreal formicine nests in hol- 
low twigs, and workers move in long, con- 
spicuous files by following a trail phero- 
mone (Wheeler 1908, Blum and Wilson 
1964). Honeydew is obtained from aphids 
and scale insects that inhabit the trees on 
which the ants live (Wolcott 1933, Smith 
1936). On laurel amarillo, the ant was as- 
sociated with three sternorrhynchans that 
colonized the foliage: a soft scale, Coccus 
moestus De Lotto (Coccidae); a mealybug, 
Nipaecoccus nipae (Maskell) (Pseudococ- 
cidae); and a whitefly (Aleyrodidae). Be- 
cause we did not collect the whitefly, its 
identity remains in doubt. In Puerto Rico, 
one aleyrodid, Aleuroplatus vinsonioides 
(Cockerell), has been recorded from laurel 
amarillo (Martorell 1976). The mirid was 
not found across the street on laurel amar- 
illo trees that appeared to lack populations 
of the ant. Trunks of laurel amarillo on both 
sides of the street had been whitewashed at 
the base, which allowed detection of ant 
trails. 

We observed six adults of B. conica on 
the abaxial surface of a leaf with ants and 
whitefly larvae. On the lower surface of an- 
other leaf, an adult was seen atop (and pos- 
sibly feeding on) whiteflies and their hon- 
eydew. We observed an adult flying a short 
distance (ca. 20 cm) from one leaf to an- 
other. A short, hopping flight was charac- 
teristic of adults held in plastic vials. 

Nymphs were bicolored and difficult to 
distinguish from the similarly colored ants 
(Fig. 5). Two late instars were on the un- 
derside of a leaf with about 12 ants that 
were tending mealybugs. Another late in- 
star was found with ants and mealybugs on 
the lower surface of a curled leaf. On an- 
other leaf, we observed an ant touching and 
displacing a late-instar mirid that was near 
a soft scale. We observed a late instar ap- 
parently feeding on crystallized honeydew 
from the scale. 

Deraeocorine mirids are largely preda- 
cious, members of the Hyaliodini perhaps 


VOLUME 107, NUMBER 1 


exclusively so (Wheeler 2001). We hypoth- 
esize that the myrmecomorphic B. conica 
will be found on foliage of other trees that 
harbor small species of ants such as M. ra- 
mulorum and will be shown to feed mainly 
on ant-attended sternorrhynchans. At the 
type locality, Maldonado (1969) collected 
the mirid on guaba, a tree on which various 
scale insects, including the mealybug N. ni- 
pae, apparently are common (Smith 1936, 
Martorell 1976). This plant bug might also 
feed on honeydew produced by Sternor- 
rhyncha, as is known for other Miridae, in- 
cluding predacious species (Wheeler 2001). 


ACKNOWLEDGMENTS 


We thank Douglass R. Miller (Systematic 
Entomology Laboratory, ARS, USDA, 
Beltsville, Maryland [SEL]) for identifying 
the Coccoidea, David R. Smith (SEL, % 
Natl. Mus. Nat. Hist., Washington, D. C.) 
for identifying the formicid, and Michele 
Touchet (SEL) for the photographs. Peter 
Adler (Clemson University, Clemson, SC), 
David R. Smith, and Steven Lingafelter 
(SEL) kindly reviewed the manuscript. 


LITERATURE CITED 


Blum, M. S. and E. O. Wilson. 1964. The anatomical 
source of trail substances in formicine ants. Psy- 
che 71: 28-31. 

China, W. E. 1931. A remarkable mirid larva from 
Cuba, apparently belonging to a new species of 
the genus Paracarnus, Dist. (Hemiptera, Miridae). 
Annals and Magazine of Natural History Ser. 10, 
8: 284-288. 

Henry, T. J. 1994. Revision of the myrmecomorphic 
plant bug genus Schaffneria Knight (Heteroptera: 
Miridae: Orthotylinae). Proceedings of the Ento- 
mological Society of Washington 96: 701—712. 

Little, E. L, Jr, R. O. Woodbury, and EF H. Wadsworth. 


1974. Trees of Puerto Rico and the Virgin Islands, 
Second Volume. United States Department of Ag- 
riculture Forest Service, Agriculture Handbook 
449: 1-1024. 

Maldonado Capriles, J. 1969. The Miridae of Puerto 
Rico (Insecta, Hemiptera). University of Puerto 
Rico Mayagiiez Campus, Agricultural Experiment 
Station Technical Paper 45: 1-133. 

Martorell, L. E 1976. Annotated Food Plant Catalog 
of the Insects of Puerto Rico. University of Puerto 
Rico, Agricultural Experiment Station, Depart- 
ment of Entomology, Rio Piedras, 303 pp. 

Mclver, J. D. and G. Stonedahl. 1993. Myrmecomor- 
phy: Morphological and behavioral mimicry of 
ants. Annual Review of Entomology 38: 351-379. 

Santiago-Blay, J. A., T. J. Henry, and A. G. Wheeler, 
Jr. 1997. Jenaro Maldonado Capriles 1919-1995: 
Biographical sketch, patronyms in his honor, and 
list of publications. Proceedings of the Entomo- 
logical Society of Washington 99: 762-771. 

Schuh, R. T. 1974. The Orthotylinae and Phylinae (He- 
miptera: Miridae) of South Africa with a phylo- 
genetic analysis of the ant-mimetic tribes of the 
two subfamilies for the world. Entomologica 
Americana 47: 1—332. 

. 1986. The influence of cladistics on Heter- 

optera classification. Annual Review of Entomol- 

ogy 31: 67-94. 

. 1991. Phylogenetic, host and biogeographic 

analyses of Pilophorini (Heteroptera: Miridae: 

Phylinae). Cladistics 7: 157-189. 

. 1995. Plant Bugs of the World (Insecta: Het- 
eroptera: Miridae): Systematic Catalog, Distribu- 
tions, Host List, and Bibliography. New York En- 
tomological Society, New York, 1329 pp. 

Smith, M. R. 1936. The ants of Puerto Rico. Journal 
of Agriculture of the University of Puerto Rico 
20: 819-875. 

Wheeler, A. G., Jr. 2001. Biology of the Plant Bugs 
(Hemiptera: Miridae): Pests, Predators, Opportun- 
ists. Cornell University Press, Ithaca, New York, 
507 pp. 

Wheeler, W. M. 1908. The ants of Porto Rico and the 
Virgin Islands. Bulletin of the American Museum 
of Natural History 24: 117-158. 

Wolcott, G. N. 1933. An Economic Entomology of the 
West Indies. Entomological Society of Puerto 
Rico, San Juan, 688 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 214-217 


A NEW SAWELY (HYMENOPTERA: PERGIDAE) FEEDING ON GUAVA, 
PSIDIUM GUAJAVA L. (MYRTACEAE), IN COSTA RICA 


DAvID R. SMITH 


Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- 
ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, 


IR@, Iworxe SOW, IMEC 
dsmith @sel.barc.usda.gov) 


168, Washington, 


DC. 20013-70125 WUsS-AG  (eaniaile 


Abstract.—Sutwanus guajavae, n. sp., from Costa Rica is described and illustrated. 
Adults were reared from larvae feeding on Psidium guajava L. (Myrtaceae). A diagnosis 
is provided for distinguishing the new species from S. nigriceps (Cameron), the only other 


species in the genus. 


Key Words: 


Specimens representing a new species of 
Sutwanus Smith reared from a cultivar of 
guava, Psidium guajava L. (Myrtaceae), 
known as Taiwanese guava in Costa Rica, 
were sent to me by Dr. Hugo Aguilar, Univ- 
ersidad de Costa Rica. They were reared 
from larvae attacking young leaves of a 
small tree. The new species, considered a 
potential pest of this important agricultural 
commodity in Costa Rica, is described to 
provide a name. 

Sutwanus was described in the Acordu- 
lecerinae by Smith (1990). Although Smith 
(1990) examined a number of undescribed 
species, he included only Sutwanus nigri- 
ceps (Cameron 1883), known from Mexico 
to Costa Rica. The new species described 
below is structurally very similar to S. ni- 
griceps but differs by its unusual color pat- 
tern and slight differences in the ovipositor 
and antennae. Sutwanus, Acorduleceridea 
Rohwer, and Busalus Smith are distin- 
guished from other Acordulecerinae by 
their 7-segmented antennae; other genera 
have 6, 8, or 9 antennal segments. Sutwanus 
is distinguished from Acorduleceridea and 
Busalus by the third antennal segment lon- 


Symphyta, guava, Taiwanese guava 


ger than the fourth, the head in dorsal view 
strongly narrowing behind the eyes, the 
lower interocular distance shorter than the 
eye length, the linear malar space, and the 
forewing with three cubital cells. Acordu- 
leceridea occurs from Mexico to Argentina, 
and Busalus occurs only in southeastern 
Brazil. Food plants are not known for either 
genus. 


Sutwanus guajavae Smith, new species 
(Figs. 1—4) 


Female.—Length, 5.0 mm. Antenna and 
head black with clypeus, labrum, and base 
of mandible white, apex of mandible red 
brown. Thorax black. Legs black with inner 
surfaces and apices of coxae, trochanters, 
and under surface of femora white. Abdo- 
men orange with basal plates, narrow an- 
terior margin of second tergum, sheath, and 
cercus black. Wings uniformly, darkly in- 
fuscated; veins and stigma black. 

Antenna 7-segmented (Fig. 3); length 
subequal to head width; third segment 1.3 x 
length of fourth segment; fourth segment 
1.4X length of fifth segment; segments 5— 
7 subequal to very slightly decreasing in 


VOLUME 107, NUMBER 1 


Figs. 1-3. Sutwanus guajavae. |, Head, front view. 2, Head, dorsal view. 3, Antenna. 


length; apical two antennal segments each 
about 2 longer than broad. Eyes large and 


slightly converging below (Fig. 1); lower 
interocular distance 0.7 eye length, upper 


interocular distance 0.9 eye length. Malar 
space linear. Head in dorsal view sharply 
narrowing behind eyes (Fig. 2). Distances 
between eye and lateral ocellus, between 


Figs. 4—5. 


lateral ocelli, and from lateral ocellus to 
hind margin of head as 1.0:1.5:1.3; posto- 
cellar area 2.2X broader than long. Hind 
basitarsus 1.4 longer than length of re- 
maining tarsal segments combined. Length 
of inner hind tibial spur 0.4 length of hind 
basitarsus and 1.2 longer than apical 
width of hind tibia. Sheath with laterally 
projecting scopae. Forewing with 3 cubital 
cells. Lancet (Fig. 4) with 12 serrulae and 
11 annuli bearing broad, blunt teeth; dorsal 
portion of basal annuli curved anteriorly; 
each serrula with numerous small posterior 
subbasal teeth. 

Male.—Length, 4.8 mm. Similar to fe- 
male except abdomen black, and femora 
and fore tibia white except blackish on dor- 
sum of femora. 

Types.—Holotype @, labeled “‘Costa 
Rica, San Jose, San Francisco Dos Rios, 
Urbanizacion La Pacifica, 1,100 m, 18/X/ 
03, H. Aguilar,” ““Guayaba taiwanesa, plan- 
tulas conseguidas Jicaral, Peninsula Nicoya, 
Puntarenas 0-500 msnm, Eclosion adultos 
28-31/X/03.”” Paratypes: 1 2, 2 d, same 
data as for holotype. Holotype and 2 ¢ 
paratypes deposited in Universidad de Cos- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


3 


Female lancets. 4, Sutwanus guajavae. 5, S. nigriceps (specimen from Mexico). 


ta Rica; | 2 paratype in National Museum 
of Natural History, Smithsonian Institution, 
Washington, DC. 

The larvae were affecting the leaves of 
commercial Taiwanese guava (Psidium 
guajava) in the central valley of Costa Rica. 
The trees originally were taken from Jicaral 
in the Peninsula of Nicoya, Puntarenas 
Province. 

Etymology.—The specific epithet is from 
the food plant, Psidium guajava. 

Discussion.— Sutwanus nigriceps was 
redescribed by Smith and Janzen (2003), 
who provided notes on biology based on 
specimens reared from larvae feeding on 
mature leaves of Psidium guajava in the 
Area de Conservacion Guanacaste at the in- 
terface of dry forest with rainforest. Larvae 
of S. guajavae were feeding on young 
leaves, which may indicate a biological dif- 
ference between the species. 

Differences in coloration separate fe- 
males of S. nigriceps and S. guajavae. The 
female of S. nigiceps is almost entirely or- 
ange to yellow orange with only the anten- 
na, head, tegula, upper part of the meso- 
pleuron, tarsi, outer surfaces of the tibiae, 


VOLUME 107, NUMBER 1 


and most of the dorsum of the abdomen 
black. Sutwanus guajavae is almost entirely 
black with a contrasting orange abdomen. 
The female lancets of the two species are 
very similar, but the basal annuli of S. gua- 
jJavae curve anteriorly at their dorsal part 
(Fig. 4) whereas those of S. nigriceps are 
straighter (Fig. 5). The apical two antennal 
segments of S. guajavae are rather stout, 
about two times longer than broad (Fig. 3), 
whereas those of S. nigriceps are more slen- 
der, about three times longer than broad 
(Smith and Janzen 2003, fig. 11). The 
sheaths of both species are similar to the 
illustration of S. nigriceps by Smith (1990, 
fig. 415). The males of both species are 
similarly colored except the tibiae are al- 
most entirely white in S. nigriceps. Males 
of many Acordulecerinae are difficult to 
separate because of their small size and 
similar coloration, and the taxonomy of this 
group is based mostly on females. The male 
genitalia of S. gudajavae appear indistin- 
guishable from the genitalia of S. nigriceps 
as illustrated by Smith (1990, fig. 418). 

I have seen a number of specimens of S. 
nigriceps from Mexico to Costa Rica, and 
there is very little color variation, except in 
the amount of black on the dorsum of the 
abdomen. Both species are very similar in 
all other structural features. The available 


217 


material strongly suggests that the two are 
distinct species rather than extreme color 
variations of the same species. There is 
probably a complex of species in Sutwanus 
with similar lancets as figured. 


ACKNOWLEDGMENTS 


I thank Dr. Hugo Aguilar, Director, Mu- 
seum of Insects CIPROC, Department of 
Agronomy, Universidad de Costa Rica, for 
bringing these specimens to my attention. 
James Coronado, Universidad de Costa 
Rica, provided a preliminary identification. 
Cathy Apgar, Systematic Entomology Lab- 
oratory (SEL), USDA, took the photos and 
arranged the plates. I appreciate the reviews 
of Nathan M. Schiff, U.S. Forest Service, 
Stoneville, MS, and Thomas J. Henry and 
John W. Brown, SEL, Washington DC. 


LITERATURE CITED 


Cameron, P. 1883. Hymenoptera, Tenthredinidae— 
Chrysididae. Jn Godman and Salvin, Biologia 
Centrali-Americana, Vol. 1, 486 pp. 

Smith, D. R. 1990. A synopsis of the sawflies (Hy- 
menoptera: Symphyta) of America south of the 
United States: Pergidae. Revista Brasileira Ento- 
mologia 34: 7—200. 

Smith, D. R. and D. H. Janzen. 2003. Food ere and 
life histories of sawflies of the families Tenthre- 
dinidae and Pergidae (Hymenoptera) in Costa 
Rica, with descriptions of four new species. Jour- 
nal of Hymenoptera Research 12: 312-332. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 218-228 


A TAXONOMIC REVIEW OF MATSUMURELLA ISHIHARA (HEMIPTERA: 
CICADELLIDAE: DELTOCEPHALINAE) FROM CHINA 


YALIN ZHANG AND WU DAI 


Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Ed- 
ucation, Entomological Museum, Northwest Sci-Tech University of Agriculture and For- 
estry, Yangling, Shaanxi 712100, China (e-mail: yalinzh@nwsuaf.edu.cn; daiwu@ 


nwsuaf.edu.cn) 


Abstract.—The nine species of the genus Matsumurella Ishihara from China are re- 
viewed and illustrations of genitalia are provided. Among them, two new species, M. 
protrudea and M. parallela, are described bringing the number of species in the world 


to twelve. A key is given to Chinese species. 


Key Words: 


The leafhopper genus Matsumurella 
(Deltocephalinae: Athysanini) was _ estab- 
lished by Ishihara (1953) for two species 
from Japan, M. kogotensis (Matsumu- 
ra,1914), the type species, and M. praesul 
(Horvath, 1899). Later, Ishihara (1958) pro- 
posed an unnecessary replacement name 
(Shonenus), for Matsumurella, |preoccu- 
pied by Matsumuraiella Enderlein, 1906 
(Psocoptera)]. Emeljanov (1962) described 
one new species M. phaea and tranfered M. 
praestans (Jacobi, 1943) to the genus from 
Allygus. Anufriev (1971) placed M. praes- 
tans (Jacobi, 1943) in synonymy with M. 
praesul (Horvath, 1899) and described two 
new species (M. curticauda and M. longi- 
cauda) from China and one (M. phaeicola) 
from Japan. Later, M. expansa Emeljanov, 
1972, and M. minor Emeljanov, 1977, were 
described from Mongolia. Recently, Cai 
and Wang (2002) described two new spe- 
cies M. rurcata and M. singularis from Chi- 
na. 

This paper is a taxonomic review of the 
genus Matsumurella Ishihara in China. All 
twelve species of the genus are listed, and 
a key is given to the nine Chinese species 
(two new). 


Homoptera, Cicadellidae, Deltocephalinae, Matsumurella, new species, China 


Type specimens and other materials ex- 
amined are deposited in the Institute of Zo- 
ology, Chinese Academy of Sciences (IZ- 
CAS), Entomological Museum of North- 
west Sci-Tech University of Agriculture 
and Forestry (NWSUAP), Shanghai Ento- 
mological Museum (SEM), and Nankai 
University (NKU), Tianjin. 


Matsumurella Ishihara 


Matsumurella Ishihara 1953a: 200. Type 
species: Jassus kogotensis Matsumura, 
1914, by original designation. 

Shonenus Ishihara 1958: 232. Unnecessary 
replacement name for Matsumurella Ishi- 
hara 1953 


Description.—Body elongate, robust. 
Head including eyes narrower than prono- 
tum. Crown longer medially than next to 
eyes, shorter than width between eyes; an- 
terior margin roundly produced anteriorly 
in dorsal view; disc somewhat flat. Transi- 
tion of vertex and frontoclypeus rounded. 
Eyes fairly large; ocellus situated on frontal 
margin of crown next to corresponding eye, 
separated from eye by distance equal to its 
diameter. Clypellus similar in width 


VOLUME 107, NUMBER 1 


throughout length. Pronotum with anterior 
margin roundly produced and posterior 
margin slightly concave. Scutellum almost 
1% times length of head, with transverse su- 
ture curved and depressed. Forewing elon- 
gate with a few extra cross veins; appendix 
wide. 

Male pygofer side tapering to a caudal 
projection, dorsoposterior margin with sev- 
eral macrosetae. Subgenital plate with sev- 
eral macrosatae laterally. Connective Y- 
shaped, shaft short and robust, arms well 
developed. Paramere with basal half wide 
and narrowing to middle, apical half nar- 
row, lateral lobe absent or weakly devel- 
oped. Aedeagus with basal apodeme well 
developed, shaft elongate, compressed dor- 
soventrally, curved dorsally with one or 
two pair of apical appendages; phallotreme 
subapical or distad from penis apex. 

Distribution.—China, Japan, Korea, 
Mongolia and Russia (Amur, Maritime Ter- 
ritory, Kurile Islands). 


WORLD CHECKLIST OF THE GENUS 
MATSUMURELLA ISHIHARA 


M. curticauda Anufriev 1971. China. 

M. expansa Emeljanov 1972. China, Mon- 
golia. 

M. kogotensis (Matsumura 1914). Japan. 

M. longicauda Anufriev 1971. China. 

M. minor Emeljanoy 1977. China, Mongo- 
lia. 

M. parallela Zhang and Dai, n. sp. China. 

M. phaea Emeljanoy 1962. Russia (Amur, 
Maritime Territory). 

M. phaeicola Anufriev 1971. Japan. 

M. praesul (Horvath 1899). China, Japan, 
Korea, Russia (Maritime Territory, Kurile 
Islands). 

M. protrudea Zhang and Dai, n. sp. China. 

M. rurcata Cai and Wang 2002. China. 

M. singularis Cai and Wang 2002. China. 


KEY TO SPECIES FROM CHINA (< ) 


1. Aedeagus with a single apical process (Fig. 65) 
er a eeu. Weiss cues s boa singularis 


— Aedeagus with one or two pair of apical pro- 
cesses (Figs. 6, 15, 23, 32, 38, 44,52,58) .. 2 


2. Aedeagus with a single pair of apical processes 
(Bigshi6; 155 235 38552455) 
— Aedeagus with two pair of apical processes 
(Pigs) 32, 44): ences «om Bete a, Seca 8 
3. Apical aedeagal processes forked (Fig. 58) 


D5 23538052) Sage eck ee ee eee - 
4. Apical processes of aedeagus directed laterally, 
perpendicular to shaft (Figs. 38,52) ....... 5 


— Apical processes of aedeagus directed antero- 
leforallky Gres, ©, 5 23) ssoanoe5ee552+5- 6 
5. Subgenital plate shorter than basal width, lat- 
eral margin strongly concave subapically, 
forming a digitate apical projection; aedeagus 
basal apodeme long, preatrium indistinct (Figs. 
35; 383039)! sh sess haere cs 2. Cuege 
— Subgenital plate longer than basal width, lat- 
eral margin weakly concave subapically; ae- 
deagus basal apodeme short, preatrium long 
(Pissv4 8 151) Eis wha meshed ue char aoe 
6. Subgenital plate with concave at apical inner 
margin (Figs. 3, 20) 


protrudea, 0. sp. 


longicauda 


— Subgenital plate without concave along inner 
margin and with apex curved dorsad (Figs. 10— 
De ered cfs scene ee eile, Sher eae eo eae mere curticauda 
7. Apical processes of aedeagus divergent, ex- 
tended anterolaterally; pygofer caudal prolon- 
gation straight in dorsal view (Figs. 2, 6) 
+ Sus co hcoise Amine oe cae ened Sen cv ee oes ore ec eaeme praesul 
— Apical processes of aedeagus nearly parallel, 
extended apically; pygofer caudal prolongation 
waved apically in dorsal view (Figs. 19, 23) 
Ee ea ret RTT IG NON Gnc. SNOT P ICY ¢ expansa 
8. Subgenital plate with lateral margin concave 
subapically, protruded at apex; aedeagus with 
subapical processes moderately long, terminat- 
ing distad of aedeagal base in lateral view 
GE1CS.29N 38) eager oreue parallela, n. sp. 
— Subgenital plate with lateral margin produced 
lobelike subapically; aedeagus with subapical 
processes long terminating near aedeagal base 
in lateral view (Figs. 40, 45) minor 


Matsumurella praesul (Horvath) 
(Figs. 1-8) 


Jassus praesul Horvath 1899: 370; Matsu- 
mura 1914: 206. 

Matsumurella praesul: Ishihara 1953b: 41; 
Metcalf 1967: 1004—1005; Anufriev 
1971: 515; Nast 1972: 369; Anufriev 
1979a: 165; Lee 1979: 463, plate 37, fig. 
186a, b, figs. 402-403, 404 (1-2); An- 
ufriev & Emeljanoy 1988: 205, plate 140, 
fig. 1, plate 141, figs. 1-7. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-8. 


Shonenus praesul: Vilbaste 1968: 143, plate 
115, figs. 1-8. 

Allygus praestans Jacobi 1943a: 29; Anuf- 
riev 1971: 515, figs. 22-23; Anufriev 
1979a: 165. 

Matsumurella praestans: Emelyanov 1962: 
162, fig. 19; Metcalf 1967: 983; Anufriev 
IMs SiS, iss, ZA—WBs INeweE IOTZs SOL. 


Material examined.—China, 146, Jilin 
Prov., 10 June 1940 (IZCAS); 16, 8@, Jilin 
Prov., Linjiang, Hongtushan Forest Farm, 
31 July 1983, coll. Wu Zheng-Liang and 
Hua Bao-Zhen (NWSUABP); 2 6,3 &, Jilin 
Prov., Linjing, Naozhi, 22—24 July 1983, 
coll. Wu Zheng-Liang and Hua Bao-Zhen 
(NWSUABF); 4 6,4 &, Jilin Prov., Linjiang, 
Dongxiaoshan Forest Farm, 3 July 1983 
coll. Wu Zheng-Liang and Hua Bao-Zhen 
(NWSUAF); | @, Heilongjiang Prov., Yuy- 


Matsumurella praesul. 1, Male pygofer side, lateral view. 2, Male pygofer side, dorsal view. 3, 
Genital valve and subgenital plate, ventral view. 4, Connective. 5, Paramere, dorsal view. 6, Aedeagus, dorsal 
view. 7, Aedeagus, lateral view. 8, Female abdominal sternum VII, ventral view. 


ing, 21 August 1978, coll. Fang San-Yang 
(NWSUAPF); 1 do, Heilongjiang Prov., Dail- 
ing, 390 m, 24 July 1957, He Zhong (IZ- 
CAS); 1 ¢@, Heilongjiang Prov., Dailing, 
390 m, 6 August 1963, coll. Bai Jiu-Wei 
(IZCAS); 1 6, Heilongjiang Prov., Yichun, 
Wuyingkou, 24 July 1980, coll. Zheng 
(NKU); 1 3, Heilongjiang Prov., Bolitong- 
tun, 29 July 1980, coll. Wang (NKU). 

Distribution.—China (Heilongjiang, Ji- 
lin), Japan, Korea, Rossia (Maritime Terri- 
tory, Kurile Islands). 


Matsumurella curticauda Anufriev 
(Figs. 9-17) 


Matsumurella curticauda Anufriev 1971: 
512, figs. 5-9; Cai and Shen 2002: 274. 


Material examined.—China: 2 6, 1 2, 
Zhejiang, Hangzhou, Zhiwuyuan (Botanic 


VOLUME 107, NUMBER 1 


Figs. 9-25. 
10, Male pygofer, caudal view. 11, Genital valve, subgenital plate, paramere and connective, dorsal view. 12 
Genital valve and subgenital plate, ventral view. 13, Connective. 14, Aedeagus, caudal view. 15, Aedeagus 
dorsal view. 16, Aedeagus, lateral view. 17, Female abdominal sternum VII, ventral view. 18-25, M. expansa 
18, Male pygofer side, lateral view. 19, Male pygofer side, dorsal view. 20, Genital valve and subgenital plate 
ventral view. 21, Paramere, dorsal view. 22, Connective. 23a, Aedeagus dorsal view. 23b, Aedeagus, caudal 
view. 24, Aedeagus, lateral view. 25, Female abdominal sternum VII, ventral view. 


9-17, Matsumurella curticauda (reproduced from Anufriev 1971). 9, Male pygofer, lateral view. 


i) 
>) 
tO 


Garden), 3 June 1982, coll. Yan Heng- 
Yuan; 1 6, Zhejiang, Hangzhou, Zhiwu- 
yuan (Botanic Garden), 24 May 1982, coll. 
Yan Hengyuan; | @, Zhejiang, Hangzhou, 
Zhiwuyuan (Botanic Garden), | June 1982, 
coll. Yan Hengyuan; 3 @, Jiangsu, Sheshan 
Mountain, 8 June 1958 (all in SEM). 

Distribution.—China (Jiangsu, Zhejiang, 
Henan). 


Matsumurella expansa Emeljanov 
(Figs. 18—25) 


Matsumurella expansa Emeljanoy 1972: 
236, figs. 63-64; Emeljanov 1977: 151; 
Anufriev & Emeljanov 1988: 205, plate 
141, fig. 13, plate 142, figs. 1-5. 


Material examined.—China, 16 3, 8 @, 
Henan Prov., Neixiang, Baotianman, 11 
July 1998, 1,300 m, coll. Hu Jian; 7 6, 3 
?, Henan Prov., Neixiang, Getiaopa, 14 
July 1998, 600-700 m, coll. Hu Jian; 1 d, 
Henan Prov., Xixia, Huangshian Forest 
Farm, 17 July 1998, 800—1,300 m, coll. Hu 
Jian; 1 @, Jilin Prov., Linjiang, Naozhi, 24 
July 1983, coll. Hua Bao-Zhen and Wu 
Zheng-Liang; 2 d, Shaanxi Prov., June 
1980, Ma Ning and Xiang Long-Cheng; | 
3,1 2, Shaanxi Prov., Huanglong, 9/10 Au- 
gust 1973, coll. Yuan Feng, MI Shun-Rong 
and Hui Jun-Rui; | 6, Shaanxi Prov., Wug- 
ong, 20 July 1987 coll. Northwestern Ag- 
riculture College; 1 ¢, Shaanxi Prov., Wug- 
ong, 24 July 1987, coll. JIA Mei-Ling; 1 d, 
2 ¢, Shaanxi Prov., Huoditang, 6 July 
1984, coll. Northwestern Agriculture Col- 
lege; 1 d, 1 2, Shaanxi Prov., Mt. Qinling, 
24 July 1995 coll. Liu Jun-Wu and Song 
Hua-Hai; 3 ¢, Shaanxi Prov., Nanwutai, 
July 1979, coll. Tian Chou and Chen Tong; 
1 2, Shaanxi Prov., Nanwutai, June 1980, 
coll. Ma Ning; | 6, Shaanxi Prov., Fe- 
ngxian Country, 18-19 July 1988, coll. Cui 
Jun-Feng; | 6, Shaanxi Proy., Yanan, June 
1980, coll. Ma Ning and Xiang Long- 
Cheng; | 2, Shaanxi Prov., Mt. Taibeishan, 
Haopingsi, 1,165 m, 19/22 July 1956, coll. 
Zhou Yao; 1 6, Shaanxi Prov., June 1980, 
coll. Ma Ning and Xiang Long-Cheng; | &, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Shanxi Prov., Mt. Hengshan, 4/6—12/15 Au- 
gust 1964, coll. Zhou Yao and Liu Shao- 
You (all above in NWSUAFP); 1 2, Gansu 
Prov., Chengxian Country, Feilongxia, 
1,020 m, 4 July 1999, coll. He Tong-Li (1Z- 
CAS). 

Distribution.—China (Henan, 
Shaanxi, Shanxi, Gansu), Mongolia. 


Jilin, 


Matsumurella parallela, Zhang and Dai, 
new species 
(Figs. 26—33) 


Male.—Length (including tegmen): 6.5 
mm. 

Yellow to fuscous. Vertex with a trans- 
verse black streak each side of midline on 
fore margin and a broad black transverse 
band between ocelli, sometimes extending 
to ocelli and sometimes interrupted medi- 
ally or fused with more anterior streaks. 
Pronotum often yellow along fore border 
with some dark spots and caudally usually 
black. Scutellum yellow, usually with black 
lateral spots and often with a black trans- 
verse band bordering transverse suture. 
Fore wing fuscous. 

External features as in generic descrip- 
tion. 

Pygofer side with caudal prolongation 
curved dorsolaterally; dorsoposterior mar- 
gin with several macrosetae. Subgenital 
plate short, lateral margin concave subapi- 
cally, with several macrosetae laterally. 
Connective Y-shaped, shaft robust and arms 
well developed. Paramere with basal half 
wider and narrowing to the middle, apical 
half narrow. Aedeagus with basal apodeme 
well produced, shaft long and curved dor- 
sad with one apical pair of short processes, 
directed dorso-laterally and a pair of longer 
subapical processes from dorsal surface, 
more or less parallel to shaft, directed ven- 
trally and terminating distad of aedeagal 
base in lateral view; phallotreme apical on 
ventral surface, short. 

Types.—Holotype: ¢, China, Gansu 
Prov., Wenxian, Qiujiaba, 2,000—2,100 m 1 
July 1998, coll. Yao Jian. Paratypes: China, 
1 d, same data as holotype, coll. Yang 


VOLUME 107, NUMBER 1 


i) 
i) 
ie) 


30 29 


33 


31 


Figs. 26-39. 26-33, Matsumurella parallela. 26, Head and thorax, dorsal view. 27, Face. 28, Male pygofer 
side, lateral view. 29, Male genital valve and subgenital plates (left plate setae omitted), ventral view. 30, 
Paramere, dorsal view. 31, Connective. 32, Aedeagus, dorsal view. 33, Aedeagus, lateral view. 34-39, M. 
protrudea. 34, Male pygofer side, lateral view. 35, Male genital valve and subgenital plate, ventral view. 36, 
Paramere, dorsal view. 37, Connective. 38, Aedeagus, dorsal view. 39, Aedeagus, lateral view. 


224 


Xing-Ke; | d, Gansu Prov., Dangchang, 
Dahebagou, 1,700—2,300 m, 1 July 1998, 
coll. Zhang Xue-Zhong (all in IZCAS). 

Notes.—This new species resembles 
Matsumurella minor, but can be distin- 
guished from the latter by: |) The pygofer 
prolongation stouter; 2) the subgenital plate 
tapered more apically; and 3) the aedeagus 
with subapical processes shorter and more 
parallel. 

Etymology.—This new species is based 
on the pair of long parallel subapical pro- 
cesses of the aedeagus. 


Matsumurella protrudea, Zhang and 
Dai, new species 
(Figs. 34-39) 


Male.—Length (including 
6.1mm. 

Brownish red, crown with a pair of black 
marks on anterior margin. Scutellum yellow 
on anterior part. 

External features as in generic descrip- 
tion. 

Pygofer side with caudal prolongation di- 
rected ventroposteriorly; dorsoposterior 
margin with several macrosetae. Subgenital 
plate shorter than broad, lateral margin 
strongly concave subapically, forming a 
digitate apical projection; lateral margin 
with several macrosatae. Connective Y- 
shaped, shaft robust and arms well devel- 
oped. Paramere with basal half wider and 
narrowing to the middle, apical half narrow. 
Aedeagus with dorsal apodeme well pro- 
duced, shaft long and curved dorsad with a 
pair of apical processes directed laterally, 
perpendicular to the shaft; phallotreme api- 
cally on ventral surface, elongate. 

Type.—Holotype: 6, China, Henan 
Prov., Xixia, Huangshian Forest Farm, 
800—1,300 m, 17 July 1998, coll. Hu Jian 
(NWSUAF). 

Notes.—This new species resembles 
Matsumurella longicauda, but can be dis- 
tinguished from the latter by: 1) The py- 
gofer caudal prolongation straighter and di- 
rected posteroventrally; 2) the subgenital 
plate shorter with lateral margin strongly 


tegmen): 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


concave subapically forming a digitate api- 
cal process; and 3) the aedeagal basal apo- 
deme more produced. 

Etymology.—This name is based on the 
subgenital plate digitate process. 


Matsumurella minor Emeljanov 
(Figs. 40—46) 


Matsumurella minor Emeljanov 1977: 151, 
figs. 5—8; Cai and Shen 2002: 275. 
Material examined.—China, | 6, Shaanxi 
Prov., Ningshan Country, Huoditang, 22 
July 2000, coll. Dai Wu and Liu Zhen- 
Jiang (NWSUAB); 1 @, Shaanxi Prov., 
Ningshan Country, 17 August 1984, 
Northwestern Agriculture College 
(NWSUAF); 1 2, Shaanxi Prov., Mt. Tai- 
baishan, Dadian, 2,300 m 30 June 1987, 
Northwestern Agriculture University 
(NWSUAF); 1 2, Shaanxi Prov., Mt. Tai- 
baishan, Zhongshansi, 17 July 1982, 
1,500 m, coll. Zhou Jing-Ruo and Liu 
Lan (NWSUAF); 1 @°, Shaanxi Prov., Mt. 
Qinling, July 1973, Zhang Xue-Zhong 
(IZCAS); 1 3, Shaanxi Prov., Yanan, 
June 1980, coll. Ma Ning (NWSUAP); 2 
3,3 2, Henan Prov., Neixiang, Getiaopa, 
14 July 1998, 66-700 m, coll. Hu Jian 
(NWSUAF); 164, Henan Prov., Longyu- 
wan, 17 July 1996, coll. Zhang Wen-Zhu 
(NWSUAF); 1 2, Henan Prov., Songxian 
Country, Mt. Baiyunshan, 17 July 1996, 
1400 m, coll. Zhang Wen-Zhu 
(NWSUABP);. 

Distribution.—China (Shaanxi, Henan), 
Mongolia. 


Matsumurella longicauda Anufriev 
(Figs. 47—53) 


Matsumurella longicauda Anufriev 1971: 
511, figs. 1-4; Cai, He and Gu 2001: 
203; Cai and Shen 2002: 274. 


Material examined.—China, 1 2, Zhe- 
jiang, Xitianmu Mountain, 21 July 1981, 
coll. Yan Heng-Yuan; 1 6d, Zhejiang, Mt. 
Tianmushan, 22 July 1981, coll. Yan Heng- 
Yuan; | 6, Zhejiang, Hangzhou, 21 May 
1983, coll. Zheng Jian-Zhong (all in SEM). 


VOLUME 107, NUMBER 1 22 


ead 


50 53 49 


Figs. 40-53. 40-46, Matsumurella minor. 40, Male pygofer side, lateral view. 41, Male genital valve and 
subgenital plate, ventral view. 42, Paramere, dorsal view. 43, Connective. 44, Aedeagus, dorsal view. 45, Ae- 


deagus, lateral view. 46, Female abdominal sternum VII, ventral view. 47-53. M. longicauda. 47, Male pygofer 
(genital capsule), lateral view. 48, Male genital valve and subgenital plate, ventral view. 49, Paramere, dorsal 
view. 50, Connective. 51, Aedeagus, lateral view. 52, Aedeagus, dorsal view. 53, Female abdominal sternum 


VII, ventral view. 


226 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 54-67. 54—60, Matsumurella rurcata (reproduced from Cai and Wang 2002). 54, Male pygofer side, 
lateral view. 55, Male genital valve and subgenital plate, ventral view. 56, Paramere, dorsal view. 57, Connective. 
58, Aedeagus, dorsal view. 59, Aedeagus, lateral view. 60, Female abdominal sternum VII, ventral view. 61— 
67, M. singularis (reproduced from Cai and Wang 2002). 61, Male pygofer side, lateral view. 62, Male genital 
valve and subgenital plate, ventral view. 63, Paramere, dorsal view. 64, Connective. 65, Aedeagus, dorsal view. 
66, Aedeagus, lateral view. 67, Female abdominal sternum VII, ventral view. 


VOLUME 107, NUMBER 1 


Distribution.—China (Jiangsu, Zhejiang, 
Henan). 


Matsumurella rurcata Cai et Wang 
(Figs. 54-60) 


Matsumurella rurcata Cai and Wang 2002: 
22, figs. 9-16; Cai and Shen 2002: 275. 
Distribution.—China (Henan). 


Matsumurella singularis Cai et Wang 
(Figs. 61—67) 


Matsumurella  singularis Cai and Wang 
2002: 21,figs. 1-8; Cai and Shen 2002: 
ZU): 

Distribution.—China (Henan). 


ACKNOWLEDGMENTS 


We acknowledge Professor Yang Xing- 
Ke (Institute of Zoology, Chinese Academy 
of Sciences, Beijing), Zhang Wei-Nian, Liu 
Xian-Wei and Yin Hai-Sheng (Shanghai 
Entomological Museum, Chinese Academy 
of Sciences, Shanghai), and Liu Guo-Qing 
and Bu Wen-Jun (Nankai University, Tian- 
jin) for lending us specimens. Sincere 
thanks also go to Mick Webb, The Natural 
History Museum, London, for his com- 
ments and revising the manuscript. The pro- 
ject was supported by The Fok Ying Tung 
Education Foundation, National Education- 
al Committee of China. 


LITERATURE CITED 


Anufriev, G. A. 1971. Study of the genus Matsumu- 
rella Ishihara 1953, (Homoptera, Auchenorrhyn- 
cha, Cicadellidae) with the description of three 
new species from China and Japan. Bulletin de 
1’ Academie Polonaise des Sciences. Serie des Sci- 
ences Biologiques 19(7&8): 511-516. 

. 1979a. Notes on some A. Jacobi’s species of 
auchenorrhynchous insects described from North- 
East China (Homoptera). Reichenbachia 17(19): 
163-170. 

Anufriev, G. A. and A. E Emeljanov. 1988. Keys to 
the Identification of Insects of the Soviet Far East, 
Vol. 2. Homoptera & Heteroptera. Nauka, Lenin- 
grad, Russia, 972 pp. [in Russian]. 

Cai, P., J. He, and X. Gu. 2001. Cicadellidae, pp. 185— 
218. In Wu, H. and Ch. Pan., eds. Insects of Tian- 
mushan National Nature Reserve. Science Press, 
Beijing, China, 764 pp. [in Chinese]. 

Cai, P. and J. Wang. 2002. Two new species of Mat- 


22 


sumurella Ishihara from Henan Province, China 
(Homoptera: Cicadellidae: Deltocephalinae), pp. 
21-24. In Shen, X. and Y. Zhao, eds. The Fauna 
and Taxonomy of Insects in Henan, Vol. 5. Insects 
of the Mountains Taihang and Tongbai Regions. 
China Agricultural Sci-Tech Press, Beijing, China, 
453 pp. [in Chinese]. 

Cai, P. and X. Shen. 2002. Homoptera: Cicadellidae, 
pp. 269-279. In Shen, X. and Y. Zhao, eds. The 
Fauna and Taxonomy of Insects in Henan, Vol. 5. 
Insects of the Mountains Taihang and Tongbai Re- 
gions. China Agricultural Sci-Tech Press, Beijing. 
China. 453 pp. [in Chinese]. 

Emeljanoy, A. FE 1962. Materials on taxonomy of Pa- 
laearctic leafhoppers (Auchenorrhyncha, Eusceli- 
nae). Trudy Zoologicheskogo Instituta Akademii 
Nauk, USSR, 30: 156—184 [in Russian]. 

. 1972. New leafhoppers (Homoptera, Auchen- 
orrhyncha) from the Mongolian People’s Repub- 
lic. Nasekomye Mongolii [Insects of Mongolia] 1: 
199—260 [in Russian]. 

1977. Leaf-hoppers (Homoptera, Auchenor- 
rhyncha) from the Mongolian People’s Republic 
based mainly on materials of the Soviet-Mongo- 
lian zoological expeditions (1967-1969). Nase- 
komye Mongolii [Insects of Mongolia] 5: 96—195 
{in Russian]. 

Horvath G. 1899. Hémipteres de Vile de Yesso (Ja- 
pon). Természetrajzi Fiizetek 22: 365-374, figs. 
1-2. 

Ishihara, T. 1953a. Some new genera including a new 
species of Japanese Deltocephalidae (Hemiptera). 
Transactions of the Shikoku Entomological Soci- 
ety 3(8): 192—200 

. 1953b. A tentative check list of the superfam- 
ily Cicadelloidea of Japan (Homoptera). The Sci- 
entific Reports of the Matsuyama Agricultural 
College, Matsuyama, Japan 11: 1—72,17 pls. 

———.. 1958. The superfamily Cicadelloidea of Nii- 
gata Prefecture, North Honshu, Japan (Hemip- 
tera). Kontyt, Tokyo, Japan 26: 225—232 [in Jap- 
anese]. 

Jacobi, A. 1943. Zur Kenntnis der Insekten von Mand- 

12. Beitrag. Eine Homopterenfaunula 

der Mandschurei. (Homoptera: Fulgoroidea, Cer- 


schukuo. 


copoidea, and Jassoidea). Arbeiten iber Morphol- 
ogische und Taxonmische Entomologie 10: 21— 
31, figs. 1-10. 

Lee, C. E. 1979. Illustrated Flora and Fauna of Korea. 
23. Insecta 7. Samhwa Publishing Co. Ltd. Seoul., 
Korea, 1,070 pp. [in Korean]. 

Matsumura, S. 1914. Die Jassinen und einige neue 
Acocephalinen Japans. Journal of the College of 
Agriculture, Tohoku Imperial University, Sapporo, 
Japan 5: 165-240, figs. 1-12. 

Metcalf, Z. P. 1967. General Catalogue of the Homop- 
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Il, Ill, Euscelidae. Agricultural Research Service, 


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(ee) 


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ington, D.C., 2,695 pp. 

Nast, J. 1972. Palaearctic Auchenorrhyncha (Homop- 
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ology, Polish Academy of Sciences, Polish Sci- 
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Gebietes. Tallin, Estonia, 180 pp. [in Russian]. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 229-234 


DESCRIPTION OF THE MALE, HOST ASSOCIATIONS, AND NEW 
DISTRIBUTION RECORDS FOR LOPHOSTIGMA CINCTA (DU BUYSSON) 
(HYMENOPTERA: MUTILLIDAE) 


ROBERTO A. CAMBRA, VICTOR H. GONZALEZ, AND WILLIAM T. WCISLO 


(RAC) Museo de Invertebrados “‘G. B. Fairchild,’ Universidad de Panama, Estafeta 
Universitaria, Panama, Republica de Panama (e-mail: reambra@ancon.up.ac.pa); (VHG) 
Department of Ecology and Evolutionary Biology, Snow Hall, 1460 Jayhawk Blvd., Uni- 
versity of Kansas, Lawrence, KS 66045, U.S.A. (e-mail: vhgonza@ku.edu); (WTW) 
Smithsonian Tropical Research Institute, Apartado 2027, Balboa, Anc6n, Reptiblica de 
Panama (e-mail: wcislow @s1.edu). Address for reprints and correspondence: W. T. Wcislo, 
Smithsonian Tropical Research Institute, Unit 0948, APO AA 34002-0948, or Apartado 
2072, Balboa, Republic of Panama 


Abstract.—The male of the mutillid wasp genus Lophostigma Mickel is described for 
the first ttme based on specimens of Lophostigma cincta (du Buysson). The sex association 
for L. cincta is based on individuals reared from nests of nocturnal sweat bees, Megalopta 
genalis (Meade-Waldo) and M. ecuadoria Friese (Hymenoptera: Halictidae). These bees 
are the first confirmed hosts for Lophostigma. We summarize the geographical distribution 


of L. cincta. 


Key Words: 


Wasps in the family Mutillidae (Hyme- 
noptera) are parasitic on the prepupae and 
pupae of other insects, including beetles 
(Coleoptera), flies (Diptera), and wasps and 
bees (Hymenoptera) (summarized in Mick- 
el 1928; Brothers 1972, 1995). Although 
mutillid wasps are often vividly colored and 
conspicuous, and therefore well-represented 
in collections, the biology of most (>95%) 
of the ~8,000 described species is unknown 
(Brothers 1972, 1995). Here we describe 
parasite-host associations for the Neotropi- 
cal sphaeropthalmine genus Lophostigma 
Mickel, based on rearing the parasites from 
nests of nocturnal sweat bees, Megalopta 
genalis (Meade-Waldo) and M. ecuadoria 
Friese (Hymenoptera: Halictidae). The 
males of this mutillid genus were unknown, 


so we describe the generic characters of 


male Lophostigma based on Lophostigma 


Sphaeropthalminae, parasitism, Halictidae, Megalopta 


cincta (du Buysson) specimens. We sum- 
marize the geographical distribution for L. 
cincta in Central America. 


MATERIALS AND METHODS 


Megalopta nests were collected periodi- 
cally in the Barro Colorado Nature Monu- 
ment [9°09'N, 79°51'W; for description of 
Barro Colorado Island (BCI), see Leigh 
1999], Republic of Panama, from January 
1999 through November 2001. Nests were 
sealed with a cotton plug and transported to 
the laboratory where they were then split 
open and the cell contents examined; the 
contents were preserved or transferred to 
plastic tissue culture trays for rearing. 

Morphological descriptions follow Mick- 
el (1928, 1952), except for the mesoscutal 
terminology that follows Menke (1993). 
Photographs were made using a scanning 


230 


electron microscope (JEOL model JSM 
5300LV). The abbreviations T and S are 
used for metasomal tergum and metasomal 
sternum, respectively. Lophostigma cincta 
specimens are deposited in the Museo de 
Invertebrados ‘“‘G. B. Fairchild,” Universi- 
dad de Panama (MIUP) and the Dry Ref- 
erence Collection of the Smithsonian Trop- 
ical Research Institute (STRI). Voucher 
specimens of the host bees are deposited at 
STRI and the Natural History Museum, 
University of Kansas. 


SYNOPSIS OF HOST BIOLOGY 


Megalopta genalis and M. ecuadoria are 
medium-sized bees that nest in dead, bro- 
ken branches in the understory of primary 
and secondary forests in central Panama 
(Wcislo et al. 2004). Females establish 
nests and provision them throughout the 
dry season (approximately mid-December 
through May) and the first half of the wet 
season. Nests contain a series of cells, usu- 
ally adjacent to a single tunnel. Nests con- 
tain 1-11 adult females per nest. If two or 
more bees live together in a nest, then that 
nest is regularly guarded by one of the bees. 
Each cell is mass provisioned with pollen, 
which females collect during foraging trips 
in the early evening and pre-dawn morning. 
After feeding, larvae develop into prepupae 
and then pupae without spinning cocoons. 


TAXONOMY 


Lophostigma Mickel 
(Figs. 1—4, 8) 


Lophostigma Mickel 1952: 146. Type spe- 
cies: Mutilla iracunda Cresson, 1902, 
original designation. 


Generic diagnosis of male.—Proboscidal 
fossa moderate in size (Fig. 1); genal carina 
absent; mandible suddenly narrowed near 
tip, angle formed on inner margin by sud- 
den narrowing, rounded (Fig. 2); mandible 
without ventral, basal tooth; scape with sin- 
gle sharp carina beneath; antennal scrobe 
not carinate above, but with small, median 
tubercle; lateral surface of pronotum with 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


anterior, strong transverse carina terminat- 
ing at humeral angle in blunt tubercle (Fig. 
3); notauli absent; parapsidal lines faint, 
present in posterior third of mesoscutum; 
scutellum flat in dorsal view; dorsum of 
propodeum rounded onto posterior surface; 
mesosternum simple, without teeth or pro- 
jections, with arcuate carina in front of me- 
socoxa; mesosternal lamellate process with 
medial, emargination at apex, ending pos- 
teriorly as spine in front of metacoxa; first 
metasomal segment slightly disciform, with 
narrow, dorsal surface and large, subcircu- 
lar anterior surface; T2 with felt lines; S2 
without felt lines; S7 (hypopygium) poste- 
riorly with broad, medial emargination (Fig. 
4); tibial spurs pale; forewing with marginal 
cell subacute at tip, two submarginal cells 
(Fig. 8); seta simple, not plumose. 


Lophostigma cincta (du Buysson) 


Mutilla cincta du Buysson 1892: 58. 


Description.—Male. Length: 10.6 mm. 
Integument black, clothed with long erect 
and recumbent white pubescence, except 
last metasomal segment with black setae. 
Head transversely ovate-rectangular, width 
slightly less than width of thorax; clypeus 
with transverse, arcuate ridge; eye strongly 
convex, shortly ovate, maximum diameter 
1.0X distance from postero-lateral ocellus 
to inner eye margin; front, vertex and gena 
with dense, close, punctures; occipital re- 
gion and postgena not coarsely punctured, 
nearly smooth; front with median, im- 
pressed, longitudinal line extending from 
anterior margin between antennal tubercles 
posteriorly about half distance to anterior 
ocellus; ocellus small, maximum diameter 
0.2X distance from postero-lateral ocellus 
to inner eye margin; anterior margin clyp- 
eus with inconspicuous tooth below anten- 
nal tubercle; pedicel short, length 0.62 
flagellomere I; flagellomeres I and II ap- 
proximately equal length. Mesosoma with 
dense, close, punctures (Fig. 3), except me- 
tapleuron, smooth; dorsum of propodeum 
reticulate throughout; scutellum with me- 


VOLUME 107, NUMBER 1 


Figs. 1-6. Lophostigma cincta, male. 1, 


Head, ventral view. 2, 


o0A% 25 kV 


Mandible (dorsal face), lateral view. 3, 


Thorax, dorsal view. 4, Apical emargination of S7. 5, Metasoma, T1—T4 (dorsal view). 6, Genitalia (dorsal 


view). 


dian, impressed, longitudinal sulcus in pos- 
terior half; mesosternum with deep, broad 
median sulcus, anterior to arcuate carina in 
front of middle coxa; axilla small, without 
tooth or spine (Fig. 3). Metasoma with TI, 
T2 (Fig. 5) and S2 with median close punc- 
tures, T2 with discal, narrow, longitudinal 


smooth area; SI mostly smooth, with dis- 
tinct, median, longitudinal carina, strongly 
elevated anteriorly into conspicuous tooth; 
S and T3-—7 with dense, small, close punc- 
tures; except 
cell, mostly clear; hindwing with basal half 


forewing infuscated, basal 


mostly clear, apical half infuscated. Para- 


DBP; 


mere as in Fig. 6, penis valve as in Fig. 7; 
cuspis, digitus, short, cuspis reduced to 
rounded lobe much shorter than digitus. 

Holotype.—Female, VENEZUELA, Car- 
abobo State: San Esteban, M. E. Simon col. 
In the Muséum National d°Histoire Natu- 
relle, Paris. 

Distribution —El Salvador, Costa Rica, 
Panama, Colombia, Venezuela, Ecuador 
(see Cambra and Quintero 1996). 

Material examined.—EL SALVADOR: 
Dpto. La Paz, campo experimental de la 
Fac. de Ciencias Agronoémicas, Universidad 
de El Salvador (CAPREX), 2 Nov 1997, J. 
Sermeno, | 2 (MIUP) [new distribution re- 
cord]. PANAMA: Veraguas Prov., Sta. Fe, 
Alto de Piedra, 23—24 Mar 1999, A. Santos, 
1 2 (MIUP); Coloén Prov., Donoso, 21—26 
Jul 2001, P. Gonzalez, 1 2 (MIUP):; Barro 
Colorado Island: 26 April 2001-4, Wcislo 
& Gonzalez, ex nest M. genalis, 5 2,3 3 
(STRI, MIUP); 10 Jan 2001-7, Wcislo & 
Gonzalez, ex nest M. genalis, 1 2 (MIUP); 
Adam Smith, emerged on May 9, into trap 
from nest code AS-20-4-2001-6, ex nest M. 
ecuadoria, 1 2 (STRI); 10 Jan 2001-2, 
Weislo & Gonzalez, ex nest M. ecuadoria, 
1 2 (STRI); 1 June 2001-7, Wcislo & Gon- 
zalez, ex nest M. ecuadoria, 1 2 (STRI): 
26 Apr 2001-3, Wcislo & Gonzalez, ex nest 
M. ecuadoria, 1 2 (STRI). 


NEST PARASITISM 


No immature or adult mutillid wasps 
were Observed in any cells from nests col- 
lected prior to January 2001 (N ~350 
nests). Between January and November 
2001, 4 mutillid wasps were reared from 4 
of 66 M. ecuadoria nests (2.5% of 160 
provisioned cells). Parasitized nests con- 
tained 2—4 sealed cells, and a single adult 
female bee; the percentage of parasitized 
cells per nest for the 4 nests ranged from 
25= 5075 OOS DON) aeSimmilarlys 
~2.1% of M. genalis cells were parasitized 
by L. cincta during this time (8 of 388 cells 
examined from 119 different nests). An 
adult female mutillid was found within a 
M. genalis nest that contained one old cell 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


and four sealed cells, each of which had a 
hole in the cell closure; subsequently adult 
mutillids emerged from sealed cells, con- 
firming the host association. Parasitized M. 
genalis nests had from 1—5 sealed cells, 
and O—2 adult female bees; the percentage 
of parasitized cells per nest for these nests 
ranged from 20—-100% OC = 2%. SD = 
0.34). Lophomutilla cincta emerged from 
a brownish paper-like cocoon that occu- 
pied almost the entire volume of the cell, 
and was attached to the wall via fine pillars 
that appeared to be extensions of the co- 
coon material. Emergence holes of L. cinc- 
ta were located near the cell entrance, and 
were roughly circular and ranged from 
2.8—4.3 mm in diameter. 


DISCUSSION 


The genera Lophostigma and Lopho- 
mutilla Mickel may be related, because in 
both genera the shape of the male mandi- 
ble (Fig. 2) is unique for any known New 
World genera. Males of Lophostigma and 
Lophomutilla key to couplet 34 (Quintero 
and Cambra 1996: 352). However, char- 
acters that they used to separate males of 
these genera are unreliable. Males of the 
two genera differ in the shape of the sev- 
enth sternite, mesosternal lamellate pro- 
cess, and scutellum (in Lophomutilla the 
mesosternal process and the S7 lacks me- 
dioapical emarginations, and the scuteilum 
is convex). 

The percentage of Megalopta cells para- 
sitized by L. cincta is extremely low (2.1— 
2.5% within a season when parasitism was 
observed, and <1% overall), as are para- 
sitism rates for Megalopta in general (Falin 
et al. 2000, Smith et al. 2003, Wcislo et al. 
2004). Nevertheless, the fact that L. cincta 
was reared from various nests of two dif- 
ferent species suggests that Megalopta bees 
are not an accidental host for this species. 
Adult males of L. cincta exhibit no mor- 
phological features typically associated 
with nocturnal activity, such as enlarged 
ocelli or pale body coloration, as known for 
genera such as Photopsis Blake, Odonto- 


VOLUME 107, NUMBER 1 


Figs. 7-8. 


photopis Viereck, and Limaytilla Casal 
(RACT, personal observation). In fact, fe- 
males of this species are frequently ob- 


served during the day on the forest floor or 


walking up plant stems (RACT and WTW, 
personal observation). No host-parasite in- 
teractions were observed, however, and fur- 
ther studies are needed to determine when 
L. cincta females attack hosts, as well as 
other details of their biology. 


ACKNOWLEDGMENTS 


We are grateful to STRI staff for out- 
standing logistical support; José M. Ser- 
meno for loan of a specimen; two anony- 


Lophostigma cincta, male. 7, Penis valve. 


8, Forewing and hindwing (scale bar = | mm). 


mous reviewers and the editor for helpful 
comments; and the Autoridad Nacional del 
Ambiente de la Republica de Panama for 
permission to export material. General re- 
search funds from STRI, as well as support 
from the Smithsonian Institution’s Baird 
Restricted Endowment to WTW, facilitated 
VHG’s participation in the field work 
through STRI’s Behind the Scenes Volun- 
teer Program; additional fieldwork by VHG 
was supported by STRI’s Internship Pro- 
gram. We are grateful to Laura Arneson, 
Adam Smith, Edgardo Garrido, and Her- 
mogenes Fernandez-Marin for their help 
with collecting nests. 


iw) 
ON) 
aS 


LITERATURE CITED 


Brothers, D. J. 1972. Biology and immature stages of 
Pseudomethoca f. frigida, with notes on other spe- 
cies (Hymenoptera: Mutillidae). University of 
Kansas Science Bulletin 50: 1—38. 

. 1995. Mutillidae, pp. 541-548. Jn Hanson, P. 
E. and I. D. Gauld, eds. The Hymenoptera of Cos- 
ta Rica. Oxford University Press, NY, xx + 893 
PP- 

Cambra, R. A. and D. Quintero. 1996. The Mexican 
and Central American species of Lophostigma 
Mickel, including a new species, new distribution 
records, and taxonomic notes for the genus (Hy- 
menoptera: Mutillidae). Pan-Pacific Entomologist 
72: 92-101. 

Cresson, E. T. 1902. Description of some Mutilla from 
Brazil. Transactions of the American Entomolog- 
ical Society 28: 1—82. 

du Buysson, R. 1892. Voyage de M. E. Simon au Ve- 
nezuela, Dec. 1887—April 1888. Hymenopteres. 
Annales de la Société Entomologique de France 
61: 55-59. 

Falin, Z. H., L. C. Arneson, and W. T. Wcislo. 2000. 
Night-flying sweat bees Megalopta genalis and 
Me. ecuadoria (Hymenoptera: Halictidae) as hosts 
of the parasitoid beetle Macrosiagon gracilis (Co- 
leoptera: Rhipiphoridae). Journal of the Kansas 
Entomological Society 73: 183-185. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Leigh, E. G., Jr. 1999. Tropical Forest Ecology. Oxford 
University Press, N.Y., xvi + 245 pp. 

Menke, A. S. 1993. Notauli and parapsidal lines: Just 
what are they? Sphecos 24: 9-11. 

Mickel, C. E. 1928. Biological and taxonomic inves- 
tigations on the mutillid wasps. Smithsonian In- 
stitution United States National Museum Bulletin 
143: 1-351. 

. 1952. The Mutillidae (wasps) of British Gui- 
ana. Zoologica: New York Zoological Society 37: 
105-150. 

Quintero, D. and R. A. Cambra. 1996. Contribucién a 
la sistematica de las mutilidas (Hymenoptera) del 
Pert, en especial las de la Estaci6n Biologica 
BIOLAT, Rio Manu, Pakitza, pp. 327-357. In Wil- 
son, D. E. and A. Sandoval, eds. Manu: The Bio- 
diversity of Southeastern Peru. Washington DC, 
Smithsonian Institution Press, 679 pp. 

Smith, A. R., W. T. Wcislo, and S. ODonnell. 2003. 
Assured fitness returns favor sociality in a mass- 
provisioning sweat bee. Behavioral Ecology and 
Sociobiology 54: 14—21. 

Weislo, W. T., L. Arneson, K. Roesch, V. H. Gonzalez, 
A. Smith, and H. Fernandez-Marin. 2004. The 
evolution of nocturnal behavior in sweat bees, 
Megalopta genalis and M. ecuadoria (Hymenop- 
tera: Halictidae): An escape from competitors and 
enemies? Biological Journal of the Linnean So- 
ciety 83:377-387. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 235-238 


NOTE 


Notes on the Distribution of Anopheles (Anopheles) sinensis Wiedemann (Diptera: 
Culicidae) in China and the Status of Some Anopheles Hyrcanus Group Type 
Specimens from China 


Anopheles (Anopheles) sinensis Wiede- 
mann is an important vector of malarial par- 
asites in China and Korea (Liu et al. 1990, 
Wilkerson et al. 2003). It has a wide distri- 
bution in Asia, including Afghanistan, 
Cambodia, China (north and south parts, 
Hong Kong), Malaysia, India (Assam), In- 
donesia, Japan (Hokkaido, Honshu, Shiko- 
ku, Kyushu, Tsushima, Ryukyu Islands), 
Korea (Korean Peninsula, Cheju Do), 
Myanmar, Singapore, Taiwan, Thailand, Vi- 
etnam (Tanaka et al. 1979, Harrison and 
Scanlon 1975), and Nepal (Darsie and Prad- 
han 1990). Several sources were used to 
document the distribution record of An. si- 
nensis in China. First, LMR examined 
about 400 pinned mosquito specimens in 
the collection of the Institute of Zoology 
Museum, Chinese Academy of Sciences, 
Beijing. These specimens are presently 
housed in more than 50 boxes (most of 
which are 29 X 21 X 5.5 cm in size). Based 
on these collections, about 27 localities 
(counties or towns) in 8 provinces and 1 
city (Beijing) were confirmed for An. sinen- 
sis (Table 1). Secondly, more than a hun- 
dred specimens located in the National Mu- 
seum of Natural History, Smithsonian I[n- 
stitution, and on loan from The Natural His- 
tory Museum, London, identified as An. 
sinensis were examined and confirmed from 
three provinces: Guandong (Guangzhou: 
Daling), Yunnan (Kaiyuan, Kunming); 
Jiangsu (Nanjing, Wuxi). Thirdly, speci- 
mens of An. sinensis deposited in the De- 
partment of Etiologic Biology, Second Mil- 
itary Medical University, Shanghai, China, 
which includes collections from Hainan 
(Sanya and Lingshui), Guandong (Zhuhai) 
and Shanghai, were confirmed. Fourth, the 
Entomology Laboratory of the Jiangsu In- 
stitute of Parasitology has on-going colo- 


nies of An. sinensis, whose origin was 
Wuxi, Jiangsu. Lastly, Ma et al. (1998) con- 
ducted molecular analyses of Chinese mos- 
quitoes and listed one city (Beijing) and 9 
provinces (county or town in parenthesis) 
of China where they collected An. sinensis, 
namely, Fujian (Jianyang), Guizhou (Sin- 
an), Hainan (Dengmai), Henan (Zheng- 
zhou), Jiangsu (Wujing, Wuxi), Liaoning 
(Faku, Shenyang), Shandong (Jining), 
Shanxi (Danfeng), Sichuan (Pixian, Pu- 
jiang), and Yunnan (Simao). The above ob- 
servations reflect a very extensive geo- 
graphical distribution of An. sinensis in 
mainland China, and we report that it is 
now known to occur in 46 locations (coun- 
ties or towns) of 16 provinces and 2 cities 
(Beijing and Shanghai) (Fig. 1). 

According to Yang et al. (1991: 84), the 
holotypes, allotypes, and paratypes of the 
Anopheles (Anopheles) changfus Ma (1981: 
65), An. (Ano.) dazhaius Ma (1981: 65—66), 
An. (Ano.) heiheensis Ma (1981: 66—67) 
and An. (Ano.) xiaokuanus Ma (1981: 67— 
68) are deposited in the Institute of Zoology 
Museum, Chinese Academy of Science. 
The type localities are Sichuan (Emai) and 
Sichuan (unknown town) of the first two 
species, respectively, and Heilongjiang (Ai- 
hui) for the last two species. During a visit 
(LMR) to the Institute of Zoology Museum 
in May 2004, the insect curators, Dr. Xiao- 
Lin Chen and Mr. Jian Yao, attempted to 
find the above types in the museum build- 
ing but could not locate them. Dr. S. EF Ma, 
who described those four species, was also 
unable to give any additional information 
about the location of the specimens (Xiao- 
Lin Chen, personal communication). If 
these specimens prove to be non-extant, 
neotype designations will be necessary to 
ensure stability in the Anopheles Hyrcanus 
Group. 


236 


Table 1. 


emy of Sciences, Beijing, China. 


Box No. Mosquitoes Location 

0269-14 35 @ Beijing: Beijing 
0269-8 I2 Oil Beijing: Pinggu 

12 9 Beijing: Daxing 

I3 OAS Beijing: Yianqing 
0013-5 5 © Zhejiang: Hangzhou 
0013-6 A OF 3 oO Zhejiang: Hangzhou 
0013-21 48 2,76 Zhejiang: Hangzhou 

992,464 Zhejiang: Hangzhou 
0013-14 2 @ Zhejiang: Hangzhou 

1 2 Zhejiang: Hangzhou 

Dey. Zhejiang: Hangzhou 
0013-7 i © Zhejiang: Hangzhou 

1¢ Zhejiang: Hangzhou 
0013-12 il 2 Guangdong: Guangzhou 

DS Guangdong: Guangzhou 
0013-1 4° Guangdong: Guangzhou 

D, QD @ Guangdong: Guangzhou 
0013-10 TY Guangdong: Guangzhou 
0269-19 lg Heilongjiang: Tahe 

DS Guangxi: Longrui (120 m eley.) 

Day © Guangxi: Xiashixiang, Pingxiang (270 m elev.) 
No label 4° Guangxi: Pingxiang (120-270 m elev.) 
0212-23 ) 2 Guangxi: Pingxiang (270 m elev.) 
No label oe Henan: Funiu 
0269-13 3 2 Anhui: Fucunniufang 

3 2 Anhui: Furen 

1 Anhui: Furenfang 

1 2 Anhui: Huangshan 

22 Anhui: Shuifutian 

il @ Anhui: Huangshan 
0212-27 21 Yunnan: Luchunxian (950-1800 m elev.) 
No label 8 2 Yunnan: Luchunxian (950—1800 m elev.) 
0212-7 4 @ Yunan: Manushu, Mengla (700 m elev. 1) 
0212-5 5 @ Yunan: Honjangu, Jiangchengxian (950 m elev.) 
0212-2 3 2 Yunan: Gejiu (1800 m elev.) 

Vg Yunan: Jiangchengxian (950 m elev.) 

3 2 Yunan: Wenshanzhen (1300-1350 m elev.) 
0212-1 2, Q Yunan: Luchunxian (1400 m elev.) 

3 Yunan: Yuanyangxian (1600 m elev.) 
0212-8 3 2 Yunan: Mengla (850—900 m elev.) 
0212-4 4° Yunan: Jiangchengxian (950 m elev.) 

2D, 2 Yunan: Lunchunxian (1400 m elev.) 
0212-22 5 © Xingjiang: Yili 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Collections of Anopheles (Anopheles) sinensis in the Institute of Zoology Museum, Chinese Acad- 


Date 


4-28 June 1957 

23 June 1970 
12-14 August 1970 
24 June 1970 

12 July 1956 

1-16 September 1956 
21-30 June 1956 
7-14 July 1956 

11, 15 June 1956 
22 August 1956 

4 September 1956 
8 July 1956 

2 September 1956 
29 September 1957 
5, 8 October 1957 
24 June 1956 

22 June 1957 
21-30 May 1957 

1 August 1971 

5 December 1986 
12 December 1986 
5, 11 December 1986 
13 December 1986 
16 September 1986 
11 July 1985 

11 July 1985 

7 July 1985 

24 June 1985 

11 July 1985 

24 June 1985 

2 July 1985 

16 September 1986 
20 August 1986 

15 September 1986 
3 August 1986 

31 July 1986 

16 September 1986 
8 August 1986 

6 August 1986 

28 August 1986 

15 August 1986 

9 August 1986 
7-17 August 1957 


We are especially grateful to the follow- 
ing staff of the Institute of Zoology Muse- 
um, Chinese Academy of Sciences, Beijing, 
China: Ai-Ping Liang for his help and hos- 
pitable arrangement of the museum visit; 
Xiao-Lin Chen, Yao Jian, and H. Liu for 
their help in curating the specimens and 


translating data labels of the specimens; and 
Ge-Xia Qiao for initially arranging the mu- 
seum visit. Thanks to Ralph Harbach (De- 
partment of Entomology, The Natural His- 
tory Museum, London, U.K.) for the loan 
of An. sinensis specimens, and James W. 
Jones and Jetsumon Prachumsri (Depart- 


VOLUME 107, NUMBER 1 


237 
75E S0E SSE SOE SE 100E 105E 410E 115E 120E 425E 130E 135E 
| [ 
— 55N 
N dhe es 
China pes 
— 50N 
i 
| 
— 45N 
— 40N 
— 36N 
— 30N 
Collection Sites = 
a Observed = 
e Published 
[__] Provinces 
— 20N 
500 (0) 500 Kilometers 
_— 
— 15N 


Province Identification: Anhui (1), Beijing (2), Fujian (3), Guandong (4), Guizhou (5), Hainan (6), 
Heilongjiang (7), Henan (8), Jiangsu (9), Liaoning (10), Shandong (11), Shanxi (12), Sichuan (13), 
Xingjiang (14), Yunnan (15), Zhejiang (16), Guangxi (17), Shanghai (18). 


Fig. 1. 
specimens). 


ment of Entomology, Armed Forces Re- 
search Institute of Medical Sciences, U.S. 
Army) for their support. Thanks also to 
Tom D. Anderson for preparing the map, 
and Richard C. Wilkerson and Yiau-Min 
Huang for reviewing the manuscript. This 
work was performed under a Memorandum 
of Understanding between the Walter Reed 
Army Institute of Research and the Smith- 
sonian Institution, with institutional support 
provided by both organizations. The opin- 
ions and assertions contained herein are 
those of the authors and are not to be con- 
strued as official or reflecting the views of 
the Department of the Army or the Depart- 
ment of Defense. 


Map of mainland China showing distribution of Anopheles sinensis (based on observed and published 


LITERATURE CITED 


Darsie, R. FE and S. P. Pradhan. 1990. The mosquitoes 
of Nepal: Their identification, distribution and bi- 
ology. Mosquito Systematics 22: 69-130. 

Harrison, B. A. and J. E. Scanlon. 1975. Medical en- 
tomology studies—II. The subgenus Anopheles in 
Thailand (Diptera: Culicidae). Contributions of 
the American Entomological Institute (Ann Ar- 
bor) 12: 1—307. 

Liu, C. EF, H. L. Quian, Z. C. Gu, J. Y. Pan, and X. 
Zheng. 1990. Comparative studies on the role of 


Anopheles anthropophagus and Anopheles sinen- 
sis in malaria transmission in China. Chinese Jour- 
nal of Epidemiology 11: 360-363. 

a, S. E 1981. Studies on the Anopheles (A.) sinensis 
group of mosquitoes in China, including four sib- 


“ 


ling species. Sinozoology 1: 59-74. 
Ma, Y., E Qu, J. Xu, and Z. Zheming. 1998. Sequence 


Nw 
o>) 
ioe) 


differences of rDNA-ITS2 and species-diagnostic 
PCR assay of Anopheles sinensis and Anopheles 
anthropophagus from China. Journal of Medical 
College, People’s Liberation Army 13: 123-128. 

Tanaka, K., K. Mizusawa, and E. S. Saugstad. 1979. 
A revision of the adult and larval mosquitoes of 
Japan (including the Ryukyu Archipelago and the 
Ogasawara Islands) and Korea (Diptera: Culici- 
dae). Contributions of the American Entomologi- 
cal Institute (Ann Arbor) 16: 1—987. 

Wilkerson, R. C., C. Li, L. M. Rueda, H. C. Kim, T. 
A. Klein, G. H. Song, and D. Strickman. 2003. 
Molecular confirmation of Anopheles (Anopheles) 
lesteri from the Republic of South Korea and its 
genetic identity with An. (Ano.) anthropophagus 
from China (Diptera: Culicidae). Zootaxa 378: 1-14. 

Yang, X., H. Sung, and G. Jiang, eds. 1991. Catalogue 
of the insect type specimens preserved in the in- 
sect collections of the Institute of Zoology, Aca- 
demia Sinica. Agriculture Publisher, Beijing. 164 pp. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Leopoldo M. Rueda, Walter Reed Bio- 
systematics Unit, Department of Entomol- 
ogy, Walter Reed Army Institute of Re- 
search, 503 Robert Grant Avenue, Silver 
Spring, MD 20910-7500, U.S.A. (email: 
ruedapol@msc.si.edu), mailing address: 
Walter Reed Biosystematics Unit, MSC 
MRC 534, Smithsonian Institution, 4210 
Silver Hill Road, Suitland, MD 20746; Ya- 
jun Ma and Guang-Hong Song, Depart- 
ments of Etiologic Biology and Parasitol- 
ogy, respectively, Second Military Medical 
University, SOO Xiangyin Road, Shanghai 
200433, People’s Republic of China; and 
Qi Gao, Institute of Parasitic Diseases, 
Meiyuan, Wuxi, Jiangsu 214064, People’s 
Republic of China. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 238 


NOTE 


Tanzaniops, Replacement Name for Certain African Baetidae (Ephemeroptera) 


Tanzaniella was proposed by Gillies 
(1991) for a genus of African Baetidae. That 
generic name, however, was originally used 
for certain African Diplopoda by Hoffman 
(1977). Unfortunately, Mick Gillies died in 
1999 so the original author cannot be given 
the opportunity to rename his homonym, 
which we feel is the ethical practice in re- 
naming homonyms. In lieu of this and as 
active workers on the Ephemeroptera of Af- 
rica, we propose the replacement name Jan- 
zaniops McCafferty and Barber-James (new 
name) [= Tanzaniella Gillies, 1991 (new 
homonym), nec Tanzaniella Hoffman, 
1977]. This baetid genus in Africa is cur- 
rently made up of Tanzaniops gorillorus 
(McCafferty) (new combination) and 7. 
spinosus (Gillies) (new combination), and 
was last treated by McCafferty (2003). 

Acknowledgments.—We thank Michelle 
Hamer, University of KwaZulu-Natal, 
South Africa, for information about the mil- 
lipede genus Tanzaniella. 


LITERATURE CITED 


Gillies, M. T. 1991. A diphlyletic origin for the two- 
tailed baetid nymphs occurring in East African 
stony streams with a description of a new genus 
and species Tanzaniella spinosa gen. nov. sp. 
nov., pp. 175-187. In Alba-Tercedor, J. and A. 
Sanchez-Ortega, eds. Overview and Strategies of 
Ephemeroptera and Plecoptera. Sandhill Crane 
Press, Gainesville. 

Hoffman, R. L. 1977. East African prepodesmid mil- 
lipeds (Chelodesmidae). Revue Zoologique Afri- 
caine 91: 69-82. 

McCafferty, W. P. 2002. Gose’s African Ephemerop- 
tera (Baetidae, Heptageniidae). Entomological 
News 113: 294-302. 


W. P. McCafferty, Department of Ento- 
mology, Purdue University, West Lafayette, 
IN 47907, U.S.A. (e-mail: mcecaffer@ 
purdue.edu), and H. M. Barber-James, De- 
partment of Freshwater Invertebrates, Al- 
bany Museum, Grahamstown 6139, South 
Africa. 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 239-240 


Book REVIEW 


Lepidoptera, Moths and Butterflies, Vol- 
ume 2. Niels P. Kristensen (editor). Hand- 
buch der Zoologie, volume IV, Arthrop- 
oda: Insecta. Series editor Maximilian Fi- 
scher. Walter de Gruyter, Berlin, New 
York, 2003. 564 pp. U.S. $334.80. cloth. 


The publication of the second volume on 
the ‘Lepidoptera, Moths and Butterflies’ 
completes a comprehensive, two volume re- 
view for this insect order, edited by Niels P. 
Kristensen. Whereas volume | (published 
1998) was concerned primarily with sys- 
tematics and evolution, the second volume 
concentrates on the morphology, physiolo- 
gy, and development of the Lepidoptera. 
Together they comprise the most authori- 
tative review ever compiled for one of the 
largest, biologically complex orders of or- 
ganisms. German has been the language tra- 
ditionally used for the volumes within the 
extensive Handbuch series. As a means to 
make the Lepidoptera volumes available to 
a broader audience, the editor, N. P. Kris- 
tensen, arranged to have both parts pub- 
lished in English. Although English was not 
the first language of most of the contribu- 
tors, surprisingly few misspellings were 
noted. A few of these (e.g., butterflie, p. 81) 
may reflect a slip into the author’s native 
tongue. The rather troublesome family 
name Roeslerstammiidae is also misspelled 
(page 547), as it was in volume 1, but is 
spelled correctly on page 550. 

Volume 2 reviews a broad range of sub- 
jects ranging from morphology to embry- 
ology, which are treated in 19 chapters au- 
thored by 28 contributors. In most chapters 
authors were listed alphabetically and not 
according to their relative contribution. The 
chapter titles and authors are: |. Integu- 
ment—Georges Chauvin & Niels P. Kris- 
tensen; 2. ‘Hairs’ and scales—Niels P. Kris- 
tensen & Thomas J. Simonsen; 3. Colora- 
tion: Patterns and morphogenesis—H. 
Frederik Nijhout; 4. Skeleton and muscles: 


adults—Niels P. Kristensen: 5. Skeleton and 
muscles: immatures—Ivar Hasenfuss & 


Niels P. Kristensen; 6. Digestive and excre- 


tory systems—Raymond YV. Barbehenn, & 
Niels P. Kristensen; 7. Respiratory sys- 
tem—Lutz Thilo Wasserthal; 8. Circulation 
and thermoregulation—Lutz Thilo Wasser- 
thal; 9. Nervous system—Jayne E. Yack & 
Uwe Homberg; 10. Sensilla and proprio- 
ceptors—Eric Hallberg, Bill S. Hansson & 
Christer L6fstedt; 11. Auditory and sound 
producing organs—Joél Minet & Annemar- 
ie Surlykke; 12. Eyes and vision—Eric 
Warrant, Almut Kelber & Niels P. Kristen- 
sen; 13. Exocrine glands: Chemical com- 
munication and chemical defense—Eric 
Hallberg and Guy Poppy; 14. Labial glands, 
silk and saliva—Hiromu Akai, Raziel S. 
Hakim, & Niels P. Kristensen; 15. Endo- 
crine glands and hormones—Sridhara Sri- 
dhara, Govindan Bhaskaran, & Karl H. 
Dahm; 16. Reproductive organs—Niels P. 
Kristensen; 17. Karyology and sex deter- 
mination—Jurate De Prins & Kazuo Saitoh; 
18. Eggs—-Heinz Fehrenbach; 19. Embry- 
ology—Yukimasa Kobayashi, Masahiro Ta- 
naka, & Hiroshi Ando. 

In addition to being the editor of this vol- 
ume, Kristensen was also the primary con- 
tributor, having authored or co-authored 
eight chapters. The two chapters reviewing 
the skeleton and muscles (4, adults, pages 
39—122 and 5, immatures, pages 123-164) 
probably best reflect Kristensen’s expertise 
and previous extensive contributions to the 
subject. Together these chapters comprise 
nearly 23%, or 125 pages, of the entire text. 
All chapters are well illustrated with de- 
tailed drawings and photographs that clear- 
ly aid in understanding the text. 

A common theme prevalent throughout 
the book is the presentation of the various 
topics in a phylogenetic context. Systema- 
tists should find such summaries, as shown 
in graphic form in chapters 18 (Eggs) and 


240 


19 (Embryology), especially informative. 
An appendix near the end of the volume, 
including a cladogram of the extant super- 
families (slightly revised from Fig. 2.2 of 
volume |) and a list of the subfamily clas- 
sification for Lepidoptera, provides a most 
useful reference in this regard. One minor 
error involving indentation of the family 
name Eriocottidae was noted on page 545. 
Although the subject index excludes taxo- 
nomic names, an index to the generic 
names used in the text, together with their 
superfamily—family—subfamily classifi- 
cation is provided in appendix 2. 

The only instance of subject duplication 
noted was in chapters 18 (Eggs), figure 18.5 
and 19 (Embryology), figure 19.2, where 
the same figure was used. In this instance 
it is probably unavoidable because of the 
inherent difficulty in discussing later egg 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


development without some mention of egg 
wall structure. 

This volume has been sorely needed for 
some time. The editor and all of the con- 
tributors are to be congratulated for provid- 
ing a well summarized update on many as- 
pects of Lepidoptera biology that will ben- 
efit both students and professional ento- 
mologists. Unfortunately the price of this 
reference ($334.80 US, amazon.com) will 
undoubtedly be beyond the budgets of most 
individuals as well as some libraries. Hope- 
fully, enough institutions can acquire this 
reference to make it readily available to a 
broad spectrum of users. 


Donald R. Davis, Department of Ento- 
mology, NHB 127, Smithsonian Institution, 
P.O. Box 37012, Washington, D.C., 20013- 
7012, U.S.A. (e-mail: davis.don@nmanh.si. 
edu) 


PROC. ENTOMOL. SOC. WASH. 
107(1), 2005, pp. 241-243 


Book REVIEW 


The Black Flies (Simuliidae) of North 
America. Peter H. Adler, Douglas C. Cur- 
rie, and D. Monty Wood. 2004. 941 pp. 
$99.95 (£57.95). Cornell University 
Press, Ithaca, NY. ISBN 0-8014-2498-4. 


The authors of “‘The Black Flies (Simu- 
liidae) of North America” have a combined 
100 years of experience working on this 
family and have harnessed their collective 
energies and expertise, as well as that of 
two excellent illustrators (Ralph M. Idema 
and Lawrence W. Zettler), in the production 
of this book. A phenomenal volume of de- 
tailed information is included in this large 
format (8.5 by I1 inch pages), 941 page 
opus magnum. The book covers 254 black 
fly species found in North America north of 
Mexico, defined by the authors as the Unit- 
ed States, Canada, and Greenland. 

The authors’ intention is to bring black 
flies into the realm of familiarity held by 
many popular insects, such as the Lepidop- 
tera. The underlying theme of the book is 
the accurate identification of black fly spe- 
cies based on morphology, cytology, distri- 
bution, and ecology. Several chapters are 
geared to general readership and intended 
to be useful for medical-veterinary ento- 
mologists, aquatic biologists, environment 
consultants, systematists, natural history en- 
thusiasts, pest management specialists, and 
students. The book is divided into four ma- 
jor parts: Background, Biology, Economic 
Aspects, and Systematics and Taxonomy, 
plus a huge reference section. 

Part 1, ““‘Background”’ includes three 
chapters, Overview, History of Research, 
and Techniques for Collection, Preparation, 
and Curation. The Overview chapter is a 
combination of information that places the 
Simuliidae in perspective for the layman and 
scientist alike. The chapter introduces black 
flies, what they are, where they live, and dif- 
ficulties encountered with identification 
based on morphology. It also presents the 


dichotomy of the family: as aquatic organ- 
isms, black fly larvae are viewed as being 
beneficial, yet as blood-feeding adults they 
are regarded as despised pests. An extensive 
table in this chapter is devoted to common 
names used for black flies in the United 
States and Canada. The History chapter cov- 
ers a 240 year period in North American si- 
muliid study beginning in the 1800’s with 
the description of the first black fly species 
in the United States by Thomas Say in 1823 
and covers other periods in the study of 
black flies up to the “Contemporary Era” 
(1970’s until the present). The remaining 
chapter in Part | provides useful technical 
information for collection of live specimens 
and their proper preservation for future ex- 
amination. It also includes a section on cy- 
totaxonomic procedures. 

Part 2, Biology, comprises three essential 
chapters for those who will study black 
flies, be it preserved specimens or living 
creatures in their natural habitats. The chap- 
ter on structure and function provides de- 
tailed illustrations and descriptions on the 
anatomy and morphology of larval, pupal, 
and adult simuliids. The chapter on cytol- 
ogy acquaints the reader with polytene 
chromosomes, chromosomal complement, 
chromosomal rearrangements, nomencla- 
ture, and how they are used in taxonomy. 
Their importance in this book is especially 
apparent. One-quarter of the 254 species 
treated in the book were revealed through 
chromosomal study. 

A chapter on behavior and ecology 
rounds out Part 2 and provides a good over- 
view of these subjects for one becoming ac- 
quainted with the family. The chapter in- 
cludes sections on distribution and habitat, 
oviposition behavior, development of im- 
mature stages, flight and longevity, labora- 
tory colonization, mating behavior, adult 
feeding behavior (with detailed tables on 
mammalian and avian hosts in North Amer- 


242 


ica), and natural enemies and symbionts 
(including nematodes, pathogens, ectopar- 
asites, and predators). Seven tables in the 
natural enemy section list the various nem- 
atodes and microbes found in larval and 
adult black flies. 

Part 3 consists of two chapters that pro- 
vide an overview of economic aspects of 
the Simuliidae in the United States and 
Canada. Chapter 7, Social and Economic 
Impact, covers factors promoting pest prob- 
lems; biting and nuisance problems for hu- 
mans and domestic and wild animals (in- 
cluding a table listing pests of humans, live- 
stock and poultry, and descriptive photo- 
graphs depicting pestiferous black flies); 
economic losses; and vector-borne diseases 
of birds and mammals. Chapter 8 provides 
a synopsis of practices employed for the 
management of larval and adult black flies. 
These include chemical control methods for 
larval abatement ranging from oils and oth- 
er early compounds, to DDT, and replace- 
ments for DDT and nontraditional com- 
pounds, especially insect growth regulators. 
A chronology of insecticide use dating from 
the employment of DDT as a larvicide and 
adulticide in the late 1940’s to the softer 
microbial insecticide, Bacillus thuringiensis 
var. israelensis from 1979 until the present 
is provided in tabular format. The chapter 
also includes sections on physical control, 
biological control, and personal protection 
(natural preventatives, synthetic repellents, 
and alternatives to repellents). 

Part 4, Systematics and Taxonomy, is the 
largest (729 pp.) and most detailed section 
of the book and contains two chapters: 
Chapter 9, Phylogeny and Classification of 
Holarctic Black Flies, and Chapter 10, Syn- 
optic List, Identification Keys, and Taxo- 
nomic Accounts of North American Black 
Flies. The chapter on phylogeny and clas- 
sification provides the logical basis for the 
classification used in the book. It comprises 
brief sections on historical context, higher 
classification of black flies, other taxonomic 
contributions, phylogenetic studies of the 
Simuliidae and molecular systematics of the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Simuliidae and a rather detailed coverage of 
the phylogeny of the Holarctic Simuliidae. 
In this section, the authors provide phylo- 
genetic trees at the species-group level and 
evidence for the monophyly and relation- 
ships of the family in the Holarctic Region. 
The phylogenetic reconstruction includes 
all Palearctic and many Neotropical taxa. In 
Chapter 10, the authors provide a variety of 
tools for the taxonomist, public health en- 
tomologist, or aquatic ecologist for the rec- 
ognition of the 254 species of black flies 
found in North America north of Mexico. 
The synoptic list contains not only the 
names of extant species, but also includes 
new nomenclatural acts which are shown in 
bold font. Forty-three new species are in- 
cluded in the list and are described by the 
authors in the Taxonomic Accounts section 
of the chapter. Comprehensive keys are fur- 
nished for the identification of adult males 
and females, pupae, and larvae of North 
American species. This section is richly il- 
lustrated and includes detailed drawings of 
major anatomical features of adults (espe- 
cially male and female terminalia), pupae, 
and larvae. Twenty-two color plates of lar- 
vae and two of adult scutal patterns provide 
additional aides for the identification of 
species. The individual taxonomic accounts 
for each species includes taxonomy, mor- 
phology, physiology, cytology, and bio- 
nomics information, especially regarding 
habitats, hosts, and economic importance. 
Some of the accounts include information 
on molecular systematics. Vernacular 
names, misidentifications, erroneous spell- 
ings, and additional details are also provid- 
ed. Distributions of the species in the Unit- 
ed States and Canada are individually 
shown on 254 maps. Capping the utility of 
this panoramic opus is a reference section 
containing over 2200 references spanning 
240 years of research. 

Richly illustrated and extensively refer- 
enced, this book will be the most definitive 
work on the North American Simuliidae 
North of Mexico for decades to come. It is 
a must for anyone working with the family 


VOLUME 107, NUMBER 1 


and will be a useful reference in the broader 
subject areas of aquatic ecology and medi- 
cal entomology. Considering the affordable 
price and sheer volume of information, it is 
a bargain. 


243 


Lawrence A. Lacey, Yakima Agricultur- 
al Research Laboratory, USDA-ARS, 
5230 Konnowac Pass Road, Wapato, WA 
98951, U.S.A. (e-mail: llacey@yarl.ars. 
usda. gov) 


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SCHEDULE OF THE REGULAR MEETINGS FOR 2004-2005 


October 7, 2004. Fred Paras (Baltimore City Community College and Maryland 
Entomological Society). A Different Olympiad: Mountain Hopping in 
Northern Greece from Mt. Gramos and Varnous across to Mt. Rhodopi 


November 4, 2004. John Lapolla (Department of Entomology, Smithsonian 
Institution). The Rise of Ranching in the Ants and the Evolution of Tro- 
phophoresy 


December 2, 2004. Sean Brady (Department of Entomology, Smithsonian In- 
stitution). Uncovering the Evolutionary History of Ants. 


January 6, 2005. Eric Grissell (Systematic Entomology Laboratory, U.S. De- 
partment of Agriculture). A Wild Rhus Chase, or How to Nearly Solve a 
Problem in 40 Years with Diversions Along the Way 


February 3, 2005. Daniel Perez (Department of Entomology, Smithsonian In- 
stitution). Survey of the Orthopteroid Insects of Hispaniola 


March 3, 2005. Edd Barrows (Department of Biology, Georgetown University). 
*‘Arthroversity”’ of the Capital Area 


April 7, 2005. Michael Engel (Department of Entomology, University of Kan- 
sas). 600,000,000 Years on Six Legs 


May 5, 2005. Irina Brake (GBIF Postdoctoral Fellow, Department of Ento- 
mology, Smithsonian Institution). Milichiid Fly Systematics and Biology 


Meetings held at: 
7:00 PM 
Cathy Kerby Seminar Room (CE-340) 
National Museum of Natural History 
10th & Constitution Ave., Washington, D.C. 


Meetings are open to the public. 
Please bring new literature, specimen demonstrations, ento-T-shirts, etc. 


Pre-meeting dinner at the R. Reagan Building Food Court, 5:30 PM. 
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David G. Furth and John Brown, Co-Program Chairs 


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 


Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae), by Alexander S. 
Konstantinov and Steven W. Lingafelter. 349 pp. 2002 


Revision of the Genus Anoplophora (Coleoptera: Cerambycidae), by Steven W. Lingafelter and E. 
Richard Hoebeke. 236 pp. 2002 


MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available. 
No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 
No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 
No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 _.. 


No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi 
Takahasi. 230 pp. 1969 


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No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 
OT DP WO SA yes 8S cae S AN ed WER ie eC eva eral Uo Aa Se eee ed RN 


No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 
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No. 15. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent 
ANG WEN GroOcane Tel OS py OOS pia ieee Ae gk NCE ie Er eae ne de 


No. 16. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 __ 


No. 17. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited 
Jay IB ABR UN arco Si aleeroval ANE rina ke alee ral ayyoye) SIS Ioye ek eve ee teh As OP eR ee 


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ATC MLCT PMNS NETO Sel OO Bites eae Ea Ly CAR aS Sey MeO Na SIE A et Ss Ca ht a 


No. 21. New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G. 
imitate trl ROM 0) olp ILSS} ee ONUeD etek anos ante Mee aD Men Leena Monn ere Se emia oo Ae TEL a 


No. 22. Systematics of the North American Species of Trichogramma Westwood (Hymenoptera: 
imehorrammaticae), by: Jobn DD, Pinto 287 pp L999 ss ne 


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INSTITUTION LIBRARIES 


wii 


01117 6104 


CONTENTS 
(Continued from front cover) 
MILLER, DOUGLASS R., GARY L. MILLER, GREG S. HODGES, and JOHN A. 
DAVIDSON—Introduced scale insects (Hemiptera: Coccoidea) of the United States and their 
impact.on (Ws:S: Aonionlliaire es eer ae oe eee SL Nv clade ae i RataMar ts aet-y ata chaverae es rete a etapa tee 


NEUNZIG, H. H. and M. A. SOLIS—Tumoriala, a new Neotropical phycitine genus 
(epidoptera:Pyralidae) ee muha elena ataevet eis\c/jolatere aie cee Ree eiey ey Teta Siete leR Tar A ete 


NOVIKOV, D. V., G. A. ANUFRIEV, and C. H. DIETRICH—New genera and species of 
leafhoppers (Hemiptera: Cicadellidae) from Kyrgyzstan ........... 0... e cece eee ee eee ect ee eee 


POINAR, GEORGE, JR.—Fossil Trigonalidae and Vespidae (Hymenoptera) in Baltic amber .... 


RANDOLPH, R. P. and W. P. McCAFFERTY—The mayflies (Ephemeroptera) of Alaska, 
InclNGIne amMew Species Ol Meptagemildae waren Me wuiwe ie nieise isch tere ec. acy fetal 2 eer ae aioe 


RIDER, DAVID A. and LE-YI ZHENG—Checklist and nomenclatural notes on the Chinese 
Pentatomidae (Heteroptera). III. Phyllocephalinae, Podopinae ...................2-.2-.00.05:- 


SCHAUFF, MICHAEL E.—Ammonoencyrtus carolinensis, n. comb. (Hymenoptera: Encyrtidae), 
a parasite of lobate lac scale Paratachardina lobata (Chamberlin) (Hemiptera: Kerriidae) .. 


SMITH, DAVID R.—Review of the genus Acordulecera Say (Hymenoptera: Pergidae) of the West 
Indies, and the first records of Symphyta from Montserrat and St. Kitts ...................... 


SMITH, DAVID R.—A new sawfly (Hymenoptera: Pergidae) feeding on guava, Psidium guajava 
i (Myrtaceae);)in' Costa Rica yan vey eye aka eerie er ila ch eth Sele eateries bet sere eae ay ete ee 


SPINELLI, GUSTAVO R., PABLO I. MARINO, and MARIA M. RONDEROS—The fourth 
instar larva and pupa of the Neotropical biting midge Forcipomyia (Forcipomyia) rioplatensis 
Marino and) Spinelli: (Diptera? Ceratoposonidac)s eee eee eee oe ner rs ste tele sheet eater 

TOGASHI, ICHIJI—Description of a new species of Eriotremex Benson (Hymenoptera: 
Sucidae) froma Japa) as 6 ae Oe Ne cs A NOIR AT UW ac eg 


WHEELER, A. G., JR. and THOMAS J. HENRY—Description of the adult and fifth instar of a 
myrmecomorphic plant bug, Bicuspidatiella conica Maldonado (Hemiptera: Miridae: 
Deracoconnas). with metesvombits abies eyes eee ee sess irs e ays oe eya yeas Hsien alc sects eu eiaene 


YANG, DING and STEPHEN D. GAIMARI—Review of the species of Elaphropeza Macquart 
(Diptera: Empididae: Tachydromiinae) from the Chinese mainland .......................... 


ZHANG, LI-JIE and XING-KE YANG—A new species of the genus Agetocera Hope 
(Coleoptera: Chrysomelidae: Galerucinae) from China ................0.. 02.00. 2 ese eee eee eee 


ZHANG, YALIN and WU DAI—A taxonomic review of Matsumurella Ishihara (Hemiptera: 
Cicadellidac; Deltocephalinae); from China sees Aa raaeeite ee eeer ieee Mee MAME TA ar 


NOTE 


RUEDA, LEOPOLDO M., YAJUN MA, QI GAO, and GUANG-HONG SONG—Notes on the 
distribution of Anopheles (Anopheles) sinensis Wiedemann (Diptera: Culicidae) in China and 
the status of some Anopheles Hyrcanus Group type specimens from China.................. 


McCAFFERTY, W. P. AND H. M. BARBER-JAMES—YTanzaniops, replacement name for 
certain African) Baetidae (Ephemeroptera) ss). s4 se eae ceases Sees nel eats oes eee 


BOOK REVIEWS 


DAVIS, DONALD R.—Lepidoptera, Moths and Butterflies, Volume 2, Niels P. Kristensen, 
ELON C6) Cay Hea Oa LES is ane UL a ater EO AR MMM RRO RM Ane AINE Aroma ean Geen os Arte 


LACEY, LAWRENCE A.—The Black Flies (Simuliidae) of North America, by Peter H. Adier, 
DouglasiC.iCurne, ands Monty? WOOd Ay 32 yee caer eae pL coat eal eag elec ae tol seen arate 


MISCELLANEOUS 
schedule of theiResular Mectines; tor 2004-2005 sie ys we ete keto etek eae te eee taleteeletebal tere Da 


i 


123 


84 


21 
55 


190 


90 


115 


99 


214 


108 


159 


209 


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119 


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235 


238 


239 


OL 
oy  PROCEEDIN 
EIt afilie 


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PUBLISHED 
QUARTERLY 


CONTENTS 


BRAILOVSKY, HARRY—A revision of the genus Acanthotyla Stal, with the description of five 
new species and synonymical note (Heteroptera: Coreidae: Colpurini)...................... 


GARCIA ALDRETE, ALFONSO N.—A new ptiloneurid genus (Psocoptera: Ptiloneuridae) 
SOMA DOLMMICA ee ee ee ear UG OUR ts ue are A RUT LB Ree, SAPS V RN telat acta ctattiajels Nalolrtavaiant oiste 


GODOY, CAROLINA—A new genus of brachypterous leafhoppers (Hemiptera: Cicadellidae: 
@readellinaeProcomiinn) irom CostaiwRican. 5): wees somes alton hoch! hei sistsls alalouece. pce 


GRISSELL, E. E. and G. F HEVEL—First report of Theocolax ingens Xiao and Huang 
(Hymenoptera: Pteromalidae) in the Western Hemisphere, with a synopsis of the genus .... 


HASTRITER, MICHAEL W. and MICHAEL F. WHITING—Records of fleas (Siphonaptera) of 
CATMIVOLES MUO LAO en vie Se Mia ae ys Peale a 898) Se ierarentl Eeyore Brava Voc tents Yalgs Rots tA SM AB cote Dacca 


HOEBEKE, E. RICHARD and A. G. WHEELER, JR.—Establishment of three European flea bee- 
tles in Nova Scotia: Longitarsus ganglbaueri Heikertinger, L. jacobaeae (Waterhouse), 
and L. rubiginosa (Foudras) (Coleoptera: Chrysomelidae: Alticinae).................... eee. 


HUANG, MIN and YALIN ZHANG—Two new leafhopper species of Bolanusoides Distant 
(Hemiptera: Cicadellidae: Typhlocybinae: Typhlocybini) from China ....................-. 


MATHIS, WAYNE N. and MANUEL A. ZUMBADO—Description of Scatella savegre, a new 
species from Costa Rica in the 7riseta Group (Diptera: Ephydridae)......................-. 


NEUNZIG, H. H. and M. A. SOLIS—A review of the Neotropical genus Difundella Dyar 
(iepidopterasibyralidaePiycittmAc) ti hire cece tce ciate cite ~ avira palate aible bia = aie arole eye's a\elcvalele(s ols ince 


PEREDO, LUIS CERVANTES and SAGRARIO GAMEZ-VIRUES—Three species of 
facultative Myodochini (Lygaeoidea: Rhyparochromidae) associated with figs in Mexico .... 
POINAR, GEORGE, JR.—A Cretaceous palm bruchid, Mesopachymerus antiqua, n. gen., n. sp. 


(Coleoptera: Bruchidae: Pachymerini) and biogeographical implications .................. 


ROBBINS, RICHARD G.—The ticks (Acari: Ixodida: Argasidae, Ixodidae) of Taiwan: 
ASSN NOUN TIT GE CHECKIRS EE oe CiN atamcniy okie verze sr NaR) entitle’, <istivlers dh Sehe Slelataieiars oe '<\e Ki¥hniee s ereet ales =) ae 


(Continued on back cover) 


APR 1 5 2009 


AR\ts 


NO. 2 


(ISSN 0013-8797) 


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PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 245-253 


THE TICKS (ACARI: IXODIDA: ARGASIDAE, IXODIDAE) OF TAIWAN: 
A SYNONYMIC CHECKLIST 


RICHARD G. ROBBINS 


Defense Pest Management Information Analysis Center, Armed Forces Pest Manage- 
ment Board, Walter Reed Army Medical Center, Washington, DC 20307-5001, U.S.A. 
(e-mail: richard.robbins @ osd.mil) 


Abstract.—Eleven checklists of Taiwanese ticks, published or privately circulated be- 
tween 1935 and 1984, are compared with specimen records, chiefly in the U.S. National 
Tick Collection, yielding a list of 32 tick species thought to definitely occur in Taiwan 
and its adjacent islets: Argas pusillus Kohls, A. robertsi Hoogstraal, Kaiser & Kohls, 
Ornithodoros capensis Neumann, Amblyomma cordiferum Neumann, A. geoemydae (Can- 
tor), A. helvolum Koch, A. testudinarium Koch, Aponomma varanense (Supino), Boophilus 
microplus (Canestrini), Dermacentor taiwanensis Sugimoto, Haemaphysalis bandicota 
Hoogstraal & Kohls, H. canestrinii (Supino), H. doenitzi Warburton & Nuttall, H. flava 
Neumann, H. formosensis Neumann, H. hystricis Supino, H. kitaokai Hoogstraal, H. ma- 
geshimaensis Saito & Hoogstraal, H. ornithophila Hoogstraal & Kohls, H. phasiana Saito, 
Hoogstraal & Wassef, H. taiwana Sugimoto, H. yeni Toumanoff, /xodes acutitarsus 
(Karsch), /. granulatus Supino, I. kuntzi Hoogstraal & Kohls, 1. nipponensis Kitaoka & 
Saito, J. ovatus Neumann, J. persulcatus Schulze, I. simplex Neumann, I. vespertilionis 
Koch, Rhipicephalus haemaphysaloides (Supino), and R. sanguineus (Latreille). An ad- 
ditional 13 tick species previously believed to occur in Taiwan are excluded from this 
list. 


Key Words: _ ticks, Ixodida, checklist, synonymy, Taiwan 


At least 11 papers and reports intended 
to serve as checklists of the ticks of Taiwan 
(formerly Formosa) and vicinity were pub- 
lished or privately circulated during the 20" 
century (Kishida 1935, Schulze 1935, Ogu- 
ra 1936, Sugimoto 1939, Elishewitz 1943, 
Anonymous 1944, Luh and Woo 1950, Shi- 
mada et al. 1961, Maa and Kuo 1966, Tseng 
1978, and Hoogstraal letter no. 251, 14 
February 1984, Smithsonian Institution Ar- 


consigned to the junior synonymy. With the 
recent emergence of Lyme borreliosis and 
human babesiosis in Taiwan (Shih and 
Chao 1998, 1999; Shih et al. 1997, 1998), 
it is imperative that these lists be reconciled 
with current taxonomic literature and avail- 
able specimen records to produce a single 
reliable roster of the Taiwan tick fauna. 
Between 1955 and 1978, teams from 
U.S. Naval Medical Research Unit Number 


chives). However, when these lists are crit- 
ically compared, it soon becomes apparent 
that there are numerous disagreements 
among them. Most are also rife with no- 
menclatural or typographical errors or cite 
species names that have long since been 


Two (NAMRU-2), Taipei, scoured Taiwan 
and its nearby islets (Lutao or Green Island, 
Lanyu or Orchid Island, and the Penghu ar- 
chipelago), collecting vertebrates and inver- 
tebrates “‘to provide the biological knowl- 
edge required for controlling the animal and 


246 


insect vectors of area diseases” (Secretary 
of the Navy Notice 5450, 9 May 1955). 
Over 1,900 tick collections, each containing 
from one to hundreds of specimens, were 
shipped to the laboratory of the late Harry 
Hoogstraal (1917-1986), then Head, Med- 
ical Zoology Department, U.S. Naval Med- 
ical Research Unit Number Three (NAM- 
RU-3), Cairo, Egypt, for identification and 
evaluation. After Hoogstraal’s death, this 
vast resource was merged with the U.S. Na- 
tional Tick Collection (USNTC), now lo- 
cated at the Institute of Arthropodology and 
Parasitology, Georgia Southern University, 
Statesboro, while the voluminous corre- 
spondence pertaining to the Taiwan collec- 
tions was archived at a Smithsonian Insti- 
tution records center in Springfield, Virgin- 
ia. 

Since 1986, I have had several opportu- 
nities to examine critical Taiwanese speci- 
mens in the USNTC. With the assistance of 
Smithsonian archivists, I have also been 
able to peruse and photocopy key portions 
of the Hoogstraal correspondence files. The 
following tabulation comprises all 30 Tai- 
wan tick species represented in the USNTC, 
plus two species known from Taiwanese 
specimens in the Bernice P. Bishop Muse- 
um (BPBM), Honolulu, Hawaii. Each spe- 
cies name appears in boldface, accompa- 
nied by 1) the total number of Taiwan-area 
collections of that species in, or recorded 
by, the USNTC or BPBM (some collections 
were not retained); 2) where applicable, a 
note on recent (and as yet controversial) 
classificatory changes advocated by Horak 
et al. (2002); and 3) a summary of all junior 
synonyms in earlier Taiwan checklists, or 
relevant remarks. I also comment on 13 tick 
species that, to my knowledge, do not occur 
in Taiwan, despite published reports or ex- 
tant specimens suggesting their presence. 


TICK SPECIES KNOWN TO OCCUR IN TAIWAN 
AND ADJACENT ISLETS 
Family Argasidae 


Argas pusillus Kohls, 1950.—13 Taiwan 
collections in USNTC. Now also classified 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


as Carios pusillus (Kohls, 1950) (Horak et 
al. 2002). 

Junior synonymy in Taiwan checklists: 
None. Published references to A. vesperti- 
lionis (Latreille, 1796) in Taiwan (Kishida 
1935, Sugimoto 1939, Shimada et al. 1961, 
Maa and Kuo 1966, Tseng 1978) may rep- 
resent A. pusillus. Hoogstraal (letter no. 
251, 14 February 1984) lists A. vesperti- 
lionis as occurring in Taiwan even though 
he earlier (letter no. 376, 14 February 1977) 
concluded that ‘‘all A. vespertilionis group 
samples from Taiwan ... represent A. pus- 
illus.’ On my copy of the Hoogstraal 
checklist, the words “‘pusillus Kohls”’ have 
been hand-entered above the typed listing 
for A. vespertilionis, but the latter name has 
not been altered. 


Argas robertsi Hoogstraal, Kaiser & 
Kohls, 1968.—1 Taiwan collection in 
USNTC. 

Junior synonymy in Taiwan checklists: 
None. Published references to A. persicus 
(Oken, 1818) from Taiwan (Kishida 1935, 
Anonymous 1944, Luh and Woo 1950, Shi- 
mada et al. 1961, Maa and Kuo 1966, Tseng 
1978) appear to be recapitulations of a mis- 
determined Argas. Tseng’s (1978) reference 
to “Argas sp. nr. arboreus Kaiser, Hoogs- 
traal & Kohls, 1964” (an African species) 
may represent the morphologically similar 
A. robertsi. See Hoogstraal et al. 1968a, 
OWS ae: 


Ornithodoros capensis Neumann, 
1901.—3 Taiwan collections in USNTC. 
Now also classified as Carios capensis 
(Neumann, 1901) (Horak et al. 2002). 

Junior synonymy in Taiwan checklists: 
None. Early references to an “‘“Ornithodoros 
sp.” in Taiwan (Kishida 1935, Maa and 
Kuo 1966) may apply to this species, but 
the host listed by these authors, domestic 
swine, would be atypical for O. capensis, 
which in Taiwan is known only from sea- 
birds. 


Family Ixodidae 


Amblyomma cordiferum Neumann, 
1899.— 31 Taiwan collections in USNTC. 


VOLUME 107, NUMBER 2 


Junior synonymy in Taiwan checklists: 
None. Listed only by Hoogstraal (letter no. 
251, 14 February 1984). Voltzit and Keir- 
ans (2002) stated that this species occurs in 
Taiwan. 


Amblyomma _ geoemydae_ (Cantor, 
1847).—3 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
Amblyomma formosanum Schulze, 1933, 
probably a junior synonym of A. geoemy- 
dae (see Camicas et al. 1998), is the name 
used for this species in the checklists of 
Schulze (1935), Elishewitz (1943), Anony- 
mous (1944), Luh and Woo (1950), and 
Shimada et al. (1961). 


Amblyomma helvolum Koch, 1844.—18 
Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Not in previous checklists, but 
Hoogstraal et al. (1968b) and Kolonin 
(1995) stated that this species occurs in Tai- 
wan. 


Amblyomma _testudinarium Koch, 
1844.—46 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
Amblyomma yajimai Kishida, 1935 (Kishi- 
da 1935, Elishewitz 1943, Anonymous 
1944, Luh and Woo 1950, Shimada et al. 
1961, Maa and Kuo 1966); A. infestum tai- 
vanicum Schulze, 1935 (Schulze 1935, Eli- 
shewitz 1943, Luh and Woo 1950, Shimada 
et al. 1961). Note that Elishewitz (1943), 
Luh and Woo (1950) and Shimada et al. 
(1961) list both junior synonyms. As well, 
Elishewitz (1943), Anonymous (1944), Luh 
and Woo (1950), Shimada et al. (1961) and 
Maa and Kuo (1966) list the senior syno- 
nym A. testudinarium. 


Aponomma varanense (Supino, 
1897).—3 Taiwan collections in USNTC. 
Now also classified as Amblyomma vara- 
nense (Supino, 1897) (Horak et al. 2002). 

Junior synonymy in Taiwan checklists: 
None. First reported from the Penghu ar- 
chipelago by Robbins (1996), who de- 
scribed this species’ convoluted taxonomic 
history. Elishewitz (1943) lists Amblyomma 


247 


serpentinum Schulze, 1936, a junior syno- 
nym of Aponomma fuscolineatum (Lucas, 
1847), from Taiwan, but A. varanense is the 
only Aponomma known from bona fide Tai- 
wan-area specimens. 


Boophilus  microplus_ (Canestrini, 
1888).—932 Taiwan collections in USNTC. 
Now also classified as Rhipicephalus (Bo- 
ophilus) microplus (Canestrini, 1888) (Hor- 
ak et al. 2002). 

Junior synonymy in Taiwan checklists: 
An abundant and widespread species in Tai- 
wan, B. microplus has been recorded under 
numerous junior synonyms: B. annulatus 
australis (Fuller, 1899) (Elishewitz 1943, 
Anonymous 1944); B. annulatus caudatus 
(Neumann, 1897) (Kishida 1935, Ogura 
1936, Elishewitz 1943, Anonymous 1944); 
B. australis (Fuller, 1899) (Luh and Woo 
1950); B. caudatus (Neumann, 1897) (Luh 
and Woo 1950, Shimada et al. 1961); B. 
(Uroboophilus) distans Minning, 1934 
(Schulze 1935, Elishewitz 1943, Anony- 
mous 1944, Luh and Woo 1950, Shimada 
et al. 1961); B. (Uroboophilus) sinensis 
Minning, 1934 (Schulze 1935, Elishewitz 
1943, Luh and Woo 1950); Margaropus an- 
nulatus australis (Fuller, 1899) (Sugimoto 
1939); and M. annulatus caudatus (Neu- 
mann, 1897) (Sugimoto 1939). Note that 
several checklist authors list more than one 
synonym. In addition, Sugimoto (1939) 
lists M. annulatus (Say, 1821), a junior syn- 
onym of B. annulatus (Say, 1821), which 
does not occur in Taiwan. 


Dermacentor taiwanensis Sugimoto, 
1935.—76 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
Indocentor bellulus Schulze, 1935 (Schulze 
1935, Elishewitz 1943, Anonymous 1944); 
Dermacentor bellulus (Schulze, 1935) (Luh 
and Woo 1950, Shimada et al. 1961, Maa 
and Kuo 1966, Tseng 1978). 


Haemaphysalis bandicota Hoogstraal 
& Kohls, 1965.—15 Taiwan collections in 
USNTC. 

Junior synonymy in Taiwan checklists: 


248 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


None. Listed only by Tseng (1978) and 
Hoogstraal (letter no. 251, 14 February 
1984). Thought to have been introduced 
with bandicoot rats, Bandicota indica 
(Bechstein) (Rodentia: Muridae), during the 
Dutch occupancy of Taiwan (1624-1662) 
(Hoogstraal and Kohls 1965, Hoogstraal 
and Wassef 1973). 


Haemaphysalis canestrinii (Supino, 
1897).—11 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Listed only by Hoogstraal (letter no. 
251, 14 February 1984), who earlier re- 
corded this species from Taiwan (Hoogs- 
traal 1971). 


Haemaphysalis doenitzi Warburton & 
Nuttall, 1909.—22 Taiwan collections in 
USNTC. 

Junior synonymy in Taiwan checklists: 
None. Not in previous checklists, but 
Hoogstraal and Wassef (1973) listed nu- 
merous collections of this species from Tai- 
wan and Lanyu. 


Haemaphysalis flava Neumann, 
1897.—4 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. This species appears in all previous 
checklists except those of Kishida (1935) 
and Schulze (1935). 


Haemaphysalis formosensis Neumann, 
1913.—27 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. This common species appears in all 
previous checklists. Recorded by Neumann 
(1913) from dogs in Taiwan. 


Haemaphysalis_ hystricis Supino, 
1897.—95 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
Haemaphysalis nishiyamai Sugimoto, 1935 
(Sugimoto 1939, Elishewitz 1943, Anony- 
mous 1944, Luh and Woo 1950, Shimada 
et al. 1961, Maa and Kuo 1966). This spe- 
cies also appears as the senior synonym H. 
hystricis in all previous checklists, reflect- 
ing its prominence in the Taiwan tick fauna 
(Hoogstraal et al. 1965, 1973). 


Haemaphysalis kitaokai Hoogstraal, 
1969.—1 Taiwan collection in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Listed only by Tseng (1978) and 
Hoogstraal (letter no. 251, 14 February 
1984). The single USNTC collection (RML 
16603) consists of two badly damaged fe- 
males obtained by Sugimoto (1937a, b) that 
were initially determined by Hoogstraal 
(1962) as H. aponommoides Warburton, 
1913 (a species of the central and eastern 
Himalayas), but later (Hoogstraal 1969) 
tentatively determined to represent H. ki- 
taokai. The abundance of this species in 
southernmost Japan (Nakao and Takada 
1997) also argues for its occurrence in Tai- 
wan. 


Haemaphysalis mageshimaensis Saito 
& Hoogstraal, 1973.—71 Taiwan collec- 
tions in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Listed only by Hoogstraal (letter no. 
251, 14 February 1984), but recorded from 
numerous hosts in Taiwan, Lutao, and Lan- 
yu (Hoogstraal and Santana 1974). 


Haemaphysalis ornithophila Hoogs- 
traal & Kohls, 1959.—No Taiwan collec- 
tions in USNTC, but 8 collections in 
BPBM. 

Junior synonymy in Taiwan checklists: 
None. Listed only by Tseng (1978). First 
reported from Taiwan by Wilson (1970). 


Haemaphysalis phasiana Saito, Hoogs- 
traal & Wassef, 1974.—16 Taiwan collec- 
tions in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Listed only by Hoogstraal (letter no. 
251, 14 February 1984). According to Saito 
et al. (1974), “the taxon phasiana may pos- 
sibly be a subspecies of doenitzi”’; these au- 
thors call for additional data to clarify the 
status of H. phasiana in Taiwan. 


Haemaphysalis taiwana Sugimoto, 
1936.—49 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
Haemaphysalis cornigera taiwana Sugi- 
moto, 1936 (Elishewitz 1943, Luh and Woo 


VOLUME 107, NUMBER 2 


1950, Shimada et al. 1961, Maa and Kuo 
1966, Tseng 1978). 


Haemaphysalis.§ yeni TYoumanoff, 
1944.—1 Taiwan collection in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Not previously listed as occurring in 
Taiwan. The single USNTC collection 
(RML 77868) consists of 15 nymphs and 2 
larvae removed from a specimen of the 
lesser coucal, Centropus bengalensis (Har- 
tlaub) (Cuculiformes: Cuculidae), on Lan- 
yu, off Taiwan’s southeastern coast. The 
presence of H. yeni in Japan’s Ryukyu Is- 
lands and in Vietnam (Saito and Hoogstraal 
1972) suggests that it also occurs in Taiwan. 


Ixodes acutitarsus (Karsch, 1880).—8 
Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. This distinctive species, said to be 
the largest Ixodes in the world (Yamaguti 
et al. 1971), appears in all previous check- 
lists except Ogura (1936). 


Ixodes granulatus Supino, 1897.—128 
Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Listed only by Tseng (1978) and 
Hoogstraal (letter no. 251, 14 February 
1984). This abundant and widespread spe- 
cies was first reported from Taiwan by Wil- 
son (1970). 


Ixodes kuntzi Hoogstraal & Kohls, 
1965.—35 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Listed by Maa and Kuo (1966), 
Tseng (1978), and Hoogstraal (letter no. 
251, 14 February 1984). This species is en- 
demic to Taiwan. 


Ixodes nipponensis Kitaoka & Saito, 
1967.—1 Taiwan collection in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Not previously listed as occurring in 
Taiwan. The single USNTC collection 
(RML 35386) consists of two females taken 
from domestic cattle in the vicinity of Tai- 
pei. Although these may have been intro- 
duced specimens, the presence of /. nippo- 


249 


nensis in southernmost Japan (Nakao and 
Takada 1997) suggests that this species also 
occurs in Taiwan. 


Ixodes ovatus Neumann, 1899.—83 Tai- 
wan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
Ixodes shinchikuensis Sugimoto, 1937 (Eli- 
shewitz 1943, Anonymous 1944, Luh and 
Woo 1950 (as “I. shinckikuensis”’), Shi- 
mada et al. 1961, Maa and Kuo 1966); /. 
taiwanensis Sugimoto, 1936 (Elishewitz 
1943, Anonymous 1944, Luh and Woo 
1950, Shimada et al. 1961, Maa and Kuo 
1966). Note that both junior synonyms are 
listed in the same five checklists. The senior 
synonym J. ovatus is listed only by Tseng 
(1978) and Hoogstraal (letter no. 251, 14 
February 1984). 


Ixodes persulcatus Schulze, 1930.—7 
Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Listed only by Hoogstraal (letter no. 
251, 14 February 1984). Published refer- 
ences to /. ricinus (Linnaeus, 1758) in Tai- 
wan (Sugimoto 1939, Elishewitz 1943, 
Anonymous 1944, Luh and Woo 1950, Shi- 
mada et al. 1961, Maa and Kuo 1966) al- 
most certainly represent /. persulcatus, an 
eastern Palearctic sister species of the large- 
ly European J. ricinus. Uncommon in Tai- 
wan tick collections, /. persulcatus is the 
principal vector of Lyme borreliosis in 
northeastern Asia and is probably respon- 
sible for the small number of cases now be- 
ing reported in Taiwan (Shih et al. 1998, 
Shih and Chao 1999). 


Ixodes simplex Neumann, 1906.—53 
Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. Listed only by Tseng (1978) and 
Hoogstraal (letter no. 251, 14 February 
1984). Wilson (1970) first reported /. sim- 
plex from Taiwan. 


Ixodes vespertilionis Koch, 1844.—No 
Taiwan collections in USNTC, but 2 col- 
lections in BPBM. 

Junior synonymy in Taiwan checklists: 


250 


None. Listed only by Tseng (1978). First 
reported from Taiwan by Wilson (1970). 


Rhipicephalus haemaphysaloides (Su- 
pino, 1897).—50 Taiwan collections in 
USNTC. 

Junior synonymy in Taiwan checklists: 
Rhipicephalus haemaphysaloides expeditus 
Neumann, 1904 (Luh and Woo 1950); R. 
expeditus Neumann, 1904 (Shimada et al. 
1961). The senior synonym R. haemaphys- 
aloides is also listed by Shimada et al. 
(1961), as well as by Maa and Kuo (1966), 
Tseng (1978), and Hoogstraal (letter no. 
251, 14 February 1984). 


Rhipicephalus sanguineus (Latreille, 
1806).—47 Taiwan collections in USNTC. 

Junior synonymy in Taiwan checklists: 
None. This ubiquitous species appears in all 
previous checklists. Hundreds of additional 
unaccessioned collections of R. sanguineus 
from Taiwan are in the USNTC. 


EXCLUDED SPECIES 


The preceding passages present argu- 
ments for excluding five tick species that 
appear in earlier checklists of the Taiwan 
fauna: Argas persicus, A. vespertilionis, 
Aponomma fuscolineatum (listed as the ju- 
nior synonym Amblyomma serpentinum), 
Boophilus annulatus (listed as the junior 
synonym Margaropus annulatus), and Ixo- 
des ricinus. Eight additional species named 
in these early checklists have either never 
occurred in Taiwan or are no longer present 
there: 

Amblyomma breviscutatum Neumann, 
1899.—Listed by Elishewitz (1943) and 
Anonymous (1944) as A. cyprium Neu- 
mann, 1899. Keirans, in Voltzit and Keirans 
(2002), determined that the types of these 
species are conspecific, and since A. brev- 
iscutatum has page priority over A. cy- 
prium, the latter falls as a junior synonym. 
There are no Taiwan collections of A. brev- 
iscutatum in the USNTC. The single Tai- 
wanese record cited in Robinson (1926) is 
from ‘“‘Kankan,” an alternate spelling of 
Kaohsiung (also formerly Koshung), Tai- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


wan’s second-largest city and its principal 
seaport, located in the tropical southwestern 
quadrant of the island. It is possible that A. 
breviscutatum either once occurred in Kao- 
hsiung (or elsewhere on Taiwan’s densely 
populated western coastal plain) and has 
since been extirpated by intense urbaniza- 
tion, or that living specimens were sporad- 
ically introduced with such hosts as cattle, 
horses, pigs and water buffalo imported 
from areas to the south and west that lie 
well within the range of this species (e.g., 
the Philippines, Indonesia). 

Dermacentor atrosignatus Neumann, 
1906.—Listed by Sugimoto (1939), Eli- 
shewitz (1943), Anonymous (1944), and 
Shimada et al. (1961). In their review of 
this Southeast Asian species, Hoogstraal 
and Wassef (1985) record specimens from 
Palawan, Busuanga, and Culion Island in 
the southwestern Philippines, localities far 
removed from Taiwan. A single supposedly 
Taiwanese collection of D. atrosignatus (1 
6, 1 2) in the USNTC (RML 15189) has 
been lost (L.A. Durden, letter of 28 Feb- 
ruary 1997 to RGR). A malformed d, also 
alleged to be D. atrosignatus, from “‘Ko- 
sempo”’ (Chia-hsien, 23.05N, 120.35E, a 
town in Kaohsiung County), Taiwan, was 
discussed by Robinson (1920). It seems un- 
likely that this relatively large and conspic- 
uously ornate Dermacentor would be over- 
looked in years of collecting by NAMRU- 
2 investigators. 

Dermacentor reticulatus (Fabricius, 
1794).—Listed by Maa and Kuo (1966) and 
by Tseng (1978). This western Palearctic 
species ranges no farther east than the Cen- 
tral Asian republics of the former Soviet 
Union (Pomerantsev 1950). 

Haemaphysalis birmaniae  Supino, 
1897.—Listed by Kishida (1935), Schulze 
(1935), Elishewitz (1943), Anonymous 
(1944), Luh and Woo (1950), and Shimada 
et al. (1961). This species, “‘the subject of 
much taxonomic confusion and many in- 
correct host and distribution records” 
(Hoogstraal 1970), is known with certainty 


VOLUME 107, NUMBER 2 


only from northeastern India, Nepal and 
Myanmar (Burma). 

Haemaphysalis bispinosa Neumann, 
1897.—Listed by Elishewitz (1943), Anon- 
ymous (1944), Luh and Woo (1950), Shi- 
mada et al. (1961), and Tseng (1978). This 
is a tropical South Asian species (Hoogs- 
traal and Trapido 1966); East Asian speci- 
mens of “H. bispinosa”’ are actually H. lon- 
gicornis Neumann, 1901, a widespread and 
widely introduced tick whose original range 
embraced temperate areas of northeastern 
China, Korea, Japan, and maritime Russia 
(Hoogstraal et al. 1968c). Significantly, H. 
longicornis appears to be absent from the 
subtropical Ryukyu Islands (Yamaguti et al. 
1971) and is therefore unlikely to occur as 
far south as Taiwan. 

Haemaphysalis inermis Birula, 1895.— 
Listed by Elishewitz (1943), Anonymous 
(1944), Shimada et al. (1961), Maa and 
Kuo (1966), and Tseng (1978). This is an- 
other western Palearctic species, ranging 
from southern Europe to Turkey, southern 
Russia, and northern Iran (Hoogstraal 
1969). 

Haemaphysalis warburtoni Nuttall, 
1912.—Listed by Kishida (1935), Schulze 
(1935), Sugimoto (1939), Elishewitz 
(1943), Luh and Woo (1950), Shimada et 
al. (1961), Maa and Kuo (1966), and Tseng 
(1978). All listings of H. warburtoni from 
Taiwan are apparently based on the single 
specimen reported by Nuttall and Warbur- 
ton (1915) from ‘“Taihoku”’ (Taipei); this 
specimen was restudied by Hoogstraal 
(1966) and found to be H. formosensis. 
True H. warburtoni is restricted to moun- 
tainous areas in southern China, Nepal, and 
northern India (Hoogstraal and Kim 1985). 

Ixodes hexagonus Leach, 1815.—Listed 
by Sugimoto (1939), Elishewitz (1943), 
Shimada et al. (1961), Maa and Kuo 
(1966), and Tseng (1978). A third western 
Palearctic species, ranging across Europe 
and North Africa to western Russia and, 
perhaps, Central Asia (Arthur 1963). 


251 


ACKNOWLEDGMENTS 


For fielding innumerable questions and 
generously making available all Taiwanese 
specimens and records in their care, I thank 
Lance A. Durden, Department of Biology, 
and James E. Keirans, Institute of Arthro- 
podology and Parasitology, Georgia South- 
ern University. Thanks also to Susan W. 
Glenn and Paul Theerman, former Smith- 
sonian archivists, for providing unlimited 
access to the meticulously organized 
Hoogstraal correspondence files. For enthu- 
siastically encouraging the compilation of 
this checklist, as the first step toward real- 
izing Harry Hoogstraal’s lifelong goal of a 
monograph on the ticks of Taiwan, I am 
grateful to Chyi-Chen Ho, Department of 
Applied Zoology, Taiwan Agricultural Re- 
search Institute, Taichung. And, as always, 
I deeply appreciate the assistance of Fu- 
Meei Yeh Robbins, who kindly prepared 
English translations of crucial paragraphs in 
Chinese papers. The opinions and _ asser- 
tions advanced herein are those of the au- 
thor and are not to be construed as official 
or reflecting the views of the U.S. Depart- 
ments of the Army, Navy, or Defense. 


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PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 254-258 


FIRST REPORT OF THEOCOLAX INGENS XIAO AND HUANG 
(HYMENOPTERA: PTEROMALIDAE) IN THE WESTERN HEMISPHERE, 
WITH A SYNOPSIS OF THE GENUS 


E. E. GRISSELL AND G. FEF HEVEL 


(EEG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, c/o National Museum of Natural History, Smithsonian Insti- 
tution, PO. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. (e-mail: 
egrissel @sel.barc.usda.gov); (GH) Department of Entomology, National Museum of Natural 
History, MRC 187, Washington, DC 20013-7012, U.S.A. (e-mail: hevel.gary @nmnhy.si.edu) 


Abstract.—Theocolax ingens Xiao and Huang is reported for the first time from the 
Western Hemisphere (Silver Spring, Maryland, USA). The known species of Theocolax 
Westwood are discussed and their host data summarized. Theecolax fasciata (Ishii 1956) 
is transferred from the genus and synonymized with Cerocephala aquila (Girault 1920) 


(n. Syn.). 


Key Words: 


Hymenoptera, Chalcidoidea, Pteromalidae, Theocolax, Cerocephala, new 


record, synonymy, Western Hemisphere 


The genus Theocolax Westwood is 
known from eight species (Noyes 2001, 
Xiao and Huang 2001, Baur 2001). One of 
these species, Theocolax fasciata (Ishii), is 
herein transferred from the genus as ex- 
plained below. The remaining seven species 
and their distributions are as follows: T. 
bakeri (Crawford), Oriental (Philippines); 
T. elegans and T. formiciformis Westwood, 
cosmopolitan; 7. frater (Girault), Australian 
(Australia, introduced into Hawaii); 7. ob- 
longa (Delucchi), Afrotropical (Zaire); T. 
phloeosini Yang and T. ingens Xiao and 
Huang, Palearctic (People’s Republic of 
China) (Bouéek 1988, Noyes 2001, Xiao 
and Huang 2001, Baur 2001). The genus is 
thought to have originated in the Eastern 
Hemisphere based on the known distribu- 
tions of its species. Only the cosmopolitan 
species T. elegans (Westwood) has been re- 
ported previously in the Nearctic (Boucéek 
and Heydon 1997). 

In this paper we report the discovery of 


Theocolax ingens Xiao and Huang in the 
Western Hemisphere (Maryland, USA) and 
summarize the known published informa- 
tion for other species of the genus. Species 
of Theocolax primarily attack small beetles, 
particularly Anobiidae and Scolytidae, in 
dead and dying trees, but 7. elegans has 
been reported from beetles in stored grains 
(Boucek 1988, Yang 1989, Xiao and Huang 
2001). 


Theocolax Westwood 


Theocolax Westwood 1832: 127. Type spe- 
cies: Theocolax formiciformis Westwood, 
by monotypy. 


There appear to be two morphological 
subgroupings of species in the genus Theo- 
colax. One group contains 7. ingens and T. 
phloeosini, in which the scrobal depression 
is bounded laterally by distinct angulate 
edges that nearly meet dorsomedially (Fig. 
1), the interantennal lamella extends % way 
to the median ocellus, and the postmarginal 


VOLUME 107, NUMBER 2 


vein is nearly subequal in length to the stig- 
mal vein (Fig. 2). The other group contains 
T. bakeri, T. elegans, T. formiciformis, and 
T. oblonga in which the scrobal depression 
is either not bounded laterally (.e., the 
junction of the face and depression is 
rounded or indefinite) or if it is then the 
edges are widely separated dorsally (Fig. 
3), the interantennal lamella extends only 4 
way to the median ocellus (Fig. 3), and the 
postmarginal vein is absent or distinctly 
shorter than the stigmal vein (Fig. 4). The 
species Theocolax frater, however, is some- 
what intermediate between the two groups, 
having the scrobal depression and interan- 
tennal lamella of the former group, and the 
shorter postmarginal vein of the latter. 


DESCRIBED SPECIES OF THEOCOLAX 
Theocolax bakeri (Crawford) 


Cerocephala bakeri Crawford 1914: 460— 
461. 

Theocolax bakeri (Crawford): Gahan 1946: 
357. Generic transfer. 


This species was described from the Phil- 
ippines. Baltazar (1966) listed the host as a 
scolytid. 


Theocolax elegans (Westwood) 
(Figs. 3, 4) 


Choetospila elegans Westwood 1874: 157. 
Theocolax elegans (Westwood): Boucéek 
1988: 339. Generic transfer. 


This is a “cosmopolitan parasite of small 
beetles developing in stored grain” (Bou- 
éek 1988). It is the only species previously 
reported from the Nearctic (Burks 1979) (as 
Choetospila elegans). 


Theocolax formiciformis Westwood 


Theocolax formiciformis Westwood 1832: 
Wie 


Although this is a ““cosmopolitan parasite 
of anobiid beetles in dead wood” (Boucek 
1988), it is not yet known from the Nearc- 
tic. 


Theocolax frater (Girault) 


Spalangiomorpha frater Girault 1913: 334. 
Theocolax frater (Girault): Bouéek 1988: 
340. Generic transfer. 


This species may belong to the genus 
Cerocephala according to Bouéek (1988), 
but little is known about it. No hosts have 
been reported. It is known from Australia 
(Boucek 1988) and Hawaii (Nishida 2002). 


Theocolax ingens Xiao and Huang 
(Figs. 1, 2) 


Theocolax ingens Xiao and Huang 2001: 
203-205. 


This species was described from China 
as a potential parasitoid of bark beetles. No 
specific host insect or host plant was given, 
but 7. ingens is nearly identical to T. 
phloeosini and would probably have similar 
hosts. The species is reported herein for the 
first ttme from the New World. In 2003, GH 
discovered several dozen specimens of T. 
ingens walking on the trunk of a dead Pru- 
nus sp. tree in his suburban backyard in Sil- 
ver Spring, Maryland, USA (specimens in 
National Museum of Natural History, 
Washington, DC). Several scolytid beetles 
were also collected from the same tree, in- 
cluding Scolytus mali (Bechstein) intro- 
duced from Europe, Africa, and Asia, and 
Xyleborinus saxeseni (Ratzeburg), intro- 
duced from Europe. Because T. ingens and 
T. phloeosini are Palearctic in origin, it ap- 
pears that 7. ingens was accidentally intro- 
duced into the United States with wood 
products exported from China. 


Theocolax oblonga (Delucchi) 


Cerocephala oblonga Delucchi 1956: 168— 
le 

Theocolax oblonga (Delucchi): Baur 2001: 
69. Generic transfer. 


This species is known only from material 
collected in Zaire (Kivu Province). It was 
reared from Mimips rugicollis Schedl (Co- 
leoptera: Scolytidae). 


256 


Figs. 1-4. 


tiny | AS 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


— 


We 
S 
SS 


WARY 


Theocolax species. 1-2, Theocolax ingens. 1, Face. 2, Forewing. 3—4, Theocolax elegans. 3, Face. 


4, Forewing marginal, postmarginal, and stigmal veins only. 


Theocolax phloeosini Yang 


Theocolax phloeosini Yang 1989: 97—99 
(Chinese), 101—102 (English). 


This species, known only from China, 
was originally reared and collected from 
trunks and branches of Sabina chinensis 
(L.) [Chinese juniper] infested with Phloeo- 
sinus aubei Perris (Coleoptera: Scolytidae) 
and Prunus persica (L.) [peach] infested 
with Scolytus japonicus Chapuis (Scolyti- 
dae). Adult wasps were reared as ectopar- 


asitoids of larvae of these two beetles. Ad- 
ditional material was collected on Ulmus 
pumila L. [elm] infested with Scolytus 
schevyrewi Seminov and S. butovitschi 
Stark (Scolytidae). Adult wasps were reared 
as ectoparasitoids of pupae and larvae col- 
lected from galleries of these bark beetles. 
Two generations were collected between 
April and July. An additional report by 
Yang (1996) added the following hosts: 
Scolytus seulensis Murayama (Scolytidae) 
on Prunus armeniaca L. and P. persica, 


VOLUME 107, NUMBER 2 


and “‘many other bark beetle species” on 
the trees Pinus tabuliformis Carriere, P. 
massoniana Lambert, and Picea spp. He 
also reported that the female enters into gal- 
leries to Oviposit. 


SPECIES REMOVED FROM THEOCOLAX 


Choetospila fasciata was described by 
Ishii (1956). The original holotype speci- 
men, a unique male collected in Tokyo, Ja- 
pan, was drawn by Ishii at an earlier date, 
but by 1956 the specimen had been “... 
devoured by a carpet beetle.” For that rea- 
son Ishii based his written description on a 
fairly detailed drawing. The species has not 
been recognized since its description, and 
nothing is known of its biology. It was 
placed in Theocolax by implication when 
Bouéek (1988) synonymized Choetopsila 
with Theocolax. The original illustration by 
Ishii (1956) clearly shows a combination of 
a sculptured propodeum and longitudinally 
striate scutellum, which is unique to Cero- 
cephala aquila (Girault), a widespread 
Indo-Australian species found also in Fiji, 
the Philippines, Cuba, and Mexico (Bouéek 
1988). The synonymy for this species ap- 
pears as follows: 


Cerocephala aquila (Girault) 


Proamotura aquila Girault 1920: 143. 
Cerocephala aquila (Girault): Gahan 1946: 
361. Generic transfer. 
Choetospila fasciata Ishii 
New Synonymy. 

Theocolax fasciata (Ishii): Bouéek 1988: 
339. Transfer by implication, Choetospila 
Westwood 1874 synonymized with Theo- 
colax Westwood 1832. 


1956: 3132. 


ACKNOWLEDGMENTS 


We thank Hui Xiao, Institute of Zoology, 
Chinese Academy of Sciences, Beijing, for 
comparing the specimens collected in Silver 
Spring with the species she coauthored with 
Da-Wei Huang. Thanks also go to Michael 
Gates, Systematic Entomology Laboratory, 
Washington, DC, for collecting additional 
material of 7. ingens from the original site. 


257 


For reviewing this manuscript and provid- 
ing helpful comments, we thank Dave 
Smith, E Christian Thompson, and Norman 
Woodley, Systematic Entomology Labora- 
tory, and Steve Heydon, Department of En- 
tomology, University of California, Davis. 
We thank Natalia Vandenberg, Systematic 
Entomology Laboratory, and Robert Ra- 
baglia, Forest Health Monitoring, Maryland 
Department of Agriculture, Annapolis, for 
identifying the scolytid beetles found at the 
same site as T. ingens. 


LITERATURE CITED 


Baltazar, C. R. 1966. A catalogue of Philippine Hy- 
menoptera. Pacific Insects Monograph 8: 1—488. 

Baur, H. 2001. The Hymenoptera (Chalcidoidea, Ich- 
neumonoidea, Platygastroidea) described by Vit- 
torio Delucchi: An annotated catalogue. Journal of 
Natural History 35: 55-125. 

Boucéek, Z. 1988. Australasian Chalcidoidea (Hyme- 
noptera). C. A. B. International, Wallingford, UK, 
832 pp. 

Bouéek, Z. and S. L. Heydon. 1997. Pteromalidae, pp. 
541-692. In Gibson, G. A. P., J. Huber, and J. B. 
Woolley, eds. Annotated Keys to the Genera of 
Nearctic Chalcidoidea (Hymenoptera). Mono- 
graph |. National Research Council, Ottawa, Can- 
ada. 

Burks, B. D. 1979. Family Pteromalidae, pp. 768-835. 
In Krombein, K. V., P. D. Hurd, Jr, D. R. Smith, 
and B. D. Burks, eds. Catalog of Hymenoptera in 
America North of Mexico, Vol. 1. Symphyta and 
Apocryta (Parasitica). Smithsonian Institution 
Press, Washington, DC. 

Crawford, J. C. 1914. New Philippine Hymenoptera. 
The Philippine Journal of Science 9: 457—464. 

Delucchi, V. 1956. Neue Chalcidier aus dem Belgisch- 
en Kongo. Revue de Zoologie et de Botanique 
Africaines 53: 158-178. 

Gahan, A. B. 1946. Review of some chalcidoid genera 
related to Cerocephala Westwood. Proceedings of 
the United States National Museum 96: 349-376, 
1 plate. 

Girault, A. A. 1913. Australian Hymenoptera Chalci- 
doidea—VI. The family Pteromalidae with the de- 
scriptions of new genera and species. Memoirs of 
the Queensland Museum 2: 303-334. 

. 1920. New genera of chalcid flies from Aus- 
tralia (Hymenoptera). Insecutor Inscitiae Men- 
struus 8: 142-146. 

Ishii, T. 1956. Some very rare wasps belonging to the 
Chalcidoidea with description of one new species. 
Kontyd 24: 31-33. 

Nishida, G. M. (Editor). 2002. Hawaiian terrestrial ar- 

checklist. 4th Edition. http://www2. 


thropod 


258 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 1874. Thesaurus Entomologicus Oxoniensis. 


bishopmuseum.org/HBS/checklist/query.asp? 


erp=Arthropod. 9 April 2002, 9724 records; last xxiv, 205 pp. Clarendon Press, Oxford, England. 
accessed 15: March 2004. Xiao, H. and D.-W. Huang. 2001. A new species of 
Noyes, J. S. 2001. Interactive Catalogue of World Theocolax Westwood (Hymenoptera: Pterom- 
Chalcidoidea (2001—second edition). CD-ROM. alidae) from China. The Raffles Bulletin of Zo- 
Taxapad and The Natural History Museum, Lon- ology 49: 203-205. 
don. Yang, Z. 1989. One new species and other pteromalids 
Westwood, J. O. 1832. Descriptions of several new parasitizing bark-beetles in Shaanxi, China. En- 
British forms amongst the parasitic hymenopter- tomotaxonomia 11: 97-103. 


ous insects. London and Edinburgh Philosophical . 1996. Parasitic Wasps on Bark Beetles in Chi- 
Magazine 2: 127-129. na. Science Press, Beijing, China, 363 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 259-266 


A NEW GENUS OF BRACHYPTEROUS LEAFHOPPERS 
(HEMIPTERA: CICADELLIDAE: CICADELLINAE: PROCONIINI) 
FROM COSTA RICA 


CAROLINA GODOY 


Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa Rica (e-mail: 


cgodoy @inbio.ac.cr) 


Abstract.—A new genus, Brevimetopia, and two new species, B. silenciosa and B. 
chusquea, are described from high altitudes in Costa Rica. These species are the first 
brachypterous leafhoppers recorded from Central America. 


Key Words: 
America, Neotropics 


Brachyptery is relatively uncommon in 
leafhoppers, and in the New World most 
records are from the Neartic Region. Some 
Neartic genera of Errhomenini (Cicadelli- 
nae) have species with brachypterous fe- 
males and macropterous males (Oman and 
Musgrave 1975, Oman 1987). In Doryce- 
phalini, some species of Aftenuipyga have 
brachypterous females and polymorphic 
males (with both macropterous and _ bra- 
chypterous individuals). In the same tribe, 
females of Neoslossonia are normally bra- 
chypterous (Oman 1985). In Lonatura (Del- 
tocephalinae), macropterous individuals of 
both sexes are rare (Kramer 1967). In the 
Neotropical Region, brachyptery in Cica- 
dellidae has been reported in four genera. 
Among Deltocephalinae, brachyptery oc- 
curs in both sexes of Faltala brachyptera 
Oman and in females (male unknown) of 
Amplicephalus papillosus  WLinnavuori 
(1959); both records are from Argentina. 
Among Proconiini, brachyptery has been 
reported in Lojata ohausi (Schmidt), which 
is known only from the female holotype 
from Ecuador, Splonia brevis (Walker) from 
Venezuela, and S. nasti Young from Ecua- 
dor (Young 1968). 


Proconiini, Cicadellinae, leafhopper, taxonomy, bamboo, Chusquea, Central 


In this paper, I describe Brevimetopia, a 
new genus of brachypterous leafhoppers 
from Costa Rica, based on two new species, 
B. silenciosa and B. chusquea. The new ge- 
nus is in the tribe Proconiini (Cicadellinae) 
and is most similar to Quichira Young. The 
two species described here, which represent 
the first records of brachypterous leafhop- 
pers from Central America, appear to be as- 
sociated with bamboos in the Talamanca 
mountain range, which extended from Cos- 
ta Rica to western Panama, although there 
are currently no records from the latter 
country. 


MATERIAL AND METHODS 


The details of preparations of genital 
structures of leafhoppers for dissections and 
study were given by Oman (1949). I have 
followed his method with some modifica- 
tions. Abdomens were removed and placed 
in 10% potassium hydroxide overnight at 
room temperature. The following day indi- 
vidual abdomens were washed in water be- 
fore examination and eventual preservation 
in glycerin in microvials. 

Specimens are deposited in the following 
collections: 


260 


CAS: California Academy of Sciences, 
San Francisco, USA. 

INBio: Instituto Nacional de Biodiver- 
sidad, Santo Domingo, Heredia, 
Costa Rica. 

INHS: Illinois Natural History Survey, 
Urbana, USA. 

NHM: The Natural History Museum, 
London, UK. 

WiGRe University of Costa Rica, San 
Pedro, San José, Costa Rica. 

USNM: National Museum of Natural 


History, Smithsonian Institution, 
Washington, D.C., USA. 


Brevimetopia Godoy, new genus 


Type species: Brevimetopia silenciosa, 
n. sp. 


Description.—Length: Male 12.0—14.8 
mm, female 13.5-15.0 mm. Head: Mod- 
erately produced, moderately rounded api- 
cally in dorsal aspect, median length of 
crown less than half interocular width and 
more than one-third transocular width, an- 
terior margin with slight depression be- 
tween crown and face, ocelli small and lo- 
cated anterad of line connecting anterior 
eye angles, each slightly closer to median 
line than to adjacent eye angle, surface of 
crown concave except for indistinct M- 
shaped elevation bordering posterior mar- 
gin, pubescence indistinct; antennal ledges 
with longitudinal fovea, anterior margins 
oblique; clypeus flattened medially, strong- 
ly convex laterally, texture of dorsomedian 
area coarsely rugose, muscle impressions 
distinct; transclypeal suture obsolete; face 
pubescent, more so below; clypellus with 
contour continuous with profile of clypeus 
(Fig: 13): 

Thorax: Pronotal width less than tran- 
socular width of head, lateral margins di- 
vergent anteriorly, disc rugose and punc- 
tate; pleura pubescent, mesepimeron inflat- 
ed as a fleshy lobe (Fig. 12); anterior por- 
tion of scutellum pubescent, inflated in the 
posterior portion (Fig. 13); metepimeron 
with shelflike projection. Forewing bra- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


chypterous, exposing at least terga VI-IX, 
with membrane including only reduced api- 
cal cells, venation absent or nearly absent, 
texture strongly coriaceous, heavily or fine- 
ly pubescent; hind wing brachypterous, 
same length as forewing; hind legs at rest 
with knees attaining or nearly attaining pos- 
terior proepimeral margins, femoral setal 
formula 2:1:1, 2:1:0, or 2:0:0, tarsomere I 
of hind leg with length nearly equal to com- 
bined length of If and III tarsomeres. 

Male genitalia: Pygofer not strongly 
produced, with numerous evenly dispersed 
microsetae over most of surface, in lateral 
view with caudal margin obliquely truncate, 
with recurved posteroventral process (Figs. 
3—4). Plates separate throughout their 
length, not extending as far posteriorly as 
apex of pygofer, each triangular, with nu- 
merous evenly dispersed microsetae (Fig. 
5). Style posteriorly extending beyond apex 
of connective. Connective Y-shaped with 
arms widely divergent, stem keeled (Fig. 6). 
Aedeagus symmetrical, in caudoventral 
view with preatrium long, shaft short, trun- 
cate apically, with long paired medial pro- 
cesses, in lateral view extending distally be- 
yond aedeagal shaft (Figs. 7-8). Paraphyses 
absent. 

Female: Ovipositor not extending be- 
yond pygofer apex. First and second val- 
vulae elongate, slightly wider subapically, 
tapered to apex; first valvulae with sculp- 
tured area striate. Second valvulae with dor- 
sal teeth, individual teeth emarginate, 
slightly elevated. 

Etymology.—The generic name is 
formed by adding the Latin brevis (in its of 
short) and the Greek metron (in its of mea- 
sure), in reference to the short wings. Gen- 
der: feminine. 

Remarks.—Brevimetopia 1s known only 
from high altitudes in Costa Rica and, with 
the exception of one specimen, the genus 
has only been collected in the Talamanca 
mountain range. The male genitalia are sim- 
ilar to those of Quichira Young, but the lat- 
ter has a more truncate head (in dorsal 
view) and the transition from the crown to 


VOLUME 107, NUMBER 2 


ate) 
H 


bine 
ui Way ma si i ‘a 


com 
4 me 


saat niin iti : 
x el a hi 
‘Ne Hi ie ibe He, ; 


ih if chu iff ik i} 
Hi tt WH ry vit 
i H ih f wily, 1 
i Wit HM! AYE Mi Ne 
4) pith ins My ie i 


Figs. 1-2. 


the face is more angulate. Brevimetopia has 
the anterior margin of the head rounded in 
dorsal view, much more body pubescence, 
brachypterous wings, inflated mesepimeron, 
and longer hind legs with knees attaining 
the posterior margin of the proepimeron. 
The unusually long hind legs, distinguish- 
ing character of Brevimetopia are clearly 
associated with brachyptery; among other 
Proconiini, this character is found only in 
Splonia (Young 1968). 


In Young’s (1968) key to the genera of 


Proconiuni, Brevimetopia would come out 


261 


Brevimetopia silenciosa. 1, Male dorsal view. 2, Apex of left hind femur. 


in the first couplet, with the other brachyp- 
terous genus, Lojata. Brevimetopia can be 
distinguished by its more rounded head, 
without a distinct carinae between the 
crown and face (in Lojata the head is pro- 
longed and obtusely angulate in dorsal 
view), as well as the lack of a pair of round- 
ed elevations bordering the posterior mar- 
gin of the pronotal disc. 


KEY TO BREVIMETOPIA SPECIES 


1. Body color mostly mustard yellow. Forewing 


densely pubescent (Fig. 1) ....... B. silenciosa 


262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


\ 


r 
ia 
~~ 


_— 


Figs. 3-9. 


Brevimetopia silenciosa. 3, Male pygofer. 4, Male pygofer lateral view. 5, One male plate, ventral 


view. 6, One style and connective, dorsal view. 7, Aedeagus, caudoventral view. 8, Aedeagus lateral view. 9, 


Female abdominal sternum VII. 


— Body black and orange. Forewing sparsely pu- 
bescent 


N 


. Forewing black with two orange spots (Fig. 10) 
B. chusquea, 3 


B. chusquea, ° 


Brevimetopia silenciosa Godoy, 
new species 
(Figs. 1-9) 


Description.—Structural characters as in 
generic description. Length: Male 14.8 mm, 
female 14.8-15.0 mm. Color of dorsum 
mustard yellow except head, pronotum, 


posterior scutellum, face, and legs, red 
brown. All parts except head, pronotum, 
and legs densely covered with mustard yel- 
low setae (Fig. 1). 

Thorax: Forewing brachypterous, expos- 
ing at least terga VI-IX. Hind legs at rest 
with knees attaining posterior proepimeral 
margins, femoral setal formula 2:1:1 (Fig. 2). 

Male genitalia: Aedeagus, with paired 
medial processes in lateral view extending. 
distally beyond aedeagal shaft, each with 
small toothlike projection (Figs. 7—8). 


VOLUME 107, NUMBER 2 


Figs. 10-12. 
arrow indicates lobelike mesepimeron. 


Female: Abdominal sternum VII broadly 
sinuate, with blunt medially produced lobe 
(Fig. 9). Ovipositor as in B. chusquea. 

Type material—-Holotype 6, COSTA 
RICA, Puntarenas, Buenos Aires, PILA- 
ACLA Est. Altamira, Sendero Valle del Sil- 
encio al Jardin, 2,400 m. 18.iv.1995 (IN- 
Bio). L. Angulo, L-S 342300-577200. Para- 
types: 3 2, same data; 2 ¢, Limon, Valle 
del Silencio, 2,420 m; 3—6.vi.1996. R. Vil- 
lalobos, L-S 342200-577400 (INBio). | 6, 
Limon P. Int. La Amistad, Send. Circular, 
2,450 m. 22.vi.-4vii.2003. R. Gonzalez, 
LS 340258-577465; 1 ¢, same data, 
27.1x.2003; 1 3, same data 20—22.1x.2003; 


Brevimetopia chusquea. 10, Male dorsal view. 11, Female lateral view. 12, Female dorsal view, 


1 3, Lim6n P. Int. La Amistad, Valle del 
Silencio, Sendero al Jardin Natural, 2,400 
Me Olen. OF De Rubies S541 2905— 
577409. 

Etymology.—The specific name refers to 
the type locality. 

Remarks.—Brevimetopia silenciosa is 
similar to B. chusquea, but the former is 
more pubescent differs in coloration, has 
the mesepimeron more inflated, and the 
basal processes of the aedeagus have tooth- 
like projections. This species is known only 
from the Talamanca mountain range. The 
host plant is Chusquea patens L. G. Clark 
(R. Rakitov, personal communication). 


264 


Figs. 13-19. 
shown. 15, One male plate, ventral view, setae not shown. 16, One style, dorsal view. 17, Connective, dorsal 
view. 18, Aedeagus, caudoventral view. 19, Aedeagus, lateral view. 


Brevimetopia chusquea Godoy, 
new species 
(Figs. 10—22) 


Description.—Length: Structural charac- 
ters as in generic description. Male, 12 mm, 
female 12.9-13.5 mm. Male. Color of dor- 
sum of head, pronotum, posterior scutel- 
lum, face and legs black; forewing black 
with large orange spot in the brachial cell 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Brevimetopia chusquea. 13, Male head and thorax, lateral view. 14, Male pygofer, setae not 


and distal area of wing, hind wing dark 
(Fig. 10). Female. Color of head orange red, 
coronal suture black, pronotum orange red, 
scutellum orange brown, anterior clypeus 
orange red and posteriorly black, clypellus 
black; legs orange, tarsus black brown; 
forewing orange red, discal cell with orange 
spots, apical portion orange, hind wing 
dark, abdomen brown red (Figs. 11—12). 


VOLUME 107, NUMBER 2 


20 
Fig. 20-22. 


22 


Brevimetopia chusquea. 20, Female genitalia (abdominal sternum VII, pygofer, ovipositor). 21, 


First valvula of ovipositor (emargination on ventral margin is the result of damage). 22, Second valvula of 


ovipositor. 


Thorax: As in generic description. Fore- 
wing very finely pubescent, rugose and 
punctate. Hind legs at rest with knees al- 
most reaching proepimeron, setal formula 
of male 2:0:0 and 2:1:0 in female. 

Male genitalia: Aedeagus, without 
toothlike medial processes. 

Female: Abdominal sternum VII pale, 
broadly sinuate with elevated truncate me- 
dian lobe shiny black (Fig. 20). First val- 
vula of ovipositor of uniform width except 
for short attenuated apex (Fig. 21). Second 
valvulae somewhat blade-shaped width 
roughly uniform for entire length, laterally 
compressed, with apex less curved than ar- 
cuate base. Apical portion narrowed and 
dorsal convex with truncate apex. Texture 
moderately to ligthly sclerotized, ventral 
hyaline area almost transparent. Dorsal 
teeth of valvulae broad and rounded (Fig. 
22). Third valvulae without setae. 

Type material—-Holotype: 6, COSTA 
RICA, San José, Cerro de la Muerte, 19 km 
S 3W Empalme, Mirador Quetzal, 2,600 m, 
xii.1999. Hanson & Godoy (UCR) (INBio). 
Paratypes: 1 3d, San José, Pavas, 1,100 m, 
20.ix.1981. A. Simons; 1 6 same data ex- 
cept vi. 2000 (CAS). 1 @, Cartago-San 
José, 20 km. SE Empalme, 2,800 m. 


V.1988. I. Gauld (INBio); 1 2, San Jose, 
Mirador de Quetzales, 2,600 m, on Chus- 
quea tonduzii, 28.V1.2003, A. Hicks 
(USNM); 1 @, same data, R. Rakitov 
(INHS)]. 

Etymology.—-The specific name refers 
to the host plant. 

Remarks.—-Brevimetopia chusquea is 
similar to B. silenciosa, but the former dif- 
fers in coloration, is less pubescent, has a 
setal formula of 2:0:0, and the basal pro- 
cesses of the aedeagus lacks toothlike pro- 
jections. Moreover, in B. chusquea the py- 
gofer is more rounded apically, the style has 
a smaller preapical lobe and narrower 
apophysis, and the connective stem is short- 
er and more slender than in B. silenciosa. 
The host plant appears to be the bamboo, 
Chusquea tonduzii Hackel (Poaceae). The 
two females collected in 2003 were placed 
in transparent plastic cages with plant cut- 
tings and were observed laying clusters of 
eggs under the lower epidermis of bamboo 
leaves (R. Rakitov, personal communica- 
tion). The Pavas (1,100 m) locality for one 
of the specimens is exceptional in that it is 
the only record of the genus outside the Tal- 
amanca mountain range. 


266 


ACKNOWLEDGMENTS 


I thank Roman Rakitov for stimulating 
discussions, permission to cite his obser- 
vations of oviposition in B. chusquea, com- 
ments on the manuscript, and the photo- 
graphs (Figs. 11-14, 20—22). I also thank 
Paul Hanson and two anonymous reviewers 
for comments on the manuscript, Alejandro 
Herrera for drawing the pygofer and plate 
of B. chusquea, and the Serrano family for 
their kind permission to collect in the Mir- 
ador de Quetzales. 


LITERATURE CITED 


Kramer, J. P. A. 1967. Taxonomic study of the bra- 
chypterous North American leafhoppers of the ge- 
nus Lonatura (Homoptera: Cicadellidae: Delto- 
cephalinae). Transactions of the American Ento- 
mological Society 93: 433-462. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Linnavuori, R. 1959. Revision of the Neotropical Del- 
tocephalinae and some related subfamilies (Ho- 
moptera). Annales Zoologici Secretaries “Vana- 
mo’ 20: 1—369. 

Oman, P. W. 1949. The Neartic Leafhoppers (Homop- 
tera: Cicadellidae). Memoirs of the Entomological 
Society of Washington, No. 3, 253 pp. 

. 1985. A synopsis of the Neartic Dorycephal- 

inae (Homoptera: Cicadellidae). Journal of the 

Kansas Entomological Society 58(2): 314-336. 

. 1987. The leafhopper genus Errhomus (Ho- 
moptera: Cicadellidae: Cicadellinae), systematics 
and biogeography. Oregon State University Sys- 
tematic Entomology Laboratory. Occasional Pub- 
lication 1, 72 pp. 

Oman, P. W. and C. A. Musgrave. 1975. The Neartic 
genera of Errhomenini (Homoptera: Cicadellidae). 
Melanderia 21: 1—14. 

Young, D. A 1968. Taxonomic study of the Cicadel- 
linae (Homoptera: Cicadellidae). Part 1. Proconi- 
ini. United States National Museum Bulletin 261: 
1-287. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 267-272 


A NEW PTILONEURID GENUS (PSOCOPTERA: PTILONEURIDAE) 
FROM DOMINICA 


ALFONSO N. GARCIA ALDRETE 


Departamento de Zoologia, Instituto de Biologia, Universidad Nacional Aut6noma de 
México. Apartado Postal 70-153, 04510 México, D. FE, México (e-mail: anga@ 
ibiologia.unam.mx) 


Abstract.—Willreevesia dominica, new genus, new species, is described from the island 
of Dominica, Lesser Antilles. It has a combination of complete labral sclerites, all distal 
labral sensilla but the central one trichoid, distinct anterior extensions of V2+3, complex 
phallosome, with triangular external parameres and mesal three pronged structure and 
hind wing M simple, which makes it unique in the family. A cladistic analysis indicates 
that it is the sister taxon to a clade consisting of Ptiloneuropsis Roesler, Ptiloneura En- 


derlein, and Loneura Navas. 


Key Words: 


The Neotropical epipsocete family Pt- 
loneuridae presently includes the genera 
Euplocania Enderlein (Brazil, Nicaragua, 
Paraguay, Peru), Triplocania Roesler (Be- 
lize, Bolivia, Brazil, Colombia, Costa Rica, 
Guatemala, México, Panama, Peru), Prilo- 
neura Enderlein (Pert), Ptiloneuropsis 
Roesler (Brazil), Loneura Navas (Argenti- 
na, Bolivia, Brazil, Costa Rica, Guatemala, 
México, Nicaragua, Peru, U.S.A., Venezue- 
la) and Perucania New & Thornton (Peru). 
All the species so far documented in the 
family are continental. Here I describe a 
new genus in the family, from specimens 
recently collected on the island of Domin- 
ica (Lesser Antilles). 

The specimens for microscopic study 
were dissected in 80% alcohol, and their 
parts (head, wings, legs, and genitals), were 
mounted on slides in Canada balsam. Color 
was recorded by observation of whole spec- 
imens illuminated with cold, white light at 
50. Measurements of parts on the slides 
(given in microns), were taken with a filar 
micrometer whose measuring unit is 1.36 


Ptiloneuridae, Willreevesia, new genus, Dominica, Lesser Antilles 


microns for wings and 0.53 microns for 
other parts. Abbreviations are as follows: 
FW = forewing; HW = hind wing; F = 
femur; T = tibia; tl, t2, t3 = hind leg tar- 
someres; cttl = number of ctenidia on tl 
of hind leg; Mx4 = fourth segment of max- 
illary palp; fl ... fn = flagellomeres fl... 
fn; IO = minimum distance between com- 
pound eyes; D = antero-posterior diameter 
of compound eye; d = transverse diameter 
of compound eye; PO = d/D. The holo- 
types and most paratypes are deposited in 
the National Insect Collection (CNIN), De- 
partamento de Zoologia, Instituto de Biol- 
ogia, Universidad Nacional Autonoma de 
México, México City. One paratype of each 
sex will be deposited in the Department of 
Entomology, Soils and Plant Sciences, 
Clemson University, Clemson, South Car- 
olina, U.S.A. 


Willreevesia Garcia Aldrete, new genus 


Diagnostic features.—Belonging in the 
Ptiloneuridae (sensu Lienhard and Smithers 
2002). Labral sclerites complete, slender, 


268 


tenuous, curved outwards at each end, not 
as well defined as in Epipsocidae, Cladiop- 
socus or Neurostigmatidae. Five distal la- 
bral sensilla, a central large placoid, and 
two smaller trichoid on each side. Forewing 
M five or six branched, branch next areola 
postica distally forked. Hindwing M simple. 
V1 long, slender; V2+3 with a distinct dis- 
tal process. Hypandrium simple. Phallo- 
some with lateral struts stout, V-shaped, ex- 
ternal parameres triangular and a mesal, 
strongly sclerotized, three pronged struc- 
ture. 

Type species.—Willreevesia dominica, 
new species. 


Willreevesia dominica Garcia Aldrete, 
new species 
(Figs. 1-11) 


Female.—Color: Ground color creamy 
white, with reddish- brown areas as indi- 
cated below. Compound eyes black, ocelli 
hyaline, with ochraceous centripetal cres- 
cents. Head pattern same as illustrated for 
male (Fig. 2). A dark brown, large, almost 
rectangular area on each gena, between 
compound eye and mandible. Maxillary 
palp creamy white. Antenna brown, flagel- 
lomeres with apices white. Thorax with ir- 
regular ochraceous areas on each pleuron. 
Tergal lobes of meso- and metathorax red- 
dish brown. Legs pale brown. Femora with 
proximal, median and distal brown spots. 
Wings almost hyaline, slightly washed with 
a brown hue; veins brown, each with a 
small brown macula distally. Forewing 
pterostigma with a proximal and a distal 
brown band; brown areas on distal ends of 
Cu2 and 1A, and between CulA and wing 
margin. Abdomen with distinct, transverse, 
ochre subcuticular bands. 

Morphology: Wacinial apex with outer 
cusp large, with 6—8 denticles (Fig. 11). La- 
bral sclerites complete, slender, outwardly 
curved at each end (Fig. 5). Five distal la- 
bral sensilla; one large central placoid, with 
two smaller trichoid ones on each side (Fig. 
6). Forewing (same as illustrated for the 
male, Fig. 1), M with 5—6 branches, branch 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


next to areola postica distally forked. Are- 
ola postica wide, high, with apex rounded. 
Pterostigma elongate, proximally narrow, 
wider in middle. Hindwing (same as illus- 
trated for male, Fig. 1), with M_ simple. 
Subgenital plate approximately triangular, 
broad, setose, with a well-defined pigment- 
ed area on each side (Fig. 7). Gonapophyses 
(Fig. 4): first valvulae (V1) long, slender, 
with a field of short spines distally on outer 
edge; V2+3 robust, extended anteriorly as 
illustrated, with a distinct, long distal pro- 
cess, armed distally with numerous micros- 
pines; third valvulae (V3) a bulge on side 
of second valvulae (V2), with 6—8 macro- 
setae. Ninth sternum distinct, with a strong- 
ly sclerotized band longitudinally, dividing 
sclerite in two symmetric halves (Fig. 4). 
Paraprocts robust, with setae as illustrated; 
sensory fields elliptical, with 22—23 tricho- 
bothria issuing from basal rosettes (Fig. 9). 
Epiproct broad, triangular, with a group of 
three mesal macrosetae near anterior border, 
other setae on distal half, as illustrated (Fig. 
)). 

Measurements: FW: 4209, HW: 2918, F: 
1224, T: 1864, tl: 871, t2: 80, t3: 145, cttl: 
PES IOS WOS)5 18 teksi5\5 ZB SIODs MOK i013), ID: 
AOS, dz 2815 lO/D= 0945 PO 0:69: 

Male.—Color: Same as female. 

Morphology: Head (Fig. 2), with large, 
bulging compound eyes. Hypandrium sim- 
ple, broad, setose, pigmented area as illus- 
trated (Fig. 8). Phallosome complex, with 
lateral struts robust, V-shaped; external par- 
ameres triangular, each internally associated 
with a stout, acuminate prong, and with an 
elongate, slender sclerite posteriorly, distal- 
ly denticulate. Mesally in area between ex- 
ternal parameres, a broad based, distinct 
sclerotized structure, posteriorly with a cen- 
tral, stout, distally rounded column, with a 
slender baculum underneath, and extended 
to form on each side an acuminate prong; 
above this three-pronged structure, posteri- 
orly, a strongly sclerotized disk (Fig. 3). 
Paraprocts broad, robust, setose; sensory 
fields elliptical, with 30—32 trichobothria is- 
suing from basal rosettes (Fig. 10). Epiproct 


VOLUME 107, NUMBER 2 269 


rei ee 
eo 


Na eee ROR Ee 
CARES oy, i [ S rie a 
EAS y 


y /; / y 
A 


Figs. 1-11. Willreevesia dominica. 1, Fore- and hind wings. d. 2, Front view of head. ¢. 3, Phallosome. 
3.4, Gonapophyses and ninth sternum. @. 5, Labrum. 2. 6, Distal labral sensilla. 2. 7, Subgenital plate. 2. 8, 
Hypandrium. d. 9, Right paraproct and epiproct. 2. 10, Epiproct and left paraproct. ¢. 11, Distal end of lacinia. 
2. Abbreviations: p, pterostigma; ap, areola postica; V1, first valvula; V2+3, valvula 2+3; Is, labral sclerite. 
Scales in mm. 


270 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Characters utilized for the phylogenetic analysis of Ptiloneuridae. 


SS 


HW M: Unbranched (0), 3—5 branched (1). 


A bBWN — 


_ EW M: 3 branched (0), 4 branched (1), 5—8 branched (2). 


FW pterostigma with a spur-vein (0), or without a spur-vein (1). 
_ FW areola postica high, rigidly triangular (0), or high, with apex rounded (1), or low, extremely long (2). 
. FW areola postica free (0), or joined to M by a crossvein (1). 


6. EW 2 A more than half the length of 1 A (0), or less than half the length of 1 A (1). 
7. Labral sclerites complete, curved outward (0), or incomplete, straight (1). 
8. Male hypandrium a single sclerite, without posterior projections (Q), or a central sclerite, with posterior 


projections, flanked by small sclerites (1). 


9. Phallosome complex, with symmetric phallic sclerites and lateral struts V shaped (O), or simple, without 


phallic sclerites and lateral struts straight (1). 


10. Phallosome with a transverse, three pronged sclerotized structure in area between external parameres (0), 
or with paired sclerites in same area (1), or without sclerites (2). 

11. External parameres broadly triangular, distally pointed (0), or elongate, distally rounded (1). 

12. Epiproct of males and females with a group of three mesal macrosetae near anterior border (0), or with- 


out mesal macrosetae near anterior border (1). 


13. Male paraprocts with mesal prongs (0), or without mesal prongs (1). 
14. Female ninth sternum with large, distinct spermapore (0), or with spermapore inconspicuous (1). 
15. Apices of female paraprocts and epiproct with stout setae (0), or without stout setae (1). 


ee 


trapezoidal, with a group of three mesal 
macrosetae near anterior border, three setae 
along posterior edge, in a field of micros- 
pines, and other setae on posterior third, as 
illustrated (Fig. 10). 

Measurements: FW: 4390, HW: 3050, F: 
DSO, Te 2Ooll, tile QS4s, ts SO, tse MSZ, eal 
Sil, IMPAS SiO, ile MOG wwe ox, Os Zilae 
De SIG, Ge FOS; WOMDs W.37/, IROs WSs. 

Types.- Holotype 6, Dominica. Parish of 
SG JOsSephy eS prneheld ae Statc als — 
20.11.2003. Malaise trap in humid forest, 
A307 mi (15222-84Ne 61205) W) 2 Ma Es I 
win, M. B. Shepard, E. Benson, G. Carnet. 
Same data as holotype, 2 ¢6¢, 1 @ para- 
types. Parish of St. Mark, 4 km N Soufriere, 
17—19.111.2003, Malaise trap in dry wash of 
deciduous forest, 70 m (15°14.3'N: 
61°22'W), same collectors, allotype 2; 7 
36,6 2 paratypes. 1 km W Pt. Guig- 
nard, 17—19.1I1.2003, Malaise trap in dry 
wash of deciduous forest (15°14.6'N: 
61°22.3’W), same collectors, 12 paratype. 

Etymology.—The genus name honors Dr. 
Will Reeves of Clemson University, South 
Carolina, for making available for study a 
collection of Psocoptera from Dominica, 
which included the specimens here de- 
scribed, and in recognition of his work on 


Diptera and on cavernicolous organisms. 
The specific name refers to the island of 
Dominica to which this taxon is endemic. 
The genus name is feminine; the species 
name is a noun in apposition. 


THE POSITION OF WILLREEVESIA IN 
PTILONEURIDAE 


To assess the phylogenetic relationships 
of the ptiloneurid genera, including Wi/lI- 
reevesia, a matrix of 15 characters was as- 
sembled (Tables 1—2). Spurostigma Eert- 
moed and Cladiopsocus Roesler were also 
included and treated as outgroups, both be- 
Jong in the family Cladiopsocidae and con- 
sidered close to the Ptiloneuridae (Yoshi- 
zawa 2002), although that author did not 
recognize Cladiopsocidae as a monophylet- 
ic taxon. The matrix was edited utilizing 
WinClada 0.9.99 (Nixon 1999) and ran in 
Nona 2.1 (Goloboff 1997). All characters 
were unordered and had the same weight. 
1,000 replicas were run in groups of 250 
sequences of random addition (Kitching 
1995, Lipscomb 1998), and retaining 20 
trees in each replica (non ambiguous opti- 
mization, heuristic search with the follow- 
ing parameters: h = 10,000; mult* = 250; 


VOLUME 107, NUMBER 2 


Table 2. Matrix of characters for the phylogenetic analysis of Ptiloneuridae. Spurostigma and Cladiopsocus are outgroups. 


15 


14 


13 


11 


10 


9 


5 


6 


nN 


0,1 


Euplocania 


0,1 


Triplocania 


AN 


Ptiloneura 


Ptiloneuropsis 


0,1 


N 


Loneura 


nN 


Perucania 


nN 


Willreevesia 


ior 


Spurostigma 


AN 


Cladiopsocus 


Dafa 


h/ = 20; the analysis was repeated ten times 
with the same parameters). 

A single tree was found, with L = 20 
steps, a Consistency Index of 0.90 and a 
Retention Index of 0.85 (Fig. 12). The clad- 
ogram indicates that Ptiloneuridae is mono- 
phyletic, supported by characters 9:0, 10:0, 
12:0, and 14:1. Two distinct clades are rec- 
ognized: one, including Perucania, Euplo- 
cania, and Triplocania, supported by char- 
acter 8:1, and the other, including Willreev- 
esia, Ptiloneuropsis, Ptiloneura, and Lo- 
neura, supported by character 1:2. 
Willreevesia is the sister group to the as- 
semblage Ptiloneuropsis-Ptiloneura-Loneu- 
ra, differing from them in having the labral 
sclerites complete (7:0) and the external 
parameres of the male phallosome distinctly 
triangular (11:0), and by having the HW M 
simple (2:0). It shares with Ptiloneuropsis 
a simple hypandrium, without posterior 
projections (8:0). 


DISCUSSION 


Willreevesia presents a combination of 
characters that makes it unique in the fam- 
ily: It is the only ptiloneurid with complete 
labral sclerites and with triangular male ex- 
ternal parameres; also, it is the only one in 
which the outermost distal labral sensilla 
are trichoid and in which the anterior ex- 
tension of the gonapophyses are distinctly 
different from those in the other genera. 
Also unique is the combination of wing ve- 
nation characters: branches of FW M as in 
Ptiloneuropsis-Ptiloneura-Loneura, com- 
bined with HW M as in the members of the 
other clade (Perucania-Euplocania-Triplo- 
cania). Other than Willreevesia, all the oth- 
er ptiloneurids are continental. Since Pti- 
loneuropsis and Ptiloneura are quite re- 
stricted geographically (only known from 
Itatiaia National Park, Rio de Janeiro, Bra- 
zil and from Callanga and Vilcanota Songo, 
Chaco, Peru, respectively), it is likely that 
Willreevesia is an offshoot of Loneura, the 
species of which occur from Arizona, USA, 
to northern Argentina, in an episode of is- 


272 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 

“a 

nm ie) 

S S < S s = S S 
5 ° SS ~ S 5} S SS < 
op aS = 8 8 co ~ ~ ~ 
> Q 3 5) Ss) Y v & = 
= S = g Q = = ~ = 
= Ss D = > = S s ca) 
a S A, Q = = aL a 5 
: j ; 55 
2 (=) 
= 2 

Fig. 12. Phylogeny of the genera of Ptiloneuridae, generated by Nona. L = 20, CI = 0.90, RI = 0.85. 


Spurostigma and Cladiopsocus are outgroups 


land colonization followed by evolution in 
isolation. 


ACKNOWLEDGMENTS 


I express thanks to José Arturo Casasola, 
Javier Garcia Figueroa, and Felipe Villegas 
(Instituto de Biologia, Universidad Nacion- 
al Autonoma de México), for technical sup- 
port in the preparation of this paper. Will 
Reeves (Clemson University, South Caro- 
lina) in addition to donating for study the 
specimens here described, obtained for me 
a bibliographical reference unobtainable in 
México, needed to understand the distribu- 
tion of Willreevesia. 


LITERATURE CITED 


Goloboff, P. A. 1997. Nona, program and documen- 
tation. Published by the author. Tucuman, Argen- 
tina, 34 pp. 


Kitching, I. J. 1995. Tree building techniques, pp. 44— 
71. In Forey, P. L., C. J. Humphries, I. J. Kitching, 
R. W. Scotland, D. J. Siebert and D. M. Williams, 
eds. Cladistics: A practical course in systematics. 
The Systematics Association. Publication 10. Ox- 
ford University Press, U.K. 

Lienhard, C. and C. N. Smithers. 2002. Psocoptera (In- 
secta). World Catalogue and Bibliography. Instru- 
menta Biodiversitatis V. Muséum d’ histoire na- 
turelle, Geneve, Switzerland, 745 pp. 

Lipscomb, D. 1998. Basics of cladistic analysis. 
George Washington University Press, Washington, 
D. C. Available at www.gwu/~clade/faculty/ 
Lipscomb/cladistics.pdf. 

Nixon, K. C. 1999. Winclada, program and docu- 
mentation. Published by the author. Ithaca, New 
York. 

Yoshizawa, K. 2002. Phylogeny and higher classifi- 
cation of suborder Psocomorpha (Insecta: Psoco- 
dea: ‘Psocoptera’). Zoological Journal of the Lin- 
nean Society 136: 371—400. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 273-278 


TWO NEW FERN-FEEDING SAWFLIES OF THE GENUS ANEUGMENUS 
HARTIG (HYMENOPTERA: TENTHREDINIDAE) FROM SOUTH AMERICA 


DAVID R. SMITH 


Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- 
ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, 
P.O. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. (e-mail: dsmith@sel. 


barc.usda.gov) 


Abstract.—Aneugmenus merida, n. sp., from Venezuela and Ecuador and A. colombia, 
n. sp., from Colombia are described. Aneugmenus merida was reared from larvae feeding 
on bracken fern, Pteridium aquilinum (L.) Kuhn. in Venezuela. These are the only two 
species of this Holarctic genus known from South America. The species are illustrated 
and separated from other New World species of Aneugmenus. Aneugmenus leucopoda 


(Cameron 1883) is a new combination. 


Key Words: 


Species of Aneugmenus Hartig are rather 
stout sawflies, around 6 mm in length, and 
are mostly black with contrasting yellow 
legs. They are distinguished from other 
New World selandriine genera by the ab- 
sence of the anal crossvein in the forewing, 
the presence of an epicnemium as a flat 
sclerite separated from the mesepisternum 
by a suture, a genal carina at least partially 
developed on the sides of the head, a cir- 
cular carina on the frons, a flat clypeus with 
the anterior margin tuncate, a sessile anal 
cell of the hind wing, and bifid tarsal claws 
with a distinct basal lobe. 

Smith (1969) revised the Nearctic species 
of Aneugmenus and included three species. It 
was later determined that A. floridella Ross 
1930 was distinct; thus, four species are listed 
in the North American catalog (Smith 1979). 
Other than A. scutellatus Smith 1969 from 
southern Arizona and northern Mexico, only 
two species have been described from the 
Neotropics, A. nigritarsis Rohwer 1911 from 
Mexico and A. leucopoda (Cameron 1883) 
from Guatemala. There are, however, a num- 


Selandriinae, ferns, Venezuela, Colombia 


ber of undescribed species from Mexico and 
Central America. 

One of the species treated here was 
reared from Pteridium aquilinum (L.) 
Kuhn. in Venezuela by Daniel Otero and 
Maria Pia Calcagno, Universidad de Los 
Andes, Mérida, Venezuela, who are work- 
ing on its life cycle, feeding habits, and 
ecology. The reared adults proved to be a 
new species of Aneugmenus Hartig. Identity 
of this species prompted this review of the 
South American species of Aneugmenus, 
and, because I am aware of only one other 
species in South America, a new species 
from Colombia also is described. Colombia, 
Ecuador, and Venezuela are the southern- 
most records for Aneugmenus in the West- 
ern Hemisphere. 


Aneugmenus merida Smith, new species 
(Figs. 1-7) 


Female.—Length, 6.0 mm. Antenna and 
head black. Thorax black with tegula and 
posterior margin of pronotum yellow. Legs 
yellow with coxae and trochanters black 


Figs. 1-3. Aneugmenus merida, female. 1, Head, 
front view. 2, Head, dorsal view. 3, Antenna. 


and tarsi blackish. Abdomen orange with 
basal plates, anterior margin of second seg- 
ment, and apical 2 to 3 segments and sheath 
black. Wings moderately uniformly infus- 
cated; veins and stigma black. 

Antennal length 1.3 head width; Ist 
segment broader than long; 2nd segment 
longer than broad; 3rd segment longer than 
4th segment (Fig. 3). Clypeus truncate. Ma- 
lar space linear. Head in front view (Fig. 1) 
with eyes strongly converging below; lower 
interocular distance slightly shorter than 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


eye length, upper interocular distance 1.4x 
eye length. Genal carina present laterally, 
absent near top of eye, on head above, and 
behind postocellar area. Head in dorsal 
view straight behind eyes; distances from 
eye to lateral ocellus, between lateral ocelli, 
and from lateral ocellus to hind margin of 
head as 1.0:1.0:0.7. Postocellar area about 
2.0 broader than long. Epicnemium pres- 
ent as a flat sclerite separated from mese- 
pisternum by a suture. Forewing with 4 cu- 
bital cells. Hind wing with anal cell sessile. 
Hind basitarsus 0.8X length of remaining 
tarsal segments combined. Tarsal claws 
with inner tooth and basal lobe. Sheath 
short and rounded in lateral view. Lancet 
(Fig. 7) with 5 serrulae, 7 broad alar spines 
with basal two small and near ventral mar- 
gin; apex truncate. 

Male.—Length, 6.0 mm. Color similar to 
female, with abdomen mostly orange (Fig. 
6) except for blackish anterior margin to an- 
terior half of basal plates and entire hypan- 
drium. Seventh tergum with a deep groove 
(sinus sexualis) and 8th tergite with broad, 
slightly concave roughened area (Figs. 4— 
5). Genitalia (Figs. 11-12) with harpe 
curved inward and apex almost truncate; 
parapenis long, narrow, gradually tapering 
to narrow rounded apex; penis valve nearly 
rectangular, apex broadly rounded. 

Types.—Holotype: 2 labeled “Cerro La 
Bandera, La Hechicera, Mérida, 2,100 m, 
Edo. Merida, Venezuela,” ‘““Julio/2003, J. 
L. Avila leg.” Deposited in Instituto de 
Zoologia Agricola, Universidad Central de 
Venezuela, Maracay, Venezuela. 

Paratypes: ECUADOR: Banos, Tungur- 
ahua, 1,820 m, II-11—55, E. I. Schlinger, E. 
S. Ross, collectors (1 2). VENEZUELA: 
Same data as for holotype (1 2, 1 d); La 
Hechicera, 1900—2100 m, Mérida, Edo Mé- 
rida, Octobre/2003, J. L. Avila, 2 ¢ reared 
from oviposition of field collected females, 
1 @ field collected (1 2, 2 ¢); Mérida; Los 
Pedregosa, in hills near Merida, 16 March 
[9S25GaE eae Hevel di oo alivs))Me= 
rida, Timotes, | June 1976, A.S. Menke & 
D. Vincent (1 2). The paratype from Ec- 


VOLUME 107, NUMBER 2 


Figs. 4-6. Aneugmenus merida, male. 4, Apex of abdomen, lateral view, showing sinus sexualis in seventh 
tergite. 5, Apex of abdomen, dorsal view. 6, Dorsal view. 


uador is deposited in the California Acad- 
emy of Sciences, San Francisco; paratypes 
from Venezuela are in the Facultad de Cien- 
cias, Universidad de Los Andes, Mérida, 
Venezuela, and the National Museum of 
Natural History, Smithsonian Institution, 
Washington, DC. 

Two larvae also associated with the ““Oc- 
tobre/2003” collection are not paratypes. 

Food plant——Larvae feed on fronds of 
bracken fern, Pteridium aquilinum (L.) 
Kuhn. 

Etymology.—The name is from the type 
locality and is a noun in apposition. 


Discussion.—This is the only known 
species of Aneugmenus with a mostly or- 
ange abdomen in both sexes. Some other 
species have the abdomen partly orange 
only in the male, but in all others the female 
abdomen is entirely black. 

Three described species are known south 
of the United States: A. leucopodus from 
Guatemala, A. nigritarsis from Mexico, and 
A. scutellatus from southern Arizona and 
northern Mexico. 

Aneugmenus leucopoda has the antenna 
and head black; the thorax black with the 
tegula, posterior margin of the pronotum, 


276 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 7-10. Female lancets. 7, Aneugmenus merida. 8, A. nigritarsis, holotype. 9, A. scutellaris. 10, A. 


colombia. 


and spot on the lower posterior margin of 
the mesepisternum yellow; the legs except 
the coxae and trochanters yellow; and the 
abdomen black with the extreme apices of 
the segments beneath broadly white. 

Aneugmenus leucopoda (Cameron 1883) is 
a new combination. Cameron described a fe- 
male as Selandria leucopoda from *“*‘Guate- 
mala, San Geronimo.” Rohwer (1912) later 
transferred it to Proselandria Forsius. I ex- 
amined the holotype, housed in The Natural 


History Museum, London, BM #1.265, la- 
beled ““S. Geronimo, Guatemala, Champion” 
and with name labels. It is a typical Aneug- 
menus. The male is unknown. 

Aneugmenus nigritarsis was described 
from “‘San Rafael, Jicoltepec, Mexico.” 
The holotype, in the National Museum of 
Natural History, Smithsonian Institution, 
was examined. I have also seen specimens 
from the Mexican states of Puebla and San 
Luis Potosi. This species is black with the 


VOLUME 107, NUMBER 2 


Si 11 12 


Figs. 11-12. Aneugmenus merida, male genitalia. 
11, Genital capsule, ventral view of left half. 12, Penis 
valve, lateral view. 


labrum, tegula, posterior margin of the 
pronotum, legs below the bases of the cox- 
ae yellow, and the apex of the clypeus 
brownish. The female lancet of A. nigritar- 
sis 18 short, with 5 serrulae and only 4—5 
distinct alar spines (Fig. 8). The serrulae are 
narrow and far apart, and the apex of the 
lancet is rounded. The male is unknown. 

Aneugmenus scutellatus is black with the 
labrum and palpi white; tegula, posterior 
angles of the pronotum and mesoscutellum 
white to yellow; legs except coxae yellow; 
and the abdomen black. The lancet has 6 
serrulae and 6—7 alar spines (Fig. 9) and the 
apex is rounded. In the male, tergites 2—6 
are reddish and sternites 1—3 are yellowish, 
and the genitalia (Smith 1969, figs. 112— 
113) have the harpe straighter, not nearly as 
curved inward as in A. merida, and the pe- 
nis valve has a narrow dorsal lobe. 

Three other species occur in North 
America. Aneugmenus flavipes (Norton 
1861) in eastern Canada and United States, 
A. floridella Ross 1930 in Florida, and A. 


DHT) 


padi (Linnaeus 1761) an introduced Pale- 
arctic species in western Canada and north- 
western United States. All are black with 
the tegula, posterior margin of the prono- 
tum, and legs yellow. Occasional specimens 
of A. flavipes and A. floridella have the 
clypeus partly yellowish, and males of A. 
floridella have the abdominal dorsum partly 
or mostly orange. Pteridium aquilinum is a 
recorded food plant for A. flavipes and A. 
padi (Smith 1969). 

The larva of A. merida is typical for spe- 
cies of Aneugmenus. The abdominal seg- 
ments are 7-annulate; prolegs are present on 
abdominal segments 2—8 and 10; the body 
is whitish (preserved specimens, probably 
green when alive) without ornamentation 
and with only the spiracles and tarsal claws 
dark brown; the head is pale without dark 
markings except black eyespots, antennal 
segments, narrow line on the clypeal suture, 
and mandible apices; and the prothorax has 
a pair of fleshy protuberances just behind 
the head capsule. The larva is very similar 
to that of A. flavipes. 

The male possesses the unusual concayv- 
ity (sinus sexualis) on the seventh tergum 
(Figs. 4—5) that appears typical for male 
Aneugmenus. This structure is present in A. 
flavipes and most other species of the ge- 
nus. Smith and Marshall (2003) reported on 
and gave photos of the female apparently 
gaining some nutritional substance from 
this structure before and/or after mating. 
Aneugmenus merida appears to have a sim- 
ilar courtship behavior, which is under in- 
vestigation by Daniel Otero and Maria Pia 
Calcagno. 


Aneugmenus colombia Smith, 
new species 
(Fig. 10) 


Female.—Length, 5.5 mm. Black with 
labrum white; apex of mandible reddish 
brown; tegula, posterior angles of prono- 
tum, and femora and tibiae yellow; tarsi 
blackish, darker apically. Wings uniformly, 
darkly infuscated; veins and stigma black. 

Antennal length 1.2 head width; oth- 


278 


erwise similar to Fig. 3. Forewing with first 
branch of Rs absent, thus with 3 cubital 
cells. Lancet (Fig. 10) short, dorsoapical 
margin protuberant, with 6 shallow serrulae 
and 6—7 alar spines. Other features similar 
to A. merida, Figs. 1—2). 

Male.—Unknown. 

Holotype.—Female, labeled ““COLOM- 
BIA: 5 mi E. Guaduas, Cundin Amarca, 
1,400 m, IHI-15—55,”" “E.I. Schlinger & E. 
S. Ross, collectors.’’ Deposited in the Cal- 
ifornia Academy of Sciences, San Francis- 
co. 

Etymology.—Named from the country of 
collection, a noun in apposition. 

Discussion.—The black coloration and 
short lancet are most similar to A. nigritar- 
sis, but in A. colombia the serrulae are shal- 
lower and broader, the apical serrulae are 
closer together, and the dorsoapical margin 
is protuberant. Examination of the color 
pattern and lancet are necessary for identi- 
fication of this species. 


ACKNOWLEDGMENTS 


I thank Daniel Otero and Maria Pia Cal- 
cagno, Universidad de Los Andes, Mérida, 
Venezuela, for bringing these specimens to 
my attention. Cathy Apgar, Systematic En- 
tomology Laboratory, U.S. Department of 
Agriculture (SEL), took the Auto-Mon- 
tage™ photos and arranged the plates. I 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


thank the curators at The Natural History 
Museum, London, and the California Acad- 
emy of Sciences, San Francisco, for allow- 
ing study of material in their possession, 
and N. M. Schiff, U.S. Forest Service, 
Stoneville, MS, and S. J. Scheffer (SEL) 
and T. J. Henry (SEL), Beltsville, MD, and 
Washingon, D.C., respectively, for review- 
ing the manuscript. 


LITERATURE CITED 


Cameron, P. 1883. Hymenoptera, Tenthredinidae— 
Chrysididae. Jn Godman and Salvin, Biologia 
Centrali-Americana, Vol. 1, 486 pp. 

Rohwer, S. A. 1911. New sawflies in the collections 
of the United States National Museum. Proceed- 
ings of the United States National Museum 41: 
377-411. 

. 1912. Sawflies from Panama, with descrip- 
tions of new genera and species. Smithsonian 
Miscellaneous Collections 59: 1-6. 

Smith, D. R. 1969. Nearctic sawflies. Il. Selandriinae: 
Adults (Hymenoptera: Tenthredinidae). United 
States Department of Agriculture Technical Bul- 
letin No. 1398, 48 pp., 10 pls. 

. 1979. Symphyta, pp. 1-132. In Krombein, K. 
V., P. D. Hurd, Jr., D. R. Smith, and B. D. Burks, 
eds. Catalog of Hymenoptera in America North of 
Mexico, Vol. 1, Symphyta and Apocrita (Parasi- 
tica). Smithsonian Institution Press, Washington, 
DC. pp. 1—-1198. 

Smith, D. R. and S. A. Marshall. 2003. First report of 
nuptial feeding in sawflies, Aneugmenus flavipes 
(Norton) (Hymenoptera: Tenthredinidae). Pro- 
ceedings of the Entomological Society of Wash- 
ington 105: 789-791. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 279-302 


A REVISION OF THE GENUS ACANTHOTYLA STAL WITH THE 
DESCRIPTION OF FIVE NEW SPECIES AND SYNONYMICAL NOTE 
(HETEROPTERA: COREIDAE: COLPURINI) 


HARRY BRAILOVSKY 


Departamento de Zoologia, Instituto de Biologia, U. N. A. M., Apdo Postal No. 70153, 
México, 04510 D.E México. (e-mail: coreidae @ servidor.unam.mx) 


Abstract.—The genus Acanthotyla Stal is revised. The following new species are de- 
scribed: A. kaloboana, A. kiungala, A. nabirenia, and A. protenta all from New Guinea 
and A. borneana from Borneo. Acanthotyla aruana Horvath is synonymized with A. 
fasciata (Walker) (n. syn.); new records for the previously known species are added. 
Habitus illustrations and drawings of the male and female genitalia, as well as head and 
pronotum, are provided, and a key to the known species is presented. 


Key Words: 


Hemiptera, Heteroptera, Coreidae, Colpurini, revision Acanthotyla, new 


species, Borneo, New Guinea, synonymical note 


The genus Acanthotyla was proposed by 
Stal (1873) to include only the species Cle- 
tus fasciatus Walker (1871) from Mysol Is- 
Jand. Cletus crassus Walker (1871), de- 
scribed in the same paper, was transferred 
to Acanthotyla by Blote (1936), but later 
Brailovsky (1995) moved it to his new ge- 
nus Brachylybastella as the type species. 
Breddin (1900) included Acanthotyla in his 
key to the genera of Colpurini. The second 
known species, A. aruana Horvath (1919), 
is synonymyzed in this paper under A. fas- 
ciata. The last group of species was de- 
scribed by Blote (1936) from New Guinea 
under the binomial names Acanthotyla dis- 
tinguenda and Brachylybas flexuosus. Brai- 
lovsky (1993) revised the Colpurini of Aus- 
tralia and recorded A. fasciata for the first 
time from that region, and later Brailovsky 
and Martinez (1994) transferred B. flexu- 
osus to Acanthotyla. 

Acanthotyla belongs to the group of Col- 
purini with abdominal sternite VII of the 
female without plica or fissura, and has not 
been previously reviewed or revised. The 


genus is characterized by having the tylus 
projecting in front of juga, upturned to form 
a horn at the apex, the mandibular plates 
with prominent tubercle, the antennal seg- 
ment III longer than IV, the antenniferous 
tubercles at outer corner projected forward, 
calli conspicuously convex, and posterior 
lobe of pronotal disk with a transverse 
wrinkle. The specific differences are chiefly 
in the development of the humeral angles 
and the intercallar space of the pronotum; 
the shape of the posteroventral edge of the 
male genital capsule, and shape of the fe- 
male genital plates. 

Members of this genus are distributed 
from Borneo, Aru Island, Mysol Island, Iri- 
an Jaya, Papua New Guinea, and Australia 
(Brailovsky 1993). 

Previously, only three species of Acan- 
thotyla, A. distinguenda, A. fasciata, and A. 


flexuosa, were known. In this contribution 


the genus is redescribed, new records for A. 
distinguenda, A. fasciata, and A. flexuosa 
are given, and five new species, from Bor- 
neo, Irian Jaya, and Papua New Guinea are 


280 


described. Acanthotyla aruana Horvath is 
synonymyzed under A. fasciata. 

The following abbreviations indicate in- 
stitutions where specimens are deposited or 
from which material was generously lent: 
The Natural History Museum, London 
(BMNH); Bernice P. Bishop Museum, Hon- 
olulu, Hawaii (BPBM); California Acade- 
my of Sciences, San Francisco, California 
(CAS); Hungarian Natural History Muse- 
um, Budapest (HNHM); Natural History 
Museum of Los Angeles County, California 
(LACM); Queensland Museum, Brisbane, 
Australia (QMBA); Rijksmuseum van Na- 
turlijke Histoire, Leiden, Netherlands 
(RNHL); Forschungsinstitut und Naturmu- 
seum Senckenberg, Frankfurt am Main, 
Germany (SMFD); Coleccion Entomol6gi- 
ca del Instituto de Biologia, Universidad 
Nacional Aut6noma de México (UNAM); 
National Museum of Natural History, 
Smithsonian Institution, Washington D.C. 
(USNM);: Zoologisches Museum, Hum- 
boldt Universitat, Berlin, Germany 
(ZMHB); Zoologische Staatssammlung 
Miinchen, Germany (ZSMC). 

All measurements are given in millime- 
ters. 


Acanthotyla Stal 


Acanthotyla Stal 1873: 68; Breddin 1900: 
194; Blote 1936: 50; Brailovsky 1995: 
93. Type species: Cletus fasciatus Walker 
1871: 196-197. Designation by Mono- 


typy. 


Diagnosis.—Like Agathyrna Stal, in this 
genus the tylus projects in front of juga, 
upturned to form a small horn at its apex, 
the mandibular plates are directly below 
apices of juga and with small prominent tu- 
bercle, buccula short, raised and angulate 
behind anterior emargination, and abdomi- 
nal sternite VII of female entire, without 
plica or fissura. In Acanthotyla, antennal 
segment IV is shorter than III, the head dor- 
sally strongly convex, antenniferous tuber- 
cles at outer apical corner projected for- 
ward, posterior lobe of pronotal disc with a 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


transverse wrinkle, calli conspicuously con- 
vex, femora unarmed or with double row of 
tiny tubercles, and male genital capsule 
usually with small to large median projec- 
tion (Figs. 11-25). In Agathyrna, antennal 
segments III and IV are subequal in length, 
the head dorsally weakly convex, the anten- 
niferous tubercles at outer apex obtuse, not 
armed, posterior lobe of pronotal disc with- 
out transverse wrinkle, calli almost flat, 
femora ventrally armed, and male genital 
capsule obtusely rounded at apex, and never 
projecting into spine. 

Generic redescription.—Head: Width 
across eyes greater than head length; qua- 
drangular, dorsally convex to moderately 
globose; tylus extending anteriorly to juga, 
apically upturned to form a sharp median 
horn; juga unarmed, thickened, shorter than 
tylus; antenniferous tubercles armed with a 
short and robust lobe; sides of head in front 
of eyes unarmed, straight; antennal segment 
I robust, thickest, slightly curved outward; 
segments II and III cylindrical, slender, and 
segment IV fusiform; antennal segment II 
longest, IV shortest, and III longer than I; 
ocelli weakly elevated, with a deep circular 
pit in front of each; eyes substylate, protu- 
berant; postocular tubercle moderately pro- 
tuberant; buccula short, elevated, angulate 
behind anterior emargination, not projected 
beyond antenniferous tubercles, with sharp 
posterior projection; rostrum reaching an- 
terior third of abdominal sternite III or IV; 
mandibular plates directly below apices of 
juga, each with prominent tubercle. 

Thorax: Pronotum wider than long, tra- 
peziform, moderately to strongly declivent, 
and bilobed; anterior lobe shorter than pos- 
terior lobe; collar wide; frontal angles pro- 
duced forward as small conical teeth some- 
times difficult to see; anterior half of an- 
terolateral margins convexly rounded, and 
posterior half obliquely straight; humeral 
angles rounded to obtuse, moderately to 
conspicuously directed upward, and hardly 
prominent; posterolateral and posterior 
margins straight; calli conspicuously con- 
vex to globose, separated along midline by 


VOLUME 107, NUMBER 2 281 


eS 
eS Z 
= = 
7 rs 
e e 
Se cep 
£ Je W 8 
& & 
mS 
5 6 S = 


Figs. 1-10. Acanthotyla spp. 1—4, Pronotum. 1, A. fasciata. 2, A. flexuosa. 3, A. kaloboana. 4, A. kiungala. 
5—10, Parameres. 5—6, A. protenta. 7-8, A. distinguenda. 9-10, A. fasciata. 


a longitudinal furrow, entirely flat, or sep- 
arated by a convex longitudinal expansion; 
posterior margin with transverse wrinkle 
(Figs. 1—4); anterior lobe of metathoracic 
peritreme reniform, posterior lobe sharp, 
small; mesosternum with longitudinal fur- 
row; propleura laterally convex to hemi- 
spheric, in dorsal view visible. Legs un- 
armed; femora usually tuberculate; tibiae 
cylindrical, sulcated. 

Scutellum: Triangular, longer than wide, 
apically rounded to subacute; disc basally 
markedly globose, and distally before the 
apex remarkably depressed. 

Hemelytron: Macropterous, almost 
reaching apex of last abdominal segment; 
apical margin sinuate to obliquely straight; 
apical angle reaching almost middle third of 
hemelytral membrane; costal margin emar- 
ginate. 

Abdomen: Connexivum higher than ter- 
ga; connexival margin entire, and posterior 
angle blunt, and not extended into spine; 
abdominal sterna with median longitudinal 
furrow running to posterior third of sternite 
IV. 

Male genital capsule: Usually with 
small to large median projection (Figs. 1 1— 
15). Parameres simple, elongate, with apical 
third slightly curved (Figs. 5—10). 

Female genitalia: Abdominal sternite 
VII complete, without plica or fissura (Figs. 
26-35). 

Integument: Body surface rather dull, 
with short, decumbent to suberect indistinct 
pubescence; abdominal sterna with few 
moderately long, and conspicuous bristle 
like hairs; pubescence of antennae and fem- 
ora short, mainly suberect; pubescence of 
tibiae and tarsi longer and rather dense; dor- 
sal and ventral surface of head, pronotum, 
scutellum, clavus, corium, propleura, me- 
sopleura, metapleura, abdominal sterna, and 
exposed parts of genital plates of both sexes 
strongly punctate, and each puncture deeply 
excavated; prosternum, mesosternum, and 
metasternum smooth; antenna and legs mi- 
nutely granulate to tuberculate. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


FEATURES IN COMMON FOR SPECIES 
DESCRIBED 


Coloration: 1, Antennal segment I yel- 
low speckled with pale brown or pale or- 
ange irregular spots. 2, Hemelytral mem- 
brane pale to dark brown with yellow veins. 
3, Connexivum reddish brown with anterior 
third and posterior margin yellow. 4, Pro- 
sternum, mesosternum, and metasternum 
black to reddish brown. 5, Anterior and 
posterior lobe of metathoracic peritreme 
pale yellow. 


KEY TO SPECIES OF ACANTHOTYLA 


1. Head in dorsal view almost entirely black to 


ineelhIa IRON ooooasanbeso5ooes000cs 2 
— Head in dorsal view yellow with punctures 

reddish brown to chestnut orange ........ 8 
Die AN alle rete mais. sieve nee ota oho tee tice ole 3) 

Femalevices sss che ee see eee 6 


3. Posteroventral edge of male genital capsule at 
middle third with a stout or slender and elon- 
gate projection obliquely directed upward 
Gigs, ISSO, 1ID=20) os cccccoocz conc 4 

—  Posteroventral edge of male genital capsule 
without stout or elongate projection (Figs. 
13-14, 21-22, 25) 

4. Posteroventral edge of genital capsule at mid- 
dle third with a stout spinous conical projec- 
tion; lateral angles of genital capsule with 
short blunt processes (Figs. 15—16); humeral 
angles broad, rounded, and not exposed 

ahi ines men ie erate ts Gra Li A. distinguenda Blote 

—  Posteroventral edge of genital capsule at mid- 
dle third with a relatively slender, elongate 
conical projection, apically acute; lateral an- 
gles of genital capsule elevated, exposed, and 
apically quadrate (Figs. 19—20); humeral an- 
gles weakly exposed, and narrowed 

SEE a EL pee STR eles eometee = A. protenta, n.sp. 

5. Posteroventral edge of genital capsule at mid- 
dle third with tiny projection directed upward, 
and sometimes hard to see (Figs. 13—14, 25); 
intercallar space broad and flat; humeral an- 
gles rounded (Fig. 1)..... A. fasciata (Walker) 

—  Posteroventral edge of genital capsule at mid- 
dle third broad, thick, without stout spinous 
projection directed upward (Figs. 21—22); in- 
tercallar space narrow, with a weakly longi- 
tudinal carina; humeral angles laminate... . 

SSI Ny ooee iecresaenen Hie Sea Skee oe A. nabirenia, n.sp. 

6. Gonocoxae I with mesial margin broadly con- 
tiguous, and not emarginate; paratergite IX at 
lower third exposed, and distally bifurcate 


VOLUME 107, NUMBER 2 283 


Figs. 11-25. Male genital capsule of Acanthotyla spp. 11, 13, 15, 17, 19, 21, 23, 25, Lateral view. 12, 14, 
16, 18, 20, 22, 24, Caudal view. 11-12, A. borneana. 13-14, 25, A. fasciata. 15-16, A. distinguenda. 17-18, A. 
kiungala. 19-20, A. protenta. 21-22, A. nabirenia. 23-24, A. kaloboana. 


284 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


aad cae ores eek A. distinguenda Blote, 

A. fasciata (Walker), and A. protenta, n.sp. 
— Gonocoxae I with the mesial margin broadly 
contiguous, and conspicuously emarginate; 
paratergite IX with lower third not exposed 

and distally truncated (Figs. 26. 31) 
7. Mesial margin of gonocoxae I with the space 
between the border and the margin strongly 
depressed throughout the entire plate (Figs. 

28, 32) A. kaloboana, n.sp. 
—  Mesial margin of gonocoxae I with the space 
between the border and the margin shallowly 


Gepresseciigencneie ne ae ene A. nabirenia, 0.sp. 
SiyIMial ay tree gen cose reece lane Menus se et eh eu. cate = 9 
= ghemaler tases sola toccitete sh emontae o oae 12 


9. Posteroventral edge of genital capsule at mid- 
dle third broad, thick, without a stout spinous 
projection directed upward (Figs. 23-24)... 
Mura Caserta se rate erica A. kaloboana, n.sp. 
—  Posteroventral edge of genital capsule at mid- 
dle third with stout and broad or elongate pro- 
jection (Figs. 11-12, 17-18) 
10. Posteroventral edge of genital capsule at mid- 
dle third with a relatively elongate spinous 
conical projection, directed posteriorly and 
upward (Figs. 17-18) 
SP SHR eee Suen iF rea A. kiungala, n.s.p. 
—  Posteroventral edge of genital capsule at mid- 
dle third with broad projection (Figs. 11—12) 
a aeCR EN aate uo. Stoo a MEA AOR Maecenas a) Smee 11 
11. Posteroventral edge of male genital capsule at 
middle third with stout quadrate projection, 
apically bifid, and clearly directed upward . . 
Da ei psi ac tna Shel esd A. flexuosa (Blote) 
—  Posteroventral edge of genital capsule at mid- 
dle third with stout, rounded tubercle, directed 
posteriorly and upward (Figs. 11-12) 
ON nu a REAROR Po rcacu on eo eater A. borneana, n.sp. 
12. Mesial margin of gonocoxae I with the space 
between the border and the margin strongly 
depressed throughout the entire plate (Figs. 
PX ESS) alot ae eee ve es A. kaloboana, n.sp. 
— Lower and mesial margin of gonocoxae I with 
the space between the border and the margin 
shallowly depressed (Figs. 27—28, 32 34) 
A. borneana, n.sp., A. flexuosa (Blote), 
and A. kiungala, n.sp. 


Acanthotyla borneana Brailovsky, 
new species 
(Figs. 11-12, 27, 34) 


Description.—Measurements: Male: 
Head length 1.50; width across eyes 1.88; 
interocular space 1.12; preocular distance 
1.22; interocellar space 0.43; length of an- 
tennal segments: I, 2.08; II, 2.96; III, 2.08; 


IV, 1.28. Pronotum: Length 2.16; maximum 
width of anterior lobe 1.84; maximum 
width of posterior lobe 3.20. Scutellar 
length 1.44; width 1.32. Body length 9.62. 
Female: Head length 1.60; width across 
eyes 1.98; interocular space 1.18; preocular 
distance 1.26; interocellar space 0.44; 
length of antennal segments: I, 2.14; II, 
3.12; Ill, 2.16; IV, 1.28. Pronotum: Length 
2.40: maximum width of anterior lobe 2.00; 
maximum width of posterior lobe 3.38. 
Scutellar length 1.60; width 1.52. Body 
length 10.48. 

Male: Dorsal coloration: Head yellow 
with punctures chestnut orange; antennal 
segments II and HI yellow, and IV dark 
chestnut orange with apex paler; anterior 
lobe of pronotal disk dark yellow, with 
transversal fascia behind calli pale yellow, 
and punctures reddish; intercallar space 
dark brown with orange reflections; poste- 
rior lobe of pronotal disk pale orange brown 
with punctures reddish; scutellum yellow 
with punctures chestnut orange and basal 
angle black; clavus and corium pale brown, 
with punctures reddish brown to chestnut 
orange, and claval and corial veins, and 
costal margin yellow; apical margin pale 
brown with yellow marks; dorsal abdominal 
segments dark orange with black marks. 
Ventral coloration: Head yellow with 
punctures chestnut orange, and middle third 
with broad longitudinal black stripe; rostral 
segment I yellow with chestnut orange re- 
flections, and segments IT to IV chestnut or- 
ange; propleura, mesopleura, and metapleu- 
ra yellow with punctures chestnut orange; 
coxae dark reddish brown; trochanters yel- 
low; fore and middle femora yellow, speck- 
led with pale brown discoidal spots, and 
with subdistal pale brown ring; hind femur 
yellow, speckled with pale brown discoidal 
spots, and with two pale brown rings, one 
near middle third and the other subdistal; 
tibiae pale brown with two yellow rings, 
one subbasal, the other near middle third; 
tarsi pale chestnut yellow; middle third of 
abdominal sterna dark reddish brown, and 
laterally yellow with punctures reddish 


VOLUME 107, NUMBER 2 285 


Figs. 26-35. Female genital plates of Acanthotyla spp. 26-30, Caudal view. 31-35, Lateral view. 26, 31, A. 
kaloboana. 27, 34, A. borneana. 28, 32, A. flexuosa. 29, 33, A. distinguenda. 30, 35, A. protenta. 


286 


brown to chestnut orange; genital capsule 
reddish brown. 

Head: Rostrum reaching anterior third of 
abdominal sternite IV. 

Thorax: Humeral angles subacute, ele- 
vated, slightly laminate, and higher than 
posterior pronotal disk; intercallar space 
narrow with prominent carina. 

Genital capsule: Posteroventral edge 
transversely concave; lateral angles straight 
not expanded; middle third with stout and 
rounded tubercle, directed posteriorly and 
clearly upward (Figs. 11—12). 

Female: Color similar to male. Connex- 
ival segments VIII and IX dark chestnut or- 
ange with reddish brown reflections; dorsal 
abdominal segments VIII and IX dark 
chestnut orange; genital plates yellow with 
punctures chestnut orange to reddish 
brown, and with middle third of gonocoxae 
I and outer margin of paratergite [X reddish 
brown. 

Genital plates: Gonocoxae I enlarged 
dorsoventrally; mesial margin broadly con- 
tiguous and weakly emarginated; parallel to 
each margin at lower and middle third shal- 
lowly depressed, with upper third overlap- 
ping; paratergite VIII small, triangular, with 
spiracle visible; paratergite [X not overlap- 
ping at middle third, and larger than para- 
tergite VIII (Figs. 27, 34). 

Type material—Holotype: d¢, Borneo 
(British N.), Sandakan Bay (SW), Sapagaya 
Lumber Camp, 2—20 m, 4 November 1957, 
J. L. Gressitt (BPBM). Paratype: 1 2, same 
data as the male holotype (BPBM). 

Distribution.—Only known from the 
type material. 

Discussion.—This species is diagnosed 
mostly on the basis of the shape of the pos- 
teroventral edge of male genital capsule 
(Figs. 11—12), the intercallar space narrow 
with prominent carina, the humeral angles 
subacute and slightly laminate, and the head 
mostly yellow with punctures chestnut or- 
ange. In Acanthotyla distinguenda and A. 
fasciata the intercallar space is broad and 
flat, the humeral angles are rounded and not 
laminate, the head is mostly black, and the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


shape of the male genital capsule is distinct 
(Figs. 13-16, 25). 

Etymology.—Named for its occurrence 
in Borneo. 


Acanthotyla distinguenda Blote 
(Figs. 7-8, 15-16, 29, 33, 36) 


Acanthotyla distinguenda Blote 1936: 50. 


Redescription.—Measurements: Male: 
Head length 1.40—144; width across eyes 
1.97-2.01; interocular space 1.24—1.34; 
preocular distance 1.08—1.12; interocellar 
space 0.38—0.44; length of antennal seg- 
ments: I, 1.62-1.64; I, 2.40—2.52; I, 
1.76-1.97; IV, 1.20—1.32. Pronotum: 
Length 1.76—1.84; maximum width of an- 
terior lobe 1.40—1.76; maximum width of 
posterior lobe 2.64—2.76. Scutellar length 
1.42-1.44; width 1.16—1.24. Body length 
8.45—9.30. Female: Head length 1.52—1.56; 
width across eyes 2.12—2.14; interocular 
space 1.33—1.36; preocular distance 1.16— 
120; interocellar space 0.40—0.46; length of 
antennal segments: I, 1.60—1.68; II, 2.48— 
2.50; Ill, 1.79—1.84; IV, 1.1.30—1.32. Pron- 
otum: Length 2.08—212; maximum width of 
anterior lobe 1.44—1.80; maximum width of 
posterior lobe 2.82—2.84. Scutellar length 
1.52-1.54; width 1.30—1.32. Body length 
10.15—10.38. 

Male: Dorsal coloration: Black with ty- 
lus and postocular tubercle yellow; antennal 
segments II and III pale yellow, and IV pale 
yellow with basal joint pale brown; jugum 
yellow with punctures reddish brown; pron- 
otum dark to pale yellow with punctures 
chestnut orange and intercallar space black; 
scutellum pale yellow with punctures chest- 
nut orange, and basal angle black; clavus 
and corium pale yellow brown with punc- 
tures reddish brown; claval and corial veins, 
and costal and apical margin pale yellow; 
dorsal abdominal segments dark orange. 
Ventral coloration: Head black with two 
short longitudinal yellow stripes lateral to 
middle line and close to eyes; buccula yel- 
low with punctures chestnut orange; rostral 
segments pale chestnut orange; propleura, 


VOLUME 107, NUMBER 2 287 


e530 


Po 


sen 
} 
[Ea 


Fig. 36. Dorsal view of Acanthotyla distinguenda, male. 


288 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mesopleura, and metapleura yellow with 
punctures chestnut orange; coxae dark red- 
dish brown with apex yellow; trochanters 
yellow; fore and middle femora yellow, 
speckled with pale brown discoidal spots, 
and with subdistal pale brown ring; hind fe- 
mur yellow, speckled with pale brown dis- 
coidal spots, and with two pale brown rings, 
one near middle third and other subdistal; 
tibiae pale brown with two yellow rings, 
one subbasal, other near middle third; tarsi 
pale chestnut yellow; middle third of ab- 
dominal sterna dark reddish brown, and lat- 
erally yellow with punctures reddish brown 
to chestnut orange; genital capsule reddish 
brown. 

Head: Rostrum reaching posterior mar- 
gin of abdominal sternite II. 

Thorax: Humeral angles rounded, slight- 
ly prominent, elevated, and higher than pos- 
terior pronotal disk; intercallar space broad 
and flat. 

Genital capsule: Posteroventral edge 
transversely straight or slightly concave; 
lateral angles with short blunt processes; 
middle third with stout spinous conical pro- 
jection, obliquely directed upward (Figs. 
15—16). Parameres in Figs. 7-8. 

Female: Color similar to male. Connex- 
ival segments VIII and IX yellow with pos- 
terior third reddish brown; dorsal abdomi- 
nal segments VIII and IX dark reddish 
brown; genital plates yellow with punc- 
tures, posterior third of paratergite VIII and 
IX, and middle third of gonocoxae I dark 
reddish brown. 

Genital plates: Gonocoxae I reniform, 
enlarged dorsoventrally, with deep depres- 
sion close to middle third; mesial margin 
broadly contiguous and not emarginate; 
paratergite VIII and IX totally exposed; 
paratergite VIII small, triangular, with spi- 
racle visible; paratergite [X not overlapping 
at middle third, larger than paratergite VIII, 
strongly carinated, exposed, and reflexed, 
with lower third bifurcate (Figs. 29, 33). 

Variation.—1, Antennal segment I pale 
chestnut orange, speckled with pale brown 
irregular spots. 2, Antennal segments II and 


III pale chestnut orange. 3, Antennal seg- 
ment IV chestnut orange with basal joint 
dark brown. 4, Hemelytral membrane dark 
brown to black with veins pale yellow. 5, 
Outer margin of calli black. 6, Trochanter 
yellow with tiny pale brown stripe. 

Distribution.—This species is known 
only from New Guinea. The only previous- 
ly known record came from the original de- 
scription in which the holotype, allotype 
and paratypes were collected: Indonesia: 
Irian Jaya (Dutch New Guinea): Manok- 
wari, Andai, Hattam, Assiki on Digul, Kop- 
stein, and Sekroe (Blote 1936). The new re- 
cords listed below show it occurs exten- 
sively in New Guinea. 

Material examined.—l1 6, INDONESIA: 
Irian Jaya (Dutch New Guinea), Manok- 
wari, J. W. van Nouhuys (holotype RNHL). 
PAPUA NEW GUINEA: 2 4, Bisianumu, 
E. of Port Moresby, 500 m, 23 September 
hOSSaeeleGressites(B EBM); siaGemlamce 
Middle Fly River, 250-300 m, July 1928, 
Pemberton (BPBM); 1 6, Morobe, D. Bul- 
log Rd., S. Slopes Yaningya, 600—1300 m, 
19-20 March 1968, Reni (BPBM); Sogeri, 
27 October 1968, Tawi and Mena (BPBM). 
Indonesia: Irian Jaya: | 36, 1 2, Jayawijaya 
Prov., Brazza River, Dekai, ca. 100 m, 21— 
22 June 1994, A. Riedel (ZSMC); 1 2, Jay- 
awijaya Proyv., Samboka, upper Koiff River, 
ca. 200 m, 10—14 October 1996, A. Riedel 
(ZSMC); 1 6, Jayawijaya Prov., Kec Wai- 
geo Sel., Gamang Isl., 0-100 m, 26 No- 
vember 1996, A. Riedel (ZSMC); Merauke 
Prov., Asmat-Patipi, Brazza River, 100 m, 
A. Riedel (USNM); i ¢, 1 2, Manokwari 
Prov., Wasior, 0-500 m, 8-12 January 
2001, A. Riedel (UNAM); 5 @, Manok- 
wari, Ransiki, Mayuby, 300 m, 26—30 Sep- 
tember 1990, A. Riedel (ZSMC); 1 3, Na- 
bire, Pusppensaat km. 60, 200 m, 15 Au- 
gust 1991, A. Riedel (ZSMC). 

Discussion.—Acanthotyla distinguenda 
is easily distinguished by the shape of the 
posteroventral edge of the male genital cap- 
sule which shows a stout conical obliquely 
erected projection, with short blunt pro- 
cesses at lateral angles (Figs. 15-16). In A. 


VOLUME 107, NUMBER 2 


289 


4) 


Fig. 37. Dorsal view of Acanthotyla kaloboana, male. 


fasciata, the most similar species, the pos- 
teroventral edge has a tiny projection, 
sometime difficult to see (Figs. 13—14, 25). 

The females of both species including the 
shape of the genital plates, are remarkably 
similar (Figs. 29, 33). 


Acanthotyla fasciata (Walker) 
(Figs. 1, 9-10, 13-14, 25) 


Cletus fasciatus Walker 1871: 196—197; 
Distant 1901: 20; Acanthotyla fasciata: 
Stal 1873: 68; Brailovsky 1993: 37. 


290 


Acanthotyla aruana Horvath 1919: 310. 
New synonym. 


Redescription.—Measurements: Male: 
Head length 1.32—1.48; width across eyes 
1.74—1.88; interocular space 1.08—1.16; 
preocular distance 1.00—1.04; interocellar 
space 0.36—0.42; length of antennal seg- 
ments.) 1) S6—1-962 sik, 1762166; 10 
PAV 1542 IV 0012038 Ss eronotum: 
Length 1.80—1.92; maximum width of an- 
terior lobe 1.64—1.74; maximum width of 
posterior lobe 2.54—3.24. Scutellar length 
1.24—1.56; width 1.04—1.44. Body length 
8.20—9.72. Female: Head length 1.36—1.50; 
width across eyes 1.84—1.92; interocular 
space 1.10—1.12; preocular distance 1.07— 
1.09; interocellar space 0.36—0.42; length of 
antennal segments: I, 1.30—2.04; II, 1.96— 
2.64; Ill, 1.51-1.54; IV, 1.04—1.08. Prono- 
tum: Length 1.88—1.96; maximum width of 
anterior lobe 1.70—1.76; maximum width of 
posterior lobe 2.80—3.22. Scutellar length 
1.36—-1.56; width 1.24—1.40. Body length 
9.20—10.30. 

Male: Dorsal coloration: Head pale yel- 
low with punctures dark reddish brown, or 
black with tylus and postocular tubercle 
yellow; antennal segments II and III pale 
chestnut yellow, and IV chestnut orange 
with basal joint pale brown; pronotum yel- 
low with punctures chestnut orange, and the 
intercallar space black; scutellum, clavus 
and corium yellow with punctures chestnut 
orange; claval and corial veins pale yellow; 
dorsal abdominal segments dark orange. 
Ventral coloration: Head black with two 
short longitudinal yellow stripes lateral to 
middle line and close to eyes; buccula yel- 
low with punctures pale chestnut orange; 
rostral segments I to IV chestnut yellow 
(apex of IV darker); propleura, mesopleura, 
and metapleura yellow with punctures 
chestnut orange; coxae dark reddish brown; 
trochanter yellow; femora yellow, speckled 
with pale brown discoidal spots; tibiae pale 
chestnut brown with two yellow rings, one 
subbasal, other near middle third; tarsi pale 
chestnut yellow; middle third of abdominal 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


sterna dark orange brown, and laterally yel- 
low with punctures chestnut orange; genital 
capsule dark orange brown. 

Head: Rostrum reaching posterior third 
of abdominal sternite III or anterior border 
of IV. 

Thorax: Humeral angles rounded, hardly 
prominent, and higher than posterior pro- 
notal disk; intercallar space broad and flat 
(Bigs 1): 

Genital capsule: Posteroventral edge 
transversely straight or slightly concave; 
lateral angles with relatively short, quadrate 
processes; middle third with small wide 
projection, apically rounded, directed up- 
ward, and sometimes hard to see (Figs. 13— 
14, 25). Parameres in Figs. 9—10. 

Female: Color similar to male. Connex- 
ival segments VIII and IX dark brown with 
anterior half yellow; dorsal abdominal seg- 
ments VIII and IX dark orange brown; gen- 
ital plates yellow with punctures and pos- 
terior third of paratergite VIII and IX chest- 
nut orange. 

Genital plates: Gonocoxae | reniform, 
enlarged dorsoventrally, with deep depres- 
sion close to middle third; mesial margin 
broadly contiguous and not emarginate; 
paratergite VIII and IX totally exposed; 
paratergite VIII small triangular with spi- 
racle visible; paratergite IX squarish, not 
overlapping at middle third, larger than par- 
atergite VIII, strongly carinate, exposed, 
and reflexed, with lower third bifurcate. 
General shape similar to A. distinguenda 
(Figs: 295 33): 

Variation.—Subdistal third of femora 
with or lacking a pale brown ring. 

Distribution.—This species was original- 
ly described from Mysol (Walker 1871), 
and later recorded from New Guinea (Stal 
1873) and Australia (Brailovsky 1993: 
North Queensland: Cape York Peninsula: 
Gordon Ck., West Claudie R., and Rocky 
River). Horvath (1919) recorded it as Acan- 
thotyla aruana from Aru Island. The new 
records listed below indicate it occurs ex- 
tensively in New Guinea. 

Material examined.—Syntypes: 


Ghaes 


VOLUME 107, NUMBER 2 


Mysol (BMNH). Holotype: d, Aru Island: 
Terangan, zwischen Erersin und Ngarangar- 
in (SMED) (Acanthotyla aruana Horvath). 
PAPUA NEW GUINEA: 6 6, 5 2, SE, 
Western District, Oriomo R, 3 m, 5—6 Au- 
gust 1964, H. Clissold (BMNH); 2 6, 1 @, 
Western District, Oriomo, Gout. Sta., 26— 
28 October 1960, J. L. Gressitt (BPBM); 5 
6, 4 2, SE, Balimo, 9 m, 7 March 1964, 
and 6-7 August 1964, H. Clissold 
(BMNH); 8 36, 5 2, SE, Western District, 
Ruka, 9 m, 12 August 1964, H. Clissold 
(BMNH); 1 6, SE, Daru Island, 3 m, 19 
July 1964, H. Clissold (BMNH); 1 d, SE, 
Brown River and Vanapa River, 16 Decem- 
ber 1964, L. and M. Gressitt (BPBM); 2 d, 
SE, Port Moresby to Brown River, 30 m, 
29 October to | November 1965, J. Sedla- 
cek (BPBM); 7 3, 3 2, Moorhead, 18 m, 
30 June 1964, and 6—14 July 1964, H. Clis- 
sold (BMNH); 4 ¢, 2 2, SE, Western Dis- 
trict, Tala, 13 July 1964, H. Clissold 
(BMNH); 1 6, 1 2, SE, Mamai, E of Port 
Glasgow, 150 m, 9-16 Febrery 1965, R. 
Straatman (BPBM); | 6,3 2, Central Dis- 
trict, Brown River, 2 March 1966 G. Mon- 
teith (QMBA); 6 6, 2 &, Port Moresby, 30 
March 1965, Balogh and Szent-Ivany 
(HNHM); 1 6, 1 2, Laloki, April 1909, E 
Muir (CAS). 

Discussion.—Like Acanthotyla distin- 
guenda with head dorsally almost black, 
humeral angles of pronotum rounded, and 
slightly prominent, gonocoxae I reniform, 
and paratergite IX with lower third exposed 
and bifurcate (Figs. 29, 33). 

In A. fasciata the posteroventral edge of 
male genital capsule at middle third has a 
tiny projection (sometimes difficult to see) 
directed upward (Figs. 13—14, 25), whereas 
in A. distinguenda the posteroventral edge 
of the genital capsule has a stout spinous 
conical projection directed obliquely up- 
ward (Figs. 15—16). 

Acanthotyla flexuosa (Bote) 
(Figs. 2, 28, 32, 41) 


Brachylybas flexuosus Bléte 1936: 31. 
Acanthotyla flexuosa: Brailovsky and Mar- 
tinez 1994: 73. 


291 


Redescription.—Measurements: Male: 
Head length 1.40; width across eyes 1.70; 
interocular space 0.98; preocular distance 
1.04; interocellar space 0.39; length of an- 
tennal segments: I, 1.52; I, 2.16; III, 1.60; 
IV, 1.08. Pronotum: Length 1.88; maximum 
width of anterior lobe 1.80; maximum 
width of posterior lobe 3.08. Scutellar 
length 1.30; width 1.24. Body length 8.26. 
Female: Head length 1.36; width across 
eyes 1.76; interocular space 1.12; preocular 
distance 1.12; interocellar space 0.42; 
length of antennal segments: I, 1.60; II, 
2.24; Ill, 1.67; IV, 1.12. Pronotum: Length 
1.96; maximum width of anterior lobe 1.24; 
maximum width of posterior lobe 3.28. 
Scutellar length 1.40; width 1.32. Body 
length 8.50. 

Male: Dorsal coloration: Head yellow to 
dark orange with punctures, frons and ver- 
tex reddish brown; antennal segments II 
and III yellow and IV yellow with basal 
joint pale brown; anterior lobe of pronotal 
disk, and transversal fascia below calli dark 
yellow with the punctures reddish brown to 
dark chestnut orange; posterior lobe of pro- 
notal disk dark chestnut orange scattered 
with dark yellow marks; intercallar space 
black; scutellum yellow with punctures red- 
dish brown, and basal angle reddish brown 
to black; clavus and corium pale brown 
with claval and corial veins, and costal mar- 
gin yellow; apical margin yellow with inner 
third pale brown; dorsal abdominal seg- 
ments dark orange. Ventral coloration: 
Head yellow to orange, with punctures red- 
dish brown, and wide black longitudinal 
stripe at middle third; rostral segment I yel- 
low, and II to IV dark brown; buccula yel- 
low with punctures chestnut orange; pro- 
pleura, mesopleura, and metapleura yellow 
with punctures chestnut orange; coxae red- 
dish brown; trochanters yellow; fore and 
middle femora yellow, speckled with pale 
brown discoidal spots, and with subdistal 
pale brown ring; hind femur yellow, speck- 
led with pale brown discoidal spots, and 
with two pale brown rings, one near middle 
third, and other subdistal; tibiae pale brown 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 38. 


Dorsal view of Acanthotyla nabirenia, male. 


VOLUME 107, NUMBER 2 


with two yellow rings, one subbasal, other 
near middle third; tarsi pale chestnut yel- 
low; middle third of abdominal sterna dark 
reddish brown, and laterally yellow with 
punctures reddish brown to dark chestnut 
orange; genital capsule yellow with punc- 
tures reddish brown to chestnut orange. 

Head: Rostrum reaching posterior bor- 
der of abdominal sternite II or anterior bor- 
der of IV. 

Thorax: WHumeral angles subacute, ele- 
vated, slightly laminate, and higher than 
posterior pronotal disk; intercallar space 
narrow, flat or tiny convex (Fig. 2). 

Genital capsule: Posteroventral edge 
transversely concave; lateral angles straight, 
and not expanded; middle third with stout 
and quadrate projection, apically slightly 
bifid, and clearly directed upward. 

Female: Color similar to male. Connex- 
ival segments VIII and IX reddish brown 
with dark yellow marks at posterior third; 
dorsal abdominal segments VIII and IX 
dark orange; genital plates yellow, with 
punctures, middle third of gonocoxae I, and 
posterior margin of paratergite IX reddish 
brown to chestnut orange. 

Genital plates: Gonocoxae I enlarged 
dorsoventrally; mesial margins broadly 
contiguous, and weakly emarginate, with 
middle third shallowly depressed parallel to 
the margin, and upper third overlapping; 
paratergite VIII small, triangular, with spi- 
racle visible; paratergite [X not overlapping 
at middle third, and larger than paratergite 
VIII (Figs. 28, 32). 

Variation.—l, Antennal segment IV dark 
orange with basal joint brown. 2. Posterior 
lobe of pronotal disk yellow with punctures 
chestnut orange. 

Distribution.—This species was de- 
scribed from two females (holotype and 
paratype) collected in Dutch New Guinea 
(Indonesia: Irian Jaya) without definite lo- 
cality, and included in the genus Brachy- 
lybas with the binomious Brachylybas flex- 
uosus. Brailovsky and Martinez (1994) re- 
vised the genus Brachylybas and transferred 


293 


that species to Acanthotyla. This species is 
now recorded from Papua New Guinea. 

Material examined—INDONESIA: 1 @, 
Irian Jaya (Dutch New Guinea), April-May 
1911, K. Gjellerup (holotype RNHL). PAP- 
UA NEW GUINEA: 3 6, 4 2, Port Mores- 
by, 30 March 1965 and 2-8 April 1965, 
Balogh and Szent Ivany (HNHM, UNAM); 
2 6, Brown River, 20 km W of Port Mores- 
by, 29 April 1960, C. W. O’ Brien (BPBM); 
1 3d, Brown River, 21 May 1956, E. J. Ford 
Jr. (BPBM). 

Discussion.—Acanthotyla flexuosa 1s dis- 
tinguished by having the humeral angles 
subacute, and laminate, the intercallar space 
narrow, and the posteroventral edge of the 
male genital capsule at middle third with 
stout and quadrate projection, apically bifid. 
The female genital plates have the gono- 
coxae I enlarged dorsoventrally with the 
middle third shallowly depressed along the 
margin, and with the upper third overlap- 
ping (Figs. 28, 32). 

In A. distinguenda and A. fasciata, the 
humeral angles are rounded, hardly promi- 
nent, and not subacute or laminate, the in- 
tercallar space is broadened, and the male 
genital capsule and female genital plates are 
quite distinct (Figs. 13—16, 25, 28-29, 32— 
33). 


Acanthotyla kaloboana Brailovsky, 
new species 
(Bigs: 3, 23-247 26; 315 37) 


Description.—Measurements: Male: 
Head length 1.48; width across eyes 1.88; 
interocular space 1.16; preocular distance 
1.06; interocellar space 0.40; length of an- 
tennal segments: I, 2.08; II, 3.00; II, 1.96: 
IV, 1.36. Pronotum: Length 2.08; maximum 
width of anterior lobe 2.08; maximum 
width of posterior lobe 3.24. Scutellar 
length 1.36; width 1.36. Body length 9.65. 
Female: Head length 1.44; width across 
eyes 1.84; interocular space 1.10; preocular 


distance 1.00; interocellar space 0.38: 
length of antennal segments: I, 1.88; II, 


2.72; Ill, 1.84; IV, 1.24. Pronotum: Length 
2.04: maximum width of anterior lobe 1.94; 


294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 39. Dorsal view of Acanthotyla kiungala, male. 


VOLUME 107, NUMBER 2 


maximum width of posterior lobe 3.04. 
Scutellar length 1.44; width 1.32. Body 
length 9.56. 

Male: Dorsal coloration: Head yellow 
with punctures reddish brown; antennal 
segment I yellow with tiny orange tuber- 
cles, segments II and III chestnut orange, 
and IV reddish brown with apex chestnut 
orange; anterior lobe of pronotal disk yel- 
low, with punctures reddish brown, and 
black spot at anterior third of intercallar 
space; posterior lobe of pronotal disk yel- 
low, with punctures reddish brown, and 
broad blackish transverse fascia near to 
posterior margin; scutellum yellow with 
punctures reddish brown, and basal angle 
black; clavus and corium pale brown, with 
claval and corial veins, costal margin, and 
apical margin dark yellow; hemelytral 
membrane dark brown with black irregular 
spots, and dark yellow veins; dorsal abdom- 
inal segments dark orange. Ventral colora- 
tion: Head yellow with punctures reddish 
brown to chestnut orange, and the middle 
third with broad black longitudinal stripe; 
rostral segments pale brown with basal and 
apical joint of rostral segment II yellow; 
propleura, mesopleura, and metapleura yel- 
low with punctures reddish brown; coxae 
dark reddish brown; trochanters yellow; 
fore and middle femora yellow, speckled 
with pale brown discoidal spots, and with 
subdistal pale brown ring; hind femur yel- 
low, speckled with pale brown discoidal 
spots, and with two pale brown rings, one 
near middle third and other subdistal; tibiae 
pale brown with two yellow rings, one sub- 
basal, other near middle third; tarsi pale 
chestnut yellow; middle third of abdominal 
sterna dark reddish brown, and laterally yel- 
low with punctures reddish brown to chest- 
nut orange; pleural margin of abdominal 
sterna HII to V yellow with punctures red- 
dish brown, and VI and VII yellow with 
black to reddish brown rectangular spots 
close to posterior third; genital capsule red- 
dish brown. 

Head: Rostrum reaching posterior mar- 
gin of abdominal sternite III. 


295 


Thorax: Humeral angles rounded or sub- 
acute, directed upward, slightly laminate, 
and higher than posterior pronotal disk: in- 
tercallar space relatively narrow, with 
weakly longitudinal carina or entirely flat 
(igs 3): 

Genital capsule: Posteroventral edge 
transversely concave; lateral angles not ex- 
panded, almost straight; middle third broad, 
thickness, without stout spinous projection 
(Figs. 23-24). 

Female: Color similar to male. Connex- 
ival segment VIII reddish brown, with an- 
terior and posterior border yellow; connex- 
ival segment IX yellow with reddish brown 
spot near posterior third; dorsal abdominal 
segments VIII and IX dark reddish brown 
to dark orange; gonocoxae I black with out- 
er margin close to middle third yellow; par- 
atergite VIII and IX reddish brown to black 
with upper margin of VIII and inner margin 
of IX yellow. 

Genital plates: Gonocoxae I reniform, 
enlarged dorsoventrally; mesial margin 
broadly contiguous and clearly emarginate; 
parallel to each margin conspicuously 
raised, and the space between the border 
and the margin strongly excavate through- 
out the entire plate; paratergite VIII and IX 
totally exposed; paratergite VIII small, tri- 
angular, with spiracle visible; paratergite [X 
not overlapping at middle third, larger than 
paratergite VIII, strongly carinated, and re- 
flexed, with lower third covered and trun- 
cated (Figs. 26, 31). 

Variation.-1, Head dorsally black with ty- 
lus, jugum and postocular tubercle yellow. 
2, Antennal segment IV pale chestnut or- 
ange with basal joint brown. 3, Basal angle 
of scutellum yellow. 4, Dorsal abdominal 
segments reddish brown to black. 5, Pleural 
margin of abdominal sternite V yellow with 
black to reddish brown spot close to middle 
third. 6, Gonocoxae I yellow with punctures 
reddish brown, and inner margin black. 

Type material.—Holotype: 4, Irian Jaya: 
Jayawijaya Prov., Salawatti Isl., Kalobo, ca. 
10-30 m, 19-22 October 1966 A. Riedel 
(ZSMC). Paratypes: Irian Jaya: 1 6, 1 9, 


296 


sama data as holotype (UNAM, ZSMC); | 
2, Jayawijaya Prov., Batanta Isl., Waylebet, 
0—100 m, 28 October 2 November 1996, A. 
Riedel (ZSMC); | @, Jayawijaya Prov., Kec 
Salawatti Kalobo, Walir Isl., 0-20 m, 20 
October 1996, A. Riedel (ZSMC); New 
Guinea: 3 6, 3 2, without data, col. Wal- 
lace (BMNH, UNAM). 

Distribution.—Only known from the 
type material. 

Discussion.—Acanthotyla kalaboana 1s 
easily recognized by the shape of the pos- 
teroventral edge of male genital capsule 
(Figs. 23-24), by the peculiar development 
of the gonocoxae I (Figs. 26, 31), by the 
laminate humeral angles (Fig. 3), the inter- 
callar space relatively narrow with weakly 
carina, and by the black transverse fascia 
running close to the posterior margin of the 
posterior lobe of the pronotal disk. In A. 
flexuosa, the most similar species, the male 
genital capsule, and the shape of the gon- 
ocoxae I (Figs. 28, 32) are distinct. 

Etymology.—Named for its occurrence 
in Kalobo (Irian Jaya). 


Acanthotyla kiungala Brailovsky, 
new species 
(Figs. 4, 17-18, 39) 


Description.—Measurements: Male: 
Head length 1.48; width across eyes 1.90; 
interocular space 1.16; preocular distance 
1.14; interocellar space 0.43; length of an- 
tennal segments: I, 1.90; I, 2.72; Ill, 1.88; 
IV, 1.26. Pronotum: Length 2.20; maximum 
width of anterior lobe 1.84; maximum 
width of posterior lobe 3.04. Scutellar 
length 1.52; width 1.20. Body length 9.25. 
Female: Head length 1.52; width across 
eyes 1.96; interocular space 1.16; preocular 
distance 1.16; interocellar space 0.47; 
length of antennal segments: I, 2.00; I, 
2.92; Ill, 2.04; IV, 1.22. Pronotum: Length 


maximum width of posterior lobe 3.28. 
Scutellar length 1.56; width 1.32. Body 
length 9.80. 

Male: Dorsal coloration: Head yellow 
with punctures reddish brown; antennal 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


segments II and HI chestnut yellow, and IV 
chestnut orange with apex yellow; anterior 
lobe of pronotal disk and scutellum yellow, 
with punctures reddish brown; intercallar 
space black; posterior lobe of pronotal disk 
pale orange brown, with punctures reddish 
brown, and area behind calli yellow; clavus 
and corium pale brown, punctures reddish 
brown, and claval and corial veins, and cos- 
tal margin yellow; dorsal abdominal seg- 
ments dark orange. Ventral coloration: 
Head yellow with punctures chestnut or- 
ange, and middle third with broad longitu- 
dinal black stripe; rostral segments pale 
brown with basal and apical joint of seg- 
ments II and II yellow; propleura, meso- 
pleura, and metapleura yellow with punc- 
tures chestnut orange; coxae dark reddish 
brown; trochanters yellow; fore and middle 
femora yellow, speckled with pale brown 
discoidal spots, and with subdistal pale 
brown ring; hind femur yellow, speckled 
with pale brown discoidal spots, and with 
two pale brown rings, one near middle third 
and other subdistal; tibiae pale brown with 
two yellow rings, one subbasal, other near 
middle third; tarsi pale chestnut yellow; 
middle third of abdominal sterna dark red- 
dish brown, and laterally yellow with 
pucntures reddish brown to chestnut or- 
ange; genital capsule reddish brown. 

Head: Rostrum reaching anterior margin 
or middle third of abdominal sternite IV. 

Thorax: Humeral angles subacute, and 
laminate, elevated, directed upward, and 
higher than posterior pronotal disk; inter- 
callar space narrow with prominent carina 
(Fig. 4). 

Genital capsule: Posteroventral edge 
transversely concave; lateral angles straight 
without projections; middle third with stout 
spinous conical projection, directed poste- 
riorly, and upward (Figs. 17-18). 

Female: Color similar to male. Connex- 
ival segment VIII reddish brown with an- 
terior and posterior margin yellow, and IX 
yellow with reddish brown spot near pos- 
terior margin; dorsal abdominal segments 


VOLUME 107, NUMBER 2 297 


| Se oY 


40 


Fig. 40. Dorsal view of Acanthotyla protenta, female. 


298 


VIII and [X dark orange; genital plates yel- 
low with punctures reddish brown. 

Genital plates: Gonocoxae I enlarged 
dorsoventrally, broadly contiguous, and 
emarginate; middle and lower third shal- 
lowly depressed along each margin; para- 
tergite VIII and IX totally exposed; para- 
tergite VIII small, triangular, with spiracle 
visible; paratergite IX squarish, and larger 
than paratergite VIII. 

Type material.—Holotype: 6, Papua 
New Guinea: Kiunga, Fly River, 14—17 Au- 
gust 1957, W. W. Brandt (BPBM). Para- 
types: Papua New Guinea: 4 d, 2 2, Kiun- 
ga, Fly River, 11-13 August 1957, W. W. 
Brandt (BPBM, UNAM); | <6, Fly River, 
Olsobip, 400—600 m, August 1969, J. and 
M. Sedlacek (BPBM); | ¢, Kiungala, 28— 
30 August 1969, J. Balogh (HNHM). 

Distribution.—Only recorded from the 
type material. 

Discussion.—Like Acanthotyla_ bor- 
neana, with the head dorsally yellow with 
punctures reddish brown to chestnut or- 
ange, the intercallar space narrow with a 
prominent carina, the humeral angles sub- 
acute and laminate, and the female genital 
plates similar. In A. kiungala, recorded only 
from Papua New Guinea, the posteroventral 
edge of the male genital capsule has at mid- 
dle third a prominent stout, spinous, conical 
projection directed posteriorly (Figs. 17— 
18), and in A. borneana, known only from 
Borneo, has at middle third of the male gen- 
ital capsule a stout, rounded, almost glo- 
bosus projection, directed posteriorly and 
weakly upward (Figs. 11—12). 

Etymology.—Named for its occurrence 
in Kiunga (Papua New Guinea). 


Acanthotyla nabirenia Brailovsky, 
new species 
(Figs. 21-22, 38) 


Description.—Measurements: Male: 
Head length 1.36; width across eyes 1.76; 
interocular space 1.00; preocular distance 
0.94; interocellar space 0.42; length of an- 
tennal segments: I, 1.68; II, 2.64; III, 1.72; 
IV, 1.24. Pronotum: Length 1.84; maximum 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


width of anterior lobe 1.72; maximum 
width of posterior lobe 2.92. Scutellar 
length 1.36; width 1.28. Body length 8.40. 
Female: Head length 1.34; width across 
eyes 1.90; interocular space 1.12; preocular 
distance 1.04; interocellar space 0.42; 
length of antennal segments: I, 1.88; II, 
2.80; III, 1.88; IV, 1.28. Pronotum: Length 
2.08; maximum width of anterior lobe 2.00; 
maximum width of posterior lobe 3.40. 
Scutellar length 1.52; width 1.44. Body 
length 9.85. 

Male: Dorsal coloration: Head reddish 
brown to black, with tylus, jugum, area ad- 
jacent to eyes and postocular tubercle yel- 
low; antennal segment I yellow speckled 
with tiny pale orange spots, segments II and 
Ill yellow and IV yellow with basal joint 
brown; anterior lobe of pronotal disk yel- 
low with punctures reddish brown, and in- 
tercallar space black; posterior lobe of pro- 
notal disk including the humeral angles 
dark brown to black with posterior margin 
and a transversal fascia behind calli yellow; 
scutellum yellow with punctures reddish 
brown, and basal angle black; clavus and 
corium pale brown with punctures reddish 
brown to chestnut orange, and claval and 
corial veins, and costal margin yellow; dor- 
sal abdominal segments dark orange. Ven- 
tral coloration: Head yellow with punc- 
tures chestnut orange and middle third with 
broad longitudinal black stripe; buccula yel- 
low; rostral segments pale brown with basal 
and apical joint of rostral segment II yel- 
low; propleura, mesopleura, and metapleura 
yellow with punctures chestnut orange; 
coxae dark reddish brown with apex yel- 
low; trochanters yellow; fore and middle 
femora yellow, speckled with pale brown 
discoidal spots, and with subdistal pale 
brown ring; hind femur yellow, speckled 
with pale brown discoidal spots, and with 
two pale brown rings, one near middle third 
and other subdistal; tibiae pale brown with 
two yellow rings, one subbasal, other near 
middle third; tarsi pale chestnut yellow; 
middle third of abdominal sterna dark red- 
dish brown, and laterally yellow with punc- 


VOLUME 107, NUMBER 2 299 


Fig. 41. Dorsal view of Acanthotyla flexuosa, female. 


Thorax: Humeral angles subacute, ele- 
vated, slightly laminate, and higher than 
posterior pronotal disk; intercallar space 
narrow, with shallow carina. 


tures reddish brown to chestnut orange; 
genital capsule reddish brown. 

Head: Rostrum reaching posterior bor- 
der of abdominal sternite HI. 


300 


Genital capsule: Posteroventral edge 
transversely concave; lateral angles straight, 
not expanded; middle third broad, thick- 
ness, rounded, without stout spinous pro- 
jection (Figs. 21—22). 

Female: Color similar to male. Connex- 
ival segment VIII reddish brown with an- 
terior and posterior border yellow, and IX 
yellow; dorsal abdominal segments VIII 
and IX dark orange; gonocoxae I yellow, 
punctures reddish brown, and inner margin 
black; paratergite VIII and IX yellow with 
outer margin of VIII and inner margin of 
IX pale brown. 

Genital plates: Gonocoxae I enlarged 
dorsoventrally; mesial margin broadly con- 
tiguous, conspicuously emarginate; parallel 
to each margin shallowly depressed; para- 
tergite VIII small, triangular, with spiracle 
visible; paratergite IX not overlapping at 
middle third, larger than paratergite VIII, 
with lower third covered and truncate. 

Variation.-1, Anterior lobe of pronotal 
disk black to reddish brown with yellowish 
marks. 2, Posterior lobe of pronotal disk 
dark to pale orange brown, with yellow 
marks behind calli. 3, Femora and tibiae 
dark to pale orange brown. 

Type material.—Holotype: 6, New 
Guinea: NW, Nabire, S. Geelvink Bay, O— 
30 m, 2—9 July 1962, J. Sedlacek (BPBM). 
Paratypes: New Guinea: 3 ¢, 2 2, NW, Na- 
bire, S. Geelvink Bay 0—40 m, 2-9 July 
1962, 1-4 September 1962, J. Sedlacek 
(BPBM, UNAM); Irian Jaya (Dutch New 
Guinea): 1 36, 2 @, Fakfak, 16-20 July 
1939, R. G. Wind (CAS); 2 2, Manokwari 
Prov., Wasior, 0-500 m, 8-15 January 
2001, A. Riedel (UNAM); 3 6, 2 @, Vo- 
gelkop, Bomberi, 700—900 m, 7—8 June 
1959, J. L. Gressitt (BPBM). 

Distribution.—Only known from the 
type material. 

Discussion.—Like Acanthotyla_ bor- 
neana, the humeral angles are slightly lam- 
inate, and the intercallar space is narrow 
with a shallow to prominent carina. In A. 
nabirenia, the first gonocoxae do not over- 
lap at the upper third and parallel to each 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


margin are entirely and shallowly excavat- 
ed; the posteroventral edge of male genital 
capsule at middle third has a stout, rounded 
processes, directed upward (Figs. 21—22); 
and the head in dorsal view is almost black 
to reddish brown. In A. borneana, the head 
is yellow with chestnut orange punctures, 
the gonocoxae I at upper third overlap, and 
parallel to each margin are shallowly ex- 
cavated at inner and middle third, and the 
posteroventral edge of male genital capsule 
has a stout and rounded tubercle directed 
posteriorly and weakly upward (Figs. 11— 
1) 

Etymology.—Named for its occurrence 
in Nabire (New Guinea). 


Acanthotyla protenta Brailovsky, 
new species 
(Figs. 5—6, 19-20, 30, 35, 40) 


Description.—Measurements: Male: 
Head length 1.42; width across eyes 1.80; 
interocular space 1.14; preocular distance 
1.08; interocellar space 0.42; length of an- 
tennal segments: I, 1.60; II, 2.40; III, 1.76; 
IV, 1.20. Pronotum: Length 1.84; maximum 
width of anterior lobe 1.82; maximum 
width of posterior lobe 2.92. Scutellar 
length 1.32; width 1.22. Body length 8.58. 
Female: Head length 1.38; width across 
eyes 1.96; interocular space 1.24; preocular 
distance 1.12; interocellar space 0.41; 
length of antennal segments: I, 1.66; II, 
2.48; Il, 1.92; IV, 1.24. Pronotum: Length 
1.96; maximum width of anterior lobe 1.96; 
maximum width of posterior lobe 3.20. 
Scutellar length 1.56; width 1.40. Body 
length 9.75. 

Male: Dorsal coloration: Head black 
with tylus, and postocular tubercle yellow; 
jugum yellow with punctures reddish 
brown; antennal segments II and III yellow 
and IV dark chestnut orange with basal 
joint brown, and apex pale chestnut orange; 
pronotum dark to pale yellow with punc- 
tures chestnut orange; intercallar space 
black; scutellum yellow with punctures 
chestnut orange, and basal angle black; cla- 
vus and corium pale yellow brown with 


VOLUME 107, NUMBER 2 


punctures reddish brown to chestnut or- 
ange, and claval and corial veins, costal 
margin, and apical margin pale yellow; dor- 
sal abdominal segments dark orange. Ven- 
tral coloration: Head black with two short 
yellow longitudinal stripes lateral to middle 
line and close to eyes and buccula; rostral 
segments pale chestnut orange; propleura, 
mesopleura, and metapleura yellow with 
punctures chestnut orange; coxae dark red- 
dish brown with apex yellow; trochanters 
yellow; fore and middle femora yellow, 
speckled with pale brown discoidal spots, 
and with subdistal pale brown ring; hind fe- 
mur yellow, speckled with pale brown dis- 
coidal spots, and with two pale brown rings, 
one near middle third and other subdistal; 
tibiae pale brown with two yellow rings, 
one subbasal, other near middle third; tarsi 
pale chestnut yellow; middle third of ab- 
dominal sterna dark reddish brown, and lat- 
erally yellow with punctures reddish brown 
to chestnut orange; genital capsule reddish 
brown. 

Head: Rostrum reaching posterior mar- 
gin of abdominal sternite III or anterior 
third of IV. 

Thorax: Humeral angles rounded, slight- 
ly prominent, elevated, and higher than pos- 
terior pronotal disk; intercallar space broad 
and flat. 

Genital capsule: Posteroventral edge 
transversely straight or slightly concave; 
lateral angles elevated, exposed, and api- 
cally quadrate; middle third with an elon- 
gate, apically acute projection, curved up- 
ward (Figs. 19-20). Parameres in Figs. 5— 
6. 

Female: Color similar to male. Connex- 
ival segments VIII and IX yellow with pos- 
terior third reddish brown; dorsal abdomi- 
nal segments VIII and IX dark reddish 
brown; genital plates yellow with punc- 
tures, posterior third of paratergite VIII and 
IX, and middle third of gonocoxae I dark 
reddish brown. 

Genital plates: Gonocoxae I reniform, 
enlarged dorsoventrally, with deep depres- 
sion close to middle third; mesial margin 


301 


broadly contiguous and not emarginate; 
paratergite VIII and IX totally exposed: 
paratergite VIII small, triangular, with spi- 
racle visible; paratergite [IX not overlapping 
at middle third, larger than paratergite VIII, 
strongly carinated, exposed, and reflexed, 
with lower third bifurcate (Figs. 30, 35). 

Variation.—1, Jugum black with anterior 
third yellow. 2, Calli laterally, humeral an- 
gles and posterior margin of pronotum red- 
dish brown to black. 3, Pronotum reddish 
brown to black with collar and transversal 
fascia behind calli yellow. 4, Hemelytral 
membrane dark brown to black with dark 
yellow veins. 5, Coxae entirely reddish 
brown. 

Type material.—Holotype: d, Papua 
New Guinea: Madang Prov., 16 km WNW 
of Sapi Forest Reserve, N of Quonona 
Creek, 160 m, 5°10’S 145°26’E, stop 89- 
68C, 8 April 1989, D. H. Kavanaugh, G. E. 
Ball, and N. D. Penny (CAS). 

Paratypes: Irian Jaya (Dutch New Guin- 
ea): 3 6, 3 2, Jayapura, Sentani, Cyclops 
Mts., 300 m, 19-21 September 1990, A. 
Riedel (UNAM, ZSMC); 1 6, Biak.L., 
Mokmer, 5—10 m, 26 May 1959, J. L. Gres- 
sitt (BPBM); 1 6, 2 @, Biak I., Mangro- 
wawa, 50-100 m, 31 May 1959, 29 Octo- 
berl9s59u ea Gressitte andes Coa Maa 
(BPBM); 2 2, Jutefa Bay, Pim, 0-100 m, 
February 1936, L. E. Cheesman (BMNH); 
1 3, Moffin Bay, September 1944, E. S. 
Ross (CAS); 9 6, 8 &, Hollandia, January 
to May 1945, B. Malkin (UNAM, USNM); 
3 6,1 %, Lae, 19 August 1944, W. Harden 
(ACM) SSiCep 2c eaverray sLopreniuss: 
14-21 April 1913, Kais Augustfl Exp., S. 
G. Burgers (ZSMC); 5 6,5 @, Biak I., Ko- 
rim, Nernu, 100-800 m, 14 February 2001, 
A. Riedel (UNAM); 4 2, Biak I., Roidilu, 
2 Febrery 2001, A. Riedel (UNAM); 4 &, 
Wasian, 9 September 1939, R. G. Wind 
(CAS). Papua New Guinea: 1 2, SE: Mu- 
tua River (Ss side)) 2m; 17) December 
1964, J. Sedlacek (BPBM); 5 6, 3 2, NE: 
Bulolo River, 680—700 m, 27 March 1969, 
23 April 1969 8 May 1969, 26 September 
1969, 26 November 1969, and 6 January 


302 


1970, J. Sedlacek (BPBM); 1, Madang 
Prov., Sapi Forest Reserve, Sapi Forest at 
confluence with Gogol River, 50 m, 15 
March 1989, stop 89-26A, D. H. Kavan- 
augh, G. E. Ball and N. D. Penny (CAS); 
1 9, Madang Prov., 14.4 km W of North 
Coast Rd, on Bunapas Rd., 10 m, 24 April 
1989, stop 89-92, D. H. Kavanaugh, G. E. 
Ball and N. D. Penny (CAS); 1 6, Madang 
Prov., 9.5 km WSW of Naru River Bridge 
on North Coast Rd, 480 m, 8 March 1989, 
stop 89-18, D. H. Kavanaugh, G. E. Ball, 
and N. D. Penny (CAS); 1 6, Popondetta 
District, Sangara, 22 March 1956, E. S. 
Brown (BMNH); 2 6, SE: Popondetta, 25— 
60 m, 1—4 September 1963, and April 1966, 
J. Sedlacek and G. Lippert (BPBM); 3 <4, 
3 2 Kokoda, 1200’, September 1933, L. E. 
Cheesman (BMNH); | ¢, NE: Upper Se- 
pik, Wagu, 180 m, 5 June 1963 (BPBM). 

Distribution.—Widespread in New Guin- 
ea. 

Discussion.—Acanthotyla distinguenda 
and A. protenta are strikingly similar in ap- 
pearance, and only the shape of the poster- 
oventral border of the male genital capsule 
distinguisheds them. 

In A. protenta, the lateral angles of the 
male genital capsule are more prominent, 
and the middle projection is more slender 
and acute than for A. distinguenda (Figs. 
16-16, 19-20). 

The female of both species are similar. 

Etymology.—Named for its narrow 
body; from the Latin protentus, meaning 
elongate. 


ACKNOWLEDGMENTS 


Thanks to the following individuals and 
institutions for loans and other assistance: 
Mick Webb (BMNH), Gordon Nishida and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Al Samuelson (BPBM), Vincent Lee and 
Keve Ribardo (CAS), Tamas Vasarhelyi 
(HNHM), Brian Harris (LACM), Geoffrey 
B. Monteith (QMBA), Jan van _ Tol 
(RNHL), Wolfgang A. Naessig (SMFD), 
Thomas J. Henry (USNM), Juergen Deck- 
ert (ZSMC), and Klaus Schonitzer (ZSMC). 
Special thanks to Ernesto Barrera (UNAM), 
Elvia Esparza (UNAM), Albino Luna 
(UNAM), Jesus Contreras (UNAM), and 
Cristina Urbina (UNAM) for the illustra- 
tions. 


LITERATURE CITED 


Bléte, H. C. 1936. Catalogue of the Coreidae in the 
Rijksmuseum van Natuurlijke Historie. Part HI, 
Coreinae, Second Part. Zoologische Mededelin- 
gen 19: 23-66. 

Brailovsky, H. 1993. A revision of the tribe Colpurini 
from Australia (Hemiptera-Heteroptera-Coreidae). 
Memoirs of the Queensland Museum 34(1): 35— 
60. 


. 1995. Un nuevo género, una nueva especie y 
algunos arreglos taxondémicos dentro de la tribu 
Colpurini (Hemiptera: Heteroptera: Coreidae). Fo- 
lia Entomol6gica Mexicana 93: 31—38. 

Brailovsky, H. and J. Martinez. 1994. Revision del gé- 
nero Brachylybas (Hemiptera-Heteroptera-Corei- 
dae-Colpurini). Publicaciones Especiales (Univ- 
ersidad Nacional Aut6noma de México) 13: 1—82. 

Breddin, G. 1900. Materiae ad cognitionem subfami- 
liae Pachycephalini (Lybantini Olim). Ex Hemip- 
teris-Heteropteris, Fam. Coreidae. Revue d’ En- 
tomologie, Caen 19: 194-217. 

Distant, W. L. 1901. Rhynchotal Notes. VIII. Heter- 
optera: Fam. Coreidae. Annals and Magazine of 
Natural History (7)7: 6—22. 

Horvath, G. 1919. Hemipteren von den Aru-und Kei 
Inseln. Abhandlungen der Senckenbergischen Na- 
turforschenden Gesellschaft 35(3): 305-314. 

Stal, C. 1873. Enumeratio Hemipterorum IiJ. Enumer- 
atio Coreidarum Africae, Asiae et Australiae. 
Kongliga Svenska Vetenskaps- Akademiens Han- 
dlingar 2(2) 33-163. 

Walker, E 1871. Catalogue of the specimens of He- 
miptera Heteroptera in the Collection of the Brit- 
ish Museum. Part IV. London. 211 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 303-314 


A REVIEW OF THE NEOTROPICAL GENUS DIFUNDELLA DYAR 
(LEPIDOPTERA: PYRALIDAE: PHYCITINAE) 


H. H. NEUNZIG AND M. A. SOLIS 


(HHN) Department of Entomology, North Carolina State University, Raleigh, NC 
27695-7613, U.S.A.; (MAS) Systematic Entomology Laboratory, PSI, Agricultural Re- 
search Service, U.S. Department of Agriculture, % National Musuem of Natural History, 
Smithsonian Institution P.O. Box 37012, MRC 168, Washington, DC, 20013-7012, U.S.A. 
(e-mail: asolis @sel.barc.usda.gov) 


Abstract.—The genus Difundella is redescribed. Seven species are recognized as be- 
longing to the genus: D. cancerella n. sp., D. corynophora Dyar, D. dumiella Neunzig 
and Dow, D. parana n. sp., D. subsutella (Schaus), D. teresina n. sp., and D. unguifera 
n. sp. A key to species based on male adults is included. Difundella is compared to the 


similar genera Nefundella Neunzig, Coptarthria Ragonot, and Rampylla Dyart. 


Key Words: 


H. G. Dyar at the United States National 
Museum had access to a large series of 
mostly previously unseen Neotropical Lep- 
idoptera as a result of a Smithsonian Insti- 
tution biological survey of the Panama Ca- 
nal Zone early in the twentieth century. 
Some of the specimens he studied were py- 
ralids belonging to the large subfamily Phy- 
citinae. In his 1914 paper, Dyar recognized 
52 genera as belonging to what he consid- 
ered to be the subfamily, one of which was 
the new genus Difundella proposed by him 
to accomodate his new species corynopho- 
ra. Heinrich (1956) included Difundella in 
his revision of the American phycitines, and 
placed three more species in the genus, 
namely, subsutella Schaus and two new 
species, distractor and tolerata. The assign- 
ment of the two new species to Difundella 
was tentative. Heinrich (1956: 62) wrote 
“The second group [distractor and tolera- 
ta| probably deserves a separate generic 
designation; but the material is too scanty 
and not in good enough condition, and the 
association of females with their proper 


Brazil, Costa Rica, Nefundella, Coptarthria, Rampylla 


males too uncertain, to permit evaluation of 
generic characters for separation at this 
time.”’ Neunzig (1986), following the ex- 
amination of specimens collected since 
Heinrich’s study, proposed the new genus 
Nefundella for distractor and tolerata, 
thereby reducing the number of species in 
Difundella to two. Subsequently, in 1993, 
Neunzig and Dow, in their treatment of the 
Phycitinae of Belize, added another species, 
dumiella, to Difundella. 

As a result of additional collecting in 
Costa Rica and Brazil, four more species of 
Difundella have come to light. The discoy- 
ery of this new material makes it appropri- 
ate to review the genus at this time. 

Specimens studied came from the follow- 
ing sources: Instituto Nacional de Biodiv- 
ersidad, Santo Domingo, Costa Rica [IN- 
Bio]; North Carolina State University Insect 
Collection, Raleigh, North Carolina, USA 
[NCSU]; National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
D.C., U.S.A. [USNM]; and the collection of 
Vitor O. Becker, Universidade de Brasilia, 
Brasilia, Brasil | VOB]. 


304 


Difundella Dyar 


Difundella Dyar 1914:327. Type species: 
Difundella corynophora Dyar, 1914, 
original designation. 


Description.—Head: Antenna of male 
slightly enlarged and flattened along basal 
half of shaft; sensilla trichodea (cilia) abun- 
dant, short (about ¥% to ¥; as long as width 
of antenna near base of shaft). Antenna of 
female simple. Labial palpus of male up- 
turned, extending above vertex; segment III 
about % as long as II (unguifera with palpus 
shorter, with segment III about % as long as 
I1). Maxillary palpus of both sexes short- 
scaled. Haustellum well developed. Ocellus 
present. Prothorax: With anteroventral, 
partially black, contrasting bib near head. 
Forewing: Upper surface with basal and 
subbasal parts and inverted triangular area 
closely following antemedial line dark red- 
dish brown, purplish brown, purple, or 
black; most of rest of wing pale (pale red- 
dish brown and ochre); antemedial and 
postmedial lines weakly developed, pale 
reddish brown and ochre; narrow, dark red- 
dish brown to black lines bordering ante- 
medial and postmedial lines (lines distally 
bordering antemedial line and proximally 
bordering postmedial line most strongly de- 
veloped); area between antemedial and 
postmedial lines usually with black or dark 
brown streaks on veins M, to CuA,, and 
fused ochre to reddish-brown discal spots; 
basal third of posterior margin of wing with 
red to black posteriorly—projecting tuft of 
scales; basal half of costa of forewing 
slightly convex (base of costa strongly pro- 
duced anteriorly in male of unguifera); un- 
derside of wing of male without contrast- 
ingly colored sex-scales; wing with 11 
veins; R;,, and R; fused for about % their 
lengths beyond cell; M, straight; M, and M, 
approximate at base; CuA, from posterod- 
istal angle of cell; CuA, from well before 
posterodistal angle of cell. Hindwing: Hy- 
aline to brown; with 8 veins (1A, 2A and 
3A treated as one vein); Sc + R, and Rs 
fused for about % distance beyond cell; M, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


and M, fused for about %4 distance beyond 
cell; CuA, fused to base of stalk of M, and 
M,; CuA, from before posterodistal angle 
of cell; 1A usually abruptly bent in basal 
half; underside of male wing with small 
cluster of thin, usually hooked scales near 
discal vein, raised, elongate-oval patch of 
sex-scales near 1A, and anal fold along 3A 
enclosing a scale pencil (no elongate-oval 
patch on unguifera, and both small cluster 
of scales and elongate-oval patch absent in 
cancerella and dumiella); cell short (4 or 
less length of wing). Male abdominal seg- 
ment 8 developed as narrow pocket, with- 
out scale tuft. Male genitalia: Uncus well 
developed with posterior margin usually 
weakly pointed (uncus distally bifurcate in 
corynophora), basolateral aspects of uncus 
usually simple (clawlike in unguifera and 
cancerella); gnathos greatly reduced, with- 
out well developed median process; tran- 
stilla absent; juxta a triangular plate with 
setiferous lobes or setal patches; valva slen- 
der basally. broadened and with spinelike 
setae distally; sacculus strongly sclerotized, 
developed into long, hooked process (pro- 
cess short in subsutella) and with strong tuft 
of scalelike setae basally; aedoeagus either 
simple, serrate, spined, or forked; vesica 
simple, scobinate, or with small cornuti; vin- 
culum slightly longer than greatest width. 
Female genitalia: Posterior part compact; 
apophysis posterioris short and slender; 
apophysis anterioris stubby (extending only 
slightly beyond ostium bursae, and usually 
more robust than apophysis posterioris); os- 
tium bursae with wide, sclerotized plate and 
many microspines (intersegmental area dor- 
sad of this plate with pair of pockets); ductus 
bursae shorter than, to about as long as, cor- 
pus bursae, usually membranous (cancerella 
with microspines in posterior half); corpus 
bursae oval with signum a patch of micros- 
pines or scobinations; ductus seminalis at- 
tached to corpus bursae near junction of duc- 
tus bursae and corpus bursae. 

Comparison to similar genera.—Species 
belonging to the genera Nefundella, Cop- 
tarthria Ragonot, and Rampylla Dyar have 


VOLUME 107, NUMBER 2 


305 


Figs. 1-4. Males, dorsal view. 1, Difundella corynophora. 2, D. dumiella. 3, D. unguifera. 4, D. cancerella. 


forewings whose color and maculation re- 
semble those of Difundella, but male geni- 
talic features separate the groups. Species 
of Nefundella differ most noticeably from 
those of Difundella in having a costal pro- 
cess on the valva (absent in Difundella). 
The male genitalia of Coptarthria and Ram- 
pylla have an unusual juxta bearing a long 
weakly attached spine, a feature absent in 
Difundella and most other phycitines. 

The immature stages of Difundella and 
their hosts are unknown. 


KEY TO SPECIES OF MALE DIFUNDELLA 


1. Uncus with apex bifurcate (Fig. 12); aedoeagus 
forked (Fig. 13) corynophora Dyar 
— Uncus with apex entire (Figs. 14, 16, 18, 20, 
22, 24); aedoeagus not forked (Figs. 15, 17, 19, 
DPI ADTE Ne DIS) stacy: oO Oe ENG Ee Ch GEERT OR OMe 2 
2. Base of uncus with medially projecting claw- 
like processes (Figs. 20, 22) 
— Base of uncus without medially projecting 
clawlike processes (Figs. 14, 16, 18, 24).... 4 


3. Each clawlike process of uncus biramous (Fig. 
2X0) Nai tts Sehr one rane cancerella Neunzig and Solis 
— Each clawlike process of uncus simple (Fig. 
DD Neth, Wee skate fa eRe unguifera Neunzig and Solis 
4. Aedoeagus with large, sclerotized process 
composed of overlapping, contiguous curved 
spines (Fig. 15) subsutella (Schaus) 
— Aedoeagus without large, sclerotized process 
composed of overlapping, contiguous spines 


5. Hooklike process of sacculus broadened dis- 
(zily (Vets, NS) ooo oo parana Neunzig and Solis 
— Hooklike process of sacculus slender distally 
(Gigs! 6324) etree einen scious enemies 6 
6. Juxta with large posteriorly projecting lobes 
(Fig. 24); process of sacculus strongly bent in 
distal half (Fig. 24) .. teresina Neunzig and Solis 
— Juxta without large posteriorly projecting lobes 
(Fig. 16); process of sacculus weakly bent in 
distal half (Fig. 16) . . dumiella Neunzig and Dow 


Difundella corynophora Dyar 
(Figs. 1, 8-13, 26) 


Difundella corynophora Dyar 1914:327. 


306 


Males of D. corynophora have the apex 
of the uncus bifurcate, the distal end of the 
arm of the sacculus very broad, and the ae- 
doeagus uniquely forked (Figs. 12—13). Fe- 
males have a wide oblong plate associated 
with the ostium bursae and the signum of 
the corpus bursae is a small, round patch of 
scobinations (Fig. 26). 

The vertex of D. corynophora is ochre 
anteriorly to pale reddish brown, or dark 
purplish brown posteriorly, and the fore- 
wing is mostly dark reddish brown and dark 
purplish brown on its basal three-fifths and 
chiefly ochre and pale reddish brown on its 
distal two-fifths. The underside of the male 
hindwing has a small cluster of thin, mostly 
hooked scales near the short discal vein at 
the distal end of the discal cell, and an elon- 
gate-oval patch of raised sex-scales near 1A 
(Fig. 8). The forewing length is 7.5—8.5 
mm. 

Material examined.—Mexico, 4 km. S. 
Estacion Palenque, Chiapas, 25 July 1984, 
H. H. Neunzig and K. M. Neunzig, geni- 
talia slide 1000 HHN (1 3) [NCSU]; Gua- 
temala, Cayuga, August, Schaus and Barnes 
(1 2) [USNM], Chejel, June, Schaus and 
Barnes, genitalia slide 102, 100 ME (1 @) 
[USNM]; Costa Rica, Sector Cerro Cocori, 
Finca de E. Rojas, 150 m., Provincia Li- 
mon, Sei, I, 18, INOS, ION ZBeouwo, 
567500, #2347, INBio CR _ 1001140900, 
1001141147, genitalia slides 5834, 5835 
HHN (2 ¢) [INBio]; Panama, La Chorrera, 
12 May 1912, A. Busck, genitalia slide 102, 
099 CH (6 holotype) [USNM]; French 
Guiana, Cayenne, [no date], [no collector’s 
name], genitalia slide 102, 101 ME (1 o) 
[USNM]. 


Difundella subsutella (Schaus) 
(Figs. 5, 14-15) 


Ulophora subsutella Schaus 1913: 248. 
Difundella subsutella: Heinrich 1956: 63. 


The most useful feature to identify D. 
subsutella is its aedoeagus, which bears dis- 
tally a large, sclerotized process composed 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


of a series of overlapping, contiguous, 
curved spines (Fig. 15). 

Difundella subsutella is known from the 
male type (Fig. 5) and one other male. The 
species appears to be most similar in color 
and maculation to D. corynophora. The 
vertex is ochre to pale reddish brown, and 
the forewing slightly darker than the fore- 
wing of D. corynophora with purplish 
brown distally. The underside of the hindw- 
ing has a small cluster of thin, hooked 
scales along the short, discal vein at the dis- 
tal end of the discal cell, and an elongate- 
oval patch of raised sex-scales near 1A. The 
forewing length is 7.0—8.0 mm. 

Material examined.—Costa Rica, Juan 
Vinas, January, [no collector’s name, prob- 
ably Schaus], genitalia slide 102, 102 CH 
(3d holotype) [USNM]; Costa Rica, Turrial- 
ba, Estacion Barbilla, P. N. Barbilla, R. E. 
Rio Pacuare, 500 m., Prov. Cartago, Ene. 
AOD, Ib, Clereiie, IG, IN|, Bits /Q=SNSAS7/, 
#66503, INBio CR INB0003417093, geni- 
talia slide 6195 HHN (1 d) [INBio]. 


Difundella dumiella Neunzig and Dow 
(Figs. 2, 16-17) 


Difundella dumiella Neunzig and Dow 
1993-32" 


Difundella dumiella has the uncus oval 
with anteriorly projecting, sinuous, blunt- 
pointed processes, and a juxta with setifer- 
ous elements completely fused into the 
sides of a triangular plate. The aedoeagus 
is elongate with a vesica bearing about five 
cornuti. Difundella dumiella also appears to 
be smaller than other Difundella, having a 
forewing length of 6.0—6.5 mm. 

The vertex of the male is white and pale 
reddish brown and the forewing mostly 
dark reddish brown and purple basally and 
mostly ochre and white distally. The un- 
derside of the hindwing lacks both the small 
cluster of hooked scales along the short, 
discal vein at the distal end of the discal 
cell, and the elongate-oval patch of raised 
sex-scales near 1A that are present in most 
Difundella. 


VOLUME 107, NUMBER 2 


307 


Figs. 5—7. 


Material examined.—Belize, San Igna- 
cio, 6 April 1989, L. C. Dow, genitalia slide 
2741 HHN 
(3 holotype) [USNM], (1 
[NCSU]. 


3 paratype) 


Difundella parana Neunzig and Solis, 
new species 
(Figs. 6, 18-19) 


Diagnosis.—The male genitalia (Fig. 18), 
in general, are more robust than in other 
Difundella. The uncus is quadrate and the 
vinculum broad and rounded. The hooklike 
process of the sacculus is strongly curved 
throughout with its distal part broadened. 

Description.—Forewing length 7.5—8.5 
mm. Head: Vertex ochre; labial palpus up- 
turned, extending above vertex, segment III 
about % as long as II, outwardly ochre and 
brownish red; maxillary palpus short-scaled 
and ochre with brownish red basally; anten- 


Males, dorsal view. 5, Difundella subsutella. 6, D. parana. 7, D. teresina. 


na with shaft of male slightly enlarged and 
flattened along basal half, with sensilla tri- 
chodea (cilia) short, about % as long as 
width of antenna near base of shaft. Tho- 
rax: Dorsum mostly pale reddish brown. 
Forewing: Dark reddish brown and brown 
basally mostly pale reddish brown, dusted 
with white distally; antemedial line weakly 
developed, pale reddish brown bordered ba- 
sally by obscure, incomplete, dark reddish 
brown and black line and bordered distally 
by more pronounced dark reddish brown 
and black line; postmedial line weakly de- 
veloped, white and pale reddish brown, bor- 
dered basally by dark reddish brown, mod- 
erately well defined line and bordered dis- 
tally by similar dark colored, but less well 
defined, line; area between antemedial line 
and postmedial line with ochre and pale 
reddish brown, fused discal spots, and with 
black or dark brown streaks on veins M, to 


308 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


discal 
scale 


cluster 


raised 


sex-scales 


anal fold 


== 


oe 
SoSs 


—2 
“~, 


> 
SS 


23 
— a 

to 
D5 


+ 
SD 
D> 


Figs. 8-13. Difundella corynophora, male. 8, Right forewing and hindwing (discal scale cluster, patch of 
raised sex-scales and anal fold are on underside of wing). 9, Left maxillary palpus, frontal view. 10, Left labial 
palpus, lateral view. 11, Right antenna, basal part, frontal view. 12, Genitalia, ventral view, without aedoeagus. 
13, Aedoeagus. 


CuA,. Hindwing: Above hyaline to pale Male genitalia (Figs. 1S—19): Uncus quad- 
brown in male; underside of male with rate, broad and acuminate distally (not al- 
small cluster of thin, hooked scales near ways apparent unless genitalia spread on 
short discal vein at distal end of discal cell, slide), simple basally; gnathos reduced, 
and raised elongate-oval patch of raised without median process; transtilla absent; 
sex-scales near 1A; anal fold along 3A. juxta in shape of inverted triangular plate 


VOLUME 107, NUMBER 2 


with low, robust, setiferous lateral arms; 
valva well developed, broadened and with 
spinelike setae distally; sacculus with large 
hooklike process strongly curved through- 
out its length and with distal part expanded, 
and with large basal tuft of slightly broad- 
ened setae; aedoeagus elongate; vesica with 
patch of many small spines; vinculum 
broad and rounded. Female unknown. 

Holotype:—¢. Curitiba, Parana, Brazil, 
12-II-1975, V. O. Becker, genitalia slide 
5739 HHN. [USNM]. 

Paratypes:—l1 d. Sete Lagoas, Minas 
Gerais, Brazil, 720 m., 25-VII-1969, V. O. 
Becker, genitalia slide 5741 HHN. [VOB]. 
1 3. Joinville, Santa Catarina, Brazil, 6- 
XII-1969, V. O. Becker, genitalia slide 5740 
HHN [NCSU]. 1 3d. Nova Teutonia, Brazil, 
VIII-1961, EF Naumann, genitalia slide 5993 
HHN [USNM]. 

Etymology.—The specific epithet is 
based on the Brazilian state (Parana) in 
which the holotype was collected. 


Difundella unguifera Neunzig and Solis, 
new species 
(Figs. 3, 22—23, 28) 


Diagnosis.—The pair of large, unira- 
mous, clawlike, medially projecting pro- 
cesses at the base of the uncus (Fig. 22) are 
peculiar to this species. Also, the costa of 
the forewing of the male is strongly pro- 
duced anteriorly at its base. The female 
genitalia have near the base of the papillae 
anales a pair of sclerotized, scobinate lateral 
lobes (Fig. 28). 

Description.—Forewing length 8.5—9.0 
mm. Head: Vertex dark brown in male, 
pale brown in female; labial palpus of both 
sexes upturned, extending above vertex, 
segment III about % as long as II in male, 
and segment III about % as long as II in 
female, outwardly mostly reddish brown in 
male, pale reddish brown in female; max- 
illary palpus short-scaled in both sexes, 
pale reddish brown with brownish red ba- 
sally; antenna with shaft of male slightly 
enlarged and flattened along basal half, with 
sensilla trichodea (cilia) short, about ¥ as 


309 


long as width of antenna near base of shaft; 
antenna of female simple. Thorax: Dorsum 
mostly purplish brown. Forewing: Mostly 
dark purplish brown basally and chiefly 
pale reddish brown distally; antemedial line 
moderately well developed, pale reddish 
brown, weakly bordered basally by a few 
dark reddish brown scales and bordered dis- 
tally by stronger line of dark reddish brown, 
and a few black, scales; postmedial line 
weakly developed, white and pale reddish 
brown bordered basally and distally with 
dark reddish brown line; area between an- 
temedial and postmedial lines with discal 
spots obscure, reddish brown, fused, and 
with dark reddish brown streaks on some 
veins. Hindwing: Above hyaline to pale 
brown in male; mostly brown in female; un- 
derside of male with cluster of mostly 
hooked scales near discal vein at distal end 
of cell (scales forming cluster longer and 
more robust than in other species in genus), 
without elongate-oval patch of raised sex- 
scales near 1A; anal fold along 3A. Male 
genitalia (Figs. 22—23): Uncus triangular in 
distal half and with pair of clawlike medi- 
ally projecting processes at base; gnathos 
reduced, without median process; transtilla 
absent; juxta an inverted triangular plate 
with very weakly developed lateroposterior, 
setiferous elements; valva well developed, 
slightly broadened, and with spinelike setae 
distally; sacculus with long, slender, slight- 
ly-curved, pointed process and with large 
basal tuft of distally broadened setae; ae- 
doeagus short; vesica with two cornuti; vin- 
culum about as long as greatest width. Fe- 
male genitalia (Fig. 28): Integment be- 
tween base of papillae analis and abdominal 
segment VIII collar produced laterally, 
sclerotized and scobinate; apophysis poster- 
ioris short and slender; apophysis anterioris 
short, about as thick as apophysis poster- 
ioris; ostium bursae with wide sclerotized, 
scobinate plate; ductus bursae about as long 
as corpus bursae, with patch of scobinations 
distally; corpus bursae membranous (unable 
to determine if signum present because of 
spermatophores); ductus bursae attached to 


310 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 14-19. Male genitalia. 14, Difundella subsutella, ventral view, without aedoeagus. 15, D. subsutella, 
aedoeagus. 16, D. dumiella, ventral view, without aedoeagus. 17, D. dumiella, aedoeagus. 18, D. parana, ventral 
view, without aedoeagus. 19, D. parana, aedoeagus. 


corpus bursae near junction of ductus bur- ando, LN 253250, 449700, INBio CR 

sae and corpus bursae. 1000895968, genitalia slide 4752 HHN [IN- 
Holotype.—<d. Estaci6n La Casona, Bio]. 

1,520 m., Res. Biol. Monteverde, Prov. Paratypes.—l @. Monte Verde, Punt. 

Puntarenas, Costa Rica, Dic. 1992, N. Ob- Prov., Costa Rica, 30—31 Jul. 1981, D. H. 


VOLUME 107, NUMBER 2 


Janzen and W. Hallwachs, INBio CR 
1002043686, genitalia slide 4753 HHN [IN- 
Bio]. 1 2, Turrialba, Sta. Cruz, 1,500 m., 
Costa Rica, VIII-1981, V. O. Becker, geni- 
talia slide 4926 HHN [USNM]. 

Etymology.—The name unguifera is de- 
rived from the Latin ungui (claw) and fera 
(to bear) in reference to the remarkable pair 
of clawlike processes at the base of the un- 
cus. 


Difundella cancerella Neunzig and Solis, 
new species 
(Figs. 4, 20-21, 27) 


Diagnosis.—A pair of biramous clawlike 
processes at the base of the uncus are 
unique to D. cancerella. Difundella cancer- 
ella males also lack the small tuft of scales 
associated with the discal vein, and the 
raised, elongate-oval patch of scales near 
1A, on the underside of the hindwing. The 
absence of both of these groups of scales is 
a feature shared only in Difundella with D. 
dumiella; the female genitalia have robust 
apophyses anteriores (about twice as thick 
as apophyses posteriores) and the signum is 
a patch of saw-toothed rows of spines. 

Description.—Forewing length 7.0—9.0 
mm. Head: Vertex white to brownish 
white; labial palpus of both sexes upturned, 
extending above vertex, outwardly pale red- 
dish brown to reddish brown; maxillary 
palpus short-scaled in both sexes, ochre; an- 
tenna with shaft of male slightly enlarged 
and flattened along basal half with sensilla 
trichodea short, about % as long as width of 
antenna near base of shaft; antenna of fe- 
male simple. Thorax: Dorsum mostly ochre 
and reddish brown. Forewing: Chiefly pur- 
plish brown basally and mostly ochre and 
pale reddish brown distally; antemedial line 
weakly developed, pale reddish brown bor- 
dered basally by obscure, incomplete red- 
dish brown line and bordered distally by 
stronger reddish brown and black line; post- 
medial line weakly developed, ochre and 
pale reddish brown, bordered basally by 
dark reddish brown, moderately well de- 
fined line and distally by a broader, less ob- 


311 


vious, mostly purplish brown line; area be- 
tween antemedial and postmedial lines with 
fused discal spots ochre and pale reddish 
brown and with streaks of purplish brown 
on some veins. Hindwing: Above hyaline 
to pale brown in male, slightly darker in 
female; underside of male without tuft of 
scales near discal vein and without elon- 
gate-oval raised patch of scales near 1A; 
with anal fold along 3A. Male genitalia 
(Figs. 20-21): Uncus broad and with pair 
of slender clawlike, biramous processes ba- 
sally; gnathos reduced, without median pro- 
cess; transtilla absent; juxta with large 
rounded, setiferous, lateral lobes; valva 
slender, slightly enlarged and with spinelike 
setae distally; sacculus with long, slender, 
untapered, blunt process and with large bas- 
al tuft of distally broadened, scalelike setae; 
aedoeagus short, slender and swollen me- 
dially; vesica with few small cornuti; vin- 
culum about as long as greatest width. Fe- 
male genitalia (Fig. 27): Apophysis 
posterioris short and slender; apophysis an- 
terioris short and robust (about 2X as wide 
as apophysis posterioris); ostium bursae 
with wide sclerotized spined plate; ductus 
bursae shorter than corpus bursae and with 
microspines in posterior half; corpus bursae 
with signum a patch of short, saw-toothed 
rows of spines; ductus seminalis attached to 
corpus bursae near junction of ductus bur- 
sae and corpus bursae. 

Holotype.—d. Sector Cerro Cocort, Fin- 
ca de E. Rojas, 150 m., Prov. Limon, Costa 
Rica, Oct. 1993, E. Rojas, LN 286000 
567500, #2395, INBio CR 1001642551, 
genitalia slide 4754 HHN [INBio]. 

Paratypes.—2 2. Same collection data as 
holotype except Ene. and Set. 1993 INBio 
CR 1001141159, 1001403076, genitalia 
slides 4755, 5833 HHN [USNM, NCSU]; 1 
6.P.N. Tapanti, A. C. Amistad, Prov. Car- 
ta., 1150 m., Costa Rica, Ene. 1994, G. 
Mora, LN 194000 559800, INBio CR 
1001830436, genitalia slide 5836 HHN 
[NCSU]; 1 ¢. Paraiso, Pque. Nat. Tapanti, 
Sect. La Represa, del Puente del Rio Porras, 
300 m. SE, 1,660 m., Prov. Cartago, Mar. 


312 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


yy yy F ii 


25 


Figs. 20-25. Male genitalia. 20, Difundella cancerella, ventral view, without aedoeagus. 21, D. cancerella, 
aedoeagus. 22, D. unguifera, ventral view, without aedoeagus. 23, D. unguifera, aedoeagus. 24, D. teresina, 
ventral view, without aedoeagus. 25, D. teresina, aedoeagus. 


2002, R. Delgado, L. N. 186550—560600, processes at the base of the uncus resemble 

#67365, INBio CR INB0003446759, geni- the claws of a crab. The specific epithet is 

talia slide 6192 HHN [INBio]. a combination of the Latin cancer (crab) 
Etymology.—The biramous’ clawlike and the diminutive Latin suffix -ella. 


VOLUME 107, NUMBER 2 


Figs. 26-28. 


Difundella teresina Neunzig and Solis, 
new species 
(Figs. 7, 24-25) 


Diagnosis.—The most distinctive feature 
of D. teresina is the juxta with its large pos- 
teriorly projecting lobes (Fig. 24). The an- 
gulate saccular spine and the serrate ae- 
doeagus also can be used to identify the 
species (Figs. 24-25). 

Description.—Forewing length 7.0 mm. 
Head: Vertex ochre and pale reddish 
brown; labial palpus upturned, extending 
above vertex, outwardly a mixture of ochre 
and reddish brown; maxillary palpus short- 


Sls} 


Female genitalia, ventral view. 26, Difundella corynophora. 27, D. cancerella. 28, D. unguifera. 


scaled, ochre with brownish red basally; an- 
tenna with shaft of male slightly enlarged 
and flattened along basal half, with sensilla 
trichodea (cilia) short, about ¥; as long as 
width of antenna near base of shaft. Tho- 
rax: Dorsum mostly reddish and purplish 
brown. Forewing: Mostly black or purplish 
black basally and chiefly ochre and pale 
reddish brown distally; antemedial line 
weakly developed, pale reddish brown, bor- 
dered basally by incomplete dark reddish 
brown line and bordered distally by a more 
complete dark reddish brown line; post- 
medial line somewhat stronger than ante- 


314 


medial line, pale reddish brown bordered 
basally by dark reddish brown line and bor- 
dered distally by less obvious dark reddish 
brown line; area between antemedial and 
postmedial line with pale reddish brown, 
fused discal spots, and streaks of dark red- 
dish brown on veins M, to CuA,. Hind- 
wing: Above hyaline to brown along mar- 
gins; underside of male with small tuft of 
scales near discal vein at distal end of discal 
cell, and with elongate-oval patch of raised 
scales near 1A; anal fold along 3A. Male 
genitalia (Figs. 24—25): Uncus elongate- 
oval, simple at base; gnathos reduced, with- 
out median process; transtilla absent; juxta 
with large, truncated, posteriorly-project- 
ing, setiferous lobes; valva slender, slightly 
enlarged and with spinelike setae distally; 
sacculus with long, slender, tapered, point- 
ed, strongly-bent process, and with large 
basal tuft of distally, slightly broadened, 
scalelike setae; aedoeagus slender, serrate 
distally; vesica scobinate; vinculum about 
as long as greatest width. Female unknown. 
Holotype.—<d. Teresina, 500 m., Goias, 
Brazil, 29-V-1994, V. O. Becker and K. S. 
Sattler, genitalia slide 5312 HHN [USNM]. 
Etymology.—The specific epithet is 
based on the type locality (Teresina). 


ACKNOWLEDGMENTS 


We thank Eugenie Phillips (Instituto Na- 
cional de Biodiversidad, Santo Domingo, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Costa Rica), and Vitor Becker (Brasilia, 
Brasil) for the opportunity to study speci- 
mens of Difundella in the INBio collection 
and in the Becker collection, respectively. 
Robert L. Blinn (North Carolina State Uni- 
versity) took the habitus photographs and 
reviewed an initial draft of the manuscript. 
Others who made useful comments to im- 
prove the manuscript were Lewis L. Deitz 
and Brian M. Wiegmann (both North Car- 
olina State University), Jay C. Shaffer 
(George Mason University), and David A. 
Nickle (Systematic Entomology Laborato- 
ry, USDA). 


LITERATURE CITED 


Dyar, H. G. 1914. Report on the Lepidoptera of the 
Smithsonian Biological Survey of the Panama Ca- 
nal Zone. Proceedings of the United States Na- 
tional Museum 47: 139-350. 

Heinrich, C. 1956. American moths of the subfamily 
Phycitinae. United States National Museum Bul- 
letin 207: 1-581. 

Neunzig, H. H. 1986. New records of Phycitinae from 
Mexico and a description of a new genus and spe- 
cies (Lepidoptera: Pyralidae). Proceedings of the 
Entomological Society of Washington 88: 122— 
126. 

Neunzig, H. H. and L. C. Dow. 1993. The Phycitinae 
of Belize (Lepidoptera: Pyralidae). North Carolina 
Agricultural Research Service Technical Bulletin 
304: 1-131. 

Schaus, W. 1913. New species of Heterocera from 
Costa Rica. Annals and Magazine of Natural His- 
tory 11: 234-262. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 315-318 


A NEW SPECIES OF THE ERIOCAMPA OVATA GROUP 
(HYMENOPTERA: TENTHREDINIDAE) FEEDING ON ALNUS MATSUMURAE 
CALL. (DICOTYLEDONEAE: FAGACEAE) FROM JAPAN 


ICHIJI TOGASHI 


1-chome, Honmachi, Tsurugi-machi, Ishikawa Prefecture, 920-2121, Japan 


Abstract.—Eriocampa alnicola, n. sp., reared from larvae feeding on leaves of Alnus 
matsumurae Call. in Mt. Hakusan, Ishikawa Prefecture, Japan, is described and illustrated. 
A key is given to the three Japanese species of the Eriocampa ovata group. 


Key Words: 


Two species of the Eriocampa ovata 
group, E. mitsukurii Rohwer, 1910, and E. 
babai Yogashi, 1980, were previously 
known from Japan (Togashi 1980). This 
group is characterized by the red median 
lobe of the mesoscutum. In 1997, I obtained 
a female belonging to the E. ovata group 
which was reared from larvae feeding on 
leaves of Alnus matsumurae Call., on Mt. 
Hakusan (alt. 2702 m), Ishikawa Prefecture. 
After comparative study, I concluded that it 
represents a new species, differing from the 
previously described species by the color 
pattern of the lateral lobes of the mesos- 
cutum and of the first tergite. Here, I de- 
scribe and illustrate this new species. 


KEY TO JAPANESE SPECIES OF THE 
ERIOCAMPA OVATA GROUP 


1. Median lobe of mesoscutum red (Figs. 1, 8); 
lateral lobe of mesoscutum mostly black; lan- 
cet with 18 or 19 serrulae  ............ 2 
— Median and lateral lobes of mesoscutum red 
(Fig. 12); lancet with 25 serrulae (apex of lance 
as in Fig. 20; 10th to 12th serrulae of lancet as 
HIM SUD) TSE te eee ss al ahs babai Togashi 
2. Lateral lobes of mesoscutum entirely black 
(Fig. 8); mesoscutellum blunt in front (Fig. 8); 
hind basitarsus slightly shorter than following 
4 segments combined (ratio about 1.0:1.1); in- 
ner spur of foretibia as in Fig. 10; apex of lance 
as in Fig. 17; lancet with 18 serrulae; 10th to 


sawflies, Tenthredinidae, Allantinae, new species, Alnus 


12th serrulae of lancet as in Fig. 19 ..... 

anit | dennte oats Oscar ig ley etal rs mitsukuriti Rohwer 
— Central, inner and lateral sides of lateral lobes 

of mesoscutum red (Fig. 1); mesoscutellum 

acute in front (Fig. 1); hind basitarsus shorter 

than following 4 segments combined (ratio 

about 1.0:1.3) (Fig. 4); inner spur of foretibia 

as in Fig. 5; apex of lance as in Fig. 14; lancet 

with 19 serrulae; 10th to 12th serrulae of lancet. 

asin Biges MGs toy siyalocentee re oe alnicola, n. sp. 


Eriocampa alnicola Togashi, new species 
(Figs. 1-7, 13-16) 


Female.—Length, 8 mm. Body black 
with following red: pronotum except for la- 
tero-ventral portion, tegula, median lobe of 
mesoscutum (Fig. 1), inner and lateral sides 
and central portion of lateral lobes of me- 
soscutum (Fig. 1), central portion of first 
tergite (Fig. 7); antero-lateral side of second 
tergite (Fig. 7), and lateral side of third ter- 
gite (Fig. 7). Antenna black with apical por- 
tion of 4th segment and apical 5 segments 
red. Wings pale yellowish tinged; stigma, 
costa, and subcosta of forewing yellowish 
red or reddish brown; other veins dark 
brown to black. Legs black; anterior 4 
knees reddish brown; inner side of foretibia 
reddish brown; fore tarsus reddish brown. 

Head: ‘Transverse; postocellar area 
slightly wider than length (ratio between 
width and length about 1.0:0.8), convex; 


316 


6 


Figs. 1-12. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


/] 


1-7, Eriocampa alnicola, holotype. 1, Mesoscutum and mesoscutellum, dorsal view. 2, Postspi- 


racular sclerite. 3, Hind wing. 4, Hind tarsus, lateral view. 5, Fore inner tibial spur, lateral view. 6, Tarsal claw, 
lateral view. 7, Ist to 4th abdominal tergites, dorsal view. 8-11, E. mitsukurii. 8, Mesoscutum and mesoscutellum, 
dorsal view. 9, Postspiracular sclerite. 10, Fore inner tibial spur, lateral view. 11, Tarsal claw, lateral view. 12, 


Mesoscutum and mesoscutellum of E. babai, dorsal view. 


OOL:POL:OCL = 1.1:1.0:1.4; interocellar 
and postocellar furrows nearly absent; lat- 
eral furrows distinct; frontal area surround- 
ed by distinct wall; area just in front of an- 
terior ocellus distinctly pitted; median fovea 
transverse in outline, distinctly concave, 
with distinct longitudinal carina; lateral fo- 
vea deep, ellipsoidal in outline; supracly- 
peal area slightly convex; clypeus rather 
flattened, frontal margin emarginate; la- 
brum rather flattened; malar space shorter 
than diameter of front ocellus (ratio about 
RO 225) 

Antenna shorter than costa of forewing 
(ratio about 1.0:1.2); relative lengths of seg- 
ments about 1.0:1.0:2.2:1.5:1.0:0.9:0.8:0.6: 
1.0; 3rd antennal segment shorter than 4th 
plus Sth segments (ratio about 1.0:1.2); 


pedicel longer than wide (ratio between 
length and width about 1.0:0.6). 

Thorax: Normal; mesoscutellum acute in 
front (Fig. 1), slightly convex; cenchrus 
small, distance between them about 3.6X as 
long as width of one; postspiracular sclerite 
small (Fig. 2); wing venation of hind wing 
as in Fig. 3, petiole of anal cell shorter than 
cu-a (ratio about 1.0:2.4); hind basitarsus 
shorter than following 4 segments com- 
bined (ratio about 1.0:1.3); inner spur of 
foretibia as in Fig. 5; tarsal claw (Fig. 6) 
with large acute basal lobe and with inner 
tooth broader and about as long as outer 
tooth. 

Abdomen: Normal; sawsheath in lateral 
view as in Fig. 13; lancet with 19 serrulae; 
apex of lance as in Fig. 14; apex of lancet 


VOLUME 107, NUMBER 2 


(ae Sena 


10th Ihikela 
Figs. 13-22. 


317 


10th 11th 12th 


13-16, Eriocampa alnicola, holotype. 13, Sawsheath, lateral view. 14, Apex of lance. 15, Apex 


of lancet. 16, 10th to 12th serrulae of lancet. 17-19, E. mitsukurii. 17, Apex of lance. 18, Apex of lancet. 19, 
10th to 12th serrulae of lancet. 20—22, E. babai. 20, Apex of lance. 21, Apex of lancet. 22, 10th to 12th serrulae 


of lancet. 


as in Fig. 15; 10th to 12th serrulae of lancet 
as in Fig. 16. 

Punctation: Vertex, frontal area, upper 
half of inner orbits, and gena covered with 
strong, craterlike punctures; lower half of 
inner orbits finely and closely punctured, 
matt; inner side of median and lateral fo- 
veae practically impunctate and shining; su- 
praclypeal area practically impunctate and 
shining; clypeus and labrum finely punc- 
tured; pronotum, meso- and metascutellum, 
postspiracular sclerite, and mesepisternum 
covered with strong, craterlike punctures; 
anterior half of median lobe of mesoscutum 
finely and rather closely punctured; poste- 
rior half of median lobe practically im- 
punctate and shining; central area of lateral 
lobe of mesoscutum distinctly and sparsely 
punctured, remainder practically impunc- 
tate and shining; sunken area of median 


lobe, mesoscutellar appendage, and metan- 
otum nearly impunctate and shining; me- 
sepimeron and mesosternum practically im- 
punctate and shining; abdominal tergites 
nearly impunctate and shining. 
Male.—Unknown. 
Food plant.—A/nus matsumurae Call. 
Distribution.—Japan (Honshu). 
Holotype.—Female, 14.1X.1997, reared 
from larva feeding on leaves of Alnus mat- 
sumurae, Ishikawa Prefecture, Mt. Haku- 
san. Deposited in the National Science Mu- 
seum (Nat. Hist.), Tokyo. 
Etymology.—The specific epithet is 
based on the genus name of the food plant. 
Remarks.—This new species is very 
closely allied to E. mitsukurii, but it is eas- 
ily distinguished from the latter by the part- 
ly red lateral lobes of the mesoscutum and 
the first tergite (both black in EF. mitsukurii, 


318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


see Figs. 1, 8), by the small postspiracular 
sclerite (large in E. mitsukurii, see Figs. 2, 
9), by the shape of the apical portion of the 
lance and lancet (see Figs. 14, 15, 17, 18), 
and by the shape of the 10th to 12th ser- 
rulae of the lancet (see Figs. 16, 19). From 
E. ovata (L.), a Palearctic species and in- 
troduced into North America, it is easily 
separated by the red tegula (black in E. ova- 
ta). From E. singularis Malaise, 1931, 
known from Siberia, it is easily distin- 
guished by the black mesoscutellum (red- 
dish yellow in E. singularis) and by the 
black legs (pale yellow in E. singularis). 


ACKNOWLEDGMENTS 


I cordially thank David R. Smith, Sys- 
tematic Entomology Laboratory, USDA, 
Washington, DC, for his kind advice and 
review of the manuscript. 


LITERATURE CITED 


Malaise, R. 1931. Blattwespen aus Wladiwostok und 
anderen Teilen Ostasiens. Entomologisk Tidskrift 
51: 97-159. 

Tohashi, I. 1980. Sawflies of the genus Eriocampa 
Hartig (Hymenoptera, Tenthredinidae) in Japan. 
Kontyt 48: 35-41. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 319-322 


ESTABLISHMENT OF THREE EUROPEAN FLEA BEETLES IN NOVA 
SCOTIA: LONGITARSUS GANGLBAUERI HEIKERTINGER, L. JACOBAEAE 
(WATERHOUSE), AND L. RUBIGINOSA (FOUDRAS) 
(COLEOPTERA: CHRYSOMELIDAE: ALTICINAE) 


E. RICHARD HOEBEKE AND A. G. WHEELER, JR. 


(ERH) Department of Entomology, Cornell University, Ithaca, NY 14853, U.S.A. (e- 
mail: erh2 @cornell.edu); (AGW) Department of Entomology, Soils, and Plant Sciences, 
Clemson University, Clemson, SC 29634, U.S.A. (e-mail: awhlr@clemson.edu) 


Abstract.—A Eurasian ragwort flea beetle, Longitarsus jacobaeae (Waterhouse), was 
released in western and eastern Canada, beginning in 1971, and in the U.S. Pacific North- 
west in 1969 to help control infestations of the noxious Eurasian weed, tansy ragwort 
(Senecio jacobaea L.; Asteraceae), in pastures and rangelands. In eastern Canada, L. 
jacobaeae previously has been recorded as established in Prince Edward Island and New 
Brunswick; we report it from 5 counties in Nova Scotia. Based on collecting from 1993 
to 2003, two other adventive species of Longitarsus are reported for the first time from 
Nova Scotia and the Canadian Maritime Provinces. Longitarsus ganglbaueri Heikertinger, 
known previously only from Oregon and Manitoba, is reported from two counties, and 
L. rubiginosa (Foudras), recorded only from Ontario and Quebec, is reported from two 
counties. 


Key Words: Chrysomelidae, biological control, new records, adventive species, tansy 


ragwort, hedge bindweed 


Tansy ragwort (Senecio jacobaea L.; As- 
teraceae), a Eurasian biennial or short-lived 
perennial, now infests pastures, rangelands, 
forest clearcuts, and disturbed areas along 
the Atlantic coast from Rhode Island to the 
Canadian Maritime Provinces and along the 
Pacific coast from northern California to 
British Columbia (ARS-USDA_ 1971, 
Coombs et al. 1991, White 1996). Senecio 
jJacobaea is especially noxious because it 
not only competes with valuable forage 
plants but also contains foliar pyrrolizidine 
alkaloids toxic to cattle, deer, horses, and 
pigs (Bain 1991). Livestock poisoning usu- 
ally occurs only with severe tansy ragwort 
infestations and when the foliage of more 
desirable plants is scarce (Turner and 
Szczawinski 1991). 


Tansy ragwort, first recorded in Califor- 
nia in 1912 and Oregon in 1922, had be- 
come an important weed on the Pacific 
coast by the mid-1950s (Pemberton and 
Turner 1990). Because of its invasive nature 
and lethal effects to livestock, this weed be- 
came a candidate for biological control in 
western North America in the late 1960s. 

The ragwort flea beetle, Longitarsus ja- 
cobaeae (Waterhouse), was deliberately in- 
troduced into western and eastern Canada 
(Julien and Griffiths 1998) from Italy and 
Switzerland. Released in Canada beginning 
in 1971, it became established in British 
Columbia, Prince Edward Island, and New 
Brunswick by 1982, but apparently failed 
to do so in Ontario or Nova Scotia (Harris 
et al. 1984, Julien and Griffiths 1998). 


320 


LeSage (1991) did not list L. jacobaeae 
from New Brunswick or Prince Edward Is- 
land, but Harris (2003) and Riley et al. 
(2003) confirmed the beetle’s establishment 
in those provinces. Releases of L. jaco- 
baeae in the United States began in 1969, 
and its establishment has been confirmed in 
California, Oregon, and Washington (Julien 
and Griffiths 1998). In combination with 
other introduced Palearctic insects, the flea 
beetle has provided successful biological 
control of tansy ragwort in Oregon (e.g., 
McEvoy et al. 1991) and northern Califor- 
nia (Piper 1985, Pemberton and Turner 
1990). 

The Old World distribution of L. jaco- 
baeae includes the British Isles, Europe, 
North Africa, Siberia, Kazakhstan, Kirgizia, 
and Dagestan (LeSage 1988, Doguet 1994). 
Known host plants in its native range in- 
clude Senecio jacobaea (British Isles, con- 
tinental Europe), S. giganteus Desf. (Mo- 
rocco), and occasionally other Senecio spe- 
cies (Frick 1970, LeSage 1988). Westcott et 
al. (1985) reported several native species of 
Senecio as adult-feeding hosts. 

During a series of survey trips to the Ca- 
nadian Maritime Provinces (1993—2003), 
we collected numerous specimens of L. ja- 
cobaeae from tansy ragwort in several lo- 
calities in Nova Scotia, a province in which 
its establishment has been considered 
doubtful. Examination of male genitalia 
verified its identity as L. jacobaeae rather 
than the morphologically similar L. flavi- 
cornis (Stephens) (Shute 1975). 

We also collected specimens of another 
Palearctic Longitarsus in Nova Scotia, L. 
ganglbaueri Heikertinger, wherever one of 
its known host plants (LeSage 1988), Se- 
necio viscosus L., was encountered. Lon- 
gitarsus ganglbaueri was detected (in vac- 
uum samples) for the first time in North 
America during studies related to the bio- 
control of tansy ragwort in Oregon (West- 
cott et al. 1985). Westcott et al. (1985) also 
reported the species from Manitoba based 
on a single specimen collected in 1978. 
LeSage (1988) considered this flea beetle to 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


have been introduced accidentally into the 
Pacific Northwest with material of S. jaco- 
baeae that was introduced intentionally 
from Italy. Westcott et al. (1985) and West- 
cott and Brown (1992), however, argued 
that it is naturally Holarctic or an early ac- 
cidental introduction that predates the bio- 
control releases of S. jacobaeae. It might 
represent an unintentional introduction in 
Nova Scotia because it appears not to have 
been introduced for biological control (Har- 
ris et al. 1984, Julien and Griffiths 1998). 
The species still is recorded only from 
Manitoba in Canada (LeSage 1991, Riley 
et al. 2003). In the Palearctic Region, L. 
ganglbaueri occurs in the British Isles, Ire- 
land, Sweden to Spain, Italy, Balkans, Cau- 
casus, and Dagestan (LeSage 1988). This 
Senecio specialist has been reported from S. 
viscosus and S. vulgaris L. (LeSage 1988) 
in Europe and from S. jacobaea, S. pseu- 
daureus Rydb., S. sylvaticus L., and S. tri- 
angularis Hooker in North America (West- 
cott et al. 1985) 

Our collecting from another invasive 
plant, hedge bindweed (Calystegia sepium 
(L.) Br; Convolvulaceae), in Nova Scotia 
yielded specimens of the Palearctic L. ru- 
biginosa (Foudras). Previous North Amer- 
ican records are limited to Ontario and Que- 
bec (LeSage 1991, Riley et al. 2003). In the 
Old World, this species is known from the 
British Isles, central Europe, Albania, Bul- 
garia, Caucasus, Siberia, Turkestan, and 
Dagestan (LeSage 1988) and northeastern 
Asia (Westcott and Brown 1992). Its hosts 
in the Palearctic Region, in addition to C. 
sepium, include other convolvulaceous 
plants (LeSage 1988). 

Here, we give the first Nova Scotian re- 
cords for L. jacobaeae, L. ganglbaueri, and 
L. rubiginosa. We collected L. jacobaeae 
on Senecio jacobaea, L. ganglbaueri on S. 
viscosus, and L. rubiginosa on Calystegia 
sepium. The number of adults collected is 
given parenthetically. Voucher specimens 
of the three species are deposited in the 
Cornell University Insect Collection (Itha- 
ca, New York) and the National Museum of 


VOLUME 107, NUMBER 2 


Natural History, Smithsonian Institution 
(Washington, D.C.). 


MATERIAL EXAMINED 


Longitarsus jacobaeae (Waterhouse).— 
CANADA: Nova Scotia: Antigonish Co., 
Antigonish, 30 July and 3 August 2003 (6). 
Cape Breton Co., Glace Bay, Renwick 
Park, | August 2003 (9); North Sydney, 24 
July 1995 (58), 2 August 2003 (3); Sydney, 
23 July 1995 (112), 31 July 2003 (11). In- 
verness Co., Port Hawksbury, 31 July 2003 
(13). Pictou Co., Pictou marine terminal, 30 
July 2003 (20); Rte. 106, Caribou, 30 July 
2003 (4). Yarmouth Co., Yarmouth, 20 July 
1994 (43), 6 August 2001 (67). 

Longitarsus ganglbaueri Heikertinger.— 
CANADA: Nova Scotia: Colchester Co., 
Truro, nr. railroad tracks, 21 July 1994 (15), 
29 July 2003 (11). Halifax Co., Dartmouth, 
3-8 August 2001 (23), 27-28 July 2003 
(3); Halifax, nr. old railroad station, 26 June 
1993 (8); 2 August 2001 (9). 

Longitarsus rubiginosa (Foudras).— 
CANADA: Nova Scotia: Annapolis Co., 
Port George, 5 August 2001 (11); Bridge- 
town, Jubilee Park, 5 August 2001 (15). 
Shelburne Co., Shelburne, 20 July 1994 (7). 


ACKNOWLEDGMENTS 


We thank Janet Ciegler (West Columbia, 
SC) for providing distribution records of 
Longitarsus listed by Riley et al. (2003), 
and Peter H. Adler (Department of Ento- 
mology, Soils, and Plant Sciences, Clem- 
son, SC) for providing helpful comments 
on a draft of the manuscript. This research 
was supported by the Cornell University 
Agricultural Experiment Station federal for- 
mula funds, Project No. NYC-139413 to 
ERH, received from Cooperative State Re- 
search, Education, and Extension Service, 
U.S. Department of Agriculture. Any opin- 
ions, findings, conclusions, or recommen- 
dations expressed in this publication are 
those of the authors and do not necessarily 
reflect the view of the U.S. Department of 
Agriculture. 


LITERATURE CITED 


ARS-USDA (Agricultural Research Service-U.S. De- 
partment of Agriculture). 1971. Common Weeds 
of the United States. Dover Publications, New 
York, 463 pp. 

Bain, J. EK 1991. The biology of Canadian weeds. Se- 
necio jacobaea L. Canadian Journal of Plant Sci- 
ence 71: 127-140. 

Coombs, E. M., T. E. Bedell, and P. B. McEvoy. 1991. 
Tansy ragwort (Senecio jacobaea): Importance, 
distribution, and control in Oregon, pp. 419-428. 
In James, L. FE, J. O. Evans, M. H. Ralphs, and 
R. D. Child, eds. Noxious Range Weeds. West- 
view Press, Boulder, Colorado. 

Doguet, S. 1994. Genre 3.—Longitarsus Berthold, pp. 
161-315. In Pericart, J. ed. Faune de France, 80. 
Coleopteres, Chrysomelidae, Vol. 2 (Alticinae). 

Frick, K.E. 1970. Longitarsus jacobaeae (Coleopera: 
Chrysomelidae), a flea beetle for the biological 
control of tansy ragwort. 1. Host plant specificity 
studies. Annals of the Entomological Society of 
America 63: 284-296. 

Harris, P. 2003. Classical biological control of weeds: 
Established biocontrol agent. Longitarsus jaco- 
baeae (Waterhouse) and L. flavicornis Stephens. 
root beetles. http://res2.agr.ca/lethbridge/weedbio/ 
agents/alongflv_e.htm (accessed 18 Jan. 2004). 

Harris, P, A. T. S. Wilkinson, and J. H. Myers. 1984. 
Senecio jacobaea L., tansy ragwort (Compositae), 
pp. 195-201. Jn Kelleher, J. S. and M. A. Hulme, 
eds. Biological Control Programmes Against -In- 
sects and Weeds in Canada, 1969-1980. Com- 
monwealth Agricultural Bureaux, Farnham Royal, 
UK. 

Julien, M. H. and M. W. Griffiths. 1998. Biological 
Control of Weeds: A World Catalogue of Agents 
and Their Target Weeds, 4" edition. CAB Inter- 
national, Wallingford, UK, 223 pp. 

LeSage, L. 1988. Notes on European Longitarsus spe- 
cies introduced in North America (Coleoptera: 
Chrysomelidae: Alticinae). Canadian Entomolo- 
gist 120: 1133-1145. 

. 1991. Family Chrysomelidae, pp. 301—323. In 
Bousquet, Y., ed. Checklist of Beetles of Canada 
and Alaska. Research Branch, Agriculture Cana- 
da, Publication 1861/E. 

McEvoy, P., C. Cox, and E. Coombs. 1991. Successful 
biological control of ragwort, Senecio jacobaea, 
by introduced insects in Oregon. Ecological Ap- 
plications 1: 430—442. 

Pemberton, R. W. and C. E. Turner. 1990. Biological 
control of Senecio jacobaea in northern Califor- 


nia, an enduring success. Entomophaga 35: 71— 
VI. 
Piper, G. L. 1985. Biological control of weeds in 


Washington: status report, pp. 817-826. /n Del- 
fosse, E.S., ed. Proceedings of the 6" International 
Symposium on the Biological Control of Weeds, 


Lo 
N 
i) 


19-25 August 1984, Vancouver, Canada. Agricul- 
ture Canada, Ottawa. 

Riley, E. G., S. M. Clark, and T. N. Seeno. 2003. Cat- 
alog of the Leaf Beetles of America North of 
Mexico (Coleoptera: Megalopodidae, Orsodacni- 
dae and Chrysomelidae, Excluding Bruchinae). 
Special Publication 1. Coleopterists Society, Sac- 
ramento, California, 290 pp. 

Shute, S. L. 1975. Longitarsus jacobaeae Waterhouse 
(Col., Chrysomelidae): Identity and distribution. 
Entomologist’s Monthly Magazine 111: 33-39. 

Turner, J. J. and A. E Szczawinski. 1991. Common 
Poisonous Plants and Mushrooms of North Amer- 
ica. Timber Press, Portland, Oregon, 311 pp. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Westscott R. L. and R. E. Brown. 1992. Comments on 
Longitarsus ganglbaueri Heikertinger in North 
America: A rebuttal (Coleoptera: Chrysomelidae). 
Coleopterists Bulletin 46: 67. 

Westcott, R. L., R. E. Brown, D. B. Sharratt, and R. 
E. White. 1985. Longitarsus: A new species from 
Oregon and a new record for North America (Co- 
leoptera: Chrysomelidae). Pan-Pacific Entomolo- 
gist 61: 323-330. 

White, R. E. 1996. Leaf beetles as biological control 
agents against injurious plants in North America, 
pp. 373-399. In Jolivet, P. H. A. and M. L. Cox, 
eds. Chrysomelidae Biology, Vol. 2. Ecological 
Studies. SPB Academic Publishing, Amsterdam, 
The Netherlands. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 323-335 


FOCUSING ON MORPHOLOGY: APPLICATIONS AND IMPLICATIONS OF 
CONFOCAL LASER SCANNING MICROSCOPY 
(DIPTERA: CAMPICHOETIDAE, CAMILLIDAE, DROSOPHILIDAE) 


VALERIE SCHAWAROCH, DAVID GRIMALDI, AND ANGELA V. KLAUS 


(VS) Natural Sciences, Baruch College, One Bernard Baruch Way Box A-0506, New 
York, NY 10010-5585, U.S.A. (e-mail: valerie-schawaroch @baruch.cuny.edu); (VS, DG) 
Division of Invertebrate Zoology, American Museum of Natural History, Central Park 
West at 79™ Street, New York, NY 10024-5192, U.S.A.; (AVK) Microscopy and Imaging 
Facility, American Museum of Natural History, Central Park West at 79" Street, New 
York, NY 10024-5192, U.S.A. 


Abstract.—Confocal laser scanning microsocopy (CLSM) produces high fidelity, three- 
dimensional digital images of complex and phylogenetically informative, diagnostic insect 
structures such as the male genitalia. This study examined terminalia, mouthparts, and 
antennae of five genera in three ephydroid fly families—Campichoetidae, Camillidae and 
Drosophilidae—by utilizing the natural fluorescence of insect tissues under visible light 
(543 nm) excitation. CLSM images better reveal the shapes and positions of individual 
structures, and particularly their connections, as compared to conventional bright field 
light microscopy where fine layers and sutures are often obscured. CLSM has important 
implications for imaging valuable insect specimens, including types, and obviates the need 


for re-examination. 


Key Words: 


Though considered quaint by some, the 
300-year tradition of studying pinned insect 
specimens is actually an ingenious method 
for preserving and observing a microscopic 
landscape of setae, spines, sensilla, append- 
ages, and microsculpture in situ. Modern 
systematic revisions, for example, empha- 
size the diversity of characters and their 
phylogenetic reliability, so it is not unusual 
now to see hundreds of anatomical struc- 
tures reported in any particular monograph, 
and the source is hardly exhausted (Gri- 
maldi 1990, Grimaldi and Nguyen 1999, 
Mathis and Zatwarnicki 2002). New mor- 
phological details are even still being dis- 
covered on what is arguably the best known 
eukaryote, Drosophila melanogaster Mei- 
gen (Ashburner 1989). Here, we present 


Ephydroidea, genitalia, systematics, pinned specimens 


confocal laser scanning microscopy 
(CLSM) as a powerful new tool in the ar- 
senal of insect morphological studies. 
Insect morphology is traditionally stud- 
ied using light microscopy, both stereomi- 
croscopy (for opaque and cleared speci- 
mens, generally 10—70), and compound 
microscopy (for transparent specimens, 
generally 60—400X). The optical con- 
straints of these two techniques comple- 
ment each other: stereomicroscopy provides 
relief and three-dimensionality, while com- 
pound microscopy provides higher resolu- 
tion and magnification. Both types of mi- 
croscopes will continue to be instrumental 
for routine morphological analysis, since 
the investigator can quickly obtain infor- 
mation about a specimen. However, record- 


324 


ing images using either stereo or compound 
microscopy is beset with the classical prob- 
lem of the interplay between resolution, il- 
lumination, and depth of field. 

Scanning electron microscopy (SEM) is 
also routinely used in insect morphology, 
although the availability of these instru- 
ments is usually very limited. SEM pro- 
vides unparalleled resolution and depth of 
field at high magnifications, but can only 
image surfaces. Insect structures are rarely 
just flat fields; they usually consist of folds, 
apodemes, and obscuring layers and lobes, 
such as in complex male genitalia, most of 
which are hidden from view in a typical 
SEM mount. Additionally, specimen prep- 
aration for conventional SEM can be de- 
structive—samples must be coated with a 
thin metal layer (usually gold/palladium) to 
render the specimen electrically conductive. 
Alternative SEM technologies exist, such as 
variable pressure (or low vacuum) SEM 
(Sammons and Marquis 1997) and cold 
field-emission SEM (Klaus 2003), that al- 
low for uncoated specimen viewing, how- 
ever, limited surface views of exposed 
structures is still a problem with these in- 
struments. 

CLSM uses a conventional compound 
microscope setup, but utilizes laser light of 
specific wavelengths for specimen illumi- 
nation rather than white light (for an intro- 
duction to CLSM, see Paddock 1999). The 
light used for illumination (excitation wave- 
length) excites fluorescent molecules pre- 
sent in the specimen, and filters are then 
used to isolate the specific wavelengths of 
light emitted by the excited molecules. Nor- 
mally, fluorescence in CLSM depends on 
the fluorescent label or tag applied to the 
specimen; however, in the case of insects 
and other arthropods, the fluorescence is 
due to naturally occurring compound(s) 
(Neff et al. 2000, Lardeux et al. 2000). The 
real power of CLSM is the ability to “op- 
tically section’ a specimen by placing a 
pinhole aperture in front of the final signal 
detector. The pinhole allows only the signal 
from the plane of focus to be collected, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


therefore excluding obscuring out-of-focus 
light. The plane of focus is changed in mi- 
nute and equal increments by the software 
controlling the microscope. The optical sec- 
tions can then be reconstructed into a three- 
dimensional (3-D) object, thus obtaining a 
clear image of the original structure without 
the blurring usually associated with images 
of thick objects obtained on a compound 
microscope. If the data from the specimen 
are collected under optimal conditions (see 
Klaus et al. 2003) a faithful 3-D rendering 
can be created and rotated in space for 
viewing at any angle. Thus, CLSM pro- 
vides the resolution of compound light mi- 
croscopy, combined with the relief and 
three-dimensionality typically seen in SEM 
images. 

There are few prior studies applying 
CLSM to image arthropods based on their 
natural fluorescence. Galassi (1997a, b) and 
Galassi et al. (1998) found CLSM to be su- 
perior to compound light microscopy for 
fine details and used the technique for tax- 
onomic descriptions of copepods. Zill et al. 
(2000) used CLSM imaging in their ex- 
amination of cockroach trochanter structure 
for biomechanical applications. Klaus et al. 
(2003) extensively explored the technique 
for imaging genitalic structures from lower 
and higher Diptera, and presented a detailed 
methodology for using CLSM in such stud- 
ies. In addition to the usefulness for imag- 
ing external structures, CLSM can also cre- 
ate images of soft internal structures such 
as muscles and ducts (Klaus et al. 2003, C. 
Chaboo, personal communication). It would 
appear that CLSM can be used for virtually 
any fluorescent insect structure small 
enough to fit within the field of view on a 
compound microscope, although there are 
real limitations on the thickness of struc- 
tures that can be imaged by CLSM (Masters 
and Farmer 1993, Skaliora and Pagakis 
2002). 

In the current work, we apply CLSM to 
various structures in the ephydroid fly fam- 
ilies of Camillidae, Campichoetidae, and 
Drosophilidae, with an emphasis on their 


VOLUME 107, NUMBER 2 


aedeagus 


paraphysis 
dorsal lobe 


paraphysis © 
ventral lobe 


epandrium 


cercus 
mon 


paraphysis 
dorsal lobe 


oe 
i, aedeagal -§ 
+ apodeme 


paraphysis 
ventral lobe 


hypandrium 
surstylus ! 


Fig. 1. CLSM maximum intensity projections for articulated male genitalia. Both ventral (A) and 
views are pictured for Campichoeta latigena; (C) posterior view 
Cladochaeta inversa. All scale bars = 


dorsal (B) 
and (D) internal view of the genitalia of 
100 microns. 


326 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Male ephydroids used in this study are from pinned specimens in the American Museum of Natural 


History collection and from VS dissertation. 


———— 


Species 


Collection Information 


— 


Cladochaeta inversa (Walker) 
Scaptodrosophila sp. 
Afrocamilla armata Barraclough 
Grimaldi 
Campichoeta latigena McAlpine 


Houghton Co. Michigan; 20 August 1959, B. and K. Driesbach 
Australia; W. B. Mather 
36°10'E, 3°50’S, Tarangire NP, Tanzania, Africa. July 1994, D. 


8.700’ Rustler Park, 7 mi. W. Portal, Cochise Co. Arizona. | June 


1991 D. Grimaldi 


Drosophila parvula Bock and Wheeler 


Drosophila Species Resource Center, Bowling Green. Stock # 


14028-0621.0; 20 miles north Kuala Lumpur, Malaysia; 22—28 
June 1962 Wasserman [type culture] 


male terminalia. Insect male terminalia are 
routinely used in insect systematics for sep- 
arating and defining species, and because of 
their complexity (Eberhard 1985) they fre- 
quently are a significant source of charac- 
ters for phylogenetic studies. The complex- 
ity of the structures has also led to many 
ambiguities regarding homology among 
male genitalia in insects, including Diptera 
(e.g., Tuxen 1978, McAlpine 1981 vs. Grif- 
fiths 1972 vs. Cumming et al. 1995). Ephy- 
droids are typical of the diverse array of 
cyclorrhaphan flies in that the male termin- 
alia are complex as well as three-dimen- 
sional (vs. the male terminalia of nemato- 
cerous flies, which are flatter), and therefore 
provide a particularly appropriate subject 
for testing applications of CLSM. 


MATERIALS AND METHODS 


Specimens.—The male genitalic dissec- 
tions are from the species listed in Table 1. 
Drosophila melanogaster females obtained 
from Carolina Biological Supply Co. were 
used for the dissections of antenna, probos- 
cis and female genitalia. 

Dissections.—Dissections were carried 
out as described by Klaus et al. (2003). 
Dried, pointed flies were relaxed in a hu- 
midity chamber prior to dissection. For gen- 
italic dissections, the distal half of the ab- 
domen was removed. For the proboscis and 
antenna, the head was disarticulated from 
the thorax at the occiput. These parts were 
cleared by placing them in a 0.6 ml Eppen- 


dorf tube containing a 10% KOH solution 
and incubated for approximately 2 h in a 
heated water bath (70°C). The dissected 
parts were rinsed, partially dehydrated and 
transferred to glycerine through successive 
wells of a spot plate. The first well con- 
tained distilled water, the next two wells 
contained 70% ethanol, and the last well 
contained glycerine. Specimens were fur- 
ther dissected in the glycerine well. 
Mounting.—Temporary slide mounts 
were made using glycerine jelly. Specimens 
were placed in a glycerine jelly droplet 
mounted between two cover slips (No. 1.5, 
nominal thickness 170 wm). Spacers mea- 
suring approximately 200 wm in thickness 
(1.e., two stacked pieces of No. 0 cover 
slips, nominal thickness 100 im each) were 
placed to either side of the glycerine jelly 
drop containing the specimen. Mounting 
the specimens between two cover slips al- 
lows the specimens to be imaged from both 
sides by simply turning over the mount, 
thereby increasing the quality of image data 
collected. This is to avoid data loss from 
the side of a specimen farthest away from 
the sources of illumination and detection. 
CLSM imaging.—Images were taken us- 
ing a Zeiss 510 confocal laser scanning mi- 
croscope (CLSM) equipped with an invert- 
ed Zeiss Axiovert microscope housed in the 
AMNH Microscopy and Imaging Facility. 
Specimens were imaged from both sides us- 
ing a 20X Fluar dry objective lens (NA = 
0.75, WD = 610 pm). In a few instances, 


VOLUME 107, NUMBER 2 


epandrium aedeagus 
paraphysis 

dorsal lobe 
paraphysis 

ventral lobe 


| Ni a hypandrium 
ventral lobe 


surstylus of epandrium anaes 
gal 
<~— "apodeme 
wt 


Cercus 


epandrium 


~~ ‘hypandrium 


Fig. 2. CLSM maximum intensity projections of disarticulated male genitalia. An exemplar Scaptodrosophila 
sp. with a posterior view of the periphallic structures in (A) plus ventral and dorsal aspects of the phallic 
structures pictured in (B) and (C) respectively. Periphallic (D) posterior view and phallic (E) ventral and (F) 


dorsal views for Afrocamilla armata. Scale bars 100 microns. 


328 


the object to be imaged (i.e., female geni- 
talia and dorso-ventral and lateral views of 
the proboscis) was so large that the zoom 
setting was reduced to 0.70. The autofluo- 
rescence of the insect structures was in- 
duced using a helium neon laser (excitation 
wavelength = 543 nm) set at full power (1 
mW). The fluorescence emission signal was 
detected using a 560 nm long pass filter; 
these are the factory preset filter settings for 
rhodamine (Cy3, Texas red) stained struc- 
tures. Due to the thickness of the speci- 
mens, series of optical slice images were 
taken through the specimen along the z- 
axis. Each z-slice was 2.2 pm thick with an 
optimal overlapping interval of approxi- 
mately 1.1 wm. Each image (or slice) was 
collected with the same parameters: pixel 
frame size of 1024 X 1024 with a unidirec- 
tional scan mode and an 8-bit pixel depth. 
The pixel scanning time was 6.40 ws. Every 
rasterized line of each frame was duplicated 
and the mean was taken. Image collection 
settings were optimized for each specimen. 
The amplitude gain was held constant at a 
value of 1.0. The amplitude offset usually 
equaled —0.06 (range of —0.05 to —0.07) 
in order to produce a maximally black 
background. For each specimen imaged, a 
single detector gain setting was used for all 
the z-slices in a single 3-D image stack. For 
each specimen the detector gain was opti- 
mized so that brightest part of the whole 
sample contained only a few red (oversat- 
uration) spots. The Zeiss 510 LSM program 
function was used for pinhole size optimi- 
zation (74 wm for all specimens). 

3-D image reconstruction.—Maximum 
intensity projections were generated using 
the Zeiss 510 LSM® software version 3.2. 
Image contrast was enhanced using Adobe 
PhotoShop® version 6.0.1. In some instanc- 
es the transparency setting for optical image 
stacks altered in the Zeiss 510 LSM® soft- 
ware to reveal internal structures in situ 
while obviating the destructiveness of ac- 
tual (physical) dissection. 

Light microscope photomicrography.— 
Bright field light microscope (BFLM) im- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ages were captured using the CLSM Zeiss 
Axiovert microscope and the software set- 
ting for transmitted light. Single frame dig- 
ital images were collected at various focal 
planes within the specimen. All the settings 
were as described above, except that no 
emission filter was used. This method of 
BFLM is similar to photomicrography us- 
ing a traditional bright field compound mi- 
croscope, except that a laser source of light 
is used for illumination instead of a white 
light flood source. BFLM images were col- 
lected using a 20 Fluar dry objective lens 
(NA = 0.75, WD = 610 um). 

Morphological terminology.—Terminol- 
ogy used follows Grimaldi (1990) and 
McAlpine (1981). 


RESULTS AND DISCUSSION 


Protocols developed at the AMNH pro- 
vide optimal CLSM images of insect cuticle 
(Klaus et al. 2003). We applied those pro- 
tocols here to complex structures of ephy- 
droid fly terminalia, proboscis and anten- 
nae. Unlike SEM, these CLSM images dif- 
ferentiate insect structures on the basis of 
degree of naturally occurring autofluores- 
cence from an undetermined molecular 
source(s) (see Klaus et al. 2003 for discus- 
sion). For male terminalia, CLSM images 
provide a refined view of structures, thus 
improving the ability to properly homolo- 
gize structures. This is particularly impor- 
tant for the periphallic structures such as the 
paraphyses and gonopods, which flank the 
aedeagus. These can be elaborately devel- 
oped, as in Cladochaeta (Fig. 1), or they 
can be reduced, or even lost in other dro- 
sophilids. The hypandrium (an internalized, 
ninth sternite) and epandrium (a capsule- 
like eighth tergite) are generally easily dis- 
cerned, though there are lobes of each de- 
veloped in various genera, and these lobes 
are sometimes articulated. The phallic and 
periphallic complex are attached to the hy- 
pandrium and lie within the epandrium, so 
these structures—which are routinely used 
for species determinations in these flies— 
are only partially observable with an SEM. 


VOLUME 107, NUMBER 2 


Cercus 


cercal spines 
cercal clasper 


surstylus 


ventral lobe of epandrium- decasternum 


aedeagus 


paraphysis 
: aa paraphysis 


dorsal lobe 


paraphysis 
ventral lobe ° » 


gonopod 


aedeagal ——___ 


apodeme 
a 


hypandrium 


Fig. 3. Maximum intensity projection of Drosophila parvula male genitalia imaged with CLSM. (A) Dorsal 
view of the periphallic structures; (B) ventral and (C) dorsal view of the phallic structures. All scale bars = 100 


microns. 


330 


Fig. 4. Bright field light microscope images of 
Drosophila parvula male phallic structures, including 
the phallus, paraphyses, gonopods and the pointed me- 
dial lobe in the upper portion of the hypandrium. The 
focal plane progresses from the ventral to the dorsal 
side of the structure—images (A) through (C). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


CLSM revealed that each paraphysis of 
Cladochaeta inversa (Walker) is composed 
of a dorsal and a ventral lobe (Fig. 1), 
which are pressed closely against each oth- 
er, but with a distinct suture between them. 
The paraphyses in Cladochaeta had previ- 
ously been interpreted as a complete struc- 
ture (Grimaldi and Nguyen 1999). In Scap- 
todrosophila sp. the paraphysis is also ar- 
ticulated, but with a small distal lobe (Fig. 
2) that is most distinct in ventral view. In 
Drosophila parvula Bock and Wheeler a bi- 
lobed paraphysis is readily seen, also with 
a dorsal one closely adpressed over a ven- 
tral one (Fig. 3), which is a structure rarely 
reported or never interpreted as such before 
in the subgenus Sophophora. The setae on 
the paraphyses of Afrocamilla, Campichoe- 
ta, and Scaptodrosophila have wide, deep 
socketed bases (Figs. 1—2), similar to those 
of trichobothria or sensilla trichodea, and 
thus are probably sensory in function. The 
gonopods in drosophilids are a pair of struc- 
tures articulating with the lateral arms of 
the hypandrium and lying against or over 
the aedeagus. Recognizing or discerning 
their structure is often difficult. Indeed, they 
are small in D. parvula, but are still clearly 
distinguished, and have scattered microtri- 
chia. Gonopods are much larger in Scap- 
todrosophila sp., but also have fine setae. 
In Campichoeta there are two pairs of per- 
iphallic structures (Fig. 1), the inner pair of 
which (immediately flanking the aedeagus) 
could be interpreted as gonopods. However, 
the deep, wide sockets of the setae on both 
pairs of lobes (similar to what is found in 
Scaptodrosophila) indicate these are pa- 
raphyses. 

Until now, scientists have had to rely on 
DIC optics and variation in illumination to 
discern subtle differences in slide-mounted, 
transparent insect structures. A comparision 
of the CLSM image for the ventral hypan- 
drium of D. parvula (Fig. 3b) with the 
bright field image for the same structure 
(Fig. 4) illustrates the limitations of trans- 
mitted light microscopy. The compound 
light microscope shows structures, albeit 


VOLUME 107, NUMBER 2 


tergite 8 


epiproct 


hypoproct 


Fig. 5. 


spermathecal capsule 


spermathecal duct 


Maximum intensity projection of CLSM image stack of internal and external female genitalic struc- 


tures of Drosophila melanogaster. Note that structures range broadly in the degree of sclerotization, from heavily 
sclerotized to completely membranous. Scale bar = 100 microns. 


blurred, simultaneously from multiple focal 
planes. Therefore, edges and depth cues can 
be confused by variation in sclerotization of 
structures. The CLSM, however, through 
the use of a pinhole, only collects in-focus 
images from one focal plane at a time, thus 
facilitating the creation of 3-D representa- 
tions of an object. 

There are few studies comparing the fe- 
male genitalia of ephydroid flies (Throck- 
morton 1962, 1966; Grimaldi 1990; Gri- 
madi and Nguyen 1999), yet female termin- 
alia possess numerous features that are 
probably important for taxonomic investi- 
gations. Among ephydriod flies the female 
genital apparatus is composed of the ter- 
minal abdominal segments (8 and 9). Fe- 
male D. melanogaster (Fig. 5) exhibit the 
apomorphic drosophilid loss of a cercus— 
a paired lobe structure located postero-ven- 
tral to the epiproct (tergite 9) and the hy- 
poproct (sternite 9) in ephydriods. The 8" 
sternite is the lobate oviscape (also referred 
to as the oviscapt). Among ephydriods, the 
Oviscape varies in overall shape. Addition- 
ally, the distribution and orientation of the 
setae along the margin varies, as does the 


type of setae (fine or pegs). The variation 
in the oviscape may corrolate with ovipo- 
sition substrate (e.g., flowers) as well as 
phylogeny. The D. melanogaster oviscape 
pictured in Fig. 5 has a broad, blunt tip used 
to deposit eggs in soft surface of rotting 
fruits and possesses pegs an apomorphic 
condition within Drosophila (Grimaldi 
1990). Among ephydriods, sperm storage 
organs are either ventral/seminal receptacle 
(e.g., Camichoetidae, Diastatidae, Ephydri- 
dae) or spermathecae. Both vary in overall 
shape, size and degree of sclerotization. The 
spermatheca of D. melanogaster is a broad 
capsule (Fig. 5) (pleisomorphic condition 
among drosophilds [Grimaldi 1990]) fairly 
well sclerotized with a single spermathecal 
duct. The duct and its articulation with the 
capsule are extremely difficult to discern 
using a light microscope, but easily visu- 
alized using CSLM (Fig. 5). 

In most higher Diptera, the proboscis is 
used for mopping surface liquids from the 
substrate, which in the case of drosophilids 
includes a suspension of yeasts, fungi, bac- 
teria and sugars, associated with rotting 
fruits or other plant parts and slime fluxes, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


lacinea 


labellum with Sept tah ec. m 
pseudotrachea _/ ()e———77, e 
a ae = —— labrum 


sieve pore 


cibarium 
posterior sensilla 


anterior sensilla 


hypopharynx 
labrum 


labellum with 
pseudotrachea 


Fig. 6. Maximum intensity projection of female Drosophila melanogaster proboscis. (A) Lateral view; (B) 
and (C) dorsal and ventral views, respectively. Notice that internal structures traditionally manually dissected 


such as the lacinia, cibarium and hypopharynx are clearly visible, allowing in situ examination. Scale bars 


100 microns. 


VOLUME 107, NUMBER 2 


scape 


pedicle 


basal | 
flagellomere 


dorsal seam 


aristomere 3 


sensilla 


Fig. 7. Medial (A) and lateral (B) views of a female Drosophila melanogaster antenna imaged using CLSM. 
Scale bars = 100 microns. The conus was made visible in (C) by changing the transparency setting in the 3-D 


rendering of the maximum intensity projection. The boxed region in (B) indicates the area of the specimen that 


was rendered transparent. 


and in a few instances pollen and nectar. 
The various components of the proboscis 
(e.g., hypopharynx, cibarium and lacinia) 
vary among ephydriod flies and have been 


used as a source of phylogenetic informa- 
tion (Grimaldi 1990). For illustrative pur- 
poses lateral and dorsal-ventral views of a 


female D. melanogaster proboscis is pre- 


334 


sented in Fig. 6. The shape of the hypo- 
pharnyx (tube used to conduct food) and 
cibarium (pump apparatus with associated 
muscles used to create the suction) vary di- 
agnostically among ephydriods. The vari- 
ous sensilla along the cibarium (anterior, 
posterior and sensilla pores) vary in number 
and arrangement (Fig. 6) these sensilla 
probably function as stretch receptors. An- 
other variable ephydriod structure is the 
paired laciniae, each laterally placed along 
the proboscis, which vary in shape and se- 
tation. Many of these minute, but phylo- 
genetically informative, structures are em- 
bedded within the proboscis and require 
difficult, destructive, and time consuming 
dissection in order to isolate them. 

Antennal surface features, especially ar- 
eas of fine setation such as the sensilla on 
the basal flagellomere, are thoroughly doc- 
umented using CLSM in Fig. 7. Addition- 
ally, by subsampling the image stack and 
manipulating the transparency setting in the 
3-D reconstruction, internal structures, such 
as the conus can be isolated. The conus pos- 
sesses structural variation at the family and 
the genus levels within drosophilids (see 
Grimaldi 1990). 

Possible applications of CLSM for insect 
morphology could be substantial, particu- 
larly for the study of obscure, intricate, 
sclerotized structures that have been diffi- 
cult to observe or interpret, including: mi- 
nute sclerites in the proboscis (e.g., the hy- 
popharynx of psocopterans and lice); the 
proventriculus and its array of internal 
spines in fleas, boreid mecopterans, Dic- 
tyoptera, and other insects; the complex of 
axillary sclerites at the base of the wing, 
and their muscular insertions; and, of 
course, terminalia. The expense of CLSM 
instruments will probably prohibit their use 
for routine imaging in systematics, but the 
implications for systematics are likewise 
highly significant. First, few illustrations 
can compare with the fidelity of a CLSM 
image. As a prolific illustrator, one of us 
(DG) acknowledges that even the best il- 
lustrations are subjective in what they do 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


not portray (as trivial information) as in 
what they do. Given that the best CSLM 
images can also be manipulated for any 
view, they provide superior renditions of 
types and other unique specimens critical 
for identification. Digitally sharing of in- 
formation with remote colleagues also ob- 
viates the need to ship types or other valu- 
able specimens. 


ACKNOWLEDGMENTS 


We thank Ronald Ochoa for a critical re- 
view and helpful suggestions. Thank you to 
Caroline Chaboo for sharing her prelimi- 
nary results of CLSM imaging. VS thanks 
Vladimir Blagoderov for thoughtful discus- 
sions concerning Diptera. We are grateful 
for generous support to DG and VS by Na- 
tional Science Foundation DEB grant 
#0075360. VS also was supported by the 
Weissman School of Arts and Sciences of 
Baruch College which provided release 
time for this research. 


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PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 336-345 


BLISSUS MINUTUS (BLATCHLEY) AND TOONGLASA UMBRATA (DISTANT): 
SELDOM-COLLECTED NATIVE CHINCH BUGS 
(HEMIPTERA: LYGAEOIDEA: BLISSIDAE) AS COLONISTS OF THE 
AFRICAN BUNCHGRASS ERAGROSTIS CURVULA, AND THEIR 
ASSOCIATION WITH OTHER GRASSES IN THE SOUTHERN UNITED STATES 


A. G. WHEELER, JR. 


Department of Entomology, Soils, and Plant Sciences, Clemson University, 
Clemson, SC 29634-0315, U.S.A. (e-mail: awhlr@clemson.edu) 


Abstract.—Weeping lovegrass (Eragrostis curvula; Poaceae), an African bunchgrass 
first introduced into the United States in the late 1920s, has acquired a diverse insect 
fauna in the southern states. Among the little-known native species that have colonized 
this chloridoid grass are the chinch bugs Blissus minutus (Blatchley) and Toonglasa um- 
brata (Distant). Known previously only from Florida, B. minutus is reported for the first 
time from Alabama, Georgia, North Carolina, Oklahoma, South Carolina, and Texas. This 
multivoltine grass generalist was collected most often on E. curvula (293 collections). 
Adults were found on 34 grasses of the subfamilies Aristoideae, Chloridoideae, and Pan- 
icoideae, with nymphs observed on 21 species, mainly chloridoids and panicoids. Toon- 
glasa umbrata was collected in Arizona, New Mexico, and Texas from seven native 
grasses and one introduced grass, in addition to weeping lovegrass (subfamilies Chlori- 
doideae, Panicoideae, and Pooideae). It was syntopic with B. minutus on E. curvula in 
parts of Oklahoma and Texas and was the only blissid found on weeping lovegrass in 
more arid regions (Arizona, New Mexico, and western Texas). 


Key Words: Insecta, Lygaeoidea, chinch bugs, weeping lovegrass, Poaceae, host expan- 


sion, novel hosts 


Weeping lovegrass, Eragrostis curvula 
(Schrad.) Nees (Poaceae), is an African 
bunchgrass that was obtained in the late 
1920s and tested in the mid-1930s for in- 
troduction into the southwestern United 
States to help reclaim rangelands depleted 
by years of drought and overgrazing (Crider 
1945). It began to be planted in southeast- 
ern states, mainly for erosion control along 
highways, in the 1940s and 1950s (Wheeler 
1999). Numerous native insects have colo- 
nized this exotic grass, especially hemipter- 
ans that develop within the extensive 
crowns. Among the little-known native in- 
sects that make up the cryptic diversity as- 


sociated with weeping lovegrass (Wheeler 
1999, 2003; Wilson and Wheeler, in press) 
are the chinch bugs Blissus minutus 
(Blatchley) and Toonglasa umbrata (Dis- 
tant). 

I first found B. minutus (May 1996) in 
crowns of a native grass, Andropogon vir- 
ginicus L., in Georgia but discovered that 
the blissid could be detected more easily by 
sampling weeping lovegrass, especially in 
or near the fall-line Sandhills from southern 
North Carolina to eastern Alabama. Era- 
grostis curvula thus was used as an “‘indi- 
cator” plant for my attempt to delimit the 
southeastern distribution of a _ species 


VOLUME 107, NUMBER 2 


known previously only from Florida. I also 
sampled native grasses to elucidate host- 
plant relationships for a blissid whose 
nymphal hosts were unknown. Southwest- 
ern states were included in my survey after 
B. minutus (1999) and T. umbrata (2000) 
were found on weeping lovegrass in 
Oklahoma. Herein I summarize the distri- 
bution records obtained from surveys of B. 
minutus 1n southern states and 7. umbrata 
in the Southwest, record the native and non- 
native grasses that serve as hosts of the blis- 
sids, and provide notes on their seasonal 
histories. 


MATERIALS AND METHODS 


Blissus minutus was sampled by holding 
a white enamel pan at the base of weeping 
lovegrass and other potential hosts, striking 
the crown with an ax handle, and noting the 
presence of dislodged nymphs and adults 
(brachypters and macropters). Specimens 
were collected from nearly every site 
(>90%) and deposited in the National Mu- 
seum of Natural History, Smithsonian In- 
stitution, Washington, D.C. (USNM). In the 
Southeast, nymphs were recorded as early 
(I-III) or late (IV—V) instars (1996-1999) 
or were sorted to instar in the field (2000— 
2004). In the Southwest, 7. umbrata was 
collected during surveys for B. minutus. 
Adults (only nymphs were present at cer- 
tain sites) of T. umbrata from each locality 
were collected and deposited in the USNM; 
when nymphs were present, instars were 
field sorted and recorded. 

Notes on the seasonality of B. minutus 
are based mainly on the sampling of weep- 
ing lovegrass in South Carolina in Richland 
County near Spears Creek Church Rd., 3.8 
km SSE of Pontiac (34°06.3'N, 80°49.5'W) 
and in Florida in Lake County along Rt. 27, 
8.5 km SSE of Clermont (28°28.7'N, 
81°43.0'W) and Polk County along Rt. 27, 


2.4 km N of jct. CR-54, 7.3 km NNW of 


Loughman (28°16.6'’N, 81°39.6'W). Notes 
on seasonality of B. minutus and T. umbra- 
ta in the Southwest are based on observa- 
tions only from late April to early June. 


337 


Although “‘chinch bugs” sometimes is 
reserved for species of the genus Blissus, | 
use this common name in a broad sense 
(e.g., Slater and Wilcox 1973, Slater 1976) 
to refer to members of the Blissidae (for- 
merly the lygaeid subfamily Blissinae; see 
Henry 1997). The names and subfamilies of 
grasses follow Soreng et al. (2004). 


Blissus minutus Blatchley 


Blissus minutus was described from Dun- 
edin (Pinellas County), Florida, by Blatch- 
ley (1925a) as Ischnodemus pusillus and 
was renamed /. minutus because of primary 
homonymy (Blatchley 1925b). Leonard 
(1968) transferred the species to Blissus 
Burmeister. All additional published re- 
cords of B. minutus are from Florida: Royal 
Palm Park [Miami-Dade Co.] (Blatchley 
1926); Bahia Honda Key [Monroe Co.] 
(Leonard 1968); and Broward, Franklin, 
Hillsborough, Levy, Palm Beach, Pasco, St. 
Lucie, and Volusia counties (Slater and 
Baranowski 1990). Previous ecological in- 
formation consists only of the collection of 
adults from “‘grass tufts” (Blatchley 1925a, 
1926) and sea oats, Uniola paniculata L. 
(Poaceae) (Leonard 1968). Siater (1976) 
noted that the blissid ““breeds” on sea oats 
and attributed the information to Leonard 
(1968); that paper, however, mentioned 
only the collection of adults. 

Distribution (Fig. 1)—Known previous- 
ly only from 11 counties in Florida (Slater 
and Baranowski 1990), B. minutus is now 
known in the southeastern United States 
from Alabama (1 county), Georgia (68), 
North Carolina (15), and South Carolina 
(24). Records also are available for fourteen 
additional counties in Florida. Although 
Fig. | shows the distribution of B. minutus 
only by county, the blissid appeared to be 
restricted to the Sandhills ecoregion of most 
counties lying within both the Sandhills and 
Piedmont (e.g., Anson, Lee, and Montgom- 
ery in North Carolina and Edgefield, Fair- 
field, and Lancaster in South Carolina). 

Attempts to collect B. minutus in the 
Piedmont of North Carolina (mainly Union 


338 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


B Record from author’s fieldwork 


Record from literature 


RY Record from author’s fieldwork and literature 


Fig. 1. 


County) were unsuccessful. The only Pied- 
mont record in South Carolina was from 
southern McCormick County near the fall 
line. My sampling of weeping lovegrass at 
multiple sites in Abbeville, Anderson, 
Chester, Greenwood, Newberry, Oconee, 
Pickens, and York counties in South Caro- 
lina was negative for the blissid. Records 
from the Piedmont of Georgia were mainly 
from counties near the fall line; extensive 
sampling of weeping lovegrass failed to 
yield specimens in more northern Piedmont 
counties: Hart, Jackson, Madison, Morgan, 
Oconee, Taliaferro, and Wilkes. An excep- 
tion to the bug’s collection from more 
southern Piedmont counties in Georgia is 
the record from Atlanta (Cobb County). 

In Alabama, B. minutus was found at 
three sites near Cottonton (Russell County) 
in sandy areas of the East Gulf Coastal 
Plain. I did not find it during sampling of 
weeping lovegrass in adjacent Lee and Ma- 
con counties to the north and west, respec- 
tively. The blissid is generally distributed in 
peninsular Florida and is known from two 
counties in the panhandle. 

In the Southwest, I collected B. minutus 


Pia 


Known distribution (by county) of Blissus minutus based on personal collecting and the literature. 


in the tall-grass prairies and mixed-grass 
plains in 16 counties of western Oklahoma. 
It was not found in sandhills of southeastern 
Oklahoma (southern Atoka County) despite 
the sampling of weeping lovegrass in April 
or May of each year from 2000 to 2003. 
The blissid was found in Texas (14 coun- 
ties) in or near the panhandle in the Rolling 
Plains and High Plains, as well as three 
counties in the Edwards Plateau. It was col- 
lected as far west as Dawson, Gaines, and 
Terry counties, where it was syntopic with 
T. umbrata, but was not found in other 
western counties (Bailey, Cochran, and 
Yoakum). 

Host plants.—Blissus minutus was col- 
lected more often (293 records) on weeping 
lovegrass (Eragrostis curvula), the princi- 
pal plant used to obtain distribution records, 
than on any other host (Tables 1—2). Other 
chloridoid grasses that served as hosts in- 
cluded two native species of Eragrostis and 
an adventive species of the genus (E. pilosa 
(L.) P. Beauv.). Among native grasses, 
broomsedge (Andropogon virginicus L.), a 
panicoid, accounted for the largest number 
of records (39; Table 1). 


VOLUME 107, NUMBER 2 


339 


Table 1. Number of collections of Blissus minutus from grasses in the southeastern United States: Florida, 
Georgia, North Carolina, and South Carolina; four collections from Eragrostis curvula in Russell County, Al- 
abama, are not listed. Species marked with an asterisk are ones from which nymphs were collected. 


— 


Grass 


Aristoideae 


Aristida beyrichiana Trin. & Rupr. 
*A. condensata Chapn. 

*A. purpurascens Poir. 

*A. tuberculosa Nutt. 


Chloridoideae 


Cynodon dactylon (L.) Pers. 
*FEragrostis curvula (Schrad.) Nees 

E. elliottii S. Wats. 
*E. hirsuta (Michx.) Nees 
*E. pilosa (L.) P. Beauv. 
*E. refracta (Muhl. ex Elliott) Scribn. 
*Muhlenbergia capillaris (Lam.) Trin. 

{including M. capillaris var. filipes (M. A. Curtis)] 
*M. expansa (Poir.) Trin. 

Spartina bakeri Merr. 

Triplasis americana P. Beauv. 


Panicoideae 


*Andropogon floridanus Scribn. 
A. glaucopsis (Elliott) Steud. 
A. glomeratus (Walter) Britton, Sterns & Poggenb. 
A. gyrans Ashe 
*A. tenuispatheus (Nash) Nash 
*A. ternarius Michx. 
*A. virginicus L. 
*Cenchrus carolinianus Hitch. 
C. echinatus L. 
*Dichanthelium scabriusculum (Elliott) Gould & 
C. A. Clark 
*Melinis repens (Willd.) Zizka 
Panicum repens L. 
*Paspalum notatum Fliiggé var. saure Parodi 
P. urvillei Steud. 
Schizachyrium scoparium (Michx.) Nash 
*Sorghastrum domingensis (Trin.) Kunth 
*S. nutans (L.) Nash 


State 


FL GA NC sc 


s) —— — — 
17 98 49 83 
| ata a =e 
_ = 
bass 2 i 8 a 
= _ Ls 2 
7 = = 2 
| a. ag a 
D) a — —— 
1 Ade a) Ao 
| ee a, He 
| st ane pe 
3 Sh mal Ase 
I got Ete oe 
1 | — oa 
—~ os —- 2 
12 3 2 22 
— iT — — 
1 oi) io vk 
6 = = — 
| ee a so 
— — — 4 
ase - — | 
ist = | 3 
A a Du fas 
2 


Adults of B. minutus were collected from 
34 species of Poaceae in the subfamilies 
Aristoideae, Chloridoideae, and Panicoi- 
deae. Nymphs were found on 21 species 
representing each of the three subfamilies. 
Chloridoid (79.8%) and panicoid (19.2%) 
grasses dominated the collection records. 
When records from the exotic E. 
are excluded, most records were from Pan- 
icoideae (72.0%), followed by Chloridoi- 


curvula 


deae (24.3%) and Aristoideae (3.7%). If 
only grasses on which nymphs were ob- 
served are considered, chloridoids account- 
ed for 81.3% of total records; panicoids, 
17.9%; and aristoids, 0.8%. With E. curvula 
omitted, panicoids (78.2%) dominated the 
nymphal hosts, followed by chloridoids 
(18.4%) and aristoids (3.4%). 
Seasonality.—In the Southeast, adults of 
B. minutus (>95% brachypters) were found 


340 


Table 2. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Number of collections of Blissus minutus from grasses in the southwestern United States: Oklahoma 


and Texas. Species marked with an asterisk are ones from which nymphs were collected. 


a ae) 


Grass 


State 


De eee 


Chloridoideae 
* Eragrostis curvula (Schrad.) Nees 
Muhlenbergia lindheimeri Hitchce. 
M. reverchonii Vasey & Scribn. 
Panicoideae 
*Bothriochloa ischaemum (L.) Keng 
*Schizachyrium scoparium (Michx.) Nash 


D — 


a 


in crowns of weeping lovegrass from mid- 
November to mid-March. First through fifth 
instars were observed in early December 
and third through fifth instars in late De- 
cember (ca. 5% of population). That second 
through fifth instars (mostly III-IV) were 
found in late February and early March 
suggests that nymphs survive the winter in 
southeastern states. The irregularity of col- 
lections and observations, coupled with the 
blissid’s overwintering as adults and 
nymphs of several instars, made it difficult 
to determine the appearance of first instars 
of the first generation and delimit the num- 
ber of annual generations. Late-developing 
cohorts of the first generation appeared to 
overlap with nymphs of the second gener- 
ation, which blurred the distinction between 
all succeeding generations. 

Adults and a mating pair were observed 
by late March; mating pairs continued to be 
beaten from weeping lovegrass from April 
to late August. First and second instars out- 
numbered late instars (presumably over- 
wintered) at the Richland County sample 
site on 1 April 2000, suggesting the begin- 
ning of a first generation. Early instars also 
were observed in late May, July, and Oc- 
tober to December. This multivoltine blissid 
appears to complete at least three genera- 
tions a year in the Southeast. 

In the Southwest, third through fifth in- 
stars (presumably overwintered), brachyp- 
ters, and an occasional macropter were pre- 
sent in late April 2000 and 2002. Mating 


pairs were observed in Wheeler County, 
Texas, on 26 April 2000. Collections in 
mid-May 2001, late May 2003, and early 
May 2004 consisted of brachypters (three 
macropters were seen). A mating pair was 
found on 25 May 2001 in Terry County, 
Texas, and another on 29 May 2003 in 
Blaine County, Oklahoma. 


Toonglasa umbrata (Distant) 


Toonglasa umbrata was described in I[s- 
chnodemus Fieber from Guatemala by Dis- 
tant (1893); it was transferred to Toonglasa 
Distant by Slater and Brailovsky (1983), 
who proposed J. macer Van Duzee, de- 
scribed from Arizona, as a synonym. Ad- 
ditional U.S. records are lowa, Kansas, Ne- 
braska, New Mexico, Oregon, and Texas 
(Slater and Brailovsky 1983). The species 
ranges through Mexico and Central Amer- 
ica to at least Brazil in South America (Slat- 
er and Brailovsky 1990). The grasses An- 
dropogon glomeratus (Walter) Britton, 
Sterns & Poggenb. and Bothriochloa blad- 
hii (Retz.) S. T. Blake (= B. intermedia (R. 
Br.) A. Camus) are hosts of 7. umbrata in 
Panama (Slater 1976); a host in Mexico is 
Hyparrhenia rufa (Nees) Stapf (Slater and 
Brailovsky 1983). The fifth instar has been 
described, based on material from Arizona 
(Slater and Brailovsky 1983), but host 
grasses have not been reported for the Unit- 
ed States. 

Distribution (Fig. 2).—Toonglasa um- 
brata was found in Arizona (1 county), 


VOLUME 107, NUMBER 2 


B Record from author’s fieldwork 
Record from literature 


BS Record from author’s fieldwork and literature 


Fig. 2. 


New Mexico (6), Oklahoma (7), and Texas 
(11); Oklahoma is a new state record. It was 
collected in western Oklahoma and Texas 
in many of the counties and the same ecore- 
gions as B. minutus. The two blissids were 
syntopic at ten sites in Oklahoma (Beck- 
ham, Blaine, Canadian, Dewey, Jackson, 
Major, and Roger Mills counties) and nine 
sites in Texas (Cottle, Dawson, Gaines, 
Gray, Hardeman, Terry, and Wheeler coun- 
ties). Toonglasa umbrata was the only blis- 
sid collected from crowns of bunchgrasses 
in southeastern New Mexico and _ south- 
eastern Arizona. 

Host plants—Nymphs and adults of 7. 
umbrata were found on one pooid, three 


341 


Known distribution (by county) of Toonglasa umbrata based on personal collecting and the literature. 


panicoid, and five chloridoid grasses (Table 
3), with records from the exotic chloridoid 
E. curvula dominating (80.0%). Another 
non-native host was a panicoid, the Old 
World bluestem Bothriochloa ischaemum 
(L.) Keng. Nymphs were found on seven 
native grasses, including the chloridoid 
sideoats grama (Bouteloua curtipendula 
(Michx.) Torr.) and big sacaton (Sporobolus 
wrightit Munroe ex Scribn.), and the pani- 
coids cane bluestem (Bothriochloa barbi- 
nodis (Lag.) Herter) and little bluestem 
(Schizachyrium scoparium (Michx.) Nash). 

Seasonality.—Late-April collections in 
2000 and 2002 consisted of apparently 
overwintered third through fifth instars 


342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 3. 


Number of collections of Toonglasa umbrata from grasses in the southwestern United States: Ari- 


zona, New Mexico, Oklahoma, and Texas. Nymphs and adults were collected from all species. 


a 


State 
Grass AZ NM OK TX 

Chloridoideae 

Bouteloua curtipendula (Michx.) Torr. 2 2 — — 

B. gracilis (Willd. ex Kunth) Lag. ex Griffiths ao | — — 

Eragrostis curvula (Schrad.) Nees 3 16 13 20 

E. intermedia A. S. Hitch. | -— — — 

Sporobolus wrightii Munro ex Scribn. —~ | — — 
Panicoideae 

Bothriochloa barbinodis (Lag.) Herter _ — 

B. ischaemum (L.) Keng — — — 

Schizachyrium scoparium (Michx.) Nash — — 1 —_— 
Pooideae 

Festuca arizonica Vasey —- ] — — 
(mostly IV-V) in western Oklahoma and DISCUSSION 


fourth and fifth instars in western Texas. 
Late instars also were found in late April in 
southeastern New Mexico except a site in 
Roosevelt County where two adults were 
present with late instars. Late instars only 
were observed in late May 2001 at all sites 
in western Oklahoma except one in Dewey 
County where teneral adults (<10) were 
present. Late-May collections in southeast- 
ern New Mexico consisted of late instars 
only or nymphs plus one or two adults, ex- 
cept the Roosevelt County site noted above, 
where adults outnumbered late instars. In 
late May 2003, adults outnumbered nymphs 
in western Oklahoma; a mating pair was 
observed on 29 May in Canadian County. 
Adults were found in southeastern New 
Mexico in late May; only fifth instars were 
observed in early June in southeastern Ar- 
izona (Santa Cruz County). In early May 
2004, only late instars were found in Daw- 
son County, Texas, whereas adults (mostly 
teneral) outnumbered fifth instars in Gaines 
County. Second through fourth instars were 
found in southeastern New Mexico in early 
May; mid-May collections in southeastern 
Arizona consisted of third and fourth in- 
stars. 


Because B. minutus has been known only 
from Florida, it probably is among lygaeoid 
species (10, not enumerated) regarded as 
precinctive (sensu Frank and McCoy 1990) 
in that state (Baranowski 1995). Its south- 
eastern range now is extended to portions 
of Alabama, Georgia, North Carolina, and 
South Carolina. The blissid can be consid- 
ered a characteristic insect of the fall-line 
Sandhills of the Carolinas and Georgia. 
Southeastern records outside the Sandhills 
also are mainly from the Coastal Plain. The 
relatively few records from the Piedmont 
are near the fail line except for one from 
Atlanta, Georgia. 

In the Southwest, B. minutus was found 
on weeping lovegrass in parts of western 
Oklahoma and western Texas, particularly 
where soils are sandy. I collected only an 
occasional adult in the Edwards Plateau of 
Texas. It was syntopic with another blissid, 
T. umbrata, at numerous sites in Oklahoma 
and Texas but was not found in the west- 
ernmost counties of Texas (except a collec- 
tion from Gaines County) or in southeastern 
Arizona and southeastern New Mexico. 
Toonglasa umbrata was the only blissid 
collected from E. curvula in these more arid 


VOLUME 107, NUMBER 2 


regions. I also did not find B. minutus on 
E. curvula in southern Arkansas, eastern 
Oklahoma, and eastern Texas. Additional 
fieldwork is needed to determine whether 
populations in the Atlantic and Gulf Coastal 
Plain are disjunct from those of the south- 
ern Great Plains. Blissus minutus might 
eventually be found in the relatively little- 
studied blackland prairies scattered in Lou- 
isiana and occurring more extensively in 
Alabama and Mississippi (Peacock and 
Schauwecker 2003). Certain other grassland 
insects are known from the Great Plains, 
the Black Belt of Alabama and Mississippi, 
and the Gulf and Atlantic Coastal Plain, 
Whereas others show a disjunct Great 
Plains-Atlantic/Gulf Coastal Plain distribu- 
tion (Brown 2003). 

Conservation and forage plantings of 
weeping lovegrass ordinarily are estab- 
lished by seeding (Crider 1945, Ahring 
1970, Voigt and Sharp 1995). Therefore, 
movement of B. minutus with commerce 
seems unlikely. Because weeping lovegrass 
can be transplanted as seedlings or crowns 
divided from older plants (Crider 1945), the 
possibility that the blissid might have be- 
come established in new areas via move- 
ment of vegetative material of E. curvula 
(or other host grasses) cannot be dismissed. 
Human-assisted spread of B. occiduus Bar- 
ber (Baxendale et al. 1999) and B. insularis 
Barber (Sweet 2000) with turfgrasses has 
been suggested. 

In analyzing host-plant relationships of 
blissids, Slater (1976) predicted that era- 
grostoid grasses “‘will prove to be widely 
used as host plants.”” Eragrostis curvula in 
South Africa, where it is native, serves as 
a host of the blissid Atrademus capeneri 
(Slater) and probably also Capodemus her- 
bosus Slater and Sweet (Slater and Wilcox 
1973, Slater 1976). After being introduced 
into the southern United States, this plant 
has been colonized by two New World blis- 
sids, B. minutus and T. umbrata. The col- 
onization of non-native hosts is common in 
the Blissidae (Slater 1976, Sweet 2000). 

Slater and Wilcox (1973) suggested that 


blissids continually test new plants, with 
most colonizations of novel hosts being un- 
successful. In other cases, they noted that a 
blissid develops on a new host but less ef- 
fectively than on its original hosts; occa- 
sionally, the new host becomes a primary 
host, as weeping lovegrass now appears to 
be for B. minutus. New plants can become 
““better”’ hosts than a blissid’s original hosts 
(Slater and Wilcox 1973). Whether nymph- 
al growth and development of B. minutus 
are increased on E. curvula relative to its 
native hosts remains to be tested. 

A South African blissid, Atrademus mar- 
itimus Slater and Wilcox, has been collected 
only from a non-native host (Slater 1976). 
When I discovered that B. minutus is com- 
mon on the introduced EF. curvula, 1 em- 
phasized collections from other grasses 
with extensive crowns in an attempt to de- 
termine its native hosts. Certain of these 
grasses, such as Andropogon species and 
Muhlenbergia capillaris, proved to be host 
plants, whereas others, such as Aristida 
beyrichiana, appeared not to be used as 
hosts. Although blissids often are host.re- 
stricted (Slater 1976), B. minutus, having 
been collected from 34 grasses (nymphs 
from 21 species) in the subfamilies Aris- 
toideae, Chloridoideae, and Panicoideae, 
can be considered a grass generalist (sensu 
Whitcomb et al. 1988). Toonglasa umbrata, 
whose nymphs were found on nine grasses 
in the Chloridoideae, Panicoideae, and 
Pooideae, might also be a grass generalist. 

Wilson and Wheeler (in press) discussed 
factors—architectural complexity and phy- 
tochemistry—that might have allowed cer- 
tain rarely collected native planthoppers to 
colonize the exotic FE. curvula. In the case 
of B. minutus, which develops in the 
crowns of its hosts, the large, dense crowns 
of E. curvula might allow larger popula- 
tions to develop than are typical on most of 
the bug’s native hosts. The large densities 
that were observed on weeping lovegrass 
might reflect the lack of resistance of this 
grass to a new herbivore. Small densities of 
pestiferous blissids, such as B. leucopterus 


344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Say, often are found on native grasses, 
whereas much larger, damaging numbers 
develop on introduced grasses (Sweet 
2000). The sandy soils to which E. curvula 
is best adapted (Crider 1945, Voigt and 
Sharp 1995) also might have facilitated the 
use of a novel host by B. minutus, which 
feeds at the base of host grasses. Blissus 
minutus is found near the soil substrate, as 
has been reported for B. leucopterus hirtus 
Montandon (Sweet 2000). 


ACKNOWLEDGMENTS 


Personnel of the USDA’s Natural Re- 
sources Conservation Service were extraor- 
dinarily helpful in providing information on 
weeping lovegrass in the Southwest: 
Oklahoma (Jim Williams), New Mexico 
(Dan Abercrombie, Wallace Cox, Joe Gib- 
son), and Texas (Fred Allison, Donald Gib- 
bons, Reggie Quiett); Donald Gibbons 
(Brownfield, TX) and Joe Gibson (Clovis, 
NM) also accompanied me in the field. I 
thank Robert Whitcomb (formerly with 
USDA, ARS, Beltsville, MD) for suggest- 
ing that I contact the NRCS and accompa- 
nying me in the field at the Audubon Re- 
search Ranch in 2004. I also am grateful to 
Paul Voigt (USDA, ARS, Beaver, WV), 
Chet Dewalt (USDA, ARS, Woodward, 
OK), and Chuck Grimes (Grasslander, Hen- 
nessey, OK) for useful information on 
weeping lovegrass, Thomas Henry (USDA, 
ARS, Systematic Entomology Laboratory, 
Washington, DC) for identifying blissids 
and accompanying me in the field in 
Oklahoma in 1999, Patrick McMillan (De- 
partment of Biological Sciences, Clemson 
University) for identifying grasses, John 
Townsend (formerly with Clemson’s De- 
partment of Biological Sciences) for obtain- 
ing herbarium records of Eragrostis curvula 
in the Southwest, David Boyd (USDA, 
ARS, Poplarville, MS) for providing the 
maps, Linda Kennedy (National Audubon 
Society Appleton-Whittell Research Ranch, 
Elgin, AZ) for issuing a collection permit 
and making my visits to the Research 
Ranch enjoyable and productive, Richard 


Brown (Department of Entomology and 
Plant Pathology, Mississippi State Univer- 
sity) for sharing his knowledge of insect 
distribution patterns, and Peter Adler (De- 
partment of Entomology, Soils, and Plant 
Sciences, Clemson University) for com- 
ments that improved the manuscript. 


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Voigt, PB. W. and W. C. Sharp. 1995. Grasses of the 
Plains and Southwest, pp. 395-408. /n Barnes, R. 
F, D. A. Miller, and C. J. Nelson, eds. Forages, 
Vol. 1. An Introduction to Grassland Agriculture, 
Sth ed. Iowa State University Press, Ames. 

Wheeler, A. G., Jr. 1999. Oncozygia clavicornis Stal 
and Allopodops mississippiensis Harris and John- 
ston: association of rarely collected Nearctic turtle 
bugs (Heteroptera: Pentatomidae: Podopinae) with 
an introduced African grass. Proceedings of the 
Entomological Society of Washington 101: 714— 
WAM 

. 2003. Rediscovery of Ligyrocoris slossoni 
(Hemiptera: Lygaeoidea: Rhyparochromidae), a 
rarely collected seed bug considered precinctive 
in Florida. Florida Entomologist 86: 219-221. 

Whitcomb, R. FE, A. L. Hicks, D. E. Lynn, H. D. 
Blocker, and J. P. Kramer. 1988. Host specificity: 
a major mechanism enhancing insect diversity in 
grasslands, Paper 11.06. Jn Davis, A. and G. Stan- 
ford, eds. The Prairie: Roots of our Culture; Foun- 
dation of our Economy. Proceedings of the Tenth 
North American Prairie Conference of Texas 
Woman’s University, Denton, Texas, June 22—26, 
1986. Native Prairie Association of Texas, Dallas. 

Wilson, S. W. and A. G. Wheeler, Jr. In press. An 
African grass recruits rarely collected native plant- 
hoppers (Hemiptera: Fulgoroidea: Dictyopharidae, 
Fulgoridae): Hidden diversity in crowns of Era- 
grostis curvula (Poaceae). Journal of the New 
York Entomological Society. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 346-356 


REVISION OF THE SOUTHEASTERN ASIAN SAWFLY GENUS BUSARBIA 
CAMERON (HYMENOPTERA: TENTHREDINIDAE) 


MALKIAT S. SAINI AND DAVID R. SMITH 


(MSS) Department of Zoology, Punjabi University, Patiala 147002, India (e-mail: 
saini20@hotmail.com); (DRS) Systematic Entomology Laboratory, PSI, Agricultural Re- 
search Service, U.S. Department of Agriculture, % National Museum of Natural History, 
Smithsonian Institution, RO. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. 
(e-mail: dsmith @sel.barc.usda.gov) 


Abstract.—Six species of Busarbia Cameron, a genus of the subfamily Selandriinae 
restricted to southeastern Asia, are known: B. viridipes Cameron, B. shinoharai, n. sp., 
B. okutanii, n. sp., and B. santokhi, n. sp., from India; B. formosana (Rohwer) from 
Taiwan, Myanmar, and India; and B. isshikii (Takeuchi) from Taiwan. Busarbia formosana 
is recorded from India for the first time. An identification key is provided, and each species 


is described and illustrated. 


Key Words: 


Busarbia was described by Cameron 
(1899) from Assam (now Megahalaya), 
Khasia Hills, India, and included a single 
species, B. viridipes Cameron. In 1902, 
Cameron added B. albipes from Shimla 
(Himalchal Pradesh). Malaise (1944) re- 
viewed this genus from Myanmar and India 
and also included Busarbia formosana 
(Rohwer 1916) and Busarbia isshikii (Yak- 
euchi 1928). Since Malaise (1944), the ge- 
nus has been only mentioned in the litera- 
ture. Togashi (1990, figs. 15—19) gave some 
additional records for B. isshikii from Tai- 
wan, briefly described the species, and il- 
lustrated the wings, top and front view of 
the head, sheath, and female lancet and 
Malaise (1963) included the genus in his 
key to world genera of Selandriinae. 

A number of specimens have been col- 
lected from India during surveys by the se- 
nior author. We treat six species, three of 
which are new, and one which is recorded 
from India for the first time. Food plants 
are not known, but most related Selandri- 


sawflies, Symphyta, Selandriinae, ferns 


inae feed on ferns and Malaise (1944) spec- 
ulated that *“*. . . without doubt the larvae 
feed on ferns, as the adults, like those of 
the genus Nesoselandria Rohwer, are al- 
ways found on or near ferns.”” Knowledge 
of fern-feeding sawflies is important be- 
cause of the interest in biological control of 
invasive ferns, especially the Old World 
climbing fern of the genus Lygodium in the 
United States. 

Terminology follows Malaise (1963) and 
Ross (1937, 1945). All holotypes are de- 
posited in the National Pusa Collections, In- 
dian Agricultural Research Institute (IARI), 
New Delhi. 

Abbreviations used are as follows: EL = 
eye length; [ATS = inner apical tibial spur; 
ICD = intercenchral distance; IDMO = in- 
terocular distance at level of median ocel- 
lus; ITD = intertegular distance; LID = 
lower interocular distance; MB = hind bas- 
itarsus; OATS = outer apical tibial spur; 
OCL = ocellar-occipital distance; OOL = 
ocular-ocellar distance; POL = postocellar 
distance. 


VOLUME 107, NUMBER 2 


Busarbia Cameron 


Busarbia Cameron 1899: 37. Type species: 
Busarbia viridipes Cameron, by mono- 
typy.—Malaise 1944: 17, 19 (genus de- 
fined, key to species).—Malaise 1963: 
174 (Gn key). 


Description.—Antenna long, slender; fi- 
liform; third antennal segment subequal in 
length to or slightly shorter than 4th seg- 
ment; pedicel twice as long as its breadth 
at apex. Inner margins of eyes straight and 
subparallel, slightly more converging below 
in males. Malar space distinct. Clypeus with 
indistinct and blunt carina along emargin- 
ated anterior margin; this margin sometimes 
with a narrow, depressed and acute edge, if 
acute edge wanting, margin appears almost 
incrassate. Mandibles roundly bent, much 
less than at a right angle; without a deep 
pit on outer surface at base. Frontal area 
surrounded by acute carina and lateral ca- 
rina extending from it almost to each eye 
(Figs. 1-4). Epicnemiun distinct, separated 
from mesepisternum by deep furrow. Fore- 
wing with veins M and Im-cu divergent, 
not parallel; with 2 radial and 4 cubital 
cells; anal cell without crossvein. Hind 
wing with 2 closed middle cells; anal cell 
petiolate. Supara-antennal pits poorly de- 
veloped and supra-antennal tubercles ab- 
sent. Tarsal claws with long subapical tooth, 
shorter than outer tooth; basal lobe absent. 
Color black, white with a greenish tinge 
when alive (whitish yellow in dried speci- 
mens) on clypeus, labrum, mandibles, palpi, 
scape, upper pronotal margin, tegula, post- 
spiracular sclerite, and triangular spot on 
posterior mesepisternum; legs except apices 
of tarsi, and most of underside of abdomen 
except for black apical 3 or 4 segments. 

Remarks.—Busarbia, Abusarbia Mal- 
aise, and Busarbidea Rohwer form a group 
of genera in the Selandriiae that share the 
sharp carinae on the frons (as in Figs. I— 
4). Abusarbia is distinguished by the tarsal 
claws which have a single tooth and a large, 
acute basal lobe, and Busarbidea by the 


347 


presence of an anal crossvein in the fore- 
wing. 

Other genera with which Busarbia may 
be confused are Anapeptamena Konow, Ar- 
busia Malaise, and Apeptamena Malaise; 
however, these genera lack the sharp ridges 
on the frontal area and have the inner mar- 
gins of the eyes distinctly converging be- 
low. Busarbia keys to near these two genera 
in Malaise (1963). For a discussion of An- 
apeptamena, see Saini et al. (2003). 

Malaise (1944) gave a key to three spe- 
cies of Busarbia, B. isshikii, B. viridipes 
Cameron, and B. formosana. He considered 
B. albipes Cameron as a color form of B. 
viridipes. These three species, plus the three 
new ones described here, are the only spe- 
cies of Busarbia that we are aware of. 


KEY TO SPECIES 


1. Antennal segment 3 subequal in length to 4; 
lateral margins of mesoprescutum, supracly- 
peal area to around median fovea, and spot on 
inner orbits whitish (from Takeuchi 1928 and 
Malaise 1944) isshikii (Takeuchi) 

— Antennal segment 3 shorter than 4 (except 
equal only in shinoharai); mesoprescutum, su-. 


praclypeal area, and inner orbits black ..... 22 
2. Scape black; head rugose with dense confluent 
PUNCHUTES |S pears eia es See See atone shinoharai, n. sp. 


— Scape yellow or fuscus; head smooth and al- 
most impunctate, punctures scattered and not 


confiuents (Gest —4) enn enennenene <= 3 
3. Hind margin of head distinctly carinate (Figs. 

1—4); medial carina of postcellar area almost 

reaching hind margin of head ........... 4 


— Hind margin of head rounded, not carinate; 
medial carina of postocellar area, if present, 
confined only to anterior half............ 5 

4. Malar space more than half diameter of front 
ocellus; tarsal claw with long inner tooth, near- 
ly as long as outer tooth (Fig. 14); lower in- 
terocular distance greater than eye length, LID: 
IDMO:EL = 1.0:1.1:0.8; postocellar area 
slightly broader than long as 5:4; hind ocelli 
closer together and farther from hind margin of 
head, OOL:POL:OCL = 1.0:0.3:0.8 (Figs. 1— 

Di) ste ky Aects oben ckchel iets formosana (Rohwer) 

— Malar space less than half diameter of front 
ocellus; tarsal claw with small inner tooth, 

much shorter than outer tooth (Fig. 11); lower 

interocular distance subequal to eye length, 

LID:IDMO:EL = 

distinctly broader than long as 7:4; hind ocelli 


1.0:1.2:1.0; postocellar area 


Nn 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


uty 


z 
Bat 
s 


9) 


Figs. 1-2. Busarbia formosana head. 1, Dorsal view. 2, Front view. Arrow indicates occipital carina. 


malar space 0.6 diameter of front ocellus 


farther apart and closer to hind margin of head, 
OOE:POL:OCE —aO:0F4:0OuGisstiSsS4) ey eee SON oes ie ees Se santokhi, n. sp. 
— Abdomen black; OOL:POL:O0CL = 1.0:0.3: 


okutanii, n. sp. 
1.0; malar space 0.4 diameter of front ocellus 
viridipes Cameron 


Abdomen pale yellow to light brownish below 
excepiatiapex:| OOP OL: OC alk 0:04 515) nenere enya eee 


349 


VOLUME 107, NUMBER 2 


al carina. 


« 


Arrow indicates occipit 


3, Dorsal view. 4, Front view. 


bia okutanii head. 


Figs. 3-4. Busar 


350 


Buarbia formosana (Rohwer) 


(Figs. 1, 2, 9, 14, 18) 


Anapeptamena formosana Rohwer 1916: 
100—101.—Forsius 1932: 53 Gn key).— 
Takeuchi 1941: 270 (in key; Taiwan re- 
cords).—Oehlke and Wudowenz 1984: 
381 (types). 

Busarbia formosana: Malaise 1944: 20. 


Female.— Color: Black with following 
pale whitish: clypeus, labrum, mandible ex- 
cept apex, scape, supraclypeal area contin- 
uous with spots on supra-antennal tuber- 
cles, posterodorsal angle of pronotum, te- 
gula, postspiracular sclerite, subtriangular 
spot on ventral half of mesepisternum, ster- 
na 1—6, and legs except somewhat rufous 
apical tarsal segments. Wings hyaline; cos- 
ta, stigma, and veins light brown to black. 

Structure: Average length, 5.5 mm. An- 
tennal length 2.9 head width; scape slight- 
ly longer than broad and almost equal to 
pedicel; pedicel length 2 apical width; 
segment 3 shorter than 4, as 5:6. Clypeus 
subconvex, roundly emarginated for one- 
quarter its medial length (Fig. 9). Labrum 
broader than long as 2:1, with rounded an- 
terior margin. Malar space 0.6 diameter 
of front ocellus. OOL:POL:OCL = 1.0:0.3: 
0.8. Supra-antennal pits only indicated. Su- 
pra-antennal tubercles slightly raised and 
sloping posteriorly. Postocellar furrow dis- 
tinct, replaced with carina; circumocellar 
furrow indistinct. Lateral furrows deep, dis- 
tinct, diverging backwards, and almost 
reaching hind margin of head. Postocellar 
area distinctly broader than long, as 5:4, 
hind margin of head distinctly carinate and 
postocellar area with medial carina reaching 
hind margin of head. Lower interocular dis- 
tance greater than eye length, LID:IDMO: 
EL = 1.0:1.1:0:9. ITD:ACD = 1:0:0.:3: Me- 
sepisternum obtusely raised without carina 
or acute apex. Subapical tooth of tarsal 
claws (Fig. 14) slightly shorter than apical 
tooth. Hind basitarsus longer than following 
tarsal segments combined, as 6:5. IATS: 
MB:OATS = 1.0:3.0:0.9. Lancet with 5 ser- 
rulae (Fig. 18). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Structure and pubescence: Head smooth, 
shining; thorax shining, almost impunctate; 
punctures on mesoscutellum anterior to its 
appendage confined to a single row. Ab- 
domen cross striated. Brownish-yellow pu- 
bescence almost negligible. 

Male.—Length, 4.0 mm. Similar to fe- 
male except for black sternites (based on 
Rohwer 1916). 

Types.—Described from 2 females (one 
type) and 1 male collected Oct. 1911, Suis- 
haryo, Formosa. Holotype and male allo- 
type in the Deutsches Entomologisches In- 
stitut, Miincheberg; female paratype in the 
National Museum of Natural History, 
Smithsonian Institution, Washington, DC 
(USNM) (Oehlke and Wudowenz 1984). 

Specimens examined.—INDIA: Aruna- 
chal Pradesh, Bomdila, 2,800 m, 26.5.1993, 
M.S. Saini (1 2); Himachal Pradesh, Ka- 
latop, 2,400 m, 25—28.7.1982, M. S. Saini 
G3 2), CE-O81GS2Y C3 2), ZIS81VSs2 
2): Uttar Pradesh, Hanuman Chatti, 2,300 
m, 5.6.1983, M.S. Saini (6 2); Kilbury, 
DEMS ii, ZLOGN9IB, IW S. Sana @ Lye 
Mandal, 2,200 m, 21.6.1985, M.S. Saini (2 
2); West Bengal, Darjeeling, 2,286 m, 
26.5.1989, M. S. Saini (1 2), 9.5.1983 C1 
2). TAIWAN: Suisharyo, Formosa, X 11, 
Sauter (Anapeptamena formosana Rohwer, 
paratype 2, USNM). 

Distribution.—Myanmar (Kambaiti at 
2,000 m; Taunggyi at 1,500 m) (Malaise 
1944); Taiwan; India (Arunachal Pradesh, 
Himachal Pradesh, Uttar Pradesh, West 
Bengal). 

Variation.—Tergites 2—5 are sometimes 
fulvous above, and the lateral sides of an- 
tennal segments 6—9 may be pale. 

Remarks.—The carinate hind margin of 
the head and coloration places this species 
close to B. okutanii, but it can be separated 
by characters given in the key and as dis- 
cussed under B. okutanii. 


Busarbia isshikii (Takeuchi) 


Anapeptamena isshikii Takeuchi 1928: 
39.—Takeuchi 1941: 270 Gn key). 


VOLUME 107, NUMBER 2 


Figs. 5—14. Clypeus and labrum and tarsal claws. 
5—9, Clypeus and labrum. 5, Busarbia shinoharai. 6, 
B. okutanii. 7, B. santokhi. 8, B. viridipes. 9, B. for- 
mosana. 10—14, Tarsal claws. 10, B. shinoharai. 11, 
B. okutanii. 12, B. santokhi. 13, B. viridipes. 14, B. 
formosana. 


Busarbia isshikii. Malaise 1944: 
gashi 1990: 180, figs. 15—19. 


Loe =To= 


Remarks.—We have not seen specimens 
of this species. According to the original 
description and Malaise (1944), it is sepa- 
rated from the other species of Busarbia by 
antennal segment 3 subequal in length to 4 
and by the whitish markings on the lateral 
margins of the mesoprescutum, supracly- 
peal area to around the median fovea, and 
spot on the inner orbits. Other species of 
Busarbia have the third antennal segment 
shorter than the fourth (execpt B. shinohar- 
at) and the above parts black. 

Takeuchi described this species from one 
female from “‘Shinsuiye, Formosa.” Toga- 
shi (1990) gave a few additional characters, 
new records from Taiwan, and illustrated 
the dorsal and frontal views of the head, 
wings, sawsheath, and lancet. 


Busarbia okutanii Saini and Smith, 
new species 
(Figs. 3—4, 6, 11, 19-20, 23) 


Female.—Color: Body black, with clyp- 
eus, labrum, dorsal pronotal margin, tegula, 
postspiracular sclerite, triangular spot on 


351 


posterior portion of mesopleuron, sternites 
1—5, and legs except apices of tarsi whitish; 
scape fuscous. Wings hyaline; costa, stig- 
ma, and veins black. 

Structure: Average length, 6.5 mm. An- 
tennal length 3 head width; scape longer 
than pedicel as 4:3; scape and pedicel 
lengths each 2X their apical widths; seg- 
ment 3 shorter than 4, as 4:5. Clypeus sub- 
convex, its extreme anterior margin arcu- 
ately emarginated up to Y; its medial length, 
with blunt lateral lobes (Fig. 6). Labrum 
broader than long, as 2:1, with rounded an- 
terior margin. Malar space less than half di- 
ameter of front ocellus. OOL:POL:OCL = 
1.0:0.4:0.9. Supra-antennal pits distinct and 
connected to antennal sockets by a very fine 
furrow. Supra-antennal tubercles indicated 
and continuous with similar frontal ridges. 
Circum- and interocellar furrows distinct: 
postocellar furrow indistinct. Lateral fur- 
rows deep, distinct, diverging backwards 
and reaching posterior margin of head. 
Postocellar area distinctly broader than 
long, as 7:4; hind margin of head distinctly 
carinate, and middle longitudinal carina of 
postocellar area reaching hind margin of 
head. Inner margins of eyes subparallel in 
middle. Lower interocular distance sube- 
qual to eye length, LID:IDMO:EL = 1.0: 
1.2:1.0. ITD:ICD = 1.0:0.3. Mesepisternum 
obtusely raised without carina or acute 
apex. Subapical tooth of tarsal claw (Fig 
11) shorter than apical tooth. Hind basitar- 
sis longer than following tarsal joints com- 
bined, as 5:4. IATS:MB:OATS = 1.0:3.3: 
0.9. Lancet with 5 serrulae (Fig. 19). 

Sculpture and pubescence: Head and 
thorax smooth, shining and impunctate ex- 
cept posterior slope of mesoscutellum with 
row of distinct punctures. Abdomen im- 
punctate, subshining. Head and thorax covy- 
ered with minute golden-yellow pubes- 
cence. 

Male.—Average length, 4.0 mm. Similar 
to female except malar space 0.3 diameter 
of front ocellus; antennal segment 3 shorter 
than 4, as 6:7, and scape and pedicel pale 
whitish. Genitalia: Penis valve (Fig. 20) 


352 


with valviceps narrow at base and broad- 
ening to truncate apex, with row of stout 
spines on ventral margin; genital capsule 
(Fig. 23) similar to B. viridipes. 

Holotype.—Female, Arunachal Pradesh, 
Bomdila, 2,800 m, 31.5.1989, M.S. Saini. 

Paratypes.—Same data as holotype (2 
2); Nagaland, Satakha, 1,600 m, 15.5.1993, 
M.S. Saini (1 6) (1 2, 1 ¢& at Punjabi Uni- 
versity; 1 2 at USNM). 

Distribution —India (Arunachal Pradesh; 
Nagaland). 

Etymology.—The species name is in 
honor of Dr. Teichi Okutani of Japan, a dis- 
tinguished worker in the field of sawfly tax- 
onomy and biology. 

Remarks.—No variation was observed. 
The carinate hind margin of the head is 
shared with B. formosana. Busarbia oku- 
tanii is separated from B. formosana by the 
short malar space less than half the diam- 
eter of the front ocellus, lower interocular 
distance subequal to the eye length, small 
inner tooth of the tarsal claw, and broad 
postocellar area. 


Busarbia santokhi Saini and Smith, 
new species 
Gigs. 75 125 7, Zl, 245) 


Female.—Color: Black, with following 
pale whitish: clypeus, labrum, mandible ex- 
cept apex, scape, upper pronotal margin, te- 
gula, postspiracular sclerite, spot on poste- 
rior portion of mesopleuron, sternites 1—6, 
and legs except infuscated tarsal claws. 
Wings hyaline; costa, stigma, and veins 
black. 

Structure: Average length, 6.0 mm. An- 
tennal length 2.5 head width; scape and 
pedicel almost equal in length, each shorter 
than 2x their apical widths; segment 3 
shorter than 4, as 8:9. Clypeus with anterior 
margin subconvex, roundly emarginate for 
Y% its medial length, with blunt lateral lobes 
(Fig. 7). Labrum broader than long as 5:2, 
with rounded anterior margin. Malar space 
0.6 diameter of front ocellus. OOL:POL: 
OCL = 1.0:0.4:1.1. Supra-antennal pits in- 
distinct. Supra-antennal tubercles only in- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


dicated and continuous with similar frontal 
ridges. Circum-, inter- and postocellar fur- 
rows distinct. Lateral furrows deep, distinct, 
diverging backwards, and ending just be- 
fore hind margin of head. Postocellar area 
subconvex, as long as broad at its maxi- 
mum width; hind margin of head rounded, 
not carinate. Lower interocular distance 
greater than eye length, LID:IDMO:EL = 
1.0:1.1:0.9. ITD:ICD = 1.0:0.3. Subapical 
tooth of tarsal claws shorter than apical 
tooth (Fig. 12). Hind basitarsus longer than 
following tarsal joints combined, as 7:6. 
IATS:MB:OATS = 1.0:3.0:0.9. Lancet with 
5 serrulae (Fig. 17). 

Sculpture and pubescence: Head and 
thorax smooth, shining, impunctate, except 
posterior slope of mesoscutellum with a 
row of distinct isolated punctures. Abdo- 
men impunctate, subshining. Body covered 
with golden-yellow pubescence. 

Male.—Average length, 5.0 mm. Similar 
to female except antennal length 3.0 head 
width, pedicel and lateral sides of third seg- 
ment pale whitish; tergites 2—5 light brown- 
ish; all sternites black; ITD:ICD = 1.0:0.4. 
Genitalia: Penis valve (Fig. 21) with val- 
viceps somewhat rectangular and of equal 
width, apex truncate, without spines on 
ventral margin; genital capsule (Fig. 24), 
with parapenis narrow and slender, separat- 
ed from gonostipes by circular notch. 

Holotype.—Female, West Bengal, Dar- 
jeclines92*3 00m =9'5511993 Masa Saini 
(IARI). 

Paratypes.—Same data as holotype (3 
2); same except 8.5.1983 (1 6d); Sikkim, 
Gangtok, 1,700 m, 18—20.9.1993, M. S. 
Saini (2 2); Himachal Pradesh, Manali, 
2,400 m, 24.8.1981, M.S. Saini (1 ¢); Uttar 
Pradesh, Chakrata, 2,300 m, 27.5.1983, 
M.S. Saini (1 d) (at Punjabi University; 1 
2, 1 6 in USNM). 

Distribution.—India (Himachal Pradesh, 
Sikkim, West Bengal, Uttar Pradesh). 

Etymology.—Named in honor of Dr. 
Santokh Singh, School of Entomology, St. 
John’s College, Agra, a reputed Indian en- 
tomologist. 


VOLUME 107, NUMBER 2 


= oc iernae 


IL SE > 
~~ 


Figs. 15-19. 
19, B. okutanii. 


Remarks.— Busarbia santokhi and B. 
viridipes both have a yellow scape, head 
smooth and almost impunctate, and the hind 
margin of the head rounded, not carinate. 
Busarbia santokhi is separated from B. vir- 
idipes by the pale yellow abdominal venter 
and broader malar space which is a little 
more than half diameter of the front ocellus. 
Busarbia viridipes has a black abdomen 
and malar space less than half the diameter 
of the front ocellus. 


ISPS LSS SEL PDE CEES SEA SIEVE NTE STATE 


Female lancets. 15, Busarbia shinoharia. 16, B. viridipes. 17, B. santokhi. 18, B. formosana. 


Busarbia shinoharai Saini and Smith, 
new species 
Gisss5selOgUS) 

Female.—Color: Body black with fol- 
lowing pale whitish: clypeus, labrum, man- 
dible at base; upper pronotal margin, tegula, 
postspiracular sclerite, triangular spot on 
mesopleuron behind; sternites 1—6, and legs 


except lightly infuscated tarsal segments of 


hind leg. Wings hyaline; costa, stigma, and 
veins black. 


354 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 20-25. 


Male genitalia. 20-23, Penis valves. 20, Busarbia okutanii. 21, B. santokhi. 22, B. viridipes. 


23-25, Genital capsule, ventral view of left half. 23, B. okutanii. 24, B. santokhi. 25, B. viridipes. 


Structure: Length, 5.5 mm. Antennal 
length 2.4 head width; scape and pedicel 
almost equal in length, each longer than 
their apical widths as 3:2; segments 3 and 
4 almost equal in length. Clypeus subcon- 
vex with anterior margin arcuately emargin- 
ate for % its medial length (Fig. 5). Labrum 
broader than long as 2:1 with rounded an- 
terior margin. Malar space 0.7 diameter 
of front ocellus. OOL:POL:OCL = 1.0:0.3: 
1.0. Supra-antennal pits only indicated. Su- 
pra-antennal tubercles moderately raised 


and continuous with similar frontal ridges. 
Circum- and postocellar furrows distinct; 
interocellar furrow indistinct. Lateral fur- 
rows distinct, straight and almost reaching 
hind margin of head. Postocellar area quad- 
rate; hind margin of head rounded. Lower 
interocular distance greater than eye length, 
LID:IDMO:EL = 1.0:1.1:0.7. ITD:ICD = 
1.0:0.3. Mesepisternum obtusely raised 
without carina or acute apex. Subapical 
tooth of tarsal claws (Fig. 10) slightly short- 
er than apical tooth. Hind basitarsus almost 


VOLUME 107, NUMBER 2 


equal to following tarsal segments com- 
bined. IATS:MB:OATS = 1.1:3.5:0.9. Lan- 
cet with 6 serrulae (Fig. 15). 

Sculpture and pubescence: Head rugose 
with dense confluent punctures. Thorax al- 
most impunctate except posterior slope of 
mesoscutellum with a row of a few large 
punctures. Head and thorax shining, abdo- 
men impuncate, subshining. Pubescence 
golden yellow. 

Male.—Unknown. 

Holotype.—Female, Arunachal Pradesh, 
Dirang, 1,500 m, 10.5.1992, M. S. Saini 
(IARI). 

Distribution.—India (Arunachal Pra- 
desh). 

Etymology.—Named in honor of Dr. 
Akihiko Shinohara, National Science Mu- 
seum (Natural History), Tokyo, a specialist 
on the family Pamphiliidae. 

Remarks.—The black scape and rugose 
head with confluent punctures separate this 
species from other species of Busarbia. 
This and B. isshikii are the only species of 
Busarbia in which the third and fouth an- 
tennal segments are subequal in length; in 
other species the third antennal segment is 
shorter than the fourth. 


Busarbia viridipes Cameron 
(rics 9 SOs 925) 


Busarbia viridipes Cameron 1899: 38— 
39.—Malaise 1944: 20 

Anapeptamena viridipes: Konow 1905: 
74.—Forsius 1932: 52 (in key). 

Busarbia albipes Cameron 1902: 445. 

Busarbia viridipes albipes: Malaise 1944: 
20 (color form). 

Anapeptamena cameroni Konow 1905: 74 
(new name for Busarbia albipes Camer- 
on, preoccupied in Anapeptamena by An- 
apeptamena albipes Konow 1898).— 
Forsius 1932: 53 (in key). 


Female.—Color: Black with following 
pale whitish: clypeus, labrum, mandible ex- 
cept apex, scape, upper pronotal margin, te- 
gula, postspiracular sclerite, subtriangular 
spot on lower half of mesepisternum, and 


S55) 


legs except tips of fore- and mid tarsal seg- 
ments and last 3 hind tarsal segments which 
are fuscous. Wings hyaline; costa, stigma, 
and veins black. 

Structure: Average length, 6.3 mm. An- 
tennal length 2.5 head width; scape longer 
than broad as 2.0:1.5; scape and pedicel 
equal in length, pedicel 2 longer than its 
apical width; segment 3 shorter than 4 as 8: 
9. Clypeus subconvex, roundly emarginated 
for 4% its medial length (Fig. 9), with ante- 
rior margin not depressed. Labrum broader 
than long as 3:2, with round anterior mar- 
gin. Malar space 0.4 diameter of front 
ocellus. OOL:POL:OCL = 1.0:0.3:1.0. Su- 
pra-antennal pits almost distinct and con- 
nected to antennal sockets by very fine fur- 
rows. Supra-antennal tubercles moderately 
raised and sloping posteriorly. Circum-, in- 
ter-, and postocellar furrows distinct. Lat- 
eral furrows distinct, diverging posteriorly 
and abruptly ending just before hind margin 
of head. Postocellar area subquadrate, 
slightly broader than long as 9:8; hind mar- 
gin of head rounded. Lower interocular dis- 
tance greater than eye length, LID:IDMO: 
EL = 1.0:1.1:0.8. ITD:ICD = 1.0:0.4. Me- 
sepisternum obtusely raised with blunt ca- 
rina. Subapical tooth of tarsal claws (Fig. 
13) slightly shorter than apical tooth. Hind 
basitarsus slightly longer than following 
tarsal segments combined, as 8:7. IATS: 
MB:OATS = 1.0:1.2:0.8. Lancet with 4 ser- 
rulae (Fig. 16). 

Sculpture and pubescence: Head and 
thorax smooth and shining, with minute 
scattered punctures; mesoscutellum punc- 
tured on a broad belt along anterior margin 
of appendage. Abdomen impunctate, sub- 
shining. Pubescence brownish yellow. 

Male.—Average length, 5.2 mm. Similar 
to female except malar space 0.2 diameter 
of front ocellus and inner margins of eyes 
faintly converging below. Genitalia: Penis 
valve (Fig. 22) with valviceps almost 
square, of equal width, truncate at apex, 
without spines on ventral margin; genital 
capsule (Fig. 25), similar to B. okutanit. 

Types.—Cameron (1899) described B. 


356 


viridipes from a female from Khasia Hills, 
and (1902) B. albipes from a female from 
‘“‘Simla.”’ The types are in The Natural His- 
tory Museum, London, U.K. 

Specimens examined.—INDIA: Sikkim, 
Chungthang, 1,500 m, 12.5.1995, M.S. Sai- 
nino) siGanetok. 1-700 mi WiS2an9935 
M.S. Saini (4 ¢d, 2 2); Phodong, 1,600 m, 
7.5.1995, M.S. Saini (5 6, 2 ¢); Singhik, 
1,440 m, 8.5.1995, M.S. Saini 9 6, 10 @); 
Sangtam, 900 m, 4.5.1995, M.S. Saini (10 
6, I Qe Akerorme, W3K00) tan, WOLS NOs), Miles). 
Saini (4 6, 1 2); Sikkim, Sangtam, 3,000 
ft., May 4, 1995, M.S. Saini; Assam, Ka- 
meng, Dirang Dzong, 10-IV-61, 5,300’, F 
Schmid (1 2); Assam, Kameng, Rupa, 3- 
IV-61, E Schmid (1 @). 

Distribution.—India (Assam, Sikkim). 

Variation.—Some males have the hind 
basitarsus and remaining hind tarsal seg- 
ments entirely fuscous. The subspecies al- 
bipes was separated by the black mesopleu- 
ron and mandibles and infuscate clypeus 
and labrum. We follow Malaise (1944) in 
considering this only a color variation of B. 
viridipes. 

Remarks.— Busarbia viridipes is similar 
to B. santokhi; see remarks under the latter 
for separation. 


ACKNOWLEDGMENTS 


We thank Cathy Apgar, Systematic En- 
tomology Laboratory (SEL), U.S. Depart- 
ment of Agriculture, Washington, DC for 
help in preparation of Figures 1—4 and for 
arrangement of plates. We appreciate the re- 
views of the following: Henri Goulet, Ag- 
riculture and Agri-Food Canada, Ottawa; 
and S. J. Scheffer and M. G. Pogue, SEL, 
Beltsville, MD, and Washington, DC, re- 
spectively. We are thankful to US-PL-480 
for funding a project under which this work 
was completed and to the authorities of 
Punjabi University, Patiala, for providing 
the basic laboratory facilities. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


LITERATURE CITED 


Cameron, P. 1899. Hymenoptera Orientalia, or contri- 
butions to a knowledge of Hymenoptera of the 
Oriental Zoological Region. Part VIII. The Hy- 
menoptera of Khasia Hills. First Paper. Memoirs 
and Proceedings of the Manchester Literary and 
Philosophical Society 43: 1-50. 

. 1902. Descriptions of new genera and species 
of Hymenoptera, collected by Maj. C. S. Nurse at 
Dessa, Ferozepur and Shimla. Part I. Journal of 
the Bombay Natural History Society 14: 433-447. 

Forsius, R. 1932. Eine neue Anapeptamena-Art aus 
Formosa (Hym., Tenthr.). Notulae Entomologicae 
12: 51-53. 

Konow, E W. 1905. Hymenoptera, Family Tenthredi- 
nidae. In Wytsman, P., ed. Genera Insectorum, 
Fasc. 29, 176 pp. 

Malaise, R. 1944. Entomological results from the 
Swedish expedition 1934 to Burma and British In- 
dia. Arikiv fér Zoologi 35: 1—58. 

. 1963. Hymenoptera Tenthredinoidea, subfam- 
ily Selandriinae, key to genera of the world. En- 
tomologisk Tijdskrift 84: 159-215. 

Oehlke, J. and J. Wudowenz. 1984. Katalog der in den 
Sammlungen der Abteilung Taxonomie der Insek- 
ten des Institutes fiir Pflanzenschutzforschung, 
Bereich Eberswalde (ehemals Deutsches Ento- 
mologisches Institut), aufbewahrten Typen—X XII 
(Hymenoptera: Symphyta). Beitrage zur Entomo- 
logie, Berlin 34: 363-320. 

Rohwer, S. A. 1916. H. Sauter’s Formosa—Ausbeute, 
Chalastogastra (Hym.). Supplementa Entomolo- 
gia, Berlin 5: 81-113. 

Ross, H. H. 1937. A generic classification of the Ne- 
arctic sawflies (Hymenoptera, Symphyta). Illinois 
Biological Monographs 34, 173 pp. 

. 1945. Sawfly genitalia, terminology and study 
techniques. Entomological News 56: 261—268. 

Saini, M.S., D. R. Smith, and T. P. Saini. 2003. Review 
of the southeastern Asian sawfly genus Anapep- 
tamena Konow (Hymenoptera: Tenthredinidae). 
Proceedings of the Entomological Society of 
Washington 105: 641—646. 

Takeuchi, K. 1928. New sawflies from Formosa—lII. 
Transactions of the Natural History Society of 
Formosa 28: 38-45. 

. 1941. A systematic study on the suborder 
Symphyta (Hymenoptera) of the Japanese Empire 
(IV). Tenthredo 3: 230—274. 

Togashi, I. 1990. Notes on Taiwanese Symphyta (Hy- 
menoptra, Siricidae, Tenthredinidae, Argidae) (II). 
Esakia, Special Issue No. 1, pp. 177—192. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 357-361 


TAXONOMIC NOTES ON SOUTH AMERICAN POGONOCEROMORPHUS PIC 
(COLEOPTERA), INCLUDING TRANSFER FROM PYROCHROIDAE 
(PY ROCHROINAE) TO ANTHICIDAE (EURYGENIINAE) 


DANIEL K. YOUNG 


Department of Entomology, University of Wisconsin, Madison, Wisconsin 53706, 
U.S.A. (email: young @entomology.wisc.edu) 


Abstract.—Pogonoceromorphus Pic is transferred from Pyrochroidae (Pyrochroinae) to 
Anthicidae (Eurygeniinae). Type material for Pogonoceromorphus gracilis Pic is dis- 
cussed and a syntype confirmed. Pogonoceromorphus lauropalui Vulcano and Pereira is 
transferred to Disema Maklin (Tenebrionidae: Lagriinae (n. comb.)). Salient anatomical 


characters supporting the proposed changes are discussed. 


Key Words: 


Pogonoceromorphus gracilis, Pogonoceromorphus lauropalui, Pyrochro- 


idae (Pyrochroinae), Anthicidae (Eurygeniinae), Disema, Tenebrionidae 


(Lagriinae), South America 


Recent years have seen activity directed 
toward better defining generic concepts 
within the pyrochroine Pyrochroidae 
(Young 1999, 2002, 2004a, b). This paper 
continues the process of reviewing generic 
relationships of Pyrochroinae in an effort to 
redefine the subfamily and genera therein as 
monophyletic taxa. This paper proposes 
taxonomic changes involving the two spe- 
cies presently comprising Pogonoceromor- 
phus Pic. 

At the time of its description (Pic 1921), 
Pogonoceromorphus was monotypic, the 
only species being P. gracilis (Figs. 1—2) 
from ‘“‘Maroni” in French Guiana. It was 
said to most closely resemble the pyro- 
chroid genus Pogonocerus Fischer (Figs. 3— 
4), differing by its more elongate, apically 
constricted prothorax with a basal sulcus, 
and by the third antennomere bearing a 
long, thin ramus. Aside from being listed in 
two catalogues (Blair 1928, Blackwelder 
1945), nothing else was published on Po- 
gonoceromorphus until Vulcano and Perei- 
ra (1972) described Pogonoceromorphus 
lauropalui from Jacareacanga, Brazil. 


METHODS AND MATERIALS 


Depositories and collection acronyms.— 
Taxonomic material came from the Musé- 
um National d’Histoire Naturelle, Paris 
(MNHN) and The Natural History Muse- 
um, London (BMNH). 

Figures.—Images (Figs. 1—4) were cap- 
tured as multiple “*.tif”’ files using a Polar- 
oid® DMC-le digital camera attached to a 
Leica® MZX75 microscope and integrated 
via Pax-It® software. Multiple files for a 
given “‘figure’’ have generally been used to 
facilitate ““building”’ a final figure that is far 
more sharply focused than any single digi- 
tal image, due to depth of focus limitations. 
All images were saved to a multi-depart- 
mental server on a Local Area Network 
(LAN) and edited using either Adobe® 
PhotoShop, version 6.0 software, or 
JASC®, Paint Shop Pro, version 7.0 soft- 
the final 
image for a given figure, the touch up pro- 


ware. In addition to “building” 


cess typically involves a number of cut/ 
paste, blend, and erase functions offered by 


358 


the software to graphically remove unwant- 
ed shadows and background. 

Material examined.—The single male of 
P. gracilis examined is certainly a syntype. 
It could be the only specimen Pic had be- 
fore him, but as there is no clear indication 
of this on the specimen labels or in the de- 
scription, it can not be considered the ho- 
lotype. Because it was cryptically labeled, 
I appended a “Syntype” label. 

Pogonoceromorphus gracilis: (1 3) [1st 
label]: Guyane, Maroni; [2nd label]: Type 
(small label appearing to be in Pic’s hand); 
[3rd label]: Pogonoceromorphus, gracilis 
Pic; [4th label]: SYNTYPE:, Pogonocero- 
morphus, gracilis 3 Pic, fide Daniel K. 
Young. 

The “‘Maroni”’ label may refer to the port 
city of St.-Laurent-du-Maroni, on the Ma- 
roni River, between Suriname and French 
Guiana; 05°29'46’N, 54°02'46”W. It also 
could refer simply to the Maroni River, or 
to the eastern State of Maroni, French Gui- 
ana. 

A second specimen of Pogonoceromor- 
phus in the Pic material of unidentified Pyr- 
ochroidae might be conspecific with P. 
gracilis. The color is not quite the same; 
more significantly, however, the vestiture of 
the prothoracic tibiae is a little different, the 
elytra are less coarsely punctured in the 
*“Maroni” specimen, and the apex of the 
fused parameres differs slightly between the 
two. For the present, however, the specimen 
is considered conspecific. 

Pogonoceromorphus gracilis? : (1 3) [1st 
label]: Teffe (Lya), Amazones; [2nd label]: 
Dendroides, probabl; [3rd label]: Pogono- 
ceromorphus, gracilis?, Pic 3d, det. Daniel 
K. Young. [Teffé (= Tefé) is located in cen- 
tral Amazonas, Brazil, along the Amazon 
River; 03°20'23"S, 64°45'17’"W.] 


DISCUSSION 


The male P. gracilis (Figs. 1-2) was dis- 
covered in the Pic material at the Muséum 
National d’Histoire Naturelle, Paris. The 
point-mounted specimen is in reasonably 
good condition and bears the locality data 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


label “‘“Guyane, Maroni’’; identical to that 
provided in the original description (Pic 
1921). The strongly, finely, pectinate anten- 
nae and nearly holoptic compound eyes are, 
as Pic noted, somewhat suggestive of the 
monotypic Russian pyrochroine, Pogono- 
cerus thoracicus Fischer (Figs. 3—4), al- 
though the eyes of the latter are not nearly 
as approximate dorsally. However, at closer 
look, there are also a number of more sig- 
nificant differences. The basal pronotal sul- 
cus described by Pic actually extends lat- 
erally onto the hypomera in P. gracilis, 
ending near the coxal cowling. This char- 
acter is suggestive of Anthicidae as char- 
acterized by Ivie (2002). The prothoracic 
coxal cavities of Pogonoceromorphus are 
widely open externally, but closed internal- 
ly. This is also more like the condition gen- 
erally seen in Anthicidae (Crowson 1955). 
The prothoracic coxal cavities are widely 
open both externally and internally in pyr- 
ochroine Pyrochroidae. The emarginate 
compound eyes are conspicuously setose 
between facets, another character fairly 
common in anthicids (Chandler 2002). The 
cranial “‘neck” in P. gracilis is about 1/3 
the width of the head and coarsely, strigo- 
sely punctate; the metathoracic coxae are 
very nearly contiguous mesally; the inter- 
coxal process of the first ventrite has the 
margins diverging at an angle of distinctly 
less than 45°. This character set clearly 
places P. gracilis in the anthicid subfamily 
Eurygeniinae. From these observations, I 
propose to remove Pogonoceromorphus 
Pic, based upon the type species, P. gracilis 
(by monotypy), from Pyrochroidae (Pyro- 
chroinae) and hereby transfer it to the an- 
thicid subfamily Eurygeniinae. 

Having examined Pic’s specimen of P. 
gracilis, it is clearly not congeneric with P. 
lauropalui Vulcano and Pereira. Presuming 
that P. lauropalui was correctly assigned to 
Pogonoceromorphus, Lawrence and New- 
ton (1995: 899) commented that, ‘‘The 
Neotropical genus Pogonoceromorphus 
...almost certainly belongs to the tenebri- 
onid subfamily Lagriinae.” Although I 


VOLUME 107, NUMBER 2 359 


r. 2 Ph. oh} 


aur 


- 


3 


Figs. 1-2. Pogonoceromorphus gracilis Pic, adult male: 1, habitus. 2, head, dorsal view. 


Figs. 3-4. Pogonocerus thoracicus Fischer, adult male: 3, habitus; 4, head, dorsal view. 


have not examined the type of P. Jauropal- genus Disema (Barsenis) sensu Champion 
ui, Vulcano and Pereira’s well-drawn habi- (1917: 141—142): “‘The chief characters of 
tus figure (1972: 30, fig. 1) is very close to the genus are—the strongly serrate (Disema 
the description of the lagriine tenebrionid _ s. str.) or flabellate (Barsenis) d-antennae 


360 


. and the presence of a depressed opaque 
area on the outer portion of the elytra... 
The eyes in the typical forms are extremely 
large and contiguous in the male ... re- 
markable modifications in the structure of 
one or more of the d- tibiae also occurring 
occasionally ...”’ In his “‘lagriid”’ contri- 
bution to the Genera Insectorum, Borch- 
mann (1936) retained Barsenis Pascoe as 
generically distinct from Disema Maklin. In 
his key to genera, Barsenis and Disema run 
together to the last couplet. He distin- 
guished the two genera using five charac- 
ters. The antennae of Barsenis were de- 
scribed as branched (= flabellate), while 
those of Disema were said to be strongly 
serrate. Specimens of Barsenis have the 
apical maxillary palpomere enlarged and 
subtriangular; it was described as small and 
cultriform in Disema. The pronotum of Di- 
sema bears scattered large punctures that 
are lacking in species of Barsenis; the elytra 
of Disema widen posteriorly as opposed to 
being cylindrical in species of Barsenis. 
Males of Barsenis were described as usu- 
ally displaying highly modified tibiae, yet 
he described those of Disema as being “‘of- 
ten sexually dimorphic.” 

Like Barsenis (sensu Borchmann) the 
specimen of P. /auropalui has flabellate an- 
tennae, an enlarged, securiform terminal 
maxillary palpomere (Vulcano and Pereira 
1972: 30, figs. 1-2), and modified protho- 
racic tibiae (Vulcano and Pereira 1972: 30, 
figs. 1, 6). However, more like Borch- 
mann’s characterization of Disema, the 
pronotum definitely has scattered large 
punctures and the elytra are widest along 
the posterior third, not cylindrical. In any 
case, P. lauropalui is not assignable to Po- 
gonoceromorphus Pic. For the present, be- 
cause of the inconsistencies noted above 
and because it is beyond the scope of this 
contribution to further examine generic lim- 
its within lagriine Tenebrionidae, I defer to 
Champion in taking a broader view of Di- 
sema. Thus, I am transferring P. lauropalui 
to Disema (Barsenis): Disema (Barsenis) 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


lauropalui (Vulcano and Pereira), new 
combination. 

It is also beyond the scope of this paper 
to comment on the validity of D. lauropalui 
as a species, since a number of species have 
been assigned to Disema and Barsenis. Di- 
sema lauropalui does appear to be related 
to material I examined at The Natural His- 
tory Museum, London, identified as Disema 
(Barsenis) fulvipes (Pascoe). Unfortunately, 
Pascoe’s (1887) figure accompanying his 
original description lacks sufficient detail to 
be useful in comparing the two. 


ACKNOWLEDGMENTS 


I am grateful to Max Barclay, Martin 
Brendell, and Malcolm Kerley (BMNH) for 
their kind assistance during my visits. I also 
extend thanks to Claude Girard (MNHN) 
for an extended loan of material critical to 
this study. Preliminary digital images and 
final figures were captured and produced by 
Michael Young, and electronic formatting 
of the plates was produced by Craig Bra- 
bant; Iam most grateful for their assistance. 
This research was supported in part by 
grants from the National Science Founda- 
tion (BSR-9006342), the University of Wis- 
consin Graduate School (900159), and the 
University of Wisconsin’s Natural History 
Museums Council Small Grants Program. 


LITERATURE CITED 


Blackwelder, R. E. 1945. Checklist of the Coleopter- 
ous insects of Mexico, Central America, the West 
Indies, and South America. Part 3. Smithsonian 
Institution. Bulletin of the United States National 
Museum 185: 343-550. 

Blair, K. G. 1928. Pars. 99: Pyrochroidae, pp. 1-14. In 
Schenkling, S., ed. Coleopterorum Catalogus, Vol. 
17, Berlin. 

Borchmann, F 1936. Coleoptera Fam. Lagriidae, pp. 
1-555. In Wytsman, P., ed. Genera Insectorum. 
Fasc. 204. L. Desmet-Verteneuil, Brussels. 

Champion, G. C. 1917. Notes on tropical American 
Lagriidae, with descriptions of new species. En- 
tomologist’s Monthly Magazine 53: 132-154. 

Chandler, D. S. 2002. Family Anthicidae, pp. 549-558. 
In Arnett, R. H. Jr., M. C. Thomas, P. E. Skelley, 
and J. H. Frank, eds. American Beetles, Vol. 2. 
CRC Press, Boca Raton, Florida. 

Crowson, R. A. 1955. The Natural Classification of the 


VOLUME 107, NUMBER 2 


Families of Coleoptera. Nathanial Lloyd, London, 
187 pp. 

Ivie, M. A. 2002. Keys to families of beetles in Amer- 
ica north of Mexico, pp. 816-835. Jn Arnett, R. 
H. Jr., M. C. Thomas, P. E. Skelley, and J. H. 
Frank, eds. American Beetles, Vol. 2. CRC Press, 
Boca Raton, Florida. 

Lawrence, J. E and A. EK Newton, Jr. 1995. Families 
and subfamilies of Coleoptera (with selected gen- 
era, notes, references and data on family-group 
names). pp. 779-1006. 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 Instytut 
Zoologil PAN, Warszawa. 

Pascoe, E P. 1887. Notes on Coleoptera, with descrip- 
tions of new genera and species.—Part VI. The 
Annals and Magazine of Natural History, includ- 
ing Zoology, Botany, and Geology (5)20: 8-20, 
plate I. 

Pic, M. 1921. Nouveautés diverses. Mélanges Exotico- 
Entomologiques 33: 1—32. 

Vulcano, M. A. and E S. Pereira. 1972. Ocorréncia da 


361 


familia Pyrochroidae Leach 1815 (Coleoptera) no 
Brasil, com descrigo de uma espécie nova. Arqui- 
vos do Instituto Biol6gico, SA4o Paulo 39: 27-34. 


Young, D. K. 1999. Transfer of the Taiwanese Pseu- 


dopyrochroa umenoi and the Japanese Pseudopyr- 
ochroa amamiana to Pseudodendroides (Coleop- 
tera: Pyrochroidae: Pyrochroinae). The Pan-Pacif- 
ic Entomologist 75: 1-7. 

. 2002. Family Pyrochroidae. pp. 540-543. Jn 
Arnett, R. H. Jr, M. C. Thomas, P. E. Skelley, and 
J. H. Frank, eds. American Beetles, Vol. 2. CRC 
Press, Boca Raton, Florida. 

. 2004a. Himalapyrochroa gibbosa, a new ge- 
nus and species of fire-colored beetle (Coleoptera: 
Pyrochroidae: Pyrochroinae) from Darjeeling, In- 
dia. Oriental Insects 38: 189-196. 

. 2004b. Dendroidopsis and Frontodendroidop- 
sis: Two new Asian genera of fire-colored beetles 
(Coleoptera: Pyrochroidae: Pyrochroinae), with 
descriptions of two new species of Dendroidopsis, 
and lectotype designations for Dendroides assa- 
mensis Blair and Pseudopyrochroa binhana Pic. 
Oriental Insects 38: 197-217. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 362-375 


THREE SPECIES OF FACULTATIVE MYODOCHINI 
(LYGAEOIDEA: RHYPAROCHROMIDAE) ASSOCIATED WITH 
FIGS IN MEXICO 


Luis CERVANTES PEREDO AND SAGRARIO GAMEZ-VIRUES 


Departamento de Entomologia, Instituto de Ecologia, A.C. km. 2.5 Antigua Carretera 
a Coatepec #351, CP 91070 Xalapa, Veracruz, México. (e-mail: cervantl@ecologia.edu.mx) 


Abstract.—The biology of the lygaeoids Myodocha intermedia Distant, M. unispinosa 
Stal, and Neopamera bilobata (Say) were studied in eastern Mexico between 2001 and 
2003. Descriptions and illustrations of their immature stages are presented for the first 
time. Their life cycles under laboratory conditions and wild fig hosts are also included. 
These lygaeoids are reported as frequent facultative, terrestrial seed predators of Ficus 


spp. 


Resumen.—La biologia de los lygaeoideos Myodocha intermedia Distant, M. unispinosa 
Stal y Neopamera bilobata (Say) fue estudiada en campo, en el Este de México, entre 
2001 y 2003. Las descripciones e ilustraciones de sus estadios inmaduros se presentan 
por primera vez. También, se incluyen datos sobre su ciclo de vida bajo condiciones de 
laboratorio e higueras silvestres hospederas. Se observ que estos lygaeoideos son depre- 


dadores facultativos terrestres frecuentes de semillas de Ficus spp. 


Key Words: 


More than 30 years ago, Slater (1972) 
recognized the association of lygaeoids 
with figs. In this work, based on species 
from Africa and the Caribbean, Slater sep- 
arated this guild of insects into four groups, 
depending on the degree of specialization 
on figs: I. ARBOREAL SEED PREDA- 
TORS. Species that live in the fig trees 
themselves and feed on the seeds, while the 
latter are still in the syconium. II. OBLIG- 
ATORY TERRESTRIAL SEED PREDA- 
TORS. Species that live in the litter layer 
below the trees and feed only, or primarily, 
on Ficus seeds. II]. FREQUENT FACUL- 
TATIVE TERRESTRIAL SEED PREDA- 
TORS. Species that live in the litter layer 
below Ficus trees and feed on the seeds, but 
feed on seeds of other plants as well; and 


Ficus, immature stages, life cycle, Lygaeoidea, Myodocha, Neopamera 


IV. ACCIDENTAL TERRESTRIAL SEED 
PREDATORS. Species that feed primarily 
on other seeds but utilize seeds of Ficus 
when they occur in the insect’s habitat. 
Since Slater (1972), there have been only 
a few publications treating the lygaeoids as- 
sociated with figs (Rodriguez 1998a, b, c; 
Brambila 2000; Cervantes and Pacheco 
2003; Cervantes et al. 2004). These studies 
deal mainly with lygaeoids that correspond 
to the first two categories proposed by Slat- 
er. The present study is the first one to con- 
sider species of Myodochini that are fre- 
quent facultative terrestrial seed predators. 
Herein we report the life cycles of Myodo- 
cha intermedia Distant, Myodocha unispi- 
nosa Stal, and Neopamera bilobata (Say) 
including descriptions and illustrations of 


VOLUME 107, NUMBER 2 


all life history stages, their biology, and 
host plants. 

This paper is the first part of a series pre- 
senting, information regarding other species 
of lygaeoids associated with figs will be 
published latter on, as well as data on the 
effect of these seed predators on seed ger- 
mination will be published later. 


MATERIALS AND METHODS 


Monthly collecting trips between 2001 
and 2003 were made to several localities in 
the Mexican states of Campeche, Puebla, 
Tamaulipas, and Veracruz. The objective 
was to collect Heteroptera associated with 
wild fruiting fig trees. About 30 fig species 
were sampled from localities occurring at 
sea level to an altitude of 1,500 m. Several 
types of vegetation were included: low 
tropical dry forest, medium tropical forest, 
high tropical rain forest, and cloud forest. 

Biological information was obtained 
mainly from two biological research sta- 
tions. Estacion Biologica La Mancha is sit- 
uated at sea level on the coast of Veracruz, 
30 km NE of Ciudad Cardel (96°22’40"W 
and 19°35'2”N); Estacion Biologica Los 
Tuxtlas (94°40'’W and 18°00'N), elevation 
varies from sea level to around 400 m is 
also situated in the state of Veracruz, but 
150 km south of Estacion Biologica La 
Mancha. The type of vegetation at La Man- 
cha is mainly medium tropical dry forest, 
and at Los Tuxtlas it is dominated by high 
tropical rain forest. 

The ground area covered by the crown of 
each fruiting tree, including leaf litter, but- 
tresses, and aerial roots, was checked. Top 
parts of each tree were searched by climb- 
ing the tree with the use of ropes or free- 
hand using their aerial root systems. Bugs 
were collected by using an aspirator or by 
cutting small branches bearing fruits. In- 
sects were kept alive and put into plastic 
containers (9 X 8 cm) covered with muslin 
cloth to avoid condensation. A dry leaf of 
the fig tree and an opened fruit were put in 
each container as well as a small damp cot- 
ton ball; these were changed every three 


363 


days. Containers were checked daily for the 
presence of eggs. Individuals were kept un- 
der laboratory conditions at about 20°C and 
70% RH. Individuals fixed in 70% alcohol 
were used for illustrations and descriptions; 
measurements are given in mm=+I1DS. De- 
scriptions of the adults are based on the 
original descriptions (Say 1831. Stal 1874, 
Distant 1882). 

Voucher specimens have been deposited 
in the Coleccion Entomologica del Instituto 
de Ecologia, A.C. Xalapa, Veracruz, Mex- 
ico (IEXA). 


DESCRIPTIONS 


Myodocha intermedia Distant 
(Figs. 1A—G) 


Egg (Fig. 1A).—Elongate, anterior pole 
rounded and posterior pole slightly pointed. 
1.22+0.04 mm long by 0.48+0.02 mm 
wide (n = 10). Yellowish white when laid, 
and turn red in four d. Chorion with very 
small projections, be coming more apparent 
on both poles. Operculum with 5 mycro- 
pilar processes. 

First instar (Fig. 1B).—Body elongate. 
Head, pronotum and antennal segment IV 
reddish brown; eyes red; antennal segment 
I pale brown, antennal segments IJ and III 
yellowish white. Rostral segment I pale 
brown, remainder yellowish white. Poste- 
rior margin of pronotum with a narrow red 
band. Meso- and metanota pale brown. Pro- 
pleuron reddish brown, meso- and meta- 
pleura pale brown. Coxae pale brown; Front 
and middle femora and tibiae, and tarsi yel- 
lowish white; hind femur and tibiae slightly 
darker. Abdomen yellowish white with ir- 
regular red areas on lateral margins of seg- 
ments I and II; with a red, cross-shape area 
on midline of segments III to VIII and 
along segment IV; tip of abdomen pale 
brown. Scent gland openings present be- 
tween segments III-IV, [V—V, and V—VI as 
small grayish areas. Abdominal venter yel- 
lowish with red areas on lateral margins of 
segments I, II, and IV. Long thick hairs sit- 
uated symmetrically on lateral margins as 


364 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Instars of Myodocha intermedia Distant. A, Egg. B, First instar. C, Second instar. D, Third instar. E, 
Fourth instar. E Fifth instar. G, Adult male. (Scale = 1 mm) 


VOLUME 107, NUMBER 2 


follows: a pair in front of eyes, one behind 
eyes, one on posterior margin of pro-, 
meso-, and metanota, two hairs on abdom- 
inal segment III, and one hair on segments 
IV to VI. Long more slender hairs also pre- 
sent on margins of head and abdomen es- 
pecially on last four segments; ventral sur- 
face of abdomen with numerous hairs. Ros- 
trum reaching metacoxae. Tibiae of all legs 
covered by numerous hairs. Measurements 
(n = 10): Body length 1.65+0.12; head 
length 0.52+0.05; width across eyes 
0.44+0.02; interocular distance 0.26+0.01:; 
postocular distance 0.150.02; antennal seg- 
noes. I O.72007, Wl Oey 0, 00 
0.340.02, IV 0.51+0.02; rostral segments: 
I 0.28+0.03, If 0.31+0.02, III 0.22+0.02, 
IV 0.27+0.02; pronotum length 0.2+0.02; 
width across humeral angles 0.43+0.01; 
width across anterior margin 0.40+0.01; 
fore leg: femur length 0.45+0.03; tibia 
length 0.44+0.02; tarsi length: I 0.13+0.02, 
HL OZaet0 (ON. 

Second instar (Fig. 1C).—Body slightly 
pyriforme. Head and pronotum brown. 
Eyes brownish red. Antennal segments I to 
III yellowish white, segment IV reddish 
brown. Rostral segment I grayish yellow, 
remainder yellowish white with tip of seg- 
ment IV brown. Propleuron brown, meso- 
and metapleura yellowish white. Legs yel- 
lowish white, although some individuals 
with darker femora. Meso- and metanota 
and abdominal segments I and II grayish; 
middle of segments III to V also grayish. 
Red areas present only on lateral margins 
of segment IV and on midline of segments 
VI to VUI. Rostrum reaching metacoxae. 
Measurements (n = 10): Body length 
2.370.18; head length 0.66+0.02; width 
across eyes 0.54+0.03; interocular distance 
0.31+£0.01; postocular distance 0.21 +0.02; 
antennal segments: I 0.24+0.02, II 
1.48+0.02, HI 0.44+0.02, IV 0.65+0.03; 
rostral segments: I 0.36+0.04, II 
0.36+0.04, I 0.29+0.02, IV 0.28+0.02; 
pronotum length 0.28+0.02; width across 
humeral angles 0.53+0.03; width across an- 
terior margin 0.44+0.01; fore leg: femur 


365 


length 0.66+0.05; tibia length 0.65+0.04: 
tarsi length: I 0.20+0.03, II 0.24+0.01. 

Third instar (Fig. 1D).—Similar to sec- 
ond instar, although meso- and metanota 
and abdomen slightly darker. Thick hairs 
less apparent. Abdomen with “‘Y”’ suture 
visible and delimited by dark brown lines. 
Red areas of abdominal segment IV of pre- 
ceding instars turning brown and red areas 
on midline of segments VI to VIII less ap- 
parent. Rostrum reaching mesocoxae. Mea- 
surements (n = 5): Body length 3.3+0.25; 
head length 0.92+0.14; width across eyes 
0.64+0.02; interocular distance 0.36+0.02; 
postocular distance 0.4+0.05; antennal seg- 
ments: 0 0253== (0104-7 tl Ol/5==O0!0 seal 
0.66+0.05, IV 0.89+0.09; rostral segments: 
I 0.52+0.07, I 0.48+0.05, Ill 0.4+0.07, IV 
0.3+0.03; pronotum length 0.42+0.04; 
width across humeral angles 0.61+0.06; 
width across anterior margin 0.46+0.01; 
fore leg: femur length 0.92+0.06; tibia 
length 0.94+0.04; tarsi length: I 0.27+0.03, 
O27 == O02: 

Fourth instar (Fig. 1E).—Pyriform, but 
with an elongated neck. Head, pro-, meso-, 
and metanota, and propleuron dark brown. 
Pro- and mesonota with a pale yellow sub- 
marginal band. Legs pale yellow; femora of 
all legs with two pale brown “rings” near 
distal ends, most apparent on middle and 
hind femora. Abdomen grayish with only a 
few pale yellow areas, with dark brown 
area along midline of segments III to VI 
and across scent gland openings. Rostrum 
slightly passing procoxae. Mesothoracic 
wing pads covering metanotum. Measure- 
ments (n = 3): Body length 4.93+0.3; head 
length 1.42+0.06; width across eyes 
0.84+0.05; interocular distance 0.41+0.01; 
postocular distance 0.65+0.05; antennal 
segments: I 0:43+0.06;,, IL 1.22=0.1, 0 
1.10, IV 1.32+0.06; rostral segments: I 
0.63+0.06, II 0.67+0.09, III 0.57+0.12, IV 
0.38+0.03; length 0.73+0.03; 
width across humeral angles 0.84+0.05; 
width across anterior margin 0.53+0.03; 
1.55+0.15; tibia 


pronotum 


fore leg: femur length 


366 


length 1.52+0.1; tarsi length: I 0.45=+0, I 
(O).235)22(0), 

Fifth instar (Fig. 1F).—Body elongate 
with neck well differentiated. Head, pron- 
otum, and scutellum brown. Eyes dark 
brown. Antennal segments I-III pale yel- 
low, distal end of segment III slightly red- 
dish, and segment IV reddish brown. Me- 
sothoracic wing pads with base pale brown 
and distal half yellowish brown. Femora 
darker than preceding and instead of hay- 
ing two pale brown rings, has one yellow- 
ish white annulus. Abdomen variegated 
with grayish and pale yellow areas. Dark 
brown area on midline of abdomen divided 
and present only around scent gland open- 
ings, red areas absent. Rostrum not reach- 
ing procoxae. Lateral margins of pronotum 
and mesothoracic wing pads emarginate. 
Front femur with two rows of small spines 
near distal end. Measurements (n = 4): 
Body length 6.85+0.68; head length 
2.04+0.14; width across eyes 1.02+0.12; 
interocular distance 0.51+0.06; postocular 
distance 1.12+0.05; antennal segments: I 
OOZAz2OLNG, IW WSO US, WO ILS2z2O, 116, IY 
1.62+0.19; rostral segments: I 0.76+0.08, 
II 0.81+0.11, Hl 0.6+0.08, IV 0.45+0.04; 
pronotum length 0.95+0.18; width across 
humeral angles 1.28+0.12; width across 
anterior margin 0.61+0.06; scutellum 
length 0.69+0.08; scutellum width 
OLOz009s tiore legs iene llengin 
2.12+0.1; tibia length 2.24+0.12; tarsi 
length: I 0.66+0.08, If 0.5+0.05. 

Adult (Fig. 1G).—Body elongate and 
narrow, posterior region of head forming a 
long neck. Posterior lobe of pronotum, scu- 
tellum, clavus, and chorion with numerous 
punctures. Head, pronotum, scutellum, and 
thoracic pleura black, head slightly shiny. 
Eyes brownish red. Antennal segment I 
dark brown, segments II and III pale brown 
with base pale yellow; segment IV brown 
with a pale yellow subasal annulus. Rostral 
segments I and IV brown, segment IV with 
yellow base; segments II and III pale yel- 
low. Anterior pronotal lobe black, some- 
times with grayish areas around calli. Cox- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ae black; front femur with proximal third 
pale yellow, and distal two-thirds brown 
with a pale yellow subapical annulus; mid- 
dle and hind femora with proximal two- 
thirds pale yellow and distal third brown 
with a pale yellow subdistal annulus. Tibiae 
and tarsi of all legs pale yellow, sometimes 
bases of tibiae slightly darker. Clavus black 
with pale yellow margins. Chorion brown- 
ish black with a few pale yellow areas on 
basal half and yellowish white area near 
apex. Membrane brownish black with veins 
and apical macula pale yellow. Abdominal 
venter brownish black, although first two 
visible sternites slightly less dark. Head 
with only a few long hairs in front of eyes, 
rest of head surface with very small, flat 
hairs. Rostrum not reaching procoxae and, 
in most individuals, only passing neck re- 
gion. Front femur with a double-ranked se- 
ries of ventral spines, longest situated on 
annulated portion; front tibiae of male with 
a small spine on proximal third, lacking in 
female. 

Male: Measurements (n = 10): Body 
length 11.04+0.5; head length 3.12+0.17; 
width across eyes 1.22+0.04; interocular 
distance 0.48+0.03; interocellar distance 
0.19+0.03; postocular distance 1.79+0.28; 
antennal segments: I 0.89+0.04, II 
DAS =0:22 238 =0 56; Ve 25 6==0 MG: 
rostral segments: I 0.93+0.07, I 
1204220: 0555 {0:92.00 55elNV 4052-01055 
pronotum length 2+0.15; width across hu- 
meral angles 2+0.12; width across anterior 
margin 0.64+0.07; scutellum length 
1.12+0.08; scutellum width 1.01+0.09; 
fore leg: femur length 3.5+0.27; tibia 
length 3.19+0.2; tarsi length: I 1.02+0.06, 
MEO == 0 MOS == OLOS: 

Female: Measurements (n = 10): Body 
length 11.29+0.58; head length 3.06+0.33; 
width across eyes 1.31+0.05; interocular 
distance 0.55+0.04; interocellar distance 
0.2+0.02; postocular distance 1.82+0.13; 
antennal segments: I 0.89+0.06, II 
Desa == O24 lly 2360.25 2 Alena: 
rostral segments: I 0.99+0.06, II 
12OS== 01085 SHI O295== 0106S IIVaIOlS 20104 


VOLUME 107, NUMBER 2 


pronotum length 1.95+0.2; width across 
humeral angles 2.22+0.09; width across an- 
terior margin 0.66+0.06; scutellum length 
1.14+0.13; scutellum width 1.18+0.11: 
fore leg: femur length 3.35+0.33; tibia 
length 3.09+0.28; tarsi length: I 1.02+0.06, 
O22 ==0!0 5 ll 03 == 003) 

Biology.—At Los Tuxtla, Myodocha in- 
termedia fed on fallen fruits of Ficus insi- 
pida Willd., it was also collected at night at 
the lights of the Field Station. In Campeche, 
this species was found under the crown of 
Ficus cotinifolia (Kunth). Nymphs and 
adults were observed running between leaf 
litter and ground vegetation feeding on the 
open fruits of this species. 

Under laboratory conditions, adults col- 
lected at light reproduced easily when they 
were fed fruits of F. cotinifolia. Adults ovi- 
posited on a cotton ball or on a leaf. Al- 
though second-instar nymphs died in great 
quantities, several nymphs reached to the 
adult stage. 

Records from material deposited in Co- 
leccion Nacional de Insectos (Instituto de 
Biologia, U.N.A.M.) showed that M. inter- 
media is often collected at light, and, al- 
though not much of its biology is known, 
it probably feeds on seeds of other plants. 


Myodocha unispinosa Stal 
(Figs. 2A—G) 


Egg (Fig. 2A).—Elongate, anterior pole 
rounded and posterior pole slightly pointed. 
1.1+0.04 mm long by 0.41+0.01 mm 
width. Yellowish white when laid and turns 
red in four d. Chorion almost smooth, with 
a few punctures near anterior pole. Oper- 
culum with 5—6 mycropilar processes. 

First instar (Fig. 2B).—Body elongate, 
evenly slender. Head, pronotum, and anten- 
nal segment IV dark brown; eyes bright red; 
antennal segment I pale brown with distal 
third yellowish white, antennal segments II 
and III yellowish white. Posterior margin of 
pronotum with a narrow red band. Meso- 
and metanota pale brown, thoracic pleura 
dark brown with a few red areas. Femora 
brown with distal ends slightly paler, tibiae 


367 


varying from grayish yellow to pale brown, 
tarsi generally yellowish white. Abdominal 
segments I to [V white, becoming yellowish 
near segment III, abdominal sternites I to 
IV yellowish white, with a reddish band 
near lateral margins. Remainder of abdo- 
men, dorsally and ventrally variegated with 
red and yellow areas. Scent glands of seg- 
ments II-IV, IV—V, and V—VI very small 
and pale brown. Posterior end of body 
brown. Head declivent, labium reaching ab- 
dominal sternum II; dorsal surface of body 
covered with numerous erect hairs, ventral 
surface with hairs only on abdomen. Mea- 
surements (n = 10): Body length 
1.65+0.21; head length 0.47+0.14; width 
across eyes 0.4+0.04; interocular distance 
0.26+0.01; postocular distance 0.17+0; an- 
tennal Sseements: Wy OstG==0) 015 al 
0.28+0.04, II 0.24+0.04, IV 0.44+0.09; 
hostraly sseements:) la Ol23==0202— 
O)-4o2e(0) Oils NNO tesss0) O27. IY OL a sae(0: 
pronotum length 0.17+0; width across hu- 
meral angles 0.36+0.06; width across an- 
terior margin 0.32+0.04; fore leg: femur 
length 0.44+0.12; tibia length 0.51+0.05; 
tarsi length: I 0.16+0.01, Il 0.16+0.01: 
Second instar (Fig. 2C).—Body slightly 
pyriform, maximum width across abdomi- 
nal segment IV. Head and pronotum dark 
brown; eyes brownish red. Antennal seg- 
ment II grayish yellow and segments III and 
IV reddish brown. Rostral segment I brown, 
II, II, and IV grayish yellow, apex of seg- 
ment IV brown. Meso- and metanota 
brown, sometimes lateral margins yellowish 
white. Femora and tibiae brown, tarsi gray- 
ish yellow. Abdominal segment I with a 
pale brown macula on each side of midline, 
segment II, anterior half of segment III, and 
posterior half of segment IV pale brown; a 
pale yellow band across segments III and 
IV; segments V to VIII with pale yellow 
areas surrounded by brown bands along lat- 
eral margins but extending toward scent 
gland openings. Scent gland opening of 
segments III-IV well defined with a rect- 
angular brown area; the ones of segments 
IV-V and V-VI smaller. Measurements (n = 


368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Instars of Myodocha unispinosa Stal. A, Egg. B, First instar. C, Second instar. D, Third instar. E, 
Fourth instar. K Fifth instar. G, Adult male. (Scale = 1 mm) 


VOLUME 107, NUMBER 2 


10). Body length 2.1+0.14; head length 
0.55+0; width across eyes 0.52+0.04; in- 
terocular distance 0.36+0.01; postocular 
distance 0.15+0; antennal segments: I 
0.26+0.01, II 0.42+0.04, II 0.32+0.04, Iv 
0.65+0; rostral segments: I 0.39+0.05, II 
O41=0:06; Tl 0352-0107, 1V 0:25+0:02: 
pronotum length 0.35+0.07; width across 
humeral angles 0.55+0; width across ante- 
rior margin 0.46+0.01; fore leg: femur 
length 0.67+0.07; tibia length 0.78+0.03; 
tarsi length: I 0.22+0.03, If 0.23+0.01. 

Third instar (Fig. 2D).—Similar to sec- 
ond instar, although head longer and neck 
Starts to appear. Antennal segments II and 
III and tibiae yellow, front tibia slightly pal- 
er. Rostrum reaching mesocoxae. Other 
characteristics as in second instar. Measure- 
ments (n = 10): Body length 2.55+0.21; 
head length 0.79+0.16; width across eyes 
0.52+0.04; interocular distance 0.37+0.1; 
postocular distance 0.18+0.1; antennal seg- 
ments: I 0.24+0.02, II 0.48+0.1, III 
0.44+0.09, IV 0.64+0.1; rostral segments: 
I 0.37+0.07, II 0.4+0.1, III 0.36+0.05, IV 
0.29+0.04; pronotum length 0.27+0.07; 
width across humeral angles 0.48+0.16; 
width across anterior margin 0.34+0.04; 
fore leg: femur length 0.72+0.11; tibia 
length 0.98+0.12; tarsi length: I 0.26+0.05, 
If 0.22+0.04. 

Fourth instar (Fig. 2E).—Body elongate, 
with a well-defined neck. Head dark brown, 
almost black. Antennal segments II and III 
yellowish red, IV reddish brown. Pro-, 
meso-, and metanota brown with a few ar- 
eas slightly darker, lateral borders of pro- 
and mesonota emarginate and yellow ochra- 
ceous. Legs mostly yellow ochraceous, with 
distal ends of femora grayish yellow. Ab- 
dominal segments with pale brown and yel- 
lowish white areas. A mesial brown band 
on segments II to V, quadrangular brown 
areas on segments IV and V similar in size 
to scent gland openings. ““Y”’ suture present 
and well defined anteriorly. Front femur 
with two rows of well-defined spines. Me- 
sothoracic wing pads almost covering me- 
tanotum. Measurements (n = 10): Body 


369 


length 5.2+0.56; head length 1.38+0.11; 
width across eyes 0.84+0.02; interocular 
distance 0.42+0.03; postocular distance 
0.28+0.2; antennal segments: I 0.42+0.03, 
IT 1.08+0.1, I 1+£0, IV 1.15+0.07; rostral 
segments jal OF62205(Fel FOWS=20) sil 
0.71+0.01, IV 0.38+0.05; pronotum length 
0.7+0.07; width across humeral angles 
0.92+0.1; width across anterior margin 
0.45+0; fore leg: femur length 1.6+0.14: 
tibia length 1.85+0.21; tarsi length: I 
0.54+0.09, II 0.37+0.04. 

Fifth instar (Fig. 2F).—Body elongate, 
with interocular distance less than posto- 
cular distance. Head with area around tylus 
and neck darker than remainder. Distal half 
of femora pale brown with pale yellow 
spots. Mesial band of abdominal segments 
Il to V better defined and slightly darker. 
Meso- and metathoracic wing pads reaching 
posterior margin of abdominal segment III. 
Other characteristics as in fourth instar. 
Measurements (n = 10): Body length 
7+0.85; head length 1.95+0.07; width 
across eyes 0.95+0; interocular distance 
0.45+0; postocular distance 1.05+0.07; an- 
tennal segments: I 0.57+0.03, Il 1.68+0.1, 
Ill 1.48+0.04, IV 1.75+0.02; rostral seg- 
ments: 1 OlS==Os Ti ==O107 iis 0 O7einy, 
0.52+0.04; pronotum length 0.95+0.07; 
width across humeral angles 1.05+0.07; 
width across anterior margin 0.48+0.04; 
scutellum length 0.8+0.1; scutellum width 
0:75=0; fore leg: femur length 2.25+0.21; 
tibia length 2.9+0.28; tarsi length: I 
0.9+0.14, IL 0.45+0.07. 

Adult (Fig. 2G).—Body narrow, elon- 
gate; posterior region of head forming a 
long neck. Head, pronotum, scutellum, tho- 
racic pleura, and abdominal venter black. 
Antennal segment I dark brown, segments 
I and III brown ochraceous, segment IV 
yellowish white with base and distal half 
dark brown. Rostrum mostly yellowish 
brown, with segment I and apex of IV dark 
brown. Coxae dark brown; femora dark 
brown with base white; tibiae and tarsi 
ochraceous. Clavus and chorion dark 
brown, margins and veins yellowish brown, 


370 


chorion also with a white subapical macula. 
Membrane dark brown, with yellowish api- 
cal area. Head and pronotum glabrous; pos- 
terior lobe of pronotum, clavus, and chorion 
with numerous punctures; scutellum with a 
few punctures along lateral margins. Ros- 
trum reaching mesocoxae. Front femur with 
two small and one large spine near distal 
end. Ventral surface with numerous short, 
silvery hairs, especially abundant on ab- 
dominal venter. 

Male: Measurements (n = 10): Body 
length 9.65+0.92; head length 2.85+0.35; 
width across eyes 1.12+0.03; interocular 
distance 0.5+0; interocellar distance 
0.22+0.02; postocular distance 1.48+0.45; 
antennal segments: I 0.78+0.04, IL 
2.85+0.78, Il 2.12£0.04, IV 2.18=0:04; 
rostral segments: I 0.88+0.1, IT 1.25+0.35, 
Il 1.25+0.49, IV 0.52+0.1; pronotum 
length 1.7+0.14; width across humeral an- 
gles 1.65+0.07; width across anterior mar- 
gin 0.45+0; scutellum length 1.05+0.07; 
scutellum width 0.88+0.1; fore leg: femur 
length 3.1+0.14; tibia length 3.95+0.49; 
tarsi length: I 1.25+0.07, II 0.180.02; Ill 
(O45) 220), 

Female: Measurements (n = 10): Body 
lene the9:s5=21F2-sheadmlensthyZ/>==0%3- 
width across eyes |.18+0.4; interocular dis- 
tance 0.55+0; interocellar distance 
0.24+0.2; postocular distance 1.38+0.25; 
antennal segments: I 0.78+0.17, II 
2.65+0.9, HI 2+0.14, IV 2.15+0.07; rostral 
segments: I 1.05+0.07 II 1.47+0.04, III 
1.55+0, IV 0.58+0.04; pronotum length 
1.65+0.21; width across humeral angles 
1.8+0.14; width across anterior margin 
0.45+0.07; scutellum length 1.15+0.14; 
scutellum width 0.75+0.14; fore leg: femur 
length 3.2+0.42; tibia length 3.9+0.42; tar- 
si length: I 1.25+0.07, I 0.180.05; III 
0.21+0.06. 

Biology.—This species is very common 
at Los Tuxtlas, Veracruz. It was found from 
January to December, although there was 
an increase in its population during April 
and May. Data from different localities 
showed that this species feeds on seeds of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Ficus albotomentosa Lundell, F. cotinifol- 
ia, F. maxima Mill., F. obtusifolia Kunth in 
eBoy F pertusa I ere renu som lel 
tecolutensis (Liebm). Miq., and on fruits of 
F. yoponensis Desv. that were already in 
the ground on a late decomposition stage. 
Adults and nymphs are active under the 
crown of the fruiting figs especially in areas 
where there was some ground vegetation. 
Its life cycle required about 48 d to com- 
plete. In the laboratory, eggs were laid 
mainly on a humid cotton ball or on fruit. 
Eggs were yellowish when laid and turned 
reddish after a few days; nymphs eclosed 
after eight days. First-instar nymphs molted 
to second instar after eight d; second instar 
lasted ten d, and third, fourth and fifth in- 
stars lasted between seven and eight d. 
Adults kept in the laboratory lived for two 
months. In Los Tuxtlas, this species also 
has been observed feeding on seeds of Cec- 
ropia obtusifolia Bertol. 


Neopamera bilobata (Say) 
(Figs. 3A—G) 


Egg (Fig. 3A).—Elongate with posterior 
pole slightly sharp and anterior pole round- 
ed. Pale yellow when laid, becoming red- 
dish with development of the embryo; 
0.86+0.03 mm long by 0.35+.00 mm wide 
(n = 10). Chorion with small, uniformly 
distributed punctures. Operculum with 4—5 
mycropilar processes. 

First instar (Fig. 3B).—Elongate, with 
abdomen only slightly wider than thorax. 
Head, pro-, and mesonota pale brown; eyes 
reddish brown; antennal segments I, II and 
distal half of IV pale yellow, segment III 
and proximal half of [V dark brown. Ros- 
trum and legs grayish yellow. Base of me- 
tanotum with pale brown and pale yellow 
areas. Abdominal segments I, II and prox- 
imal half of IV pale yellow, remainder of 
segment IV and segment V red; last three 
abdominal segments pale yellow with a few 
ochraceous areas along lateral margins. 
Scent gland openings between segments 
IIJ-IV, IV—V, and V—VI pale brown, elon- 
gated, narrowing caudally. Tylus reaching 


VOLUME 107, NUMBER 2 371 


Fig. 3. Instars of Neopamera bilobata Say. A, Egg. B, First instar. C, Second instar. D, Third instar. E, 
Fourth instar. E Fifth instar. G, Adult male. (Scale = | mm) 


three-fourths of first antennal segment; width across eyes 0.34+0.01; interocular 
rostrum reaching metacoxae. Head with a_ distance 0.20+0.01; postocular  dis- 
few disperse hairs especially around apex tance 0.03+0.01; antennal segments: | 
of tylus. Measurements (n = 10): Body 0.1+0.00, II 0.19+0.01, II 0.18+0.01, IV 
length 1.26+0.57; head length 0.29+0.06; 0.30+0.01; rostral segments: I 0.14+0.03, 


372 


II 0.16+0.00, II 0.13+0.02, IV 0.14+0.02; 
pronotum length 0.14+0.04; width across 
humeral angles 0.29+0.01; width across an- 
terior margin 0.29+0.02; fore leg: femur 
length 0.34+0.01; tibia length 0.38+0.02; 
tarsi length: I 0.12+0.0, II 0.15+0.01. 
Second instar (Fig. 3C).—Body elongate, 
becoming antlike because of narrower me- 
sonotum. Head and pronotum dark brown; 
antennal segment I grayish yellow, segments 
Il and II pale yellow, segment IV reddish 
brown. Rostral segment I and apex of IV 
brown, segments II and III, and rest of IV 
pale yellow. Lateral margins of meso- and 
metanota varying from pale yellow to red, 
midline pale yellow. Femora and _ tibiae 
brown, tarsi grayish yellow. Thoracic pleura 
mostly brown, with shiny red and yellow re- 
flections dorsally. Abdominal segments I 
and II with a pale brown macula on each 
side of midline, midline and lateral margins 
yellowish white; anterior three-fourths of 
segment III pale brown with a subapical red 
band transverse to union with segment IV. 
Scent gland between segments III-I'V encir- 
cled by a white band delimited in red on 
outer margins, this band also extending 
down along lateral margins of segments IV 
to VI; segments IV to VIII reddish yellow; 
rectangular scent gland openings dark brown 
with a similar rectangular red area located 
anterior to glands IV—V and V—VI. Head 
slightly declivent; dorsal surface of head, 
thorax, and abdominal segment III with a 
few long and erect hairs; antenna covered by 
numerous small hairs; abdominal venter 
with numerous caudally directed hairs. Ros- 
trum just reaching metacoxae. Measure- 
ments (n = 10): Body length 1.8+0.18; 
head length 0.38+0.02; width across eyes 
0.43+0.02; interocular distance 0.29+0.02; 
postocular distance 0.05+0.05; antennal seg- 
moemss J OMWsz20.Ol, Wl O.A75220 0), IU 
0.23+0.02, ITV 0.40+0.02; rostral segments: 
IO 22501025 O22 == 0102 Siig ON ss 0208 
IV 0.18+0.02; pronotum length 0.24+0.01; 
width across humeral angles 0.27+0.06; 
width across anterior margin 0.28+0.02; 
fore leg: femur length 0.46+0.02; tibia 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


length 0.50+0.2; tarsi length: I 0.16+0.02, 
II 0.16+0.02. 

Third instar (Fig. 3D).—Antlike, with a 
rounded head and protruding eyes. Head 
ochraceus brown; antennal segments I and 
III yellowish white, segment IV reddish 
brown. Pronotum with anterior, posterior, 
and lateral margins ochraceous, rest pale 
brown. Meso- and metanotum ocrhaceous 
with a few brown areas delimiting what are 
going to be the wings and scutellum. Fem- 
ora and tibiae ocrhaceous, tarsi pale yellow. 
Pale brown areas of abdomen turns ochra- 
ceous; white band between segments III 
and IV becomes more apparent and well de- 
fined. Rostrum reaching mesocoxae. Erect 
hairs of abdominal dorsum are bent cau- 
dally. Front femora wider than midle and 
hind femora. Measurements (n = 10): 
Body length 2.33+0.17; head length 
0.40+0.07; width across eyes 0.55+0.03; 
interocular distance 0.37+0.02; postocular 
distance 0.08+0.08; antennal segments: I 
0.16+0.02, II 0.34+0.02, Il 0.32+0.03, IV 
0.45+0.03; rostral segments: I 0.28+0.03, 
II 0.31+0.02, Ml 0.20+£0.03, IV 0.23=0.01; 
pronotum length 0.36+0.03; width across 
humeral angles 0.40+0.03; width across an- 
terior margin 0.38+0.02; fore leg: femur 
length 0.64+0.4; tibia length 0.76+0.04; 
tarsi length: I 0.21+0.03, II 0.21+0.03. 

Fourth instar (Fig. 3E).—Body elongat- 
ed, ant like shape. Head, pro-, meso-, and 
metanotum ochraceous; lateral margins of 
pro- and mesonotum dark brown. Yellowish 
white banda along union between abdomi- 
nal segments III and IV delimited frontally 
and caudally by a red band; dark brown 
area infront of scent gland of segments IV- 
V dissapear almost totally and the one in- 
front gland V-VI turns reddish; scent gland 
openning III-IV turns pale brown. Head 
slightly declivente with pop out eyes; ros- 
trum reaching mesocoxae. Mesothoracic 
wing pads covering almost totally metano- 
tum. Front femora with two rows of dark 
brown spines in number of 2 or 3. Mea- 
surements (n = 10). Body length 3.02+0.23 
head length 0.57+0.07; width across eyes 


VOLUME 107, NUMBER 2 


0.74+0.03; interocular distance 0.47+0.02: 
postocular distance 0.07+0.07; antennal 
segments: I 0.24+0.01, II 0.53+0.02, III 
0.49+0.01, IV 0.64+0.03; rostral segments: 
I 0.39+0.03, Il 0.40+0.03, HI 0.24+0.03, 
IV 0.27+0.03; pronotum length 0.57+0.03; 
width across humeral angles 0.68+0.06; 
width across anterior margin 0.53+0.03; 
scutellum length 0.31+0.03; scutellum 
width 0.42+0.02; fore leg: femur length 
0.79+0.03; tibia length 1.10+0.05; tarsi 
length: I 0.30+0.02, II 0.29+0.02. 

Fifth instar (Fig. 3F).—Elongated, an 
ochraceous general coloration, with osten- 
sible scent gland openings and white band 
between segments III and IV. Similar to 
fourth instar, although sometimes hind tibia 
are brown, as well as tarsi Il. Scent gland 
of segments III-IV turns ochraceus. Ocelli 
appear as small red spots on moulting su- 
ture. Spines of front femora turn larger and 
more numerous. Meso- and metathoracic 
wing pads reaching middle of abdominal 
segment III. Measurements (n = 10): Body 
length 4.62+0.25; head length 0.72+0.07; 
width across eyes 0.92+0.03; interocular 
distance 0.55+0.04; interocellar distance 
0.26+0.26; postocular distance 0.07+0.02; 
antennal segments: I 0.36+0.03, II 
0.74+0.04, Ill 0.72+0.03, IV 0.84+0.05; 
ROStralewseoments ll TOSOEO105%) IT 
OSs OS FSO 5220104 IVs OS 5220103; 
pronotum length 0.86+0.04; width across 
humeral angles 1.04+0.11; width across an- 
terior margin 0.66+0.06; scutellum length 
0.64+0.02; scutellum width 0.71 +0.06; 
fore leg: femur length 1.32+0.09; tibia 
length 1.61+0.11; tarsi length: I 0.51+0.02, 
II 0.34+0.04. 

Adult (Fig. 3G).—Body elongated, ant- 
like shape. Head pale brown with numerous 
silvery hairs. Antennal segments I to III 
yellowish brown, segment IV brown. Ros- 
trum yellowish brown. Pronotum varying 
from brown to yellowish brown, generally 
with lateral margins of posterior lobe dark 
brown, covered by long hairs. Propleura 
pale brown, meso-, and metapleura dark 
brown; acetabula yellowish white. Proximal 


337/53} 


half of femora yellowish brown, distal half 
dark brown; tibiae and tarsi yellowish 
brown, apices of tibiae sometimes dark 
brown. Scutellum dark brown, only lateral 
margins pale brown, surface covered by nu- 
merous silvery hairs. Chorion whitish yel- 
low with numerous brown hairs, with a 
brown band along midline and a small 
brown macula near apex. Membrane tran- 
slucid with some brown areas between 
veins. Head slightly declivent, pop out and 
large eyes. Rostrum reaching mesocoxas. 
Front femur with two rows of ventral 
spines. 

Male: Measurements (n = 10): Body 
length 5.17+0.49; head length 0.71+0.09; 
width across eyes 0.97+0.05; interocular 
distance 0.51+0.02; interocellar distance 
0.26+0.26; postocular distance 0.06+0.02; 
antennal segments: I 0.42+0.04, II 
LP US002" WLeO9S4= 0107 IWene0G= Ost 
rostral segments: I 0.53+0.05, II 0.62+0.09, 
Il 0.46+0.09, IV 0.37+0.03; pronotum 
length 1.26+0.15; width across humeral an- 
gles 1.25+0.35; width across anterior mar- 
gin 0.87+0.07; scutellum length 0.97=+0.10; 
scutellum width 0.71+0.07; fore leg: femur 
length 1.51+0.21; tibia length 1.98+0.27; 
tarsi length: I 0.65+0.03, Il 0.15+0.01; Il 
0.19+0.02. 

Female: Measurements (n = 10): Body 
length 5.15+0.36; head length 0.72+0.07; 
width across eyes 0.97+0.07; interocular 
distance 0.55+0.02; interocellar distance 
0.29+0.29; postocular distance 0.09+0.02; 
antennal segments: I 0.4+0.04, I 
0.097+0.08, III 0.83+0.08, [IV 0.96+0.06; 
rostral segments: I 0.54+0.02, Il 0.63+0.04, 
I 0.43+0.03, IV 0.37+0.02; pronotum 
length 1.14+0.06; width across humeral an- 
gles 1.23+0.17; width across anterior mar- 
gin 0.7+0.16; scutellum length 0.93+0.08; 
scutellum width 1.7+0.10; fore leg: femur 
length 1.58+0.15; tibia length 1.8620.18 
tarsi length: I 0.56+0.06, I 0.147+0.00; III 
Onl7 0102. 

Biology.—Neopamera bilobata was pre- 
sent all year around, although in Los Tux- 
tlas it was very abundant between March 


374 


and April. In the field both adults and 
nymphs were found gregariously on the leaf 
litter under fruiting fig trees. They moved 
actively, feeding on seeds of fresh and dry 
fruits. Nymphs of fourth and fifth instar and 
adults also were found between weeds. UN. 
bilobata was collected in Los Tuxtlas as- 
sociated with Ficus perforata L., F. pertu- 
sa, F. tecolutensis, and F. yoponensis, spe- 
cies that were frequently found in grazing 
areas, used as fences or for shadow for the 
cattle. In La Mancha it was found only in 
a few occasions associated with figs, al- 
though it was collected associated mainly 
with Euphorbia sp. and other weeds. In Ta- 
maulipas it was commonly found feeding 
on fig seeds contained in racoon excrement 
deposited near fig trees. Other fig species 
host of N. bilobata were: F. albotomentosa, 
F. aurea Nutt, F. calyculata P. Miller, F. 
cookii Standl., F. cotinifolia, F. maxima, F. 
obtusifolia, F. retusa. The life cycle of N. 
bilobata took around 36 d, the egg hatched 
after 7 d, it was white for a day, then turns 
yellowish and finally reddish with the de- 
velopment of the embryo. First instar 
nymphs lasted 8 d, second instar 6 d, third 
5 d, and fourth and fifth instar nymphs only 
5 d. Adults kept in laboratory lasted 22 d. 

Adults and last instar nymphs are very 
similar to ants due to the shape of their 
body, size, color, and movement. It is also 
considered a terrestrial facultative species 
according to Slater’s (1972) classification. 
It has been reported feeding on Chenopo- 
dium ambrosoides, Croton sp., Richardia 
sp., Solidago sp., Euphorbia maculata, and 
has been considered as a pest of strawber- 
ries in Florida (Slater and Baranows- 
ki,1990). Rodriguez (1998b) reported N. bi- 
lobata associted with Ficus padifolia 
H.B.K. in Costa Rica. 


DISCUSSION 


These three species of Myodochini are 
considered multivoltine due to: their po- 
lyphagous habit, the asyncronous fruting of 
their fig hosts, which provide resources al! 
year around, and regarding to their short 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


life cycles varying from 40 to 50 d. As 
some species of Ozophorini (Cervantes et 
al, in press) these Myodochini probably 
move from one fruting fig to another or to 
another host other than figs. N. bilobata 
seems to be the more polyphagous species, 
feeding on 12 species of figs and on fig 
seeds contained in racoon excrement; M. 
unispinosa feeds on eigth species; and fi- 
nally M. intermedia which was only found 
feeding on two species of figs. It was com- 
mon to found these species living sympat- 
rically, and it was also common to found 
them associated with Ozophora baranows- 
kii Slater and O’Donnell and O. maculata 
Slater and O’ Donnell; no predators of par- 
asitoids have been recorded for these My- 
odochini species. 

Nymphs of IV and V instars of the two 
Myodocha species can be differentiated 
morphologically by the coloration of the 
legs, M. intermedia nymphs have the an- 
nulated femora characteristic of the adult 
stage, while M. unispinosa femora are un- 
icolorous or have the distal two-thirds 
brown. Mesothoracic wing pads of fifth in- 
star nymphs of M. intermedia are lighter 
than in M. unispinosa 


ACKNOWLEDGMENTS 


We are very grateful to Aaron Sanchez 
for the drawings of Myodocha unispinosa 
and Neopamera bilobata and to Iliana Pa- 
checo for also recording some of the bio- 
logical information. Financial support was 
granted by a CONACYT project (34238-V) 
assigned to the first author. 


LITERATURE CITED 


Brambila, J. 2000. A review of Cligenes with the de- 
scription of a new genus, Valeris (Hemiptera: 
Rhyparochromidae: Antillocorini). Florida Ento- 
mologist 83(3): 303-315. 

Cervantes, P. L. and R. I. Pacheco. 2003. Biology and 
description of a new species of Cholula (Rhypar- 
ochromidae: Myodochini) associated with a fig in 
Mexico. Journal of the New York Entomological 
Society 111(1): 41-47. 

Cervantes, P. L., R. I. Pacheco, and P. A. Sanchez, (in 
press). Immature stages and life cycles of five spe- 
cies of Ozophora (Heteroptera: Rhyparochromi- 


VOLUME 107, NUMBER 2 


dae: Ozophorini) associated with figs in Mexico. 
Proceedings of the Entomological Society of 
Washington. 

Distant, W. L. 1880-1893. Insecta. Rhynchota. He- 
miptera-Heteroptera, Vol 1. Biol. Cent. 
London. 203 pp. 

Rodriguez, S. R. L. 1998a. Notes on the natural history 
of two Ozophora bugs (Heteroptera: Lygaeidae) 
in Costa Rica. Journal of the Kansas Entomolog- 
ical Society. 70: 203-206. 

. 1998b. Copulation and fighting behavior, and 

life cycle of Neopamera bilobata (Heteroptera: 

Lygaeidae). Revista de Biologia Tropical. 46: 

837-840. 

. 1998c. Possible female choice during copu- 

lation in Ozophora baranowskii (Heteroptera: Ly- 


Amer. 


375 


gaeidae): Female behavior, multiple copulations, 
and sperm transfer. Journal of Insect Behavior. 
11(5): 725-741. 

Say, T. 1831. Descriptions of new species of Heter- 
opterous Hemiptera of North America. New Har- 
mony, Indiana. pp 334-335. 

Slater, J. A. 1972. Lygaeid Bugs (Hemiptera: Lygaei- 
dae) as seed predators of figs. Biotropica 4(3): 
145-151. 

Slater, J. A. and R. M. Baranowski, 1990. Arthropods 
of Florida and neighboring land areas. Lygaeidae 
of Florida (Hemiptera: Heteroptera). Florida De- 
partment of Agriculture and Consumer Services. 
Doyle Conner, Commissioner. Gainesville, Flori- 
da, 211 pp. 

Stal, C. 1874. Enumeration Hemipterorum Pt. 4. K. 
Svenska Vetensk. Acad. Handl. 147 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 376-381 


TWO NEW LEAFHOPPER GENERA, DIRENAIA AND XANIONA 
(HEMIPTERA: CICADELLIDAE: TYPHLOCYBINAE: TYPHLOCYBINI) 
FROM CHINA 


YALIN ZHANG AND MIN HUANG 


Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Edu- 
cation, Entomological Museum, Northwest Sci-Tech University of Agriculture and Forestry, 
Yangling, Shaanxi 712100, China (e-mail: yalinzh@nwsuaf.edu.cn; huangmin4399 @sina. 


com) 


Abstract.—The following new taxa are described from China: Direnaia, n. gen., with 
one new species, D. quadripunctata, n. sp. and Xaniona, n. gen., with two new species, 
Xaniona galacta, n. sp. (type species) and X. cerina, n. sp. 


Key Words: 


Auchenorrhyncha, Cicadellidae, Typhlocybinae, Typhlocybini, Direnaia, 


Xaniona, new genus, new species, China 


The Typhlocybini from China were re- 
viewed by Zhang (1990) with additional 
studies by Dworakowska (1970, 1980, 
1982). This paper deals with two new ty- 
phlocybine genera of this tribe from China. 
Three new species are described and illus- 
trated: Direnaia quadripunctata, Xaniona 
galacta (type species of Xaniona), and X. 
cerina. 

All type specimens are deposited in the 
collections of the Entomological Museum, 
Northwest Sci-Tech University of Agricul- 
ture and Forestry, Yangling, China. 


Direnaia Zhang and Huang, new genus 


Type species.—Direnaia quadripunctata, 
n. sp. 

Diagnosis.—The new genus belongs to 
the Typhlocyba complex and is distin- 
guished from the similar Agnesiella Dwor- 
akowska by 1) the light body color, yellow- 
ish to yellowish white; 2) basal half of sub- 
genital plate broadened; 3) gracile macro- 
setae absent at apical part of subgenital 


plate; and 4) aedeagus without apical ven- 
tral ledge. 

Description.—Yellowish white with 1ir- 
regular yellowish patches on vertex and 
pronotum; forewing with indistinct fuscous 
pattern. Head slightiy narrower than pron- 
otum; vertex rounded to face, fore and hind 
margins subparallel, coronal suture visible. 
Forewing rather long, first apical cell small, 
second apical cell largest, widening towards 
apex, third apical cell petiolate and sub- 
triangular, stem short. Hind wing with veins 
R and M confluent apically; apical submar- 
ginal vein absent. Abdominal apodemes 
well developed. 

Posterior part of pygofer side covered 
with minute teeth at top and middle and an 
oblique ledge bridging ventral and dorsal 
margins; setosity consisting of some rigid 
microsetae at posterior margin and feeble 
microsetae scattered at ventral part. Sub- 
genital plate extending well beyond poste- 
rior margin of pygofer with basal half 
broadened and sculptured, and with a stout 
subbasal macroseta near inner side and a 


VOLUME 107, NUMBER 2 


Siill 


Figs. 1-28. 
Abdominal apodeme. 5, Male genital capsule and anal tube, lateral view. 6, Male pygofer side, lateral view. 
Paramere, connective and subgenital plate, dorsal view. 8, Subgenital plate, ventral view. 9, Connective. 10, 
Paramere. 11, Aedeagus, posterior view. 12, Aedeagus, lateral view. 


protrusion at middle of outer margin; apical 
half narrow with sparse short microsetae 
roughly in rows. Paramere long with broad 
base, distal part narrowing to apex with two 
apical processes, a row of sensorial pits at 
inner margin and several scattered short and 
rigid microsetae. Connective stem broad 
basally with a longitudinal lamellate dorsal 
ledge medially. Aedeagus with indistinct 
preatrium and well-developed dorsal apo- 
deme; shaft elongate, tubular with apical 
gonopore, a single dorsal process at apex 
and subapical posterior process. 

Etymology.—The new genus name is a 
combination of “Dr. Irena” to honor Dr. Ir- 
ena Dworakowska’s contribution to the 
knowledge of the Chinese fauna of Typhlo- 
cybinae. The gender is feminine. 


1-12, Direnaia quadripunctata. 1, Head and thorax, dorsal view. 2, Forewing. 3, Hindwing. 


SU sS 


Direnaia quadripunctata Zhang and 
Huang, new species 
(Figs. 1-12) 


Description—Length: ¢ 3.84 mm; & 
3.60 mm. Vertex and pronotum yellowish 
white with irregular yellowish patches; pair 
of blackish spots on each side of pronotum. 
Face and scutellum yellowish. Forewing 
usually with pale fuscous patches on clavus 
and at end of corium. Abdominal apodemes 
reaching to end of abdominal sternite V. Py- 
gofer side with long digitate process near 
dorsocaudal corner, directed dorsad; several 
rigid microsetae below process. Paramere 
with one long sinuate process directed me- 
dially and another short straight process di- 
rected laterally. Aedeagus with apex of dor- 
sal apodeme extending laterally on each 


378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


side; shaft evenly curved dorsally in lateral 
view, similar width throughout length ba- 
sally, slightly narrower distally, a straight 
posterior process at apical ’4, directed to- 
wards apex. 

Types.—Holotype: ¢, China, Mt. Emei 
(29.6°N, 103.4°E), Sichuan Province, alt. 
(00) tan, in avis, Now, Il, ISLS), coll, Il 
Dworakowska. Paratypes: China, 1 d, 2 &, 
Mt. Gongga (29.7°N,101.9°E), Sichuan 
Province, alt. 2,650 m, Nov. 5, 1999; 1 6, 
Lijiang (26.9°N, 100.3°E), Xinzhu, Yunnan 
Province, alt. 2,300 m, Nov. 15, 1999; 1°, 
Lijiang (26.9°N, 100.3°E), Xinzhu, Yunnan 
Province, on Juglans, Nov. 14, 1999; 2 @, 
Tengchong (25.1°N, 98.5°E), Yunnan Prov- 
ince, alt. 2,000 m, on Pinus, Rosa and 
Quercus, Nov. 24, 1999; 1 2, Tengchong 
(25.1°N, 98.5°E),Yunnan Province, alt. 
1,700 m, Nov.22, 1999; all collected by I. 
Dworakowska. 

Etymology.—The specific name is de- 
rived from the Latin words “‘quadri”’ and 
‘“‘punctata”’ referring to the four spots on 
the pronotum. 


Xaniona Zhang and Huang, new genus 


Type species.—Xaniona galacta, n. sp. 

Diagnosis.—The new genus belongs to 
the Farynala complex and is related to the 
genus Warodia but can be distinguished 
from the latter genus by 1) pygofer side 
without posterior marginal ledge; 2) pygo- 
fer side with one microseta subbasally near 
ventral margin; 3) caudal part of paramere 
shorter than the latter; and 4) apex of sub- 
genital plate curved and expanded with 
short stout setae. 

Description.—Uniformly — yellowish 
white. Head wider than pronotum; vertex 
rounded to face, fore and hind margins sub- 
parallel; coronal suture distinct, extending 
to near anterior margin. Forewing with first 
apical cell small, second apical cell largest, 
widening towards apex, third apical cell 
subtriangular, petiolate with stem short. 
Hind wing with R and M veins confluent 
apically; apical submarginal vein absent. 
Abdominal apodemes well developed. 


Genital valve long, %—% length of sub- 
genital plate. Pygofer long, sides with an 
inner middle transverse ledge in basal part 
(Figs. 20, 32) with a rigid microseta near 
ventral margin subbasally, and some short, 
rigid microsetae at lower part of posterior 
margin. Subgenital plate long, narrowing to 
apex, subtriangular with expanded tip, row 
of fine microsetae along inner margin and 
a row of short stout microsetae along outer 
margin from basal %4 to apex, a row of sharp 
peglike microsetae at outer apex and a rigid 
macroseta subbasally near outer margin. 
Paramere with elongate basal part, broad 
middle part, and a very long and stellate 
caudal part; sensorial pits distinct and scat- 
tered in a row from inner middle to sub- 
apical outer margin; some fairly long fine 
setae laterally at midlength. Connective 
large and lamellate with incomplete poorly- 
sclerotized central ridge. Aedeagus with 
basal apodeme well developed, shaft elon- 
gate with apical gonopore, pair of elongate 
apical processes strongly curved laterally to 
opposite side, crossed basally. 

Etymology.—The new genus name is de- 
rived from the Latin “‘xanion-” which 
means comblike and refers to the shape of 
apical part of subgenital plate. The gender 
is feminine. 


Xaniona galacta Zhang and Huang, 
new species 
(Figs. 13-28) 


Diagnosis.—Distinguished from the sim- 
ilar X. cerina by 1) pygofer side with an 
acute process near dorsocaudal angle; and 
2) aedeagal shaft with only one pair of pro- 
cess. 

Description.—Length: ¢ 3.45 mm; @ 
3.50 mm. Uniformly yellowish white. Ab- 
dominal apodemes strongly rounded apical- 
ly, reaching to midlength of abdominal ster- 
nite V. Genital valve about 4% length of sub- 
genital plate. Pygofer side with a spinelike 
process at dorsocaudal angle. Aedeagal 
shaft in lateral view of similar width 
throughout length and straight from near 
base to apex, with pair of elongate apical 


VOLUME 107, NUMBER 2 


379 


Figs. 13-28. 
view. 16, Forewing. 17, hindwing. 18, Abdominal apodemes. 19, Male genital capsule and anal tube, lateral 
view. 20, Male pygofer side, lateral view. 21, Posterior part of d pygofer side, lateral view. 22, Paramere, 
connective, subgenital plate and valve, dorsal view. 23, Connective. 24, Subgenital plate, ventral view. 25, Apical 
part of subgenital plate, ventral view. 26, Paramere. 27, Aedeagus, posterior view. 28, Aedeagus, lateral view. 


process, strongly curved laterally to oppo- 
site side, crossed basally. 

Types.—Holotype: d, China, Yingxiu 
(30.1°N, 101.3°E), Sichuan Province, alt. 
1,000 m, Oct. 25, 1999, coll. I. Dwora- 
kowska. Paratypes: China, 3 6, 2 2, Mt. 
Emei (29.6°N, 103.4°E), Sichuan Province, 
alt. 600 m, on Debregeasia, Nov. 2, 1999, 
coll. I. Dworakowska; 2 6, 1 2, Mt. Emei 
(29.6°N, 103.4°E), Sichuan Province, alt. 
600 m, on shrub, Oct.29, 1999, coll. I. 
Dworakowska. 

Etymology.—The specific name is de- 
rived from the Latin “‘galact-"’ which 
means milky white referring to its body col- 
or. 


Xaniona galacta. 13, Head and thorax, dorsal view. 14, Face. 15, Head and pronotum, lateral 


Xaniona cerina Zhang and Huang, 
new species 
(Figs. 29-38) 


Diagnosis.—Distinguished from the sim- 
ilar X. galacta sp. nov. by 1) pygofer side 
without a process, and 2) aedeagal shaft 
with two pairs of processes. 

Description—Length: ¢ 3.45 mm; ° 
3.60 mm. Body yellowish white. Vertex 
rounded to face, slightly produced anteri- 
orly. Abdominal apodemes_ shallowly 
rounded posteriorly, reaching to 5th abdom- 
inal sternite. Genital valve about half length 
of subgenital plate. Pygofer side without 
process. Aedeagal shaft in lateral view ta- 


380 
WSS) 
31 
30 
35 
Figs. 29-38. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Xaniona cerina. 29, Forewing. 30, Hindwing. 31, Abdominal apodeme. 32, Male pygofer side, 


lateral view. 33, Paramere, connective, subgenital plate and valve, dorsal view. 34, Subgenital plate, dorsal view. 
35, Apical part of subgenital plate, dorsal view. 36, Paramere. 37, Aedeagus, posterior view. 38, Aedeagus, 


lateral view. 


pered from base to apex, evenly curved 
dorsally, with a pair of elongate apical pro- 
cesses, strongly curved laterally to other 
side, crossed basally, and a subapical short 
straight process on each side, directed dor- 
sally. 

Types.—Holotype: ¢, Mt. Emei (29.6°N, 
103.4°E), Sichuan Province, alt. 550 m, 
Nov.2, 1999, coll. I. Dworakowska. Para- 
types: | d, 1 2, alt. 950 m, on Hortensia; 
2 3,2 2, alt. 1,500 m, Oct. 31, 1999, other 
data same as holotype. 

Etymology.—The specific name is de- 
rived from the Latin “‘cerin-’’ which 
means yellowish white referring to its 
body color. 


ACKNOWLEDGMENTS 


We give our sincere thanks to Dr. Irena 
Dworakowska (retired, Canada) for her 


help and contribution to the knowledge of 
Chinese Typhlocybinae during her visit to 
the Entomology Museum of Northwest Sci- 
Tech University of Agriculture and Forestry 
(supported by CIDA), and also to Mr. Mick 
Webb, The Natural History Museum, Lon- 
don, for revising the manuscript. This pro- 
ject was supported by “The Fok Ying Tung 
Education Foundation, National Education- 
al Committee of China.” 


LITERATURE CITED 


Dworakowska, I. 1970. On some East Palaearctic and 
Oriental Typhlocybini (Homoptera, Cicadellidae, 
Typhlocybinae). Bulletin de 1’ Academie Polonaise 
des Sciences, Serie des Sciences Biologiques 
18(4): 211-217. 

. 1980. On some Typhlocybinae from India 

(Auchenorrhyncha, Cicadellidae, Typhlocybinae). 

Entomologische Abhandlungen und Berichte aus 


VOLUME 107, NUMBER 2 381 


dem Staatlichen Museum fiir Tierkunde in Dres- lichen Museum fiir Tierkunde in Dresden 45(6): 
den 43(8): 151-201. 99-181. 

Dworakowska, I. 1982. Typhlocybini of Asia(Homop- Zhang Y. 1990. A Taxonomic Study of Chinese Ci- 
tera, Auchenorrhyncha, Cicadellidae). Entomolo- cadellidae (Homoptera). Tianze Press, Shaanxi. 


gische Abhandlungen und Berichte aus dem Staat- 218 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 382-385 


DESCRIPTION OF A NEW SPECIES OF THE GENUS APETHYMUS BENSON 
(HYMENOPTERA: TENTHREDINIDAE) FEEDING ON QUERCUS 
ACUTISSIMA CARRUTHERS (FAGACEAE) IN JAPAN 


ICH] TOGASHI 


1-chome, Honmachi, Tsurugi-machi, Ishikawa Prefecture, 920-2121, Japan 


Abstract.—Apethymus kunugi, n. sp., from Japan is described and illustrated. It was 
reared from a larva feeding on Quercus acutissima Carruthers (Fagaceae). A key is pro- 
vided for the six Japanese species of Apethymus. 


Key Words: 
Quercus acutissima 


Apethymus Benson, a genus of the sub- 
family Allantinae, is Palearctic and includes 
about 15 species (Koch 1988). Five species 
are listed for Japan by Abe and Togashi 
(1989): A. hakusanensis Togashi, 1976, A. 
hisamatsui Togashi, 1978, A. kaiensis To- 
gashi and Shinohara, 1975, A. kuri Takeu- 
chi, 1952, and A. quercivorus Togashi, 
1980. 

I had the opportunity to examine a spec- 
imen of Apethymus which was reared from 
a larva feeding on the leaves of Quercus 
acutissima Carruthers from Tochigi Prefec- 
ture, through the courtesy of Dr. K. Tanaka. 
According to examination of the Japanese 
species and the European literature (Koch 
1988), this specimen resembles A. hakusa- 
nensis and A. cereus (Klug) from Japan and 
Europe, respectively. However, this speci- 
men is separated by the coloration and the 
characters of the clypeus and the sawsheath. 
Thus, I concluded that the specimen repre- 
sents a new species, and I describe it here 
and give a key to the Japanese species. 


KEY TO JAPANESE SPECIES OF APETHYMUS 


(Females) 


[FaAntennayentinelvalblackan isin near ntrn 2 
— Apical three segments of antenna milky white 


Symphyta, Tenthredinidae, Allantinae, Apethymus, new species, food plant, 


2. Posterior margin of pronotum yellow; second 
tergite entirely black; postocellar area rather 
pentagonal; postorbital groove distinct 
SpiArarin Oras Fiat etn) Soe CONS hakusanensis Togashi 

— Pronotum black; lateral side of second tergite 
milky white (Fig. 11); postocellar area sub- 
quadrate; postorbital groove indistinct .... 

kunugi, 0. sp. 

3. Most of sixth antennal segment milky white; 
second to fifth tergites reddish yellow .... 


hisamatsui Togashi 
— Sixth antennal segment black; second to fifth 
tergites black 
4. Lateral side of third tergite milky white 
ea ene kaiensis Togashi and Shinohara 
— Third tergite black 
5. Second tergite black; apical three antennal seg- 
ments milky white; postocellar area without 
median longitudinal suture ...... kuri Takeuchi 
— Lateral side of second tergite milky white; api- 
cal portion of seventh antennal segment, apical 
half of eighth antennal segment, and ninth an- 
tennal segment except ventrally milky white; 
apical half of postocellar area with a median 
longitudinal suture ....... quercivorus Togashi 


Apethymus kunugi Togashi, new species 
(Figs. 1-13) 


Female.—Length, 12 mm. Body includ- 
ing antenna black with following from al- 
most white to milky white: labrum, tegula, 
cenchrus, lateral side of second tergite (Fig. 
11), very narrow posterior margin of third 


VOLUME 107, NUMBER 2 


Nl" ft 


2 


SS Drs 


Nag aes 4 


Figs. 1-10. Apethymus kunugi, holotype. 1, Head, dorsal view. 2, Head, lateral view. 3, Clypeus and labrum, 
front view. 4, Antenna, lateral view. 5, Mesoscutellum, dorsal view. 6, Forewing. 7, Hindwing. 8, Hind tarsus, 
lateral view. 9, Front inner tibial spur, lateral view. 10, Tarsal claw, lateral view. 


to eighth tergites, and last tergite (Fig. 11). 
Wings hyaline, stigma black with basal por- 
tion milky white; costa pale yellow with 
apical portion dark brown; other veins dark 
brown to black. Legs black with following 
milky white: hind trochanter, fore- and mid- 
tibiae except for apical third, hind tibia ex- 
cept for apical half, and all tarsi except for 
apical segment. 

Head: Transverse, slightly enlarged be- 
hind eyes (Fig. 1); interocellar furrow dis- 
tinct; postocellar furrow slightly depressed; 
lateral furrows distinct and deep; postocel- 
lar area slightly convex, subquadrate (Fig. 
1); OOL:POL:OCL = 1.2:1.0:1.7; frontal 
area nearly flattened; median fovea deep 
and elongate; lateral fovea deep and elon- 
gate; antenno-ocular distance nearly as long 
as distance between antennal sockets; su- 
praclypeal area slightly convex; basal half 
of clypeus with a median carina, anterior 
margin emarginate (Fig. 3); malar space 


narrow, nearly half as long as diameter of 
front ocellus; postorbital groove indistinct 
(Fig. 2); occipital carina distinct laterally 
(Fig. 2). Antenna nearly as long as costa of 
forewing; relative lengths of segments 1—9 
about 1.7 3150:4. 124. 122.8:2-4:2 03> lee 
pedicel longer than breadth (ratio between 
length and breadth as 1.0:0.7). 

Thorax: Median lobe of mesoscutum 
with a distinct median furrow; mesoscutel- 
lum gently convex, apical margin acute 
(Fig. 5); mesoscutellar appendage very 
short (Fig. 5); cenchrus small, distance be- 
tween them twice as long as each. Wing 
venation as in Figs. 6, 7; hind wing without 
middle cell; petiole of anal cell of hind 
wing very short. Legs with fore inner tibial 
spur as in Fig. 9; tarsal claw with lobe and 
large preapical tooth (Fig. 10); inner hind 
tibial spur slightly shorter than breadth of 
hind tibia (ratio between length of spur and 
breadth of tibia as 1.0:1.3); hind basitarsus 


11 


Figs. 11-13. 
sheath, dorsal view. 


slightly longer than following three seg- 
ments combined (Fig. 8). 

Abdomen: Sawsheath as in Figs. 12, 13; 
cercus short, half as long as sawsheath 
along dorsal margin. 

Punctation: Head and thorax distinctly, 
moderately, and closely punctured, spaces 
between punctures nearly impunctate; su- 
praclypeal area and clypeus coarsely and 
reticulately sculptured; labrum nearly im- 
punctate, shining; lower half of mesopleu- 
ron rather sparsely and finely punctured, 
spaces between punctures nearly impunc- 
tate, shining; katepimeron finely and dense- 
ly punctured, matt; abdominal tergites sha- 
greened. 

Male.—Unknown. 

Distribution.—Japan (Honshu). 

Food plant.—Quercus acutissima Car- 
ruthers (Japanese name: Kunugi). 

Holotype.—Female, emerged on 
31.X.1976 from larva feeding on leaves of 
Quercus acutissima,; Mt. Matsugamine, 
Utsunomiya City, H. Watanabe leg. Depos- 
ited in the collection of the National Sci- 
ence Museum (Nat. Hist.), Tokyo. 

Etymology.—Named after the Japanese 
name of the host plant, Kunugi, a noun in 
apposition. 

Remarks.—This new species is closely 
allied to Apethymus hakusanensis, but it is 
easily distinguished from the latter by the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


12 13 


Apethymus kunugi, holotype. 11, Abdomen, dorsal view. 12, Sawsheath, lateral view. 13, Saw- 


black pronotum (posterior margin milky 
white in A. hakusanensis), by the milky 
white macula on the second tergite (Fig. 11) 
(second tergite entirely black in A. haku- 
sanensis), by the absence of the postorbital 
groove (distinct in A. hakusanensis) and by 
the shape of the sawsheath (see fig. 5 in 
Togashi 1980 and Fig. 12). From A. cereus, 
the new species is easily distinguished by 
the black coxae (lateral side of midcoxa 
yellow in A. cereus), by the milky white 
hind trochanters (all trochanters black in A. 
cereus), and by the shape of the sawsheath 
(see fig. 3-1 in Koch 1988 and Fig. 12). 
From A. proceratis Lee and Ryu, 1996, 
from Korea, A. kunugi is easily separated 
by the mostly black abdomen and legs 
(both reddish brown in A. proceratis). 


ACKNOWLEDGMENTS 


I thank David R. Smith, USDA, Wash- 
ington, DC, for reviewing this manuscript 
and giving me copies of the valuable liter- 
ature. I also thank Dr. K. Tanaka for lending 
me the valuable specimen. 


LITERATURE CITED 


Abe, M. and I. Togashi. 1989. Tenthredinidae, pp. 
545-558. In Hirashima, Y., ed. supervisor. A 
Check List of Japanese Insects, Vol. I, pp. 541— 
1088. Entomological Laboratory, Faculty of Ag- 
riculture, Kyushu University, Fukuoka (in Japa- 
nese). 


VOLUME 107, NUMBER 2 


Koch, FE 1988. Die palaearctischen Arten der Gattung 
Apethymus Benson, 1939 (Hymenoptera, Sym- 
phyta, Allantinae). Mitteilungen der Miinchner 
Entomologische Gesellchaften 78: 155-178. 


Lee, J. W. and S. M. Ryu. 1996. A systematic study on 
the Tenthredinidae (Hymenoptera: Symphyta) from 


Korea II. Ten new species of the Tenthredinidae. 
Entomological Research Bulletin (KEI) 22: 17—34. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 386-391 


DESCRIPTION OF SCATELLA SAVEGRE, A NEW SPECIES FROM 
COSTA RICA IN THE TRISETA GROUP (DIPTERA: EPHYDRIDAE) 


WAYNE N. MATHIS AND MANUEL A. ZUMBADO 


(WNM) Department of Entomology, P.O. Box 37012, National Museum of Natural 
History, CE 619, MRC 169, Smithsonian Institution, Washington, DC 20013-7012, U.S.A. 
(e-mail: mathis.wayne@nmnh.si.edu); (MAZ) Instituto Nacional de Biodiversidad, 
A.P. 22-3100, Santo Domingo, Heredia, Costa Rica (e-mail: mzumbado @inbio.ac.cr) 


Abstract.—A new species of Scatella belonging to the triseta group, S. savegre, is 
described from specimens collected in Costa Rica (San José: Rio Savegre, San Gerardo 
de Dota (9°39.5'N, 83°51'W; 2,180 m)). For perspective, diagnoses for Scatella and the 
triseta group are provided, and to facilitate identification a key to species of the triseta 
group as well as figures of the head and wings (male and female) are provided. 


Key Words: 


Scatella Robineau-Desvoidy is among 
the most speciose genera of shore flies with 
135 species in six subgenera worldwide 
(Mathis and Zatwarnicki 1995). Of the six 
subgenera, the nominate subgenus, Scatella, 
comprises most of the diversity with 76 
species that collectively occur essentially 
worldwide in temperate and tropical zones. 
Within the subgenus Scatella, the triseta 
group has been recognized (Mathis and 
Shewell 1978) for three New World species 
(S. triseta Coquillett, S. marinensis (Cres- 
son), S. melanderi (Cresson)) that: (1) form 
a monophyletic group, (2) are easily distin- 
guished as a group from congeners, and (3) 
occur in the western United States (British 
Columbia south to Baja California Norte). 
While conducting field work in Costa Rica, 
we discovered a fourth species, which is the 
subject of this paper. This fourth species 
substantially extends the range of the triseta 
group and represents another connection 
between the shore-fly faunas occurring in 
montane habitats of Central America and 
western North America (Clausen 1987). 


Scatella, the triseta group, Ephydridae, shore flies, Costa Rica 


MATERIALS AND METHODS 


The descriptive terminology, with the ex- 
ceptions noted in Mathis (1986) and Mathis 
and Zatwarnicki (1990), follows that pub- 
lished in the Manual of Nearctic Diptera 
(McAlpine 1981). Because specimens of 
the new species are small, less than 3.5 mm 
in length, study and illustration of the male 
terminalia required use of a compound mi- 
croscope. The species’ description is com- 
posite and not based solely on the holotype. 
One head and two venational ratios that are 
used in the descriptions are defined below 
(all ratios are based on three specimens: the 
largest, smallest, and one other). Gena-to- 
eye ratio is the genal height measured at the 
maximum eye height/eye height. Costal 
vein ratio is the straight-line distance be- 
tween 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/dis- 
tance apicad of dm-cu. 

Although most specimens are in the Na- 
tional Museum of Natural History, Smith- 


VOLUME 107, NUMBER 2 


sonian Institution, Washington, DC 
(USNM), we also borrowed and studied nu- 
merous specimens from the Instituto Na- 
cional de Biodiversidad (INBio), Santo Do- 
mingo, Heredia, Costa Rica. 


Genus Scatella Robineau-Desvoidy 


Scatella Robineau-Desvoidy 1830: 801. 
Type species: Scatella buccata Robineau- 
Desvoidy, 1830 (= Ephydra_ stagnalis 
Fallén, 1813), subsequent designation of 
Coquillett 1910: 603.—Wirth 1965: 757 
[Nearctic catalog]; 1968: 24-26 [Neo- 
tropical catalog].—Mathis and Zatwarni- 
cki 1995: 262—281 [world catalog]. 


Diagnosis.—Scatella is distinguished 
from other genera of the tribe Scatellini by 
the following combination of characters: 
minute to small shore flies, body length 
0.8—3.5 mm, 2 lateroclinate fronto-orbital 
setae; face protrudent with an interfoveal 
dorsal hump; mesonotum generally unico- 
lorous or with inconspicuous longitudinal 
stripes; wings lightly to darkly infuscate or 
gray with white spots; costa long, extended 
to vein M; gonal arch divided ventrally, in 
most cases separated into 3 parts: 2 lateral 
gonites and a ventral bandlike neohypan- 
drium may be reduced. 

Description.—Head: Frons dull, usually 
with distinct subshiny to shiny mesofrons; 
lateroclinate fronto-orbital setae 2. Antenna 
short, dark; pedicel with strong seta ven- 
trally; 1st flagellomere round; arista macro- 
pubescent to at most bearing short, hairlike, 
dorsal branches. Face conspicuously pro- 
truding, with an interfoveal dorsal hump, 
uniformly sclerotized, lacking processes; 
facial setae conspicuous, usually 1—2 por- 
rect to slightly dorsoclinate setae, 4—8 ven- 
troclinate setae along oral margin. Eye 
nearly round. Gena short to moderately 
high, usually bearing a large seta; palpus 
elongate, mostly dark, occasionally yellow. 

Thorax: Mesonotum generally dark col- 
ored, microtomentose, density of microto- 
mentum varying, generally unicolorous or 
with inconspicuous longitudinal stripes 


387 


with pattern of bands and/or spots; dorso- 
central setae usually 2 (0+2), sometimes 3 
(1+2, subgenus Neoscatella); scutellum 
flat, disc bare, bearing 2 pairs of marginal 
setae; pleural region generally gray, lighter 
than mesonotum; legs typical, usually with- 
out distinct setae (ventral row of spinulae 
on midtibiae of males of some species); col- 
or of tarsi variable; stem of halter short, 
knob oval, white. Wing generally with pale 
to conspicuous white spots, especially in 
cells R,,3, R4,;, and discal cell but only oc- 
casionally in cell R,; costa long, extended 
to vein M; wing rarely brachypterous. 

Abdomen: Tergites gray to brown, mi- 
crotomentose, sometimes with lighter pos- 
terior margins, or mostly shiny, blackish 
brown. Male terminalia as follows: epan- 
drium a closed plate, narrowed ventrally; 
cerci completely round, rarely separated an- 
teriorly; gonites elongate, sharply terminat- 
ed, sometimes bearing setae on dorsal mar- 
gin of anterior portion; neohypandrium as 
a more or less sinuous band; phallapodeme 
reduced; ejaculatory apodeme large, band- 
like, broad at marginal connection to dorsal 
aedeagal opening; aedeagus shoe-shaped in 
lateral view (broader and roundedly angu- 
late basally, tapered to apex), in most spe- 
cies bearing narrow, single or paired sinu- 
ous ventral process that originates from 
ventral side of distal aedeagal margin. 

Discussion.—There are six subgenera in 
Scatella, and the triseta group is in the 
nominate subgenus Scatella, which is dis- 
tinguished from other subgenera by the fol- 
lowing combination of characters: setae of 
pedicel, if present, short, not more than 
length of arista; dorsocentral setae 2, both 
postsutural (O+2); 1 prominent, presutural 
acrostichal seta; postsutural supra-alar seta 
reduced, 2 length of postalar seta; scutellar 
disc bare; wing generally infuscate with ev- 
ident pattern of white spots; setae of mid- 
coxa short; male midfemur lacking row of 
setae; pulvilli evident, normally developed, 
tarsal claws normally developed, not con- 
spicuously elongate. 

The Neotropical fauna of the subgenus 


388 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-2. 


Scatella comprises 35 species (Mathis and 
Zatwarnicki 1995), and of these, six occur 
in Mesoamerica. Most of the known Neo- 
tropical species occur in Argentina and 
Chile, including the Juan Ferandez Islands, 
where there is considerable endemicity. 


The triseta group 


Diagnosis.—Species of the triseta group 
are distinguished by the following combi- 
nation of characters: usually larger than 
other Scatella, body length 2.20—4.00 mm. 

Description.—Head: Ocelli arranged in 
an equilateral triangle; dorsalmost postoc- 
ular setae weakly developed, subequal to 
pseudopostocellar setae; arista longer than 
combined length of scape, pedicel, and Ist 
flagellomere, micropectinate dorsally; facial 
setae extended from interfoveal hump to 
posteroventral corner of face, well devel- 
oped, subequal to length of setae along oral 
margin, often semiporrect. 

Thorax: Acrostichal setae small, hair- 
like, lacking distinctly larger pair at trans- 
verse suture; dorsocentral setae 3 (1+2); su- 
pra-alar seta rudimentary, at most % length 
of postalar seta; scutellum with 2 lateral se- 


Scatella savegre (COSTA RICA. San José: Rio Savegre (9°35'N, 83°48'W; 2,450 m). 1, Head, 
anterior view. 2, Same, lateral view. 


tae; costal margin with interspersed, slightly 
larger setulae, length of these not more than 
width of costal vein. 

Abdomen: Phallapodeme flattened dor- 
soventrally; surstylus fused indistinguish- 
ably with venter of epandrium. 

Remarks.—Mathis and Shewell (1978) 
first recognized and characterized the tri- 
seta group, which then included three Ne- 
arctic species that occur in the western 
United States and Canada. He also docu- 
mented the species group’s monophyly, 
which is based on two synapomorphies 
(their numbers 24 and 25). 


KEY TO SPECIES OF THE TRISETA 
GROUP OF SCATELLA 


1. Face uniformly dark brown; wing spots con- 
spicuous, well developed (west coast of North 
AMeriCa)\tne. 6 Ae ee S. melanderi (Cresson) 

— Face bicolored, dorsum of interfoveal hump 
brown, contrasted with grayer ventral portion; 
wing mostly hyaline to very lightly infuscate, 
white spots weakly to moderately evident ... 2 


i) 


. Wing spots not evident, male wing completely 
hyaline except for an apical, narrow brown 
spot at apex of veins R,,, and R,,; (Costa 


IRI Ga))i7s aaa Becks Glee chs S. savegre, new species 


VOLUME 107, NUMBER 2 


389 


Figs. 3-4. Scatella savegre (COSTA RICA. San José: Rio Savegre (9°35'N, 83°48'W; 2,450 m).3, Wing of 


male, dorsal view. 4, Wing of female, dorsal view. 


— Wing spots moderately evident, lacking apical 
brown spot 

3. Mesonotum and scutellum dull, microtomen- 
tose, brown with some weak golden microto- 
mentum; dorsum of scutellum flat (western 
United States and Canada) S. triseta Coquillett 

— Mesonotum and scutellum subshiny to shiny, 
very thinly microtomentose, dark brown; scu- 
tellum weakly convex (western United States 
and Canada)......... S. marinensis (Cresson) 


Scatella savegre Mathis and Zumbado, 
new species 
(Figs. 1—4) 


Description.—Moderately small to me- 
dium-sized shore flies, body length 2.20— 
3.35 mm; generally dark colored. 

Head (Figs. 1—2): Mesofrons shiny, dark 


brown, mostly bare of microtomentum ex- 
cept for narrow area immediately anterior 
of anteromedial ocellus and extended to 
frontal suture; parafrons dark brown, dense- 
ly microtomentose; fronto-orbits slightly 
darker than parafrons; ocelli arranged in al- 
most equilateral triangle with distance be- 
tween posterior pair slightly less than that 
between either posterior ocellus and antero- 
medial ocellus. Face bicolored, dorsum of 
interfoveal hump brown, contrasted with 
grayer ventral portion. Gena concolorous 
with ventral portion of face, slightly more 
than %4 eye height; gena-to-eye ratio 0.26— 
0.28. 

Thorax: Mesonotum mostly dark brown, 


390 


subshiny, thinly microtomentose, becoming 
lighter brown and more microtomentose lat- 
erally; scutellum weakly convex; postpro- 
notum very light tan to gray; notopleuron 
brown, densely microtomentose; pleural 
area generally gray, densely microtomen- 
tose, except for brown dorsal % of anepis- 
ternum and faintly brown area at ventral 
margin of anepisternum. Wing of male (Fig. 
3) mostly hyaline, no evident spots except 
for an apical, narrow, brown linear spot at 
apex of vein R,,, and R,,;; wing of female 
(Fig. 4) generally faintly infuscate with 
white markings as follows: cell r,,, with a 
linear spot at mid length and subapically; 
on either side of crossvein rm; cell r,,; with 
2 additional spots apicad of level of cross- 
vein dm-cu; |—2 irregular spots subapically 
in discal cell, cell M with a basal, C-shaped 
spot and a faint subapical spot; cell cua, 
with a spot just basad of level of crossvein 
dm-cu; crossvein rm and area immediately 
surrounding in female more darkly infus- 
cate than remainder of wing; costal vein ra- 
tio 0.17—0.19; M vein ratio 0.59—0.61. Legs 
with femora gray, slightly darker posteroap- 
ically; hindfemora shiny blackish brown 
posteriorly; tibiae blackish brown to black; 
tarsi black. 

Abdomen: Dorsum of tergites dark 
brown, subshiny; basal tergites slightly 
lighter, especially laterally; ventral portion 
of tergites becoming gray. Structures of 
male terminalia not described or illustrated 
as they are similar to other congeners of the 
triseta group. 

Type material.—The holotype male is la- 
beled “COSTA RICA. San José: Rio Sav- 
egre (9°35’N, 83°48’W; 2,450 m); 29 Jun 
2001, W. N. Mathis/USNM ENT 00187058 
[plastic bar code labelJ//HOLOTYPE 3d 
Scatella savegre W.N.Mathis & M.A. Zum- 
bado [red].”” The holotype is double mount- 
ed (minuten in a small block of plastic), is 
in excellent condition, and is deposited in 
INBio. Paratypes are as follows: bearing the 
same locality data as the holotype (13 6, 6 
2; INBio, USNM). COSTA RICA. San 
José: Rio Savegre, San Gerardo de Dota 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(9°39.5'N, 83°51’W; 2,180 m), 29-30 Jun 
2001, W. N. Mathis 9 do, 8 &; INBio, 
USNM); Rio Savegre, Cabinas Quetzal 
(933.9'N, 83°48'’W; 2,270 m), 7-8 Aug 
2001, D. and W. N. Mathis (10 6, 4 9; 
INBio, USNM); Rio Savegre (9°33'N, 
83°48.5'W; 2,180 m), 7-8 Aug 2001, D. 
and W. N. Mathis (2 6, 1 2: USNM). 

Distribution.—Neotropical. Costa Rica 
(San José). 

Natural history.—This species occurs 
along fast flowing, montane streamlets and 
rivers where there are sandy to muddy 
shorelines or where there are pockets of less 
turbulent water. We also found specimens 
at the head of a small spring or seepage area 
that was filled with emergent but low-lying 
vegetation. All specimens were netted in 
the mountains of the central cordilleras of 
Costa Rica at higher elevations between 
2,180—2,450 m. 

Etymology.—The species epithet, saveg- 
re, is to recognize the locality and drainage 
system where this species occurs and is a 
noun in apposition. 

Remarks.—Although this species sub- 
stantially extends the range of the triseta 
group and is somewhat disjunct from the 
other included species, collection of addi- 
tional species in other mountainous regions 
of Mesoamerica is anticipated. 


ACKNOWLEDGMENTS 


For reviewing a draft of this paper, we 
thank Dr. James EK Edmiston. We are grate- 
ful to Mr. Ross Simons, former Associate 
Director for Science, National Museum of 
Natural History, Smithsonian Institution, 
and to Mr. Michael A. Lang, Director of the 
Smithsonian Institution’s Marine Science 
Network, for financial support to conduct 
field work in Costa Rica. We express sin- 
cere thanks also to Mr. Jonathan M. Eibl, 
who helped prepare the head and wing pho- 
tographs. 


LITERATURE CITED 


Clausen, P. J. 1987. A new species of Pelina (Diptera: 
Ephydridae) from the Neotropical Region. Ento- 
mological News 98(1): 10-12. 


VOLUME 107, NUMBER 2 


Coquillett, D. W. 1910. The type-species of the North 
American genera of Diptera. Proceedings of the 
United States National Museum 37: 499-647. 

Mathis, W. N. 1986. Studies of Psilopinae (Diptera: 
Ephydridae), I: A Revision of the shore fly genus 
Placopsidella Kertész. Smithsonian Contributions 
to Zoology 430: iv+30 pp. 

Mathis, W. N. and G. E. Shewell. 1978. Studies of 
Ephydrinae (Diptera: Ephydridae), I: Revisions of 
Parascatella Cresson and the triseta group of Sca- 
tella Robineau-Desvoidy. Smithsonian Contribu- 
tions to Zoology 285: iv+44 pp. 

Mathis, W. N. and T. Zatwarnicki. 1990. A revision of 
the western Palearctic species of Athyroglossa 
(Diptera: Ephydridae). Transactions of the Amer- 
ican Entomological Society 116(1): 103-133. 

. 1995. A world catalog of the shore flies (Dip- 

tera: Ephydridae). Memoirs on Entomology, In- 


391 


ternational, Associated Publishers, 
Florida, 4: vit+423 pp. 

McAlpine, J. EF 1981. Morphology and terminology- 
adults, pp. 9-63. In McAlpine, J. E, B. V. Peter- 
son, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, 
and D. M. Wood, eds. Manual of Nearctic Diptera, 
Vol. 1, vit+674 pp. Agriculture Canada Mono- 
graph 27, Ottawa. 

Wirth, W. W. 1965. Family Ephydridae, pp. 734-759. 
In Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. 
Foote, and J. R. Coulson, eds. A Catalog of the 
Diptera of America north of Mexico. United 
States Departament of Agriculture, Agriculture 
Handbook 276: iv+ 1696 pp. 

. 1968. 77. Family Ephydridae, pp. 1—43. Jn 

Papavero, N., ed. A Catalogue of the Diptera of 

the Americas South of the United States. Depar- 

tamento de Zoologia, Secretaria da Agricultura. 

Sao Paulo. 


Gainesville, 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 392-397 


A CRETACEOUS PALM BRUCHID, MESOPACHYMERUS ANTIQUA, N. GEN., 
N. SP. (COLEOPTERA: BRUCHIDAE: PACHYMERINI) AND 
BIOGEOGRAPHICAL IMPLICATIONS 


GEORGE POINAR, JR. 


Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. email: 
poinarg @science.oregonstate.edu) 


Abstract.—Mesopachymerus antiqua (Coleoptera: Bruchidae), a new genus and species 
of palm seed beetles, is described from Cretaceous Canadian amber. The new genus is 
characterized by its small size (under 3 mm in length with head deflexed), head prolonged 
into a short beak, coarse eye facets, non-existent ocular sinus, complete pronotal carina, 
pro- and metatarsi segment | well expanded at apices, metafemur incrassate, pecten with 
6 denticles, prepectenal ridge with 8 spines and with the denticles and spines offset when 
the leg is flexed and metatibia positioned on the lateral side of the pecten and on the 
mesal side of the prepectenal spines. Based on this fossil, it is proposed that the Bruchidae 
arose in the Nearctic during the Jurassic or Early Cretaceous and then migrated to the 
Palearctic over the Beringia land bridge before the Oligocene. Movement into South 
America could have occurred at the end of the Cretaceous when the Proto-Greater Antilles 
formed a land bridge connecting North and South America. Palm seeds are suggested to 
be the ancestral hosts of the Bruchidae. 


Key Words: Mesopachymerus, new genus, Mesopachymerus antiqua, new species, Bru- 


chidae, Canadian amber, Cretaceous 


A beetle in Canadian amber that was 
originally considered to be a sagrine chrys- 
omelid (Poinar 1999a) was re-examined 
and determined to be a pachymerine bru- 
chid. The amber piece containing the fossil 
bruchid originated from deposits of subbi- 
tuminous coals and associated shales in 
Grassy Lake, Alberta, Canada. The amber 
sediments belong to the Foremost Forma- 
tion (Judith River Formation) within the 
Campanian of the Upper Cretaceous. These 
deposits were originally radiometrically 
dated between 70 and 80 million years (Fol- 


Bruchidae and the original host plants of 
this group. 


MATERIALS AND METHODS 


The specimen has been embedded in bio- 
plastic for stabilization and mounted on a 
microscope slide. While embedding speci- 
mens in bioplastic may stabile the amber, it 
often restricts observations. This specimen, 
which is essentially complete except for 
missing portions of some legs, was mount- 
ed facing the microscope slide and not the 
cover slip. Since the bioplastic covers all 


insbee et al. 1964) and more recently by 
Eberth and Hamilton (1993) at 79 million 
years. The present study describes the fossil 
in a new genus and discusses its signifi- 
cance regarding the origin of the family 


edges of the slide, the specimen could not 
be observed from the dorsal, ventral, ante- 
rior or posterior sides, which made viewing 
many important taxonomic characters im- 
possible. Also a milky deposit almost total- 


VOLUME 107, NUMBER 2 


ly covers the cover glass side as well as 
much of the abdomen and elytra on the 
viewing left side (Fig. 1). The description 
is based on characters observed on the 
head, antennae, pronotum and legs. 

Observations and photographs were 
made with a Nikon stereoscopic microscope 
SMZ-1tOR and Nikon Optiphot micro- 
scope™ at magnifications up to 600. All 
measurements are in microns unless other- 
wise noted. 


Mesopachymerus Poinar, new genus 


Description.—Small, under 3 mm in 
length with head deflexed; head short, pro- 
longed into a short beak; eye facets coarse; 
ocular sinus apparently non-existent; 11 an- 
tennal segments serrate (slide must be tilted 
to see this character) except scape; pronotal 
carina complete; protarsus and metatarsus 
segment | well expanded at apex; metafe- 
mur incrassate; pecten with 6 blunt tipped 
denticles; prepectenal ridge with 8 spines; 
denticles and spines offset when leg flexed; 
tibia positioned on lateral side of pecten and 
on mesal side of prepecten; metatibia with 
at least two carina; metatibia with well-de- 
veloped mucro and small spur; calcaria ab- 
sent. 

Etymology.—““Meso”’ is from Mesozoic. 
The gender is masculine. 

Diagnosis.—The non-metallic body and 
shape of the metafemur separate the speci- 
men from the Rhaebinae. The absence of 
calcaria and presence of a mucro separates 
it from the Amblycerinae. The presence of 
a pecten and a marginal line on the disk of 
the pronotum separate it from the Bruchi- 
nae and place the fossil in the Pachymeri- 
nae (Nilsson and Johnston 1993). The ab- 
sence of deep ocular sinuses and a varie- 
gated vestiture separate the specimen from 
the Caryopemini (Nilsson and Johnson 
1993). While the Pachymerini and Carye- 
dontini are obviously closely related, char- 
acters on the fossil such as the strongly ex- 
panded metatarsal segment 1, the short ros- 
trum, the coarse eye facets, the nonexistent 
ocular sinus, the structure of the pecten and 


prepecten, and the complete pronotal carina 
show its affinity with the Pachymerini. 

The shape and size of the denticles on 
the pecten and the spines on the prepecten, 
the serrate nature of antennal segments 2— 
11, the offset nature of the pectenal denti- 
cles and prepectenal spines, and the small 
size separate the new genus from all others 
in the Pachymerini. 


Mesopachymerus antigua Poinar, 
new species 
(Figs. 1—5) 


Description.—With characters listed un- 
der generic description; small species with 
body length of 2.77 mm (from frons to tip 
of abdomen; with head deflexed; if head 
porrect, then body length of 3.26 mm). In- 
tegument dull black; most of body with 
moderate to heavy pubescence. 

Head: Length = 0.60 mm, eye spherical, 
diameter = 0.23 mm; postocular lobe ab- 
sent; left antenna outstretched over thorax, 
antennal segments covered with short hairs 
and a few long hairs near apices of each 
segment, antenna 1.45 mm in length, 11 
segments serrate except linear scape: right 
antenna curved under body; maxillary palp 
3-segmented, terminal segment long, sec- 
ond segment short; labial palp 2-segmented, 
both segments short, subequal; short hairs 
on labrum and clypeus; tuft of recurved, 
scalelike hairs protruding from tip of la- 
brum. 

Thorax: Length = 0.63 mm: pronotum 
with marginal carina, covered with short 
hairs; legs brown, covered with short, thick 
pubescence; left proleg bent under thorax; 
apex of left femur and base of left tibia 
missing; apex of protibia with cluster of 11 
terminal spines; protarsus with second and 
third segments bilobed; claws paired, apical 
half curved, 52 long, with swollen base; 
metafemur incrassate, 940 in greatest length 
and 470 in greatest width; ventral side of 
pecten armed with 6 blunt-tipped denticles, 
denticle one slightly larger than remainder, 
closer to femur apex than base, following 4 
denticles subequal, denticle 6 smallest; pre- 


394 


sd 
by 


Fig. 1. 


pectenal ridge longer than pecten, with se- 
ries of 8 spines, subequal in length, proxi- 
mal two wide; metatibia evenly arcuate, 
with 2 carinae (possibly more). 

Abdomen: Length = 2.03 mm, dark, el- 
ytra with short hairs, elytra tips cover base 
of pygidium, length elytra = 1.91 mm, el- 
ytra with striations containing punctures; 
number of striations and punctures ob- 
scured by milky deposit; venter with 5 seg- 
ments. 

Female.—unknown 

Material examined.—Holotype male in 
Canadian amber, deposited at the Royal 
Tyrrell Museum in Drumheller, Alberta, 
Canada (accession number TMP 
91.148.771). 

Type locality—Deposits of subbitumi- 
nous coals and associated shales in Grassy 
Lake, Alberta, Canada. 

Etymology.—‘“‘Antiqua”’ refers to an- 
tique or old. 

Comments.—According to Nilsson and 
Johnson (1993), a widened protarsus 1 is 
indicative of male palm bruchids. In M. an- 
tiqua, protarsus | is widened and both pro- 
tarsus 2 and 3 are bilobed and heavily se- 
tose (mesotarsus 2 and metatarsus 2 are lin- 


é 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 
"44 hae 


Lateral view of Mesopachymerus antiqua in Canadian amber. Bar = 554 wm. 


ear and less setose). The specimen is con- 
sidered a male and these _ protarsal 
modifications would probably have been 
used for clasping the female. 

Diagnosis.—This is the first description 
of a Mesozoic member of the family Bru- 
chidae and represents the smallest known 
Pachymerini (Nilsson and Johnson 1993). 
Previously described fossil palm bruchids 
include several taxa from the Florissant 
shales in North America (approximately 35 
mya) (Wickham 1914) that were subse- 
quently placed in the fossil genus Oligo- 
bruchus Kingsolver (1965) and assigned to 
the tribe Caryopemini (Nilsson 1992) and 
the pachymerini, Caryobruchus dominican- 
us Poinar (1999b) from Dominican amber. 
An undescribed specimen from Lower Eo- 
cene beds in British Columbia tentatively 
has been placed in the Caryopemini (Ar- 
chibald and Mathewes 2000). 


DISCUSSION 


The present find disputes various theories 
on the origin of the Bruchidae. The sug- 
gestion by Lawrence and Newton (1982) 
that the Bruchidae evolved during the Ter- 
tiary is no longer valid. The hypothesis that 


VOLUME 107, NUMBER 2 


ee) 
\O 
Nn 


Figs. 2-5. Mesopachymerus antiqua. 2, Lateral view of entire specimen. Note that when flexed, the metatibia 
is positioned on the lateral side of the pectin and on the mesal side of the prepectenal ridge. Bar = 260 kum. 3, 
Reconstruction of left hind leg with metatibia slightly opened to show pecten and prepectenal ridge. Bar = 243 


wm. 4, Left protarsus showing bilobed condition of second and third segments. Bar = 52 xm. 5, Left metatarsus 
showing basitarsus greatly expanded at apex. Bar = 49 jm. 


396 


the Pachymerinae originated in Gondwan- 
aland (Nilsson 1992) is also challenged. An 
alternative hypothesis is that the Pachymer- 
inae arose in what is now the Nearctic Re- 
gion in the Late Jurassic or Early Creta- 
ceous when conditions were tropical- sub- 
tropical (Boucot et al., in press). During this 
period, the beetles dispersed into the Pale- 
arctic and Asia across the Beringia land 
bridge, which was present by the Late Cre- 
taceous and continued to be open intermit- 
tently throughout the Eocene and into the 
Oligocene (Lillegraven et al. 1979). The 
beetles eventually reached Africa in the 
Miocene when the African plate made con- 
tact with Southern Europe (Smith et al. 
1994). Another possible migratory route to 
the Old World could have been via the 
DeGreer and Thulean land bridges. Migra- 
tion from North America to South America 
could have occurred when the Caribbean 
Plate (Proto-Greater Antilles) made contact 
with the Chortes terrane (southern Guate- 
mala, Honduras, northern El Salvador and 
part of Nicaragua) and the Maya terrane 
(southern Mexico, Belize, and northern 
Guatemala) some 65 mya (Donnelley 
1992). Origin of the Bruchidae in North 
America is supported by an undescribed 
specimen from Lower Eocene beds in Brit- 
ish Columbia (Archibald and Mathewes 
2000) as well as fossil species of Oligobru- 
chus Kingsolver from the Florissant shales. 
The above taxa have been assigned to the 
tribe Caryopemini that is restricted to the 
Old World today. The presence of both Pa- 
chymerini and Caryopemini in Nearctic 
Cretaceous and Eocene deposits indicates 
that several types of seed beetles occurred 
in the New World by the Eocene. Migration 
of the Caryopemini into the Old World 
probably occurred by the end of the Eocene 
since northern land masses would have 
been hostile for these forms during the Ol- 
igocene when the climate cooled signifi- 
cantly (Prothero 1994). 

Since, with one possible spurious excep- 
tion, the Pachymerini are restricted to de- 
veloping in palm seeds (Johnson et al. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


1995), and it is likely that M. antiqua de- 
veloped in palm seeds. It is proposed here 
that palm seeds, and not legume seeds 
(Nilsson 1992), were the original hosts of 
bruchids. In western North America, there 
are no records of Cretaceous legumes; how- 
ever, Upper Cretaceous palm fossils belong- 
ing to several genera, including Sabal, exist 
(Tidwell 1998). Many species of Sabal are 
hosts to Caryebruchus Schonherr and Pa- 
chymerus Thunberg species today (Johnson 
et al. 1995) and a species of Sabal could 
have been the host plant of M. antiqua. 


ACKNOWLEDGMENTS 


I thank Dr. Paul Johnston at the Tyrell 
Museum in Drumheller, Alberta, Canada, 
for loaning the bruchid specimen for study 
and Roberta Poinar for comments on earlier 
drafts of this paper. 


LITERATURE CITED 


Archibald, S. B. and R. W. Mathewes. 2000. Early 
Eocene Insects from Quilchena, British Columbia, 
and their paleoclimatic implications. Canadian 
Journal of Zoology 78: 1141-1462. 

Boucot, A. J., Xu Chen, and C. R. Scotese, In press. 
Preliminary compilation of Cambrian through 
Miocene climatically sensitive deposits. Memoirs 
of the Society of Economic Paleontologists and 
Mineralogists. 

Donnelly, T. W. 1992. Geological setting and tectonic 
history of Mesoamerica, pp. 1-13. Jn Quintero, D. 
and A. Aiello, eds. Insects of Panama and Me- 
soamerica, selected studies, Oxford University 
Press, Oxford. 

Eberth, D. A. and A. P. Hamblin, 1993. Tectonic, strati- 
graphic and sedimentological significance of a re- 
gional discontinuity in the Judith River Group 
(Belly River Wedge) of southern Alberta, Sas- 
katchewan and northern Montana. Canadian Jour- 
nal of Earth Science 30: 174—200. 

Folinsbee, R., G. Baadsgaard, G. Cumming and J. Na- 
simbene, 1964. Radiometric dating of the Bear- 
paw Sea. Bulletin of the American Association of 
Petroleum Geology 48: 525. 

Johnson, C. D., S. Zona, and J. A. Nilsson, 1995. Bru- 
chid beetles and palm seeds: Recorded relation- 
ships. Principes 39: 25-35. 

Kingsolver, J. M. 1965. A new fossil bruchid genus 
and its relationships to modern genera (Coleop- 
tera: Bruchidae: Pachymerinae). Coleopterists’ 
Bulletin 19: 25-30. 

Lawrence, J. EK and A. EK Newton, Jr, 1982. Evolution 


VOLUME 107, NUMBER 2 


and classification of beetles. Annual Review of 
Systematics and Ecology 13: 261—290. 

Lillegraven, J. A., M. J. Kraus, and T. M. Brown, 1979. 
Paleogeography of the world of the Mesozoic. pp. 
277-308. In Lillegraven, J. A., Z. Kielan-Jawo- 
rowska, and W. A. Clemens, eds. Mesozoic Mam- 
mals, University of California Press, Berkeley. 

Nilsson, J. A. 1992. A taxonomic revision of the palm 
bruchids (Platymerini) and a preliminary phylo- 
genetic analysis of the world genera of Pachy- 
merinae (Coleoptera: Bruchidae), Ph.D Disserta- 
tion, Northern Arizona University, Flagstaff, Ari- 
zona, 276 pp. 

Nilsson, J. A. and C. D. Johnson, 1993. A taxonomic 
revision of the palm bruchids (Platymerini) and a 
description of the world genera of Pachymerinae. 
Memoirs of the American Entomological Society, 
No. 41, 104 pp. 

Poinar, Jr, G. O. 1999a. Chrysomelidae in fossilized 


397 


resin: behavioral inferences, pp. 1-16. In Cox, M. 
L. ed. Advances in Chrysomelidae Biology 1, Lei- 
den, Backhuys Pub. 

. 1999b. A fossil palm bruchid, Caryobruchus 
dominicanus sp. n. (Pachymerini:Bruchidae) in 
Dominican amber. Entomologica Scandinavica 
30: 210-224. 

Prothero, D. R. 1994. The Eocene-Oligocene transi- 
tion. Columbia University Press, New York. 291 
PPp- 

Smith, A. C., D. G. Smith, and B. M. Funnell, 1994. 
Atlas of Mesozoic and Cenozoic Coastlines. Cam- 
bridge University Press, Cambridge. 99 pp. 

Tidwell, W. D. 1998. Common fossil plants of Western 
North America. Second Edition. Smithsonian In- 
stitution Press, Washington, DC. 299 pp. 

Wickham, H. FE 1914. New Miocene Coleoptera from 
Florissant. Bulletin of the Museum of Compara- 
tive Zoology 58: 423-494. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 398-416 


PHYLOGENETIC ANALYSIS OF CYANOPHRYS CLENCH, A SYNOPSIS OF 
ITS SPECIES, AND THE POTENTIALLY THREATENED C. BERTHA (JONES) 
(LYCAENIDAE: THECLINAE: EUMAEIND) 


ROBERT K. ROBBINS AND MARCELO DUARTE 


(RKR) Department of Entomology, P.O. Box 37012, NHB Stop 127, Smithsonian In- 
stitution, Washington, DC 20013-7012 U.S.A. (e-mail: RobbinsR @s1.edu); (MD) Colegao 
de Lepidoptera, Museu de Zoologia, Universidade de Sao Paulo, Av. Nazaré 481, 
04263-000 Sao Paulo SP Brasil (e-mail: mduartes @usp.br) 


Abstract.—A species level phylogenetic analysis of Cyanophrys Clench was performed 
using 14 characters of the frons, wing venation, wing shape, androconia, and genitalia. 
There were 15 most parsimonious cladograms, and a successive weighting iteration re- 
sulted in three of these 15. Cyanophrys is characterized in this paper by two hypothesized 
synapomorphies (an expanded hindwing anal lobe that is tan-brown and a pair of dorsal 
brush organs), and adults can be readily identified in the field. Cyanophrys has been 
divided into Plesiocyanophrys K. Johnson, Eisele and MacPherson, Apophrys K. Johnson 
and Le Crom, Antephrys K. Johnson, Eisele and MacPherson, Mesocyanophrys K. John- 
son, and Cyanophrys (as delimited in Johnson and Le Crom 1997a), but only the mono- 
typic Antephrys is monophyletic on the most parsimonious cladograms. A synopsis of 
Cyanophrys species includes notes on their distribution, habitat, identification, nomencla- 
ture, larval food plants, and male behavior. Cyanophrys bertha, which occurs in moist 
evergreen and seasonally deciduous forests in the mountains of southern Brazil from 800 
to 1,400 m elevation, has been proposed for threatened status. It is “‘cladistically distinct”’ 
(sensu Vane-Wright et al. 1991) in that its sister is a lineage of five montane species or 
seven primarily lowland species. 


Key Words: Callophrys, phylogeny, venation, genitalia, androconia, brush organs 


Generic taxonomy of the primarily Neo- 
tropical Cyanophrys Clench (Theclinae: 
Eumaeini) has been unstable (Robbins 
2004b). Clench (1961) described it as a 
subgenus of the Holarctic Callophrys West- 
wood, but since 1993, Cyanophrys has been 
divided into five genera and subgenera, pri- 
marily on the basis of differences in geni- 
talia and color of the frons (Johnson et al. 
1993, Johnson and Le Crom 1997a). 

While most of the 16 Cyanophrys species 
are common and widespread, the Brazilian 
C. bertha (Jones) is exceedingly rare and 
has been proposed for “‘vulnerable”’ status 


(Brown 1993, Brown and Freitas 2000 and 
references therein, Otero et al. 2000). More 
recently, it has been listed as ‘‘almost 
threatened” (Mielke and Casagrande 2004). 
Little is known about C. bertha, and it is 
unclear which species, or group of species, 
is its closest relative. Such information 
might provide clues to its biology. Further, 
phylogenetic position is a factor to be con- 
sidered in assessing the status of threatened 
species (Vane-Wright et al 1991). 

The first purpose of this paper is to assess 
the monophyly of the genera and subgenera 
into which Cyanophrys has been parti- 


VOLUME 107, NUMBER 2 


tioned. To accomplish this goal, we code 
interspecific morphological variation to in- 
fer phylogenetic relations among Cyano- 
phrys species. Besides frons color and gen- 
italia, we code characters of the male fore- 
wing venation, androconia, brush organs 
(sensu Eliot 1973), and wing shape. 

Johnson and co-authors have proposed 
433 new species names in the Eumaeini 
during the past three decades, including 31 
in Cyanophrys (Robbins 2004a,b), but have 
not been careful. The adults and genitalia 
of many types are different sexes or species 
(Robbins and Lamas 2002). Data labels on 
types appear to have been switched (Rob- 
bins and Nicolay 1999, Robbins and Lamas 
2002). In one case, the two adult types of 
one species from different collections were 
similarly glued parts of species in different 
genera (Robbins and Lamas 2002). Finally, 
a number of types cannot be found (Rob- 
bins and Nicolay 2002, G. Lamas, personal 
communication). 

The publications of Johnson and co-au- 
thors on Cyanophrys contain similar prob- 
lems. For example, Johnson and Le Crom 
(1997b:23) designated a neotype for Papi- 
lio amyntor Cramer 1775 from Surinam and 
deposited it in the Natural History Museum 
(London). However, the labeled neotype de- 
posited in this collection has data labels 
from French Guiana. Another specimen 
with their neotype label in the American 
Museum (New York) has a Surinam label, 
but is a different species than the designated 
neotype in London. A second representative 
example of their lack of care is outlined in 
the synopsis below under Cyanophrys ro- 
raimiensis K. Johnson and D. S. Smith. Al- 
though we plan to address the neotype des- 
ignation elsewhere, correcting all problems 
created by Johnson and co-authors in Cy- 
anophrys would be a Sisyphean task that 
could delay publication of our current re- 
sults for years. We follow the classification 
in Robbins (2004b). 

The second purpose of this paper is to 
present a synopsis of Cyanophrys species, 
with special emphasis on C. bertha. A\l- 


399 


though identification of Cyanophrys species 
was not a goal of this study, the coded char- 
acters can be used to identify most species. 
And in the species synopsis, we note infor- 
mation on the distribution, habitat, identi- 
fication, nomenclature, behavior, larval 
food plants, and published illustrations of 
adult Cyanophrys species. We summarize 
and assess what we know about C. bertha 
in light of the phylogenetic results. 


MATERIALS AND METHODS 


Genitalia, androconia, and venation were 
examined by standard techniques (Robbins 
1991) using the approximately 1,075 spec- 
imens of Cyanophrys in the National Mu- 
seum of Natural History (USNM) as well 
as others borrowed from the American Mu- 
seum of Natural History (AMNH) in New 
York, the Illinois Natural History Survey 
(INHS) in Champaign—Urbana, The Natural 
History Museum (BMNH) in London, and 
the Universidade Federal do Parana (UFPR) 
in Curitiba. Wing venation illustrations 
were made by digital scanning of wing 
slides. We examined 82 male and 62 female 
genitalic preparations. Genitalic terms fol- 
low those in Klots (1970), wing vein names 
follow Nicolay (1971, 1977), and andro- 
conial terminology follows Robbins (1991). 

The terminal taxa are C. goodsoni 
(Clench), C. argentinensis (Clench), C. ber- 
tha, C. acaste (Prittwitz), C. amyntor (Cra- 
mer), C. fusius (Godman and Salvin), C. 
herodotus (Fabricius), C. miserabilis (Hew- 
itson), C. velezi K. Johnson and Kruse, C. 
crethona (Hewitson), C. longula (Hewit- 
son), C. pseudolongula (Clench), C. agri- 
color (Butler and H. Druce), C. banosensis 
(Clench), and C. remus (Hewitson). Cyano- 
phrys roraimiensis K. Johnson and D.S. 
Smith is omitted because it is known only 
from the holotype female, but we discuss 
its likely phylogenetic placement on the ba- 
sis of structures in the female type. As out- 
groups, we chose three species from three 
other subgenera of Callophrys recognized 
by Clench (1961); Callophrys (Callophrys) 


400 


Table 1. 
and their states are detailed in the text. 


Species | 2 3 4 5 
Callophrys rubi | O 0) O 0) 
C. gryneus 0) 0) O 0) 0) 
C. niphon 0) O 0 0 0) 
Cyanophrys goodsoni | 0) 0) 0) O 
C. argentinensis 0) | O | 
C. bertha 0) O 0) 0) I 
C. acaste | 0 | l ] 
C. amyntor | 0) O | ] 
C. fusius | 1 0 2 u 
C. herodotus | 1 0) 1 
C. miserabilis 0) 1 0) ] ] 
C. velezi O ] 0) 1 
C. crethona 0) ] 0) ] ] 
C. longula 0) O 0) 0) ] 
C. pseudolongula 0) ? O 0) 1 
C. remus 0) 1 0) O 1 
C. banosensis O 1 0) O 1 
C. agricolor 0) 1 0) O i 


OQOOoQoooqg Ss mia aw oe © 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Data matrix for Cyanophrys. The three Callophrys species are the outgroups, and the characters 


Characters 
7 8 9 10 1] 12 13 14 
0) O 0) 0) z ? u ? 
0) 0) 0) 0) 2 7 ub 2 
O O | 0) a zy ? ? 
| l 0) 0) 0) 0) O 0) 
] ] 0) 0) 0) 0) O 0) 
| 1 2 O O l 0) 0) 
| ] y 0) 0) | 0) 0) 
| 1 Dy 0) 0) | 0 O 
| 1 a 0) 0) ] 0) O 
] ] 2 1 O 0) 0) 0) 
| 1 2 ] O 0) 0) O 
] 1 2 i 0) 0) O O 
] 1 2 1 0) 0) 0) 0) 
| ] 2 0) ] 0) 1 1 
] ] 2 0) 1 0) 1 1 
i 1 2) 0) 1 0) 1 1 
] 1 2 0) 1 0) 1 1 
1 ] 1 O ] O 1 ] 


rubi (L.), C. (Mitoura) gryneus (Hiibner), 
and C. (Incisalia) niphon (Hiibner). 

We coded 14 characters (Table 1) and 
used a question mark (?) for inapplicable 
states and for one case of an intraspecific 
dimorphism. All multi-state characters were 
treated non-additively (unordered). We used 
the implicit enumeration option of Hen- 
nig86 software to derive most parsimonious 
cladograms. A strict consensus tree was de- 
termined. To test the assumption of equally 
weighted characters, a successive weighting 
iteration was performed (Farris 1969), and 
a consensus of the resulting trees was de- 
termined. Mapping of characters on trees 
was done with Winclada software (Nixon 
2002) using the “unambiguous changes 
only” optimization option. Jackknife sup- 
ports were determined in Winclada using 
Nona (1000 replications with mult*10, 
memory 1000 trees). 

The synopsis contains additional detail 
for C. bertha because of its conservation 
interest. The distribution of C. bertha is 
based on 13 museum specimens (Appen- 
dix). Its morphology is based on 6 individ- 
uals in the USNM collection (Appendix). 


Distributional information for the other spe- 
cies is based on specimens from many mu- 
seums and from the literature. Records of 
larval food plants without citation are based 
on Janzen and Hallwachs (2004) or data 
compiled by Robbins. Plant family names 
follow those in Willis (1973). Records of 
“territorial’’ behavior are based on unpub- 
lished data compiled by Robbins. Type lo- 
calities are listed in Robbins (2004b), and 
citations for original descriptions can be 
found in Lamas (1995). 


MORPHOLOGY AND CODED CHARACTERS 


Head.—Presence or absence of iridescent 
green scales on the frons is intraspecifically 
consistent. 

Character 1: Iridescent green scales on 
frons (O) absent, (1) present. 

This character needs to be used with cau- 
tion because exposure to humidity, partic- 
ularly during preparation, or to physical 
abrasion can change scale color from green 
to brown. Iridescent butterfly scales return 
to their original color after drying (Nijhout 
1991), but for reasons currently unknown, 
this is generally untrue for hairstreaks 


VOLUME 107, NUMBER 2 


401 


Fig. 1. 


(Robbins, unpublished). Also, green scales 
on the frons may be rubbed off of worn 
individuals. 

Wing pattern.—We were unable to code 
wing pattern variation in Cyanophrys for a 
number of reasons. Presence of a postme- 
dian line on the ventral forewing varies in- 
terspecifically and intraspecifically among 
species. A dark maroon band along the sub- 
margin of the ventral hindwing and dark 
brown scales at the base of the ventral 
hindwing are conspicuous wing pattern el- 
ements in some Cyanophrys species, but the 
dark maroon and brown scales occur in all 
Cyanophrys and the extent of their expres- 
sion shows almost continuous variation 
among species and sometimes within a spe- 
cies. 

Male wing venation.—Male forewing 
venation varies within Cyanophrys, and we 
code it as two characters. 

Character 2: Male forewing veins R3 


Dorsal (top) and ventral wings of male (left) and female Cyanophrys bertha. Scale | cm. 


and M1 (OQ) connected by vein UDC, which 
may be poorly developed (Figs. 3, 5—7), (1) 
fused at the origin of vein M1 (vein UDC 
absent) (Fig. 4). Males of C. pseudolongula 
are coded with a question mark because 
they are geographically variable for fore- 
wing venation, with both states occurring in 
males from Ecuador and northern Peru east 
of the Andes. 

Character 3: Male forewing veins R2 
and R3 (Q) arise from the discal cell (Figs. 
3—4, 6), (1) are stalked (Fig. 5). 

Androconia.—There are three major an- 
droconial patterns. Males of some Cyano- 


phrys species have two clusters of andro- 
conia; one at the base of veins R2, R3, and 
M1 and one along the base of vein M3 
(Figs. 4—5). Other species have androconia 
only at the base of veins R2, R3, and M1 
(Figs. 3, 6—7), although this cluster is some- 
times small, as in C. remus. Finally, males 
of C. fusius are unique among Cyanophrys 


402 


60° 50° 


Fig. 2. 


in having brown dorsal forewings (irides- 
cent blue in the others) that lack androcon- 
ia. Because it lacks androconia, C. fusius 
was coded ? (inapplicable) for characters 
5-6. 

Character 4: Male dorsal forewing (0) 
with androconia only at the base of veins 
R2, R3, and M1 (Figs. 3, 6-7), (1) with 
androconia at the base of veins R2, R3, and 
M1 and along the base of vein M3 (Figs. 
4—5), (2) lacking androconia. 

Character 5: Dorsal forewing androcon- 
ial cluster at the base of veins R2, R3, and 
M1 (Q) extends into cell R2-R3 (Figs. 6-7), 
(1) absent from cell R2-R3 (Fig. 3-5). 

Character 6: Basal edge of dorsal fore- 
wing androconial cluster at the base of 
veins R2, R3, and M1 (QO) located at, or just 
basal to, the origin of vein R2 (Figs. 3, 5— 
7), (1) extending basally well beyond the 
origin of vein R2 to, or almost to, the origin 
of vein R1 (Fig. 4). 

Wing shape and hindwing tails.-Despite 
variability in wing shape and number of 
hindwing tails, we code only one such char- 
acter. We did not code number of tails be- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


10° 


20° 


40° 30° 


Distribution of Cyanophrys bertha in southern Brazil from 13 museum specimens (Appendix). 


cause it can vary between males and fe- 
males (C. longula), geographically (C. her- 
odotus, C. acaste), or at one locality (C. 
amyntor, C. herodotus). We also did not 
code wing shape because there appears to 
be a continuum of variation from the round- 
ed wings of C. goodsoni to the angular 
wings of C. agricolor. The character below, 
however, 1s distinct and unambiguously 
scored. 

Character 7: Hindwing anal lobe (O) not 
produced posteriorly (<0O.1 mm beyond the 
outer margin of cell Cu2—2A) (Fig. 7), (1) 
produced posteriorly, at least 0.25 mm be- 
yond the outer margin of cell Cu2—2A 
(Figs. 3, 6). 

Male genitalia——We found considerable 
interspecific variation in the male genitalia 
and its associated dorsal brush organs. Var- 
iation of some structures, such as length of 
the saccus (Table 2), phallobase (internal 
part of the penis), entire penis, and tegu- 
men, was not phylogenetically useful be- 
cause differences in means between species 
were small (typically less than one standard 
deviation), which made it difficult to rec- 


VOLUME 107, NUMBER 2 403 


3 


Figs. 3-5. Forewing and hindwing venation with detail of androconial clusters. 3, Cyanophrys bertha. Arrow 
at top points to the upper discocellular vein (UDC), which arises distal of the origin of vein R2. Arrow at bottom 
points to the posteriorly produced hindwing anal lobe. 4, C. herodotus. Detail shows anastomosis of veins R3 
and M1; there is no upper discocellular vein. Arrow points to androconia along the base of vein M3. 5, C. 
acaste. Veins R2 and R3 are stalked. Arrow points to androconia along the base of vein M3. The upper 
discocellular vein arises basal of the origin of vein R2. 


ognize distinct states. We were able to code Character 8: Dorsal male genitalia brush 
other aspects of interspecific variation. Be- organs (0) absent, (1) present (Figs. 8-10). 
cause they lack brush organs, the outgroup Character 9: Male genitalia valvae (0) 


species were coded? (inapplicable) for char- with a terminal thickening that extends 
acters 11—14. along the inner margin of the valva (Fig. 3 


404 


Table 2. Mean length in mm of the saccus mea- 
sured in ventral aspect along the sagittal plane. Species 
are listed in ascending rank order with differences be- 
tween succeeding species typically less than a standard 
deviation. 


Standard 
Species Mean Length Deviation Sample Size 
C. acaste 0.53 0.050 7 
C. remus 0.54 0.036 5 
C. amyntor 0.61 0.086 5 
C. agricolor 0.64 0.042 4 
C. pseudolongula 0.68 0.036 3 
C. longula 0.69 0.035 4 
C. bertha 0.72 0) 2 
C. herodotus 0.74 0.062 7 
C. velezi 0.79 0.051 D; 
C. goodsoni 0.79 0.204 2 
C. fusius 0.81 0.091 4 
C. argentinensis 0.82 0.062 4 
C. banosensis 0.83 0) 1 
C. miserabilis 0.86 0.030 5) 
C. crethona 1.35 0.040 D} 


in Warren and Robbins 1993), (1) with a 
strongly demarcated terminal thickening 
that does not extend along the inner margin 
of the valva (Fig. 10, Fig. 3 in Warren and 
Robbins 1993), (2) with no terminal thick- 
ening (“‘not capped”’ in Clench 1961) (Figs. 
8-9). 

Clench (1961) reported valva tips with a 
terminal thickening in subgenera /ncisalia, 
Sandia Clench and Ehrlich, and Xamia 
Clench. This valva tip was illustrated by 
Warren and Robbins (1993), and also oc- 
curs in C. agricolor. A less prominent 
thickening that extends along the inner mar- 
gin of the valvae occurs in subgenera Cal- 
lophrys and Mitoura (Warren and Robbins 
1993) as well as C. goodsoni and C. argen- 
tinensis. 

Character 10: Posterior-ventral process 
of vinculum-tegumen (0) less than 0.15 mm 
in length (Figs. 8, 10), (1) with a long (> 
0.15 mm) pointed process (Fig. 9). 

Character 11: Anterior end of the setae 
comprising the dorsal male genitalia brush 
organs oriented (0) dorso-posteriorly (Figs. 
8, 9), (1) horizontally (Fig. 10). 

Character 12: Distance from anterior to 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


posterior ends of the dorsal male genitalia 
brush organs in lateral aspect (0) less than 
1.3 mm long (Figs. 9-10), (1) more than 
1.4 mm long (Fig. 8). 

Character 13: Anterior end of the dorsal 
male genitalia brush organs originates in 
lateral aspect (0) near or below the origin 
of the vinculum strut (Figs. 8—9), (1) pri- 
marily above the origin of the vinculum 
strut (Fig. 10). 

Character 14: Width of the dorsal male 
genitalia brush organs (dorsal aspect) at its 
widest point (0) less than 0.5 mm (Figs. 8— 
9), (1) more than 0.5 mm (Fig. 10). 

Female genitalia——Although length and 
width of the ductus bursae, development of 
sclerotized plates surrounding the ostium 
bursae, and length of the spines of the signa 
vary in Cyanophrys, we did not code this 
variation. For the ductus bursae, mean 
lengths and widths for species were similar, 
which made it difficult to delineate distinct 
states. Similarly, we were unable to char- 
acterize distinct states among species in de- 
velopment of the lamellae vaginalis or 
spines of the signa. In brief, interspecific 
variation in the female genitalia was not 
phylogenetically informative even though it 
was sometimes useful for distinguishing 
species, such as C. herodotus and C. amyn- 
tor. The bursa copulatrix of C. bertha is 
illustrated (Fig. 11). 


PHYLOGENETIC ANALYSES AND RESULTS 


Analysis of the character matrix with the 
Hennig&86 “‘ie*”’ option, which searches ex- 
haustively for the most parsimonious clad- 
ograms, yielded 15 equally parsimonious 
23-step trees with a consistency index of 
0.69 and retention index of 0.86. The equal- 
ly parsimonious trees differed primarily in 
the placement of C. bertha and C. fusius, 
with C. bertha being the sister of either the 
C. longula group (C. longula, C. pseudo- 
longula, C. agricolor, C. banosensis, and C. 
remus) Or C. herodotus group (C. acaste, 
C. amyntor, C. fusius, C. herodotus, C. mis- 
erabilis, C. velezi, and C. crethona). This 
result is represented in the consensus tree 


VOLUME 107, NUMBER 2 


(‘‘nelsen” option) as a trichotomy among 
C. bertha, the C. longula group, and the C. 
herodotus group, but the latter group is not 
strongly supported (Fig. 12 with jackknife 
values). The consensus tree is not one of 
the 15 most parsimonious trees. 

A successive weighting iteration (Farris 
1969) produced three trees, each of which 
was one of the 15 original most parsimo- 
nious ones. Wing venation and frons color 
(characters 1—3) were weighted four or less 
out of 10 while the other characters (4—14) 
were fully weighted (10 out of 10) except 
for “capped” valvae (character 9) and 
brush organ length (character 12). In other 
words, androconia and male genitalia char- 
acters were less likely to be homoplastic 
than those of wing venation and frons color. 
Carpenter (1988) argued that these trees are 
the best phylogenetic hypotheses. The con- 
sensus of the three successively weighted 
trees (24 steps) shows a sister relationship 
between C. bertha and the C. herodotus lin- 
eage, and there is greater resolution within 
the C. herodotus group (Fig. 13) than in the 
original consensus tree (Fig. 12). 


MONOPHYLY OF GENERA AND SUBGENERA 


Hypothesized synapomorphies for Cy- 
anophrys are (1) paired dorsal brush organs 
on the intersegmental membrane between 
the male genitalia vinculum and the 8" ab- 
dominal tergum (Character 8, Figs. 8—10) 
and (2) an elongate hindwing anal lobe with 
tan-brown coloration (Character 7, Figs. 3, 
6). These synapomorphies are unique in the 
Callophrys Section (Robbins 2004b), but 
occur in other sections of the Eumaeini. 
However, the green underside of the wings 
coupled with the elongate hindwing anal 
lobe allow adult Cyanophrys to be identi- 
fied in the field. These results do not pro- 
vide data on whether the Holarctic mem- 
bers of Callophrys (sensu Clench 1961) 
form a monophyletic group. 

Cyanophrys as characterized in this paper 
has been partitioned into subgenera (some- 
times treated as genera) Cyanophrys, Ple- 
siocyanophrys KK. Johnson, Eisele and 


405 


MacPherson, Antephrys K. Johnson, Eisele 
and MacPherson, Apophrys K. Johnson and 
Le Crom, and Mesocyanophrys K. Johnson 
(Johnson et al. 1993, Johnson and Le Crom 
1997a). The species that belong to each is 
noted (Fig. 12). Subgenus Cyanoprhys con- 
tains the C. longula group (as denoted 
above), C. miserabilis, C. velezi, C. cre- 
thona, and C. roraimiensis (the latter omit- 
ted from the phylogenetic analysis) (John- 
son and Le Crom 1997a). This grouping is 
polyphyletic in the 15 original most parsi- 
monious cladograms, as summarized by the 
consensus tree (Fig. 12). Subgenus Plesio- 
cyanophrys contains C. goodsoni and C. ar- 
gentinensis (Johnson et al. 1993), a group- 
ing that is paraphyletic in the 15 most par- 
simonious trees. Subgenus Antephrys con- 
tains only C. fusius (Johnson et al. 1993, 
Johnson and Le Crom 1997a, Johnson and 
Amarillo 1997, Robbins 2004b), making it 
a monotypic genus. Subgenus Mesocyano- 
phrys contains C. acaste and C. bertha 
(Johnson and Le Crom 1997a), a grouping 
that is paraphyletic or polyphyletic in the 
original 15 most parsimonious trees. Final- 
ly, subgenus Apophrys contains C. herod- 
otus and C. amyntor (Johnson and Le Crom 
1997a), a grouping that is not monophyletic 
in the 15 most parsimonious cladograms. 


SYNOPSIS OF SPECIES 
Cyanophrys goodsoni (Clench, 1946) 


Distribution, habitat, and abundance.— 
Usually uncommon in seasonally dry hab- 
itats from southern Texas (United States) to 
Guanacaste (Costa Rica). 

Larval food plants.—Blossom buds of 
Rivina (Phytolaccaceae) in Texas and on 
Vernonia (Compositae) in Costa Rica. 

Illustrations of adults.—Both sexes in 
Scott (1986). 


Cyanophrys argentinensis (Clench, 1946) 


Distribution, habitat, and abundance.— 
Uncommon to rare in both dry and wet for- 
ests from sea level to over 1,000 m eleva- 


tion from eastern Panama (Darien) and 


406 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 6-7. Forewing and hindwing venation with detail of androconial clusters. 6, Cyanophrys goodsoni. 
Arrow at top points to the upper discocellular vein. Arrow at bottom points to the posteriorly produced hindwing 
anal lobe. 7, Callophrys rubi. Arrow at top points to the upper discocellular vein. Arrow at bottom points to 
the hindwing anal lobe that is not posteriorly produced. 


northern Venezuela (Aragua) south along (Panama, Venezuela) to 0.25 mm (Peru) to 
the Andes to northwestern Argentina, then 0.20 mm (Argentina). 


eastward to Paraguay and southern Brazil. Male behavior.—A “‘territorial”” male was 
Variation—Width of the ductus bursae in recorded at about 1400 hours on a hilltop. 
the middle in ventral aspect appears to vary Illustrations of adults—Both sexes in 


geographically, from more than 0.30 mm _ D’Abrera (1995). 


VOLUME 107, NUMBER 2 


407 


Figs. 8-9. 


Male genitalia in lateral (left) and ventral aspect. 8, Cyanophrys bertha. B-brush organs, P-penis, 


S-saccus, T-tegumen, V-valvae, Vi-vinculum (which is fused with the tegumen in all eumaeines). 9, C. herodotus. 
The arrow points to the ventro-lateral process of the tegumen, which is foreshortened in this aspect. 


Fig. 10. 


Male genitalia of Cyanophrys agricolor in 
lateral (left) and ventral aspect. The arrow (right fig- 
ure) points to terminal thickening of valvae. ViS-vin- 
culum strut, which is an internal ridge. 


Cyanophrys bertha (Jones, 1912) . 


Distribution.—Coastal mountains of 
southern Brazil from about 800 to 1,400 m 
elevation in the states of Minas Gerais, Sao 
Paulo, Parana, and Santa Catarina (Fig. 2, 
Appendix). There are literature records 
from Rio de Janeiro (Brown 1993) and Rio 
Grande do Sul (Draudt 1919-1920), but we 
know of no extant specimens to verify the 
occurrence of C. bertha in these states. The 
elevation of the locality where one speci- 
men was supposedly collected is 75 m, but 
it is probably mislabeled (discussed in Ap- 
pendix). 

Habitat.—The habitat of C. bertha is var- 
ied. Holdridge’s (1947) ecological life 
zones of the localities in the Appendix in- 
clude subtropical lower montane moist for- 
est, subtropical moist forest, and warm tem- 
perate moist forest (Tosi 1983). Vegetation 
zones include dense evergreen forest, 
mixed evergreen forest, and seasonally de- 
ciduous forest (IBGE 1993). Although C. 
bertha appears to be primarily a resident of 


408 


ale 


Fig. 11. 


moist coastal mountains, three specimens 
found at Barbacena (Minas Gerais) indicate 
that it also occurs, at least on occasion, in 
semi-deciduous forest and may possibly be 
more widespread inland than is currently 
realized. 

Identification.—The unique ventral wing 
pattern of C. bertha (Fig. 1) has distin- 
guished this species since it was described. 

Larval food plants.—The larval food 
plant and immature stages of C. bertha are 
unknown. Larvae of other Cyanophrys have 
been reared on flowers, fruits, or young 
leaves of plants in families Anacardiaceae, 
Boraginaceae, Dipsacaceae, Compositae, 
Leguminosae, Malvaceae, Phytolaccaceae, 
Sambucaceae, Sterculiaceae, Ulmaceae, and 
Verbenaceae. 

Male behavior.—Brown (1993) observed 
males of C. bertha setting up mating terri- 
tories in the crowns of trees on hilltops at 
Serra do Japi (cf. Appendix) in the early 
afternoon. Other than C. velezi, whose 
males set up mating territories on hilltops 
before 0930 hours, the afternoon male “‘ter- 
ritorial” behavior of C. bertha is typical for 
the genus 

Illustrations of adults.—Fig. 1. 


Cyanophrys acaste (Prittwitz, 1865) 


Distribution, habitat, and abundance.—A 
very Common species in a variety of habi- 
tats and elevations from southern Brazil to 
eastern Bolivia, south to central Argentina 
and Uruguay. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


\ Y 
1mm 


l J 


Female genitalia bursa copulatrix of Cyanophrys bertha in ventral aspect. 


Variation.—Variation of the ventral wing 
pattern, particularly expression of the post- 
median line and a white hindwing discal 
bar, accounts for this species being named 
repeatedly, usually as a subspecies. Individ- 
uals from the southern temperate parts of 
the range are tailed (C. acastoides pheno- 
type). 

Larval food plants.—Chuquiraga (Com- 
positae) in Brazil. 

Male behavior.—Males “‘perch”’ on hill- 
tops from about 1100 to 1500 hours. 

Illustrations of adults——Both sexes in 
D’ Abrera (1995). 


Cyanophrys amyntor (Cramer, 1775) 


Distribution, habitat, and abundance.— 
Widespread and common from sea level to 
about 1,100 m from northern Mexico to 
southern Brazil. Recorded from Texas, 
United States (Kendall and McGuire 1984), 
based on a female in the Illinois Natural 
History Survey. Johnson and Le Crom 
(1997b) stated that it was a misidentified 
female of C. herodotus. Robbins (unpubl.) 
confirmed their identification. 

Variation.—Individuals in the southern 
parts of its range may have or lack tails, but 
do not differ otherwise. The name Cyano- 
Phrys caramba (Clench) refers to tailless 
individuals. 

Identification.—Slightly larger on aver- 
age than sympatric C. herodotus, but males 
are distinguished by different androconial 


VOLUME 107, NUMBER 2 


Callophrys (Cailophrys) rubi 


409 


1 Callophrys (Mitoura) gryneus 
O 9 
0 Callophrys (Incisalia) niphon 
C. (Plesiocyanophrys) goodsoni 
3 
i eaWwas, C. (Plesiocyanophrys) argentinensis 
1 4 C. (Mesocyanophrys) bertha 
Cyanophrys 2 
5 (54) O C. (Cyanophrys) longula 
1 C. (Cyanophrys) pseudolongula 
7 ih 13 9 (64) 
O C. (Cyanophrys) remus 
9 AP ssi pelt a4 
C. (Cyanophrys) banosensis 
2 9 
O C. (Cyanophrys) agricolor 
3 1 
O C. (Mesocyanophrys) acaste 
1 
C. (Apophrys) amyntor 
4 ama 
O C. (Antephrys) fusius 
i 2 
C. (Apophrys) herodotus 
2} 0 12 (63) 
O O C. (Cyanophrys) miserabilis 
1b har ss Le i(0) { 
O C. (Cyanophrys) velezi 
0 
C. (Cyanophrys) crethona 
Fig. 12. Strict consensus of the fifteen equally most parsimonious cladograms (23 steps, ci = 0.69, ri = 


0.86) for Cyanophrys species. Callophrys rubi, C. gryneus, and C. niphon (top) are outgroups. Character numbers 
are placed above nodes and character state numbers below nodes. Open circles represent reversal or convergence 
of the character state at that node. Jackknife values are noted in parentheses. Subgeneric placements for Cyano- 
phrys from Johnson et al. (1993), Johnson and Le Crom (1997a), and Johnson and Amarillo (1997) 


patches on the dorsal forewing. Females of 
the two have different genitalia. 

Larval food plants.—Reared from UI- 
maceae (Celtis, Trema) in Mexico, Ecuador, 
and Brazil (Kendall 1975, Hoffmann 1937), 
and from Verbenaceae (Clerodendron) in 
Trinidad. 

Male behavior.—Males set up mating ter- 
ritories along trail edges and on hilltops 
from about 1200 to 1600 hours. 
Illustrations of adults.—Male in D’Abrera 
(1995), but identity of the female that he 
illustrated as this species is unclear. 


Cyanophrys fusius 
(Godman and Salvin, 1887) 

Distribution, habitat, and abundance.— 
Seasonally dry habitats from northern Mex- 
ico to Colombia and Venezuela (Clench 
1946). 

Identification.—Males are brown above, 
not blue, and lack forewing androconia, 
both unique traits in Cyanophrys. Clench 
(1946) noted the similarity of females of 
this and the previous species, but female C. 
fusius have a short white-tipped tail at the 
end of vein Cul while Central American 


410 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
rubi 
1 ryneus 
O 9 g 
0 O f 
niphon 
i 
3 
() . . 
aye argentinensis ; 
ae longula 
6 pseudolongula 
21 8 14 
% remus 
leslie 3} 
9 9 
() . 
: agricolor 
3 
12 
acaste 
1 
4 amyntor 
4 
1 . 
fusius 
D) 2 
herodotus 
1 10 12 
niserabilis 
velezi 
crethona 
Fig. 13. Strict consensus of the three most parsimonious successively weighted trees for Cyanophrys species. 


Characters of wing venation and frons color were under-weighted because they were homoplastic. 


females of C. amyntor have only a black 
stub at the end of vein Cul. 

Larval food plants.—Reared from Arra- 
bidaea (Bignoniaceae) in Costa Rica. 

Male behavior.—A “‘territorial’’ male 
was collected on a ridge top at 1500 hours. 

Illustrations of adults.—Male in 
D’ Abrera (1995), but identity of the female 
that he illustrated as this species is unclear. 


Cyanophrys herodotus (Fabricius, 1793) 


Distribution, habitat, and abundance.— 
The most common, widespread, variable, 
and weedy species in Cyanophrys, occur- 
ring in almost all habitats under 1,500 m 
throughout the Neotropics. 


Variation.—Size and wing pattern of this 
species are highly variable, with the largest 
individuals being most frequent in the 
southern part of the range. Individuals from 
northwestern Peru, western Ecuador and 
Colombia, and northern Colombia lack a 
tail (to which the name C. detesta refers), 
but do not otherwise differ. Specimens with 
and without a tail occur sympatrically and 
synchronically in El Salvador and southern 
Brazil, but again, do not otherwise differ. 
Despite substantive genitalic variation, in- 
cluding clinal differences, we find no con- 
vincing evidence of sibling species. 

Larval food plants.—A polyphagous spe- 
cies that has been reared from plants in the 


VOLUME 107, NUMBER 2 


Anacardiaceae, Boraginaceae, Compositae, 
Dipsacaceae, Malvaceae, Sambucaceae, and 
Verbenaceae (Lima 1928, 1930, 1936: San- 
tos 1933; Monte 1934; Biezanko et al. 
1974; Robbins and Aiello 1982). 

Male behavior.—Males set up mating ter- 
ritories on hilltops and ridges in the middle 
of the afternoon from about 1300 to 1600 
hours at slightly greater heights above the 
ground, on average, than males of C. amyn- 
tor. 

Illustrations of adults.—Male_ in 
D’ Abrera (1995), but identity of the female 
that he illustrated as this species is unclear. 


Cyanophrys miserabilis (Clench, 1946) 


Distribution, habitat, and abundance.— 
Occurs in seasonally dry habitats from sea 
level to about 2,000 m elevation from 
southern United States (Texas) to the Pacif- 
ic side of Costa Rica. 

Identification.—It is easily distinguished 
from the sympatric C. herodotus by the 
brown frons (green in C. herodotus) and by 
the greater amount of dark maroon scaling 
along the outer margin of the ventral 
hindwing. Cyanophrys miserabilis appears 
to form a superspecies (monophyletic line- 
age in which no two species are sympatric) 
with the next two species (Figs. 12, 13). 
Besides the presumed evolution of a brown 
frons in the ancestor of these species, they 
also possess “thin, stringy” brush organs. 

Larval food plants.—Recorded larval 
food plants are Compositae (Eupatorium) 
and Leguminosae (Caesalpinia, Parkinson- 
ia). 

Illustrations of adults.—Both sexes in 
D’ Abrera (1995). 


Cyanophrys velezi 
Johnson and Kruse 1997 


Distribution, habitat, and abundance.— 
Occurs in Panama, where it has been re- 
corded only during the dry season, to west- 
ern Ecuador in seasonally dry habitats. 

Identification. —Unlike C. miserabilis, it 
lacks a hindwing tail and dark maroon scal- 


411 


ing along the outer margin of the ventral 
hindwing. 

Male behavior.—Males “hilltop” in the 
morning before 0930 hours, which is 
unique in the genus, so far as is known, but 
which may be shared by the preceding and 
succeeding species. 

Illustrations of adults—This species has 
been illustrated, so far as we are aware, 
only in the original description. 


Cyanophrys crethona (Hewitson, 1874) 

Distribution, habitat, and abundance.—A 
Jamaican endemic. 

Identification.—The male genitalia of C. 
crethona is similar to the preceding two 
species, but is distinguished by its larger 
size (1.e., Table 2) and a very lightly scler- 
otized ventral cornutus. 

Nomenclature.—Cyanophrys hartii 
Turner and J. Y. Miller differs from C. cre- 
thona by ventral brown coloration. Because 
exposure to humidity or physical abrasion 
can change scale color from green to 
brown, as mentioned in the discussion of 
Character 1, this name was synonymized 
with C. crethona (Robbins 2004b). 

Illustrations of adults —A male is illus- 
trated in D’ Abrera (1995). 


Cyanophrys roraimiensis Johnson and 
Smith, 1993 


Distribution, habitat, and abundance.— 
Known only from the holotype female, 
which was collected on the Brazilian side 
of Mt. Roraima. We have seen a picture of 
a female collected in the adjoining fepui re- 
gion of Venezuela that may be this species, 
but have not had the opportunity to exam- 
ine it. 

Identification.—The ventral hindwing 
pattern and size of this species is exceed- 
ingly similar to that of C. crethona 
(Huntington 1933, Comstock = and 
Huntington 1943), with which it also shares 
a brown frons, for which reason we suspect 
that it is closely related to the preceding 
three species. 

Nomenclature.—Johnson 


and Smith 


412 


(1993) wrote that Comstock and 
Huntington (1943) mentioned other speci- 
mens of Cyanophrys roraimiensis. Johnson 
and Smith (1993) then noted that these 
specimens were missing and speculated at 
length on the reasons why they had been 
‘“*stolen.’’ However, neither Huntington 
(1933) nor Comstock and Huntington 
(1943) mentioned any specimens of this 
species other than the one that was subse- 
quently designated the holotype. 

Illustrations of adults.—This species has 
not been illustrated other than in the origi- 
nal description. 


Cyanophrys longula (Hewitson, 1868) 


Distribution, habitat, and abundance.—A 
common species that occurs above 800 m 
from northern Mexico to western Panama. 
This species has been recorded as a stray in 
Arizona, United States (Bailowitz and 
Brock 1991), but no voucher specimen or 
photograph exists, so far as we are aware. 

Identification.—Males are tailless and fe- 
males have a single tail. This and the fol- 
lowing 4 species appear to form a mono- 
phyletic lineage of montane species that is 
uniquely characterized by brush organ 
structure, as described in the text (Fig. 10). 

Larval food plants.—Larvae have been 
reared from Compositae (Eupatorium) and 
Verbenaceae (Lantana, Stachytarpheta). 

Male behavior.—“‘Territorial’’ males 
have been recorded during the late morning 
(1000-1130 hours) on trees at the edge of 
a road, but the hours of such activity may 
depend largely upon good weather. 

Illustrations of adults——Both sexes in 
D’ Abrera (1995). 


Cyanophrys pseudolongula (Clench, 1944) 


Distribution, habitat, and abundance.—A 
common species above 700 m elevation in 
the Andes from Venezuela to Argentina and 
east through Paraguay to the mountains of 
southern Brazil. In the southern subtropical 
parts of its range, it may occur at lower el- 
evations. 

Variation.—Males in the northern part of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


the range (Venezuela and Colombia to 
western Ecuador and Peru) have relatively 
small androconial clusters and forewing 
veins R3 and M1 fused in contrast to those 
from the south (central Peru to southern 
Brazil, for which the name C. longuloides 
Clench was proposed) with larger andro- 
conial clusters and forewing veins R3 and 
M1 connected by vein UDC. However, 
some males in Ecuador and northern Peru 
east of the Andes have intermediate vena- 
tion (veins R3 and M1 “‘touch,”? a small 
vein UDC) as well as intermediate sized an- 
droconial clusters (although closer to the 
northern phenotype). 

Larval food plants.—Calliandra (Legu- 
minosae), Abutilon (Malvaceae), Pavonia 
(Malvaceae), and Buettneria (Sterculiaceae) 
(Biezanko et al. 1974). 

Male behavior.—‘*Hilltopping”’ males 
have been recorded from 1340 to 1415 
hours. 

Illustrations of adults——Both sexes in 
D’ Abrera (1995). 


Cyanophrys agricolor 
(Butler and Druce, 1872) 


Distribution, habitat, and abundance.— 
Occurs commonly in the mountains of Cen- 
tral America above 1,000 m elevation. 

Variation.—A short series of males col- 
lected at 1,400 m on the wet Atlantic side 
of Panama (Veraguas) are considerably 
brighter blue above, but do not differ in any 
other way. 

Male behavior.—Males have been ob- 
served setting up mating territories on trees 
lining a dirt road from 1000 to 1430 hours. 

Illustrations of adults——Both sexes in 
D’ Abrera (1995). 


Cyanophrys banosensis (Clench, 1944) 


Distribution, habitat, and abundance.—A 
montane species that usually occurs above 
1500 m from Venezuela to southern Peru. 
Although rare to uncommon in museum 
collections, it has recently been found com- 
monly in both wet and dry montane habitats 
in Ecuador (Busby and Hall, pers. comm.). 


VOLUME 107, NUMBER 2 


It probably occurs in Bolivia, but we know 
of no records. 

Illustrations of adults—A female is il- 
lustrated in D’ Abrera (1995). 


Cyanophrys remus (Hewitson, 1868) 


Distribution, habitat, and abundance.—A 
very common species in the mountains of 
southern Brazil, Argentina (Misiones), Uru- 
guay, and Paraguay. In the subtropical parts 
of its range, it occurs at lower elevations. 

Identification.—Males have brilliant dor- 
sal blue color, similar to that in C. longula 
and C. pseudolongula, but the underside 
wing pattern has extensive brown markings, 
similar to the preceding two species. 

Larval food plants.—It has been reared 
from Calliandra (Leguminosae) in Uruguay 
and from Abutilon and Pavonia (Malva- 
ceae) in Brazil (Zikan 1956, Biezanko et al. 
1966, Silva et al. 1968). 

Male behavior.—Males “‘hilltop”’ in the 
early afternoon, with records from 1145 to 
1500 hours. 

Illustrations of adults——Both sexes in 
D’ Abrera (1995). 


DISCUSSION 


Building on the work of Clench (1961), 
Robbins (2004a) distinguished the Callo- 
phrys Section of the Eumaeini by valve tips 
that are flattened and without setae, but 
sometimes with a terminal thickening (il- 
lustrated in Warren and Robbins 1993). The 
form of the anal lobe (Character 7) and 
presence of brush organs (Character 8) dis- 
tinguish Cyanophrys from other members 
of the Callophrys Section. Although the ge- 
nus could be split into smaller genera on 
the basis of the phylogenetic results (Figs. 
12 and 13), the anal lobe character allows 
individuals to be recognized in the field. It 
is unclear whether the remaining Holarctic 
members of Callophrys, or the many genera 
into which they have been divided, are 
monophyletic. 

The subgeneric nomenclature used by 
Johnson and colleagues (Johnson et al. 
1993, Johnson and Le Crom 1997a, John- 


413 


son and Amarillo 1997) is inconsistent with 
the phylogenetic results (Fig. 12). Four of 
these subgenera are not monophyletic and 
One is monotypic. In some cases, such as 
the aptly named Plesiocyanophrys, they ap- 
pear to have been characterized with sym- 
plesiomorphies. In others, such as the char- 
acterization of Cyanophrys (a brown frons), 
they were delimited by a homoplastic char- 
acter. 

The C. longula and C. herodotus groups 
are monophyletic on the 15 most parsimo- 
nious trees. The C. Jongula group consists 
of species with a brown frons (Character 1, 
which is homoplastic), brush organs wider 
than 0.05 mm (Character 14, Fig. 10), brush 
organs that are oriented horizontally at the 
anterior end (Character 11, Fig. 10), and 
brush organs that arise primarily above the 
origin of the vinculum strut (Character 13, 
Fig. 10). The latter three character states are 
unique in the Callophrys Section. These 
species occur in montane habitats (above 
800 m) except in the subtropical parts of 
southern South America. The C. herodotus 
group consists of species with two andro- 
conial clusters on the dorsal forewing 
(Character 4, Figs. 4—5, both lost in C. fu- 
sius according to our results). All seven 
species occur in the lowlands, unlike the 
previous lineage, but most are also found in 
montane habitats. 

Cyanophrys bertha is the sister of the 
seven-species C. herodotus lineage in the 
successively weighted cladogram (Fig. 13), 
but in the equally weighted original most 
parsimonious trees, it was sometimes the 
sister of the five-species C. /ongula lineage. 
The genitalia of C. bertha are nearly iden- 
tical to those of C. amynitor, but C. bertha 
shares frons color, androconial structure, 
and restriction to montane habitats with C. 
longula (Table 1). 

Cyanophrys bertha has been proposed 
for “‘vulnerable’’ status (Brown 1993, 
Brown and Freitas 2000 and references 
therein, Otero et al. 2000) and listed as *‘al- 
most threatened”’ (Mielke and Casagrande 
2004). Its relatively basal position in the 


414 


cladogram increases its priority as a species 
of conservation concern (Atkinson 1989, 
Vane-Wright et al. 1991). 


ACKNOWLEDGMENTS 


We are grateful to Keith Brown for his 
kind hospitality, for generously sharing his 
knowledge of C. bertha, and for providing 
us with study specimens. We thank Mary 
Sangrey and the NMNH Research Training 
Program for providing an internship to 
Duarte; the Women’s Committee of the 
Smithsonian Institution and the National 
Science Foundation (award #BIR-953 1331) 
for supporting this program; Olaf Mielke 
and Mirna Casagrande for obtaining CNPq 
support for Robbins to curate the UFPR 
collection in Curitiba; Karl Ebert for allow- 
ing us to examine his father’s collection; the 
curators and collections managers at the 
AMNH, BMNH, UFPR, INHS for allowing 
us to examine specimens under their care; 
the many biologists who shared larval food 
plant information with us; Carl Hansen for 
technical help in digitizing wing venation 
directly from slides; George Venable for 
making the figures; Astrid Caldas for pro- 
viding a smooth interface between English 
and Portuguese; and especially Andrew 
Brower, John Brown, Keith Brown, Astrid 
Caldas, Larry Gall, Jason Hall, Donald Har- 
vey, Gerardo Lamas, James Miller, Carla 
Penz, J. B. Sullivan, and Andy Warren for 
commenting, sometimes extensively, on the 
manuscript. The junior author has been fi- 
nancially supported by the State of Sao 
Paulo Research Foundation—FAPESP (02/ 
13898-0 and 03/05895-4). 


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doptera, Vol. 5A. Association for Tropical Lepi- 

doptera, Scientific Publishers, Gainesville, xxxvi 

+ 439 pp. 

. 2004b. Lycaenidae. Theclinae. Tribe Euma- 
eini, pp. 118-137. In Lamas, G., ed. Checklist: 
Part 4A. Hesperioidea—Papilionoidea. /n He- 
ppner, J. B., ed. Atlas of Neotropical Lepidoptera, 
Vol. 5A. Association for Tropical Lepidoptera, 
Scientific Publishers, Gainesville, xxxvi + 439 pp. 

Robbins, R. K. and A. Aiello. 1982. Foodplant and 
oviposition records for Panamanian Lycaenidae 
and Riodinidae. Journal of the Lepidopterists’ So- 
ciety 36: 65-75. 

Robbins, R. K. and G. Lamas. 2002. Nomenclatural 
Changes in the Neotropical Eumaeini (Lepidop- 
tera: Lycaenidae: Theclinae). Revista brasileira de 
Zoologia 19: 197-214. 

Robbins, R.K. and S.S. Nicolay. 1999. Taxonomy of 


Strymon toussainti, S. andrewi, S. amonensis, and 


S. rhaptos (Lycaenidae: Theclinae). Journal of the 
Lepidopterists’ Society 52: 328-334. 
. 2002. An overview of Strymon Hiibner (Ly- 


caenidae: Theclinae: Eumaeini). Journal of the 
Lepidopterists’ Society 55: 85-100. 

Santos, E. 1933. Inimigos e Doengas das Fruteiras. Rio 
de Janeiro, Biblioteca Agricola d?O Campo. 81 
PP: 

Scott, J. A. 1986. The butterflies of North America. A 


416 


Natural History and Field Guide. Stanford Uni- 
versity Press, Stanford, xv + 583p. 

Silva, A. G. d’A. e, C. R. Gongalves, D. M. Galvao, 
A. J. L. Gongalves, J. Gomes, M. do N. Silva, and 
L. de Simoni. 1968. Quarto catalogo dos insetos 
que vivem nas plantas do Brasil seus parasitos e 
predadores. Edigaéo ampliada do ‘*3° catalogo dos 
insetos que vivem nas plantas do Brasil’ de au- 
toria do Prof. A. M. da Costa Lima. Parte II. In- 
setos, hospedeiros e inimigos naturais. Indice de 
insetos e indice de plantas. Rio de Janeiro, Min- 
istério da Agricultura. 1: xxvii + 622 pp.; 2: [viii] 
+ 265 pp. 

Tosi, J. Jr. 1983. Provisional life zone map of Brazil 
at 1:5,000,000 scale. Publication of the Institute 
of Tropical Forestry, Rio Piedras, Puerto Rico, 16 
pp. + map. 

Vane-Wright, R. I., Humphries, C. J. and P. H. Wil- 
liams. 1991. What to protect?—-Systematics and 
the agony of choice. Biological Conservation 55: 
235-254. 

Warren, A. D. and R. K. Robbins. 1993. A natural 
hybrid between Callophrys (Callophrys) sherida- 
nit and C. (Uncisalia) augustinus (Lycaenidae). 
Journal of the Lepidopterists’ Society 47: 236— 
240. 

Willis, J. C. 1973. A Dictionary of the Flowering 
Plants and Ferns. 8th edition revised by H. K. 
Airy Shaw. University Press, Cambridge, 1,245 
Pp. 

Zikan, J. F 1956. Beitrag zur Biologie von 12 Thecli- 
nen-arten. Dusenia 7: 139-148. 


APPENDIX 


The data for the 13 museum specimens 
of C. bertha from 7 localities in southern 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Brazil that we have examined are listed be- 
low (museum acronyms listed in Methods). 
If the elevation was not recorded on a spec- 
imen’s data label, we parenthetically note 
the elevation for that site as it is listed in 
gazetteers. So far as we are aware, there are 
no other known specimens of C. bertha. 


1. Minas Gerais, Barbacena, 900 m. 12 
(Private collection of Karl Ebert, Santa 
Clara, Sao Paulo, Brazil); 1 2, 11 May 
1969 (USNM); 12, 27 May 1986 
(USNM). 

2. Minas Gerais, Pocgos de Caldas, 1,300— 
1,400 m. 1 3d and 1 &, March (Ebert). 

3. Sao Paulo, Serra do Japi, 1,050—1,250 
m. 1 36, 24 March 1990 (USNM); 1 6, 
25 March 1990 (USNM); 1 6, 14 May 
1990 (USNM); 1 6, 2 June 1990 
(USNM). 

4. Parana, Curitiba, (934 m). 1d, Decem- 
ber 1945 (UFPR). 

5. Parana, Ponta Grossa, (969 m). 

March 1948 (UFPR). 

Parana, Castro, (999 m). 12 (BMNH) 

7. Santa Catarina, Corupaé, 19 October 
1975. 12 (UFPR). (Corupa is located at 
75 m elevation at the base of the moun- 
tains on the road leading to Sao Bento 
do Sul, a well-known butterfly locality 
above 800 m elevation that is the likely 
collection place for this specimen.) 


[oe 


oN 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 417-427 


RECORDS OF FLEAS (SIPHONAPTERA) OF CARNIVORES FROM IDAHO 
MICHAEL W. HASTRITER AND MICHAEL E WHITING 


(MWH) Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, 
P.O. Box 20200, Provo, UT 84602-0200, U.S.A. (e-mail: mwhastriter@sprintmail.com); 
(MFW) Department of Integrated Biology, 693 Widsoe Building, Brigham Young Uni- 
versity, Provo, UT 84602, U.S.A. 


Abstract. Records of fleas from carnivores trapped at six localities in Valley county, 
Idaho, during November through February, 2002—2004, are presented. Additional records 
collected from small mammals by the senior author over a period of 26 years are also 
included. Twenty-eight species of fleas representing 24 genera and 6 families are docu- 
mented. Three new state records [Delotelis telgoni (Rothschild, 1905) from Idaho County, 
Nearctopsylla hyrtaci (Rothschild, 1904) from Valley County, and Peromyscopsylla ham- 
ifer vigens (Jordan, 1937) from Valley and Idaho counties] and 18 additional new county 


records are reported (15 for Valley County and 3 for Idaho County). 


Key Words: 


Few studies have been conducted to as- 
sess the flea fauna in Idaho. Most notewor- 
thy are those of Allred (1968, 1971) from 
the National Reactor Testing Station (cur- 
rently the National Engineering Laboratory 
located in Bingham, Bonneville, Butte, 
Clark, and Jefferson counties), Hubbard 
(1943, 1947), Lewis et al. (1988), and Yen- 
sen et al. (1996). Saunders (1978) provided 
a preliminary checklist, while Baird and 
Saunders (1992) published a comprehen- 
sive checklist. In the latter, Baird and Saun- 
ders report descriptions of nine currently 
valid taxa from Latah (seven), Elmore 
(one), and Fremont (one) counties and list- 
ed ten species from Valley County. 

Eads et al. (1979) demonstrated the ex- 
treme diversity (18 species) encountered on 
only three species of carnivores [Lynx rufus 
(Schreber, 1777), Bassaricus astutus (Lich- 
tenstein, 1830) and Martes americana (Tur- 
ton, 1806)] in Larimer County, Colorado. 
Flea species listed by Eads et al. are all ac- 
cidental associations reflecting carnivore- 


Siphonaptera, carnivore fleas, [Idaho 


prey relationships, e.g., bobcats [L. rufus] 
acquiring the fleas (Cediopsylla inaequalis 
ssp.) off the animals they routinely feed 
upon (lagomorphs). Although a well known 
concept, few studies have focused on the 
diversity of fleas found on carnivores to as- 
sess specific flea fauna. We report similar 
diversity and cite new county and state rec- 
ords for Idaho. The senior author opportu- 
nistically collected in Valley County spo- 
radically over 26 years. These records are 
also included. 


MATERIALS AND METHODS 


During the winter months of November 
through February 2002—2004, James Ba- 
con, a resident fur trapper of McCall, Valley 
County, Idaho, trapped fur-bearing mam- 
mals. Larger animals, e.g., bobcats, foxes, 
and coyotes, were trapped with No. 1% 
jump traps and American martens (MM. 
americana) were trapped with No. | long 
spring traps. Traps were baited with car- 
casses of animals skinned from previous 


418 


catches and were checked within 48 hours 
after sets. Captured animals were skinned, 
and their pelts were immediately rolled up, 
placed in plastic bags, and put into a freezer 
for later processing. All pelts, except those 
of martens, were stored in individual bags. 
Marten pelts were bagged singly or as mul- 
tiples when the number captured in a day 
exceeded more than one. Following the 
trapping season, processing of pelts was 
completed and the pelage and bags were ex- 
amined for fleas. Fleas were preserved in 
80 percent ethanol, processed by conven- 
tional mounting techniques, and identified 
by the senior author. 

Carnivore trapping localities in Valley 
County included an area between Payette 
Lake and Upper Payette Lake, 45°05'N, 
116°03'W (1), vicinity of Brundage Moun- 
tain, 45°01’N, 116°07'’ (2), Lake Fork 
Drainage on the North Fork of Lake Fork, 
44°07'N, 115°57'W (3), Lake Fork Drain- 
age, North Fork of Payette River, 44°51’'N, 
116°04'W (4), South Fork of the Salmon 
River, Forest Service Trail #50674, 
44°53'N, 115°42'W (5), and South Fork of 
the Salmon River near Warm Lake, 
44°41’N, 115°42'W (6). Fleas were also 
taken from two cougars [Puma concolor 
(Linnaeus 1771)] from the Gold Fork area, 
near Donnelly 44°41'N, 115°56’W (7) and 
from the West Mountains, ~44°30'N, 
116°O5’W (8) by Craig Derrick, a local 
taxidermist (Cascade, Idaho). These locali- 
ties (numerals in parentheses) precede each 
record under “‘Carnivore Records.’ New 
county and state records are indicated by 
single and double asterisks, respectively. 
Specific records collected by the senior au- 
thor are listed under the category “‘Miscel- 
laneous Records.” All localities are from 
Valley County except for a few noted from 
Idaho County under Miscellaneous Rec- 
ords. Mammal synonymies follow those of 
Wilson and Reeder (1993). Fleas are re- 
tained in the personal collection of the se- 
nior author. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


RESULTS AND DISCUSSION 
Ceratophyllidae 
Aetheca wagneri (Baker, 1904)* 


Miscellaneous records.—Payette Lake, 
(45°N, 116°05’W), ex Peromyscus mani- 
culatus (Wagner, 1845), 11 September 
1997, 2 3; Upper Payette Lake, (45°08'N, 
116°O1’W), ex P. maniculatus, 13 August 
1997, 5 6. 


Amardix bitterrootensis bitterrootensis 
(Dunn, 1923)* 


Carnivore records.—(1), ex M. ameri- 
cana, 20 December 2002, | 2. (4), ex Vul- 
pes vulpes Linnaeus, 1758, 3 November 
2003, 1 &. (5), ex M. americana, 28 Jan- 
uary 2004, 1 do. 

Remarks.—This flea is rare in collec- 
tions, possibly because it is a winter flea 
found at high elevations occurring primarily 
on pikas [Ochotona princeps Richardson, 
1828)] whose habitats are snow-covered 
during most of the winter months. Hubbard 
(1941) reported 17 3 and 13 @ from 100 
pikas, seldom obtaining more than one on 
any individual animal. There are a number 
of records also from the bushy-tailed wood- 
rat [Neotoma cinerea (Ord, 1815)], which 
often lives in close association with pikas 
in rocky slopes. Only one report (Holland, 
1985) indicates a carnivore host (Mustela 
erminea invicta = M. erminea Linnaeus, 
1758). It 1s not surprising to find this flea 
during the winter months from carnivores 
that feed on both pikas and N. cinerea in 
our trapping areas. This is noted by the 
large numbers of woodrat fleas (see Orcho- 
peas agilis) and one pika flea (see Cteno- 
phyllus armatus armatus) collected from M. 
americana. A curious record of 10 ¢ and 
13 2 of A. b. bitterrootensis from a single 
Mus musculus Linnaeus, 1758 was noted by 
Holland (1985) (season not listed). 


Amonopsyllus ciliatus kincaidi 
(Hubbard, 1947)* 
Miscellaneous records.—Wagon Wheel 
Road (44°58’'N, 116°06’W), ex Tamiasciu- 
rus hudsonicus 4 August 1998, 2 @. 


VOLUME 107, NUMBER 2 


Ceratophyllus niger C. Fox, 1908* 


Miscellaneous records.—Little Payette 
Lake (44°56'N, 116°02’W), ex Woodpecker 
nest cavity (1.25 inch opening), 27 May 
1996, 2 2; ex Woodpecker nest cavity (1.5 
inch opening), 28 May 1996, 1 36,1 9. 


Eumolpianus eumolpi (Rothschild, 1905) 


Miscellaneous records.—Wagon Wheel 
Road (44°58'N, 116°06'W), ex Spermophi- 
lus lateralis (Say, 1823), 4 August 1998, 3 
6; ex T. hudsonicus, 3 C. 


Foxella ignota ignota (Baker, 1895) 


Carnivore records.—(1), ex M. ameri- 
cana, 20 December 2002, 5 @ and ex V. 
vulpes, 5 December 2002, 1 2. (2), ex M. 
americana, 6 November 2003, | @. (5), ex 
M. americana, 14 December 2003, 2 9; and 
ex L. rufus, 19 December 2003, 1 &. (6), ex 
M. americana, | January 2003, 2 @. (7), ex 
P. concolor, 20 December 2002, 1 . 

Miscellaneous records.—North edge of 
Little Payette Lake (44°56'N, 116°02’W), 
1,555 m, ex Thomomys bottae (Eydoux and 
Gervais, 1836), 27 and 29 May 1996, 3 6, 
2 2; McCall (44°53'N, 116°06’W), 1,520 
m, ex JT. bottae, 13 and 14 July 2000, 2 3, 
One 

Remarks.—The common host for this 
flea is the pocket gopher (Thomomys sp.). 
The senior author has trapped this plentiful 
host species in several of the trapping areas. 
It is smaller than varieties occurring at low- 
er elevations and thus creates burrows with 
very small diameters that would preclude 
entry of an animal as large as M. ameri- 
cana. The data suggest that martens prey on 
pocket gophers frequently. During the win- 
ter months, pocket gophers frequently leave 
their subterranean burrows extending their 
tunnel systems into the strata of snow (ev- 
idenced by tubes of earth left behind on the 
ground’s surface following spring thaws). 
This perhaps provides an added opportunity 
for martens as well as other carnivores to 
prey upon them. 

Baird and Saunders (1992) reported a 


419 


single male of F. ignota recula (Jordan and 
Rothschild, 1915) from Valley County (not 
examined). A single male of 13 specimens 
from carnivores and 5 males of 15 speci- 
mens from 7. bottae represents F. i. ignota 
(not F. 7. recula) based on diagnostic cri- 
teria noted in Holland (1985). Female F. 
ignota subspecies are indistinguishable and 
are only tentatively assigned in our study to 
the nominate subspecies. 


Megabothris abantis (Rothschild, 1905) 


Miscellaneous records.—Josephine Lake, 
2,287 m, Idaho Co. (45°13’N, 115°58’W), 
ex Clethrionomys gapperi (Vigors, 1830), 
23 October 1999, 3 6, 1 9; ex Microtus 
sp ld, See sUpper Payette. ake 
(45°08'N, 116°01’W), ex Microtus sp., 13 
August 1997, 1 9. 

Remarks.—Baird and Saunders (1992) 
recorded this species in Valley County from 
American mink (Mustela vison Schreber, 
1777); however, it had not been reported 
previously from Idaho County. 


Monopsyllus vison Baker, 1904 
(Fig. 1) 


Carnivore records.—(3), ex M. ameri- 
cana, 10 November 2003, 1 2 (1 d without 
host, date, or locality data). 

Miscellaneous records.—NW of Wagon 
Wheel Road, (44°58'N, 116°06’W), ex T. 
hudsonicus, 27 May 1996, 1 6,5 2, 4 Au- 
gust 1998, | 2, and ex woodpecker nest 
cavity (1.5 inch hole), 28 May 1996, 1 @. 

Remarks.—Haas and Wilson (1982) 
found M. vison to be the dominant flea in- 
festing red squirrel (7. hudsonicus) nests in 
Alaska. Holland (1985) also listed exten- 
sive records (some inclusive of Haas and 
Wilson’s work) of M. vison on red squirrels. 
The red squirrel is the main staple of M. 
americana, but only two specimens of M. 
vison were recovered in our study. This 
may be explained in Fig. 1, which is assem- 
bled from host records compiled in Holland 
(1985). Only specimens of M. vison taken 
from 7. hudsonicus are included in Fig. 1, 
while those from nests, predators, and mis- 


420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 
200 
Ej Males 
180 Females 
160 : 
140 
n 
5 120 
ae 
xs 
mn 0)0) 
1) 
Oo 
5 
F 80 
60 
40 
20 
0 TLE oe 
| HP TE WA WE SWAT WANT ANI (PDS OXI = AVL 
Months 
Fig. 1. Seasonal abundance of males and females of Monopsyllus vision north of the U.S. and Canadian 


border (raw data from Holland 1985). 


cellaneous other hosts are excluded because 
they are considered accidental associations. 
Populations of M. vison are minimal during 
the winter months. Among Holland’s rec- 
ords, M. americana was cited as a host to 
M. vison only four times. These were dur- 
ing the months of June (1 6d), October (4 
3, 1 2), and November (3 6, 1 2). The 
occurrence of M. vison on martens primar- 
ily during the winter months might be re- 
flective of: 1) increased trapping by fur 


trappers during winter months; 2) the mar- 
ten’s diet of red squirrels increases during 
the heavy snow-cover of winter (even 
though flea numbers are low) when other 
prey are scarce; or 3) a combination of 
both. The sex ratio (male/female) of 1,025 
adults from Holland’s records (Fig. 1) is 1: 
1.29. Valley County records in Baird and 
Saunders (1992) were collected from an 
American mink (M. vison) and T. hudson- 
icus. 


VOLUME 107, NUMBER 2 


Opisodasys keeni (Baker, 1896)* 


Miscellaneous records.—McCall 
(44°53'’N, 116°06’W), ex P. maniculatus, 
11 September 1997, 1 2; Upper Payette 
Lake (45°08’N, 116°01’W), ex P. manicu- 
latus, 13 August 1997, 1 6. 


Orchopeas agilis (Rothschild, 1905)* 


Carnivore records.—(1), ex M. ameri- 
cana, 20 December 2002, 1 2; 4 November 
2003, 3 6, 13 &. (2), ex M. americana, 6 
November 2003, 1 3. (3), ex M. ameri- 
cana, 10 November 2003, 1 @. (4), ex M. 
americana, 10 November 2003, 1 <o. (5), 
ex L. rufus, 10 January 2004, 1 2, 4 Feb- 
ruary 2004, 1 d, 1 2; ex M. americana, 20 
December 2003, 5 2, 14 December 2003, 
5 6, 13 2, 6 danny ZOO, i 6, 3 2, 2B 
January 2004, 13 ¢, 19 &. (6), ex M. amer- 
icana, | January 2003, 1 ¢ (1 6, 1 2 with- 
out host, date, or locality data). 

Remarks.—Orchopeas agilis was given 
full specific status by Lewis (2000). This 
flea is a common and widespread species 
occurring on woodrats. Neotoma cinerea, a 
high elevation species, is the sole species 
occurring in the trapping areas of this study. 
Holland (1985) also found this species 
commonly on N. cinerea in southern British 
Columbia and Alberta. The distribution of 
this species extends from these two Cana- 
dian provinces to western Texas, and the 
southern borders of New Mexico and Ari- 
zona. Neotoma cinerea is the primary host 
throughout its range, but at lower elevations 
it readily parasitises other species of wood- 
rats (Morlan, 1955). The bushy-tailed 
woodrat appears to be a significant part of 
the diet of M. americana. 


Orchopeas caedens (Jordan, 1925) 


Carnivore records.—(1), ex V. vulpes, 5 
December 2002, 2 6, 1 2; ex M. ameri- 
cana, 4 November 2003, 1 @. (2), ex M. 
americana, 6 November 2003, 2 &. (3), ex 
M. americana, 10—12 November 2003, | d, 
2 &. (4), ex V. vulpes, 3 November 2003, 
1 3d. G), ex EL. rufus, 19 December 2003, 2 


421 


2; 30 December 2003, 4 9, 4 February 
2004, 1 2; ex M. americana, 14 December 
2003, 5 2, 20 December 2003, 1 6, 3 @, 
6 January 2004, 1 2, 28 January 2004, 1 
2. (6), ex M. americana, 1 January 2003, 
3 2 (1 3,4 & without host, date, or locality 
data). 

Miscellaneous records.—Wagon Wheel 
Road, (44°58'N, 116°06’W), ex T. hudson- 
icus, 4 August 1998, 1 6; McCall, 
(44°53’N, 116°06’W), ex T. hudsonicus, 27 
May sl 996sIe Sas 

Remarks.—Lewis (2000) synonymized 
Orchopeas caedens durus Jordan, 1929, 
with O. caedens. Haas and Wilson (1982) 
listed this flea second only to M. vison in 
red squirrel nests in Alaska. Females of O. 
agilis and O. caedens are difficult to distin- 
guish and, based on males, both were pre- 
sent simultaneously on M. americana 
throughout the trapping areas. Based on the 
key (couplet 4) provided in Lewis (2000), 
the frontal setal row has one or no setae in 
O. caedens, while O. agilis always has two 
or more. In the latter, there were seldom 
more than two in Valley County popula- 
tions, and the second more ventral seta was 
often minute. The caudal margin of st. VII 
(seventh sternite) is highly variable in both 
species; however, based on the dicotomy 
created by the frontal setal row, specimens 
attributed to O. caedens usually have an 
acutely pointed dorsal lobe on the caudal 
margin of the st. VII. 


Oropsylla idahoensis (Baker, 1904)* 


Miscellaneous records.—Silver Creek, 
near Silver Creek community hot springs 
plunge (44°20'N, 115°47'W), ex Spermo- 
philus columbianus (Ord, 1815), 13 and 15 
hubby nl S74 Seely oe 


Oropsylla tuberculata tuberculata 
(Baker, 1904) 


Miscellaneous records.—Silver Creek, 
near Silver Creek community hot springs 
plunge (44°20'N, 115°47'W), ex S. colum- 
bianus, 15 July 1974, 2 o. 

Remarks.—Baird and Saunders (1992) 


422 


recorded two pairs of this flea from S. col- 
umbianus in Valley County. 


Thrassis pandorae pandorae 
Jellison, 1937 


Miscellaneous records.—Silver Creek, 
near Silver Creek community hot springs 
plunge (44°20'N, 115°47'W), ex S. colum- 
bianus, 13 and 15 July 1974, 1 6,7 &. 

Remarks.—Stark (1970) recorded 7. p. 
pandorae from Valley County, and Baird 
and Saunders (1992) repeated Stark’s rec- 
ord. 


CTENOPHTHALMIDAE 


Catallagia decipiens Rothschild, 1915* 
(Figs. 2A—C, G, H) 


Miscellaneous records.—Josephine Lake, 
2,287 m, Idaho Co. (45°13’N, 115°58W), ex 
Microtus sp., 23 October 1999, 1 3. Upper 
Payette Lake (45°08N, 116°01'W), ex P. 
maniculatus, 13 August 1997, 1 @. 

Remarks.—These are the first records for 
both Idaho and Valley counties. This female 
has an aberrant spermatheca, which resem- 
bles that of Delotelis telgoni (Rothschild, 
1905) but is smaller (Figs. 2C, E). The hilla 
does not project into the bulga, as in all 
species of Catallagia Rothschild, 1915 
(Fig. 2B). The spiracle on t. VIII is char- 
acteristic of Catallagia (compare spiracle of 
Catallagia and Delotelis Figs. 2F—H) and 
there are only two rows of frontal setae ver- 
sus the typical three in Delotelis (see Figs. 
2A, D). Similar observations of an aberrant 
spermatheca were noted by Hubbard (1947) 
and these were illustrated in Hopkins and 
Rothschild (1962) for Catallagia moneris 
Jordan, 1937 and Catallagia motei Hub- 
bard, 1940. Hubbard (1947) synonymized 
each of these species with C. decipiens and 
C. charlottensis (Baker, 1898), respectively. 
Hubbard termed these anomalous speci- 
mens, whose characters resulted in descrip- 
tions of distinct species, ‘“‘dimorphic 
forms” of valid taxa. Delotelis is remark- 
ably similar to the genus Catallagia (it may 
eventually prove to represent a subgenus of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Catallagia as may also the Mexican genus 
Strepsylla Traub, 1950). Perhaps these three 
aberrant specimens are merely freak genetic 
expressions of genes common to both Ca- 
tallagia and Delotelis. 


Delotelis telgoni (Rothschild, 1905)** 
(Figs. 2D—F) 


Miscellaneous records.—Josephine Lake, 
2,287 m, Idaho Co. (45°13'N, 115°58’W), 
ex C. gapperi, 23 October 1999, 1 2. 

Remarks.—This is a new state record. 
The flea has a broad distribution from Brit- 
ish Columbia to New Mexico; however, it 
is extremely rare in collections. Rarely are 
more than one or two specimens collected 
from a host. Jellison and Senger (1973) list- 
ed 8 6 and 20 2 from Missoula County, 
Montana, also from C. gapperi. 


Epitedia wenmanni (Rothschild, 1904)* 


Carnivore records.—(2), ex M. ameri- 
cana, 6 November 2003, 1 @. 


Megarthroglossus divisus (Baker, 1898)* 


Carnivore records.—1 2 without host, 
date or locality data. 


Nearctopsylla hyrtaci 
(Rothschild, 1904)** 


Carnivore records.—(1), ex M. ameri- 
cana, 20 December 2002, 1 °, 4 and 10 
November 2003, 2 @. 

Remarks.—Although Holland (1985) re- 
ported this species numerous times from 
various species of mustelids, it is primarily 
a parasite of shrews (Soricidae). It has been 
reported in Montana (Jellison and Senger 
1973, Kohls 1950, Senger 1966), Utah (Ku- 
cera 1995, Tipton and Allred 1952), and 
Wyoming (Wiseman 1955) but is seemingly 
more prevalent to the north in British Co- 
lumbia (Holland 1985). It appears from the 
records that small mustelids, such as wea- 
sels, prey on soricids more frequently than 
do pine martens. Two females of Nearctop- 
sylla princei Holland and Jameson, 1950, 
reported in Larimer County, Colorado, by 
Eads, et al. (1979) may represent N. hyrtaci. 


VOLUME 107, NUMBER 2 423 


Fig. 2. A-C, Catallagia decipiens, female. A, Head, arrows depicting two preantennal frontal rows. B, 
Normal sperm theca. C, Aberrant spermatheca. D—F, Delotelis telgoni, female. D, Head, arrows depicting three 
frontal rows. E, Spermatheca. F Eighth spiracular fossa. G-H, C. decipiens, female. G, Eighth spiracular fossa. 
H, Eighth spiracular fossa of female with aberrant spermatheca. Scale: A and D = 100 microns: B, C, and E— 
H = 200 microns. 


These specimens could not be located for Nevadas considering the isolating barrier of 
comparison. Females of these two species the Great Basin. 

are very similar (if not indistinguishable) 
and it is unlikely that the distribution of N. 
princei would be so disjunct as to occur in Miscellaneous records.—Silver Creek, 
both the Rocky Mountains and the Sierra near Silver Creek community hot springs 


Neopsylla inopina Rothschild, 1915* 


424 


plunge (44°20'N, 115°47'W), ex S. colum- 
bianus, 13 and 15 July 1974, 4 @. 

Remarks.—Neopsylla inopina is the only 
species of this large genus in North Amer- 
ica. Holland (1985) reported S. columbi- 
anus and §. richardsoni (Sabine, 1822) as 
its principle hosts. 


Rhadinopsylla (Actenophthalmus) difficilis 
Smit, 1957* 


Carnivore records.—(1), ex V. vulpes, 5 
December 2002, 1 2. (2), ex M. americana, 
6 November 2003, 1 3, 2 &. (5), ex M. 
americana, 6 January 2004, 1 @. (6), ex M. 
americana, | January 2003, 1 2 (1 @ with- 
out host, date, or locality data). 

Miscellaneous records.—Josephine Lake, 
2,287 m, Idaho Co. (44°53’N, 115°58’W), 
ex Microtus or Clethrionomys nest, 12 Sep- 
tember 19977 12 

Remarks.—This is a new record for Ida- 
ho County. Our material belongs to the sub- 
genus Actenophthalmus and is tentatively 
assigned to R. difficilis. Holland (1985) re- 
ported similar difficulty in placing Canadi- 
an specimens. Smit (1957) described R. ar- 
borea, R. difficilis, R. linta, and R. media 
(the latter three each based only on 2-3 
specimens), and he re-described the male of 
R. fraterna (Baker, 1895). Each of these 
species (if valid) potentially occurs in Ida- 
ho. Based on the presence of only one seta 
below the level of spiracles on t. II-VI, our 
specimens are clearly not R. fraterna. The 
diagnostic characters that Smit used were of 
little benefit in identifying our material 
(shape of basimere/telomere and st. IX of 
males, the caudal margin of st. VII of fe- 
males, chaetotaxy of tergum VIII, shape of 
8" spiracular fossa, and arrangement of ge- 
nal spines). Tentative assignment to R. dif- 
ficilis is based primarily on Smit’s declara- 
tion of geographic distribution, although the 
atrium of the 8" spiracle was very narrow 
in all specimens which is most similar to 
that of R. linta. Baird and Saunders (1992) 
recorded a single female of R. difficilis from 
M. americana, taken in Valley County but 
there is no indication of who identified the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


specimen. North American material belong- 
ing to this subgenus is in need of revision, 
which surely will result in several syno- 
nyms. 


HyYSTRICHOPSYLLIDAE 


Hystrichopsylla dippiei dippiei 
Rothschild, 1902* 


Miscellaneous records.—Josephine Lake, 
2,287 m, Idaho Co. (44°53'N, 115°58'W), 
ex C. gapperi, 23 October 1999, 1 6, 2 9; 
Microtus or Clethrionomys nest, 1 3. 

Remarks.—The only other record in Ida- 
ho was reported by Baird and Saunder 
(1992) from Boise County. 


LEPTOPSYLLIDAE 


Ctenophyllus armatus armatus 
(Wagner, 1901) 


Carnivore records.—(1), ex M. ameri- 
cana, 20 December 2002, 1 @. 

Remarks.—Crenophyllus a. armatus is a 
well established parasite of pikas. 


Peromyscopsylla hamifer vigens 
(Jordan, 1937)** 


Carnivore records.—(4), ex V. vulpes, 3 
November 2003, 1 6. (1 @ without host, 
date, or locality data). 

Miscellaneous records.—Josephine Lake, 
2,287 m, Idaho Co. (45°13'N, 115°58’W), 
ex Microtus sp.?, 23 October 1999, 3 6. 

Remarks.—These are new records for 
Idaho. Although the distribution of this flea 
is broad, extending from Montana south to 
the Jemez Mountains of New Mexico, it is 
collected infrequently. An exception is the 
collection from the Jemez Mountains, New 
Mexico, where Haas (1973) reported 71 6 
and 88 @. He did not specify the source of 
these specimens but virtually all were col- 
lected from animals (personal communica- 
tion, G.E. Haas). He indicated that he has 
rarely collected any species of Peromyscop- 
sylla from nests, and that they spend most 
of their time in the fur of their host. Hol- 
denried and Morlan (1956) reported 10 
specimens taken from six of 41 voles ex- 


VOLUME 107, NUMBER 2 


amined in Sante Fe County, New Mexico. 
The preferred host of this flea is species of 
Microtus. 


Peromyscopsylla selenis 
(Rothschild, 1906)* 


Miscellaneous records.—Upper Payette 
Lake (45°08'N, 116°01'’W), 13 August 
1997, ex P. maniculatus, 1 3, ex Microtus 
Spal la.oe 

Remarks.—This species is widely dis- 
tributed in western North America from 
Alaska to New Mexico, but is not common 
in collections. 


PULICIDAE 


Euhoplopsyllus glacialis lynx 
(Baker, 1904)* 


Carnivore records.—(5), ex L. rufus, 19 
December 2003, 3 3, 10 2, 30 December 
2003 ING yee atOManuany, 20042556), 110 
2, 20 January 2004, 2 5d, 3 2, 4 February 
2004, 9 5d, 12 2: ex M. americana, 28 Jan- 
uary 2004, 2 2; V. fulva, 4 February 2004, 
8) 

Remarks.—This rabbit flea occurs pri- 
marily on the snowshoe hare (Lepus amer- 
icanus Erxleben, 1777) according to Hol- 
land (1985) who also suggested that it 
might be established on the Canada lynx 
(Lynx canadensis Kerr, 1792) because of 
the large numbers frequently found on it. 
Baird and Saunders (1992) reported one 
record of this species in Latah County, Ida- 
ho, from Felis lynx = L. rufus. Jellison and 
Senger (1973), in their synopsis of the fleas 
of Montana, recorded only two records in 
Missoula County, one each from a bobcat 
and the Canada lynx. Of the 14 bobcats that 
were trapped in trapping area (5), only five 
harbored this flea. Bobcats were not cap- 
tured in other areas. Records from three car- 
nivorous species, plus significant total num- 
bers of fleas (20 6, 45 2), suggest that this 
flea might be widespread in Idaho com- 
mensurate with the range of L. americanus. 
This can be validated only with additional 
collections from larger carnivores and from 
L. americanus. 


Pulex irritans Linnaeus, 1758* 


Carnivore records.—(1), ex V. vulpes, 5 
December 2002, 6 6, 12 9. (2), ex Canis 
latrans Say, 1823, 3 November 2003, 3 6, 
3) Ds 

Remarks.—This is a cosmopolitan spe- 
cies commonly found on these hosts. 


VERMIPSYLLIDAE 
Chaetopsylla setosa Rothschild, 1906* 


Carnivore records.—(3), ex M. ameri- 
cana, 12 November 2003, 1 2. (S), ex ZL. 
rufus, 19 December 2003, 1 6, 2 9, 30 
December 2003, 4 6, 1 2, 20 January 
2004, 1 3; M. americana, 20 December 
2003, 1 2, 6 January 2004, 1 @. (7), ex P. 
concolor, 20 December 2002, 1 d, 1 &. (8), 
ex P. concolor, 19 January 2004, 1 3 (1 2 
without host, date, or locality data). 

Remarks.—Chaetopsylla setosa is con- 
sidered a parasite of larger carnivores, i.e., 
black bears and wolverines, while Chaetop- 
sylla floridensis (1. Fox, 1939), a closely al- 
lied species, occurs on small mustelids, e.g., 
martens, mink, and ermine (Baird ~ and 
Saunders 1992, Haas and Wilson 1982, 
Haas et al. 1978, 1979, 1980, 1982, 1989, 
Holland 1985, Hopla 1965, and Lewis and 
Lewis 1994). McDaniel and Easton (1986) 
reported an interesting record of C. setosa 
on porcupines from South Dakota where 
bears and wolverines are not known to oc- 
cur. Jellison et al. (1943) reported this flea 
from a cougar from Mineral County, Mon- 
tana (two d and II 2 annotated from cou- 
gar in Mineral County in Jellison and Sen- 
ger (1973) are assumed to be the same rec- 
ord). Three specimens of C. setosa collect- 
ed from two different cougars are new for 
this host in Idaho. Martens and cougars are 
accidental hosts for this flea. Although the 
precise interaction between martens, cou- 
gars, and black bears is unknown, occa- 
sional intermittent occupation of black bear 
dens by martens and cougars because of se- 
vere Winter weather may provide an expla- 
nation of how these two animals become 
infested with C. setosa. The low incidence 


426 


of marten flea infestations suggest that cou- 
gars do not become infested from eating 
martens. 

Our specimens of C. setosa were com- 
pared with the syntypes of C. floridensis (I. 
Fox, 1939) collected from “‘Garden truck 
leaf mold”’ in Florida and with a series (7 
3d, 2 2) reported from M. americana by 
Eads, Campos and Barnes (1979) (all de- 
posited in the NMNH). The aedeagus of 
these two species is virtually indistinguish- 
able but displays significant variation from 
one specimen to another within the same 
species. The flexibility of bilateral membra- 
nous lobes associated with the median dor- 
sal lobe creates these variations (compare 
illustrations of apex of aedeagus Wagner 
(1936, fig. 4) and Hopkins and Rothschild 
(1956, fig. 140). The crochet, as illustrated 
in Johnson (1955, figs. 6—8, is sometimes 
extended dorso-caudally, uncurling an oth- 
erwise obvious “loop.” These two morpho- 
logical variables portray quite different fea- 
tures. Contrary to couplet 5 in the key to 
the Chaetopsylla in Hopkins and Roths- 
child (1956), both species bear a single long 
seta at the posterior margin of the eye. This 
character cannot be used as a distinguishing 
feature, nor can the morphology of the 
clasper (basimere and telomere) which also 
shows considerable variation. Chaetopsylla 
setosa may be distinguished from the close- 
ly allied species C. floridensis by: 1) its 
larger size; 2) labial palpi that extend to the 
middle of the trochanter, or well beyond (7— 
10 segments versus < 7); and 3) the pres- 
ence of several anterior rows of setae on t. 
VIII of the male (exclusive of the primary 
row of 3—5 large setae). 


ACKNOWLEDGMENTS 


Without the help of diligent collectors, 
the advancement of our knowledge of ec- 
toparasites would not be possible. Such is 
the cooperative support of James Bacon 
who unselfishly took considerable time to 
collect and record the data for the fleas from 
the fur-bearers that he trapped for two sea- 
sons. We are most grateful for his dedica- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


tion and efforts and to Craig Derrick for 
contributing specimens from cougars. We 
also express our thanks to Nancy Adams, 
National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 
(NMNH) for the loan of specimens essen- 
tial for comparative work and to Duke S. 
Rogers, College of Integrative Biology, 
Brigham Young University, Provo, Utah for 
providing identifications for some of the 
small mammals. Publication and laboratory 
costs were provided in part by National Sci- 
ence Foundation, Grant Number DEB- 
hea) II). 


LITERATURE CITED 


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. 1971. Mammalian ectoparasite consortism at 
the National Reactor Testing Station. Great Basin 
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Baird, C. R. and R. C. Saunders. 1992. An annotated 
checklist of the fleas of Idaho. University of Idaho 
Research Bulletin No. 148, pp. 3-34. 

Eads, R. B., E. G. Campos, and A. M. Barnes. 1979. 
New Records for several flea (Siphonaptera) spe- 
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species parasitizing carnivores in the Rocky 
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Haas, G. E. 1973. Morphological notes on some Si- 
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Haas, G. E., R. E. Barrett, and N. Wilson. 1978. Si- 
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1822-1825. 

Haas, G. E., L. Johnson, and N. Wilson. 1980. Si- 
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Haas, G. E., L. Johnson, and R. E. Wood. 1982. Si- 
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Haas, G. E. and N. Wilson. 1982. Fleas (Siphonaptera) 
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Haas, G. E., N. Wilson, T. O. Osborne, R. L. Zarnke, 
L. Johnson, and J. O. Wolff. 1989. Mammal fleas 
(Siphonaptera) of Alaska and Yukon Territory. 
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Holdenried, R. and H. B. Morlan. 1956. A field study 
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55: 369-381. 

Holland, G. P. 1985. The fleas of Canada, Alaska and 
Greenland (Siphonaptera). Memoirs of the Ento- 
mological Society of Canada, No. 130, 631 pp. 

Hopkins, G. H. E. and M. Rothschild. 1956. An illus- 
trated catalogue of the Rothschild collection of 
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ural History), Vol. I, Coptopsyllidae, Vermipsyl- 
lidae, Stephanocircidae, Ischnopsyllidae, Hypso- 
phthalmidae, and Xiphiopsyllidae, London, 32 
plates, 445 pp. 

. 1962. An illustrated catalogue of the Roths- 
child collection of fleas (Siphonaptera) in the Brit- 
ish Museum (Natural History), Vol. III, Hystri- 
chopsyllidae, London, 10 plates, 560 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. 

Hubbard, C. A. 1941. Ectoparasites of western Lago- 
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. 1943. The fleas of California with checklists 

of the fleas of Oregon, Washington, British Co- 

lumbia, Alaska, Idaho, Nevada, and Arizona. Pa- 

cific University Bulletin 39: 1—12. 

. 1947. Fleas of western North America, their 
relation to public health. lowa State College Press, 
Ames, Iowa, 533 pp. 

Jellison, W. L., G. M. Kohls, and H. B. Mills. 1943. 
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fleas. Miscellaneous Publication No. 2, Montana 
State Board of Entomology, 22 pp. 

Jellison, W. L. and S. Senger. 1973. Fleas of Montana. 
Montana Agricultural Experiment Station, Re- 
search Report No. 29, Montana State University, 
Bozeman, 78 pp. 

Johnson, P. T. 1955. The genus Chaetopsylla Kohaut, 
1903 in North America, with the description of a 
new species. The Pan-Pacific Entomologist 31: 
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Kohls, G. M. 1950. Note on the occurrence of the flea 
Nearctopsylla hyrtaci (Rothschild) in the United 
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92-94. 

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phonaptera: Ceratophyllidae: Ceratophyllinae). 
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North America north of Mexico: Vermipsyllidae 
and Rhopalopsyllidae. Journal of Medical Ento- 
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fleas of the Pacific Northwest. Oregon State Uni- 
versity Press, Corvallis, 296 pp. 

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flea (Siphonaptera) associated with porcupines 
collected in South Dakota. Proceedings of the En- 
tomological Society of Washington 88: 265-267. 

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ty, New Mexico. Texas Reports on Biology and 
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Siphonaptera. Journal of the Idaho Academy of 
Science 14: 26-28. 

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Montana. Journal of the Kansas Entomological 
Society 39: 105-109. 

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aptera. Bulletin of the British Museum (Natural 
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Entomology 53: 1—184. 

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tribution records of Utah Siphonaptera with the 
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Jordan and Rothschild, 1915. Great Basin Natu- 
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harten. Zeitschrift fiir Parasitenkunde 8: 333-352. 

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cies of the world, a taxonomic and geographic ref- 
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PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 428-431 


TWO NEW LEAFHOPPER SPECIES OF BOLANUSOIDES DISTANT 
(HEMIPTERA: CICADELLIDAE: TYPHLOCYBINAE: TYPHLOCYBINI) 
FROM CHINA 


MIN HUANG AND Y ALIN ZHANG 


Key Laboratory of Plant Protection Resources and Pest Management, Ministry of Ed- 
ucation, Entomological Museum, Northwest Sci-Tech University of Agriculture and For- 
estry, Yangling, Shaanxi 712100, China (e-mail: huangmin4399@sina.com; yalinzh@ 


nwsuaf.edu.cn) 


Abstract.—The leafhopper genus Bolanusoides Distant, 1918, is reported for the first 
time from China. Two new species, B. yunnanensis and B. shaanxiensis from China are 


described and illustrated. 


Key Words: 
species, China 


The leafhopper genus Bolanusoides Dis- 
tant, 1918 (Typhlocybinae: Typhlocybin1) 
was reviewed by Dworakowska (1993), and 
the 11 species, mainly from the Oriental 
and Australian regions, were divided into 
the B. heros and B. bohater groups. This is 
the first report of Bolanusoides from China, 
and two new species of the B. heros group 
are described in this paper. 

Type specimens are deposited in the col- 
lections of the Entomological Museum, 
Northwest Sci-Tech University of Agricul- 
ture and Forestry, China (NWSUAP), and 
China Agricultural University, Beijing, Chi- 
na (CAU). 


Bolanusoides Distant 


Bolanusoides Distant 1918: 90, type-spe- 
cies: B. heros Distant; Dworakowska 
Uhsts32 55 IGOBe So), 

Camulus Distant 1918: 97, type species: C. 
ornatus Distant; McAtee 1934: 104. 


Diagnosis.—Bolanusoides differs from 
other genera in this tribe by the following 
combined characters: body robust and flat, 
body color with rich brown to dark brown 


Hemiptera, Cicadellidae, Typhlocybinae, Bolanusoides, new record, new 


fasciae on head, pronotum, and fore wing; 
RP, MP’ and MP”+CuA’ of fore wing with 
conjuncted stem at their base; connective 
lamellate without longitudinal ridge in cen- 
tral part; aedeagal shaft tubular with round- 
ed and swollen apex and paired processes. 


KEY TO SPECIES OF CHINA (MALES) 


1. Pygofer with grouped microsetae on ventro- 
basal part (Fig. 6); lateral aedeagal process 
very slim and straight in lateral view (Fig. 11) 

Sana rae shh ed hap eta Nene tte de ey ee pere d B. yunnanensis 

— Pygofer without grouped microsetae on ventro- 
basal part (Fig. 15); lateral aedeagal process 
strong and slightly sinuated in lateral view 


(Fig. 20) B. shaanxiensis 


Bolanusoides yunnanensis Huang and 
Zhang, new species 
(Figs. 1-11) 


Diagnosis.—This species resembles B. 
pahlawan Dworakowska in external color- 
ation and male genitalia but can be distin- 
guished from the latter by the configuration 
of the aedeagal processes. Its lateroventral 
processes smooth with apex curved later- 
ally, lateral processes long and extending 
beyond the tip of lateroventral ones. 


VOLUME 107, NUMBER 2 429 


2, Head, pronotum and scutellum, lateral 


1-11, Bolanusoides yunnanensis. 1, Adult, dorsal view. 
view. 3, Forewing. 4, Hindwing. 5, Abdominal apodeme. 6, ¢ pygofer side, lateral view. 7, Paramere, connective, 
subgenital plate and valve, dorsal view. 8, Apical part of subgenital plat 


posterior view. 11, Aedeagus, lateral view. 12-20, Bolanusoides shaanxiensts. 


Figs. 1—20. 
e. 9, Paramere, dorsal view. 10, Aedeagus, 
12, Adult, dorsal view. 13, Forewing. 
14, Hindwing. 15, 3 pygofer side, lateral view. 16, Paramere, connective and subgenital plate, dorsal view. 17, Apical 


part of subgenital plate. 18, Paramere, dorsal view. 19, Aedeagus, posterior view. 20, Aedeagus, lateral view. 


430 


Description—Length: ¢ 3.84 mm, 2 
4.02 mm. Head with anterior margin of ver- 
tex silvery white, delimited posteriorly by 
brown transverse line, remaining part light 
yellowish ochre. Face light yellow with nar- 
row undulating transverse brown line dor- 
sally. Pronotum orange ochre except lateral 
band, transverse line at hind margin and 
patch on discal area, brownish. Scutellum 
orange ochre, two small spots against me- 
dial suture brownish, middle of lateral mar- 
gin and very tip of scutellum blackish. 
Forewing mostly yellow ochre with brown 
pattern (Fig. 3); patch at proximal end, 
streak from distal end of wax field to outer 
apical cell, and a spot at base of third apical 
cell, blackish brown. 

Male pygofer sides narrowing posteriorly 
with several macrosetae dorsoposteriorly; 
ventral process short and stout. Subgenital 
plate with one basal macroseta, apex curved 
dorsally with subapical lateral margin with 
pigmented peglike setae. Paramere elongate 
with numerous setae in middle part, inner 
margin serrate over distal half. Connective 
lamellate without central lobe. Aedeagal 
shaft cylindrical, expanded in medial part 
in lateral and posterior views, gonopore api- 
cal; two stout processes on each side one- 
third distance from base to apex of shaft, 
one curved ventrally and other curved dor- 
solaterally and a slender straight process on 
each side, situated more basally, directed 
dorsally. Male basal abdominal apodemes 
reaching to base of 5th abdominal sternite. 

Types.—Holotype d, China, Yingjiang 
(24.7°N, 97.9°E), Yunnan Province, alt. 380 
m, April 29, 1981, coll. Li Fa-sheng 
(CAU). Paratypes: China, 22, same data as 
holotype; 12, Tengchong (25.1°N, 98.5°E), 
Yunnan Province, alt. 1,650 m, April 28, 
1981, coll. Li Fa-sheng (all CAU). 

Etymology.—The specific name is de- 
rived from the type locality, Yunnan Prov- 
ince. 


B. shaanxiensis Huang and Zhang, 
new species 
(Figs. 12—20) 
Diagnosis.—This species is similar to B. 
yunnanensis. n. sp., and B. pahlawan in ex- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ternal coloration and male genitalia but can 
be distinguished from both latter species by 
configuration of the aedeagal processes. Its 
aedeagus with smooth and fairly strong la- 
teroventral processes, two pairs of lateral 
processes arising from one point and upper 
ones, which are almost parallel to the later- 
oventral process, are the strongest. More- 
over, the new species does not have anterior 
fascia on the vertex. 

Description.—Length: ¢ 4.02 mm, 2 
4.11 mm. Head with anterior part of vertex 
silvery white, remaining part ivory; face 
light yellow. Pronotum orange ochre except 
lateral margin, central area, and transverse 
posterior band, brownish. Scutellum orange 
ochre, two spots on front of medial suture 
and patch at inside of basal triangles, mid- 
dle part of lateral margin, and very tip of 
scutellum blackish brown. Forewing mostly 
yellow ochre with brown pattern as shown 
in Fig. 13; patch at proximal end, streak 
from distal end of wax field to outer apical 
cell, and spot at base of 3rd apical cell 
blackish brown. 

Male pygofer, paramere and connective 
as in B. yunnanensis but pygofer process 
elongate and subgenital plate with one to 
three pigmented peglike setae. Aedeagal 
shaft expanded apically in lateral and pos- 
terior views, two processes on each side of 
shaft near midlength directed dorsolaterally, 
lower process with small basal process di- 
rected ventrally. 

Types.—Holotype ¢6, China, Hanzhong 
(33.1°N, 107.1°E), Shaanxi Province, Au- 
gust 12, 1984, coll. Zhang Ya-lin 
(NWSUAF). Paratypes: 1d, 12, same data 
as holotype (NWSUAF). 

Etymology.—The specific name is de- 
rived from the type locality, Shaanxi Prov- 
ince. 


ACKNOWLEDGMENTS 


We thank Dr. I. Dworakowska (retired, 
Canada) for her contribution to the knowl- 
edge of Chinese Typhlocybinae in cooper- 
ation with the Entomology Museum of 
Northwest Sci-Tech University of Agricul- 


VOLUME 107, NUMBER 2 


ture and Forestry (supported by the CIDA). 
Our sincere thanks also go to Mr. Mick 
Webb, The Natural History Museum, Lon- 
don, for revising the manuscript. This proj- 
ect was supported by ““The Fok Ying Tung 
Education Foundation, National Education- 
al Committee of China.” 


LITERATURE CITED 


Distant, W. L. 1918. Rhynchota. Homoptera: Appen- 
dix. Heteroptera: Addenda. The Fauna of British 
India, including Ceylon and Burma. Published un- 
der the authority of the Secretary of State for India 


431 


in Council. Ed. By A. E. Shipley, assisted by Guy 
A. K. Marshall, Vol. 7, vii + 210 pp. 

Dworakowska, I. 1988. Bolanusoides bohater sp.n. 

from New Guinea (Insecta, Homoptera, Auchen- 

orrhyncha, Cicadellidae: Typhlocybinae). Rei- 
chenbachia Staatlichen Museum fuer Tierkunde 

Dresden 26: 5-7. 

. 1993. Contribution to the knowledge of Bo- 
lanusoides Dist. (Auchenorrhyncha, Cicadellidae, 
Typhlocybinae, Typhlocibini). Bulletin de LIn- 
stitut Royal des Sciences Naturelles de Belgique, 
Entomologie 63: 59-69. 

McAtee, W. L. 1934. Genera and subgenera of Eup- 
terygidae (Homoptera; Jassidae). Proceedings of 
the Zoological Society of London 1934: 93-117. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 432—435 


THE GENUS [SCHNOCEROS GRAVENHORST 
(HYMENOPTERA: ICHNEUMONIDAE) IN CHINA 


MAo-LING SHENG 


Forest Resource and Environment College, Beijing Forestry University, Beijing, 
100083, China; General Station of Forest Pest Management, State Forestry Administra- 
tion, Shenyang, Liaoning, 110034, China (e-mail: smlzylx @mail.sy.In.cn) 


Abstract.—Four species of the genus /schnoceros Gravenhorst, 1829, collected in China 
are reported. One species is described, Ischnoceros pedipullus, n. sp., and two species, 
I. rusticus (Geoffroy, 1785) reared from Acanthocinus aedilis (Linnaeus) (Cerambycidae) 
in Pinus, and I. sapporensis Uchida, 1928, reared from Callidium sp. (Cerambycidae) in 
Quercus, are new records for China. A key to the four species known in China is given. 


Key Words: 


Ischnoceros Gravenhorst, 1829, is a 
small genus belonging to the subfamily 
Xoridinae which are important parasitoids 
of wood borers (Cerambycidae). Species 
are distributed in the Oriental, Neotropical, 
and Holarctic regions. Ten world species 
are known (Wang et al. 1997, Yu and Horst- 
mann 1997). The genus has not been stud- 
ied thoroughly in the Oriental and Palearc- 
tic regions of China (Zhang and Zhao 
1996). Only one species, /. sanxiaensis 
Wang, 1997, previously has been recorded. 
In this study, one new species is described 
and two species are newly recorded for Chi- 
na. 

Identifications of /. rusticus (Geoffroy) 
and /. sapporensis Uchida are based on de- 
termined specimens loaned from K. Sch6n- 
itzer, J. Hilszczanski, and R. Matsumoto. 
Identification of /. sanxianesis Wang is 
based on the study of the type. Specimens 
from this study are deposited in the collec- 
tions of the General Station of Forest Pest 
Management, State Forestry Administra- 
tion, Shenyang, Liaoning, China. 

Ischnoceros is distinguished from other 
genera of Xoridinae by the two subequal 


Hymenoptera, Ichneumonidae, /schnoceros, new species, China 


teeth at the apex of the mandible, frons with 
a strong median horn or tubercle, and hind 
femur without a median ventral tooth. 


Ischnoceros pedipullus Sheng, 
new species 
(lisse Ie 2s 3a) 


Female.—Body 9.5 mm long. Forewing 
7.4 mm long. Face 2.0 wider than long, 
strongly convex centrally, densely punctate 
with puncture diameters 3.0 to 4.0X dis- 
tance between punctures, lateral margins 
with faint punctures; upper portion below 
antennal sockets with oblique wrinkles. 
Clypeal suture distinct and deep. Clypeus 
smooth, without punctures; basal portion 
transversely convex; lateral sides with short 
transverse wrinkles; median portion flat; 
apical portion weakly concave. Mandible 
with two teeth, lower tooth distinctly short- 
er than upper tooth. Malar space 0.4X as 
long as basal width of mandible. Temple 
smooth, with fine punctures, diameter of 
punctures about same as distance between 
punctures; median portion weakly convex 
longitudinally; posterior portion converging 
backward; 1.3 as long as eye in dorsal 


VOLUME 107, NUMBER 2 


Figs. 1-3. 


Ischnoceros species. 1, I. pedipullus, propodeum; arrow points to semicircular notch. 2, /. pedi- 


pullus, lateral view of metasomal apex. 3, Apex of ovipositor of /. pedipullus (a), I. rusticus (b), and I. sappo- 


rensis (C). 


view. Vertex smooth, with fine, very sparse 
punctures, diameter of punctures 0.3x— 
0.8 distance between punctures; distance 
between lateral ocellus and occipital carina 
very wide, lateral ocellus distinctly located 
anterior to line connecting hind margins of 
eyes. Outside of stemmaticum with dense 
transverse wrinkles. Distance between lat- 
eral ocelli 0.8 distance between lateral 
ocellus and compound eye. Frons with 
dense transverse wrinkles, lateral margins 
with unclear and shallow punctures, its cen- 
ter with a strong, very deeply hollowed pro- 
tuberance, bifurcated at upper portion. An- 
tenna filiform with 33 flagellomeres: first 
flagellomere 1.1 longer than second; sec- 


ond, third, and fourth about same length; 
remaining flagellomeres gradually decreas- 
ing in length. Occipital carina complete and 
strong. Front and hind portion of pronotum 
with dense longitudinal wrinkles, dense 
transverse wrinkles at center of lateral side. 
Epomia indistinct. Front portion of middle 
lobe of mesonotum with distinct punctures; 
hind portion strongly rough; outside of lat- 
eral lobe with dense and large punctures, 
diameter of punctures about 4.0% distance 
between punctures; sublaterally with dense 
and oblique longitudinal wrinkles. Notaulus 
reaching center of mesoscutum. Scutellum 
nearly flat, with dense punctures, diameter 
of punctures about 3.0% distance between 


434 


punctures. Postscutellum strongly convex, 
with a triangular concavity in front. Meso- 
pleuron with dense longitudinal wrinkles. 
Speculum smooth. Upper end of epicnemial 
carina reaching front margin of mesopleu- 
ron. Epicnimium rough. Mesosternum with 
dense punctures, diameter of punctures 
3.0X—-5.0 distance between punctures; 
median groove of mesosternum deep and 
broad a hind portion. Metapleuron rough, 
with rough and irregular wrinkles. Wings 
brownish hyaline. Forewing vein Icu-a dis- 
tinctly basad of 1M; 3rs-m absent; 2rs-m 
distinct, 0.3 as long as distance between 
it to 2m-cu; base of Im-cu separated from 
cula by 2.0 length of culb. Hind wing 
veins 1Cu and cu-a approximately equal. 
Median part of base of propodeum descend- 
ing strongly to forma a semicircular notch 
(Fig. 1). Carinae of propodeum strong, lat- 
eral carinae of basal area parallel; areola 
confluent with basal area and separated 
from petiolar area by a distinct carina. Cos- 
tula distinct. Propodeal apophysis large and 
compressed, about 1.5 diameter of second 
antennal flagellomere. Areola and pleural 
areas with transverse wrinkles. Petiolar area 
with oblique longitudinal wrinkles. First 
lateral areas with punctures. Second lateral 
areas with longitudinal wrinkles. Metasoma 
comparatively short and glossy. First tergite 
1.7X longer than its apical width, strongly 
bent at spiracle, with irregular and fine 
lines, fine oblique transverse lines at middle 
portion of subapex. Second to fourth ter- 
gites with dense and fine transverse lines. 
Fifth and sixth tergites with very weak 
transverse lines, each about 0.3 longer 
than their basal width. Ovipositor sheath 
5.0 mm long, about 1.6 longer than hind 
tibia. Apical portion of lower valve of ovi- 
positor (Fig. 3a) with 6 distinct ridges, basal 
ridge weakly oblique, rest vertical; rough- 
ened area basad of basal ridge. 

Black. Clypeus dark brown. Palpi brown- 
ish black. Front leg brown to dark brown. 
Middle and hind legs, except for black cox- 
ae and hind femur, blackish brown to 
brownish black. Bases of tibiae and first tar- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


sal segments white. Stigmata darkish 
brown. Veins brownish black. 
Holotype.—®, China: Baishilazi Natural 
Reserve, 400 m, Kuandian County, Liaon- 
ing Prov., June 1, 2001, Mao-Ling Sheng. 
Host.—Unknown. 
Etymology.—The species name is based 
on the color of the legs. 
Diagnosis.—Ischnoceros pedipullus is 
similar to 1. sapporensis but can be distin- 
guished from the latter by the brownish 
black palpi, median part of the base of the 
propodeum with a semicircular notch (Fig. 
1), lateral carinae of basal area parallel, 
lower valve of ovipositor with a roughened 
area basad of the basal ridge (Fig. 3a), most 
of the middle legs brownish black, and the 
middle and hind coxae black. /schnoceros 
sapporensis has the palpi white; basal por- 
tion of the propodeum without a notch, lat- 
eral carinae of the basal area strongly di- 
vergent anteriorly, lower valve of the ovi- 
positor without a roughened area (Fig. 3c), 
most of the middle legs light brown to 
brown, and most of the middle and hind 
coxae brown to darkish brown. 


Ischnoceros rusticus (Geoffroy, 1785) 
(Fig. 3b) 


New record for China. 

Specimens examined.—1 @, China: Dong- 
cun, Mudanjiang, Heilongjiang Prov., June 
25, 1980, Jing-Wen Shao; 1 @, China: 
Tonghua, Jilin Prov., June 30, 1992, Mao- 
Ling Sheng. 

Host.—I reared this species from Acan- 
thocinus aedilis (Linnaeus) (Cerambycidae) 
in Pinus from China: Tonghua, Jilin Prov- 
ince. 


Ischnoceros sapporensis Uchida, 1928 
(Fig. 3c) 


New record for China. 

Specimens examined.—2 ?, China: Xin- 
bin, Liaoning Prov., June 10, 1999, Mao- 
Ling Sheng; | 2, Kuandian (Baishilazi Nat- 
ural Reserve), 400 m, Liaoning Prov., June 
1, 2001, Mao-Ling Sheng. 

Host.—I reared this species from Calli- 


VOLUME 107, NUMBER 2 


dium sp. (Cerambycidae) in Quercus from 
China: Xinbin, Liaoning Province. 


KEY TO SPECIES OF [SCHNOCEROS IN CHINA 


1. Median part of base of propodeum without 
notch; lateral carinae of basal area strongly di- 
WETYOME AMWETIOTY 5 2soc0c0c00s000bs 000% 2 

— Median part of base of propodeum descending 
strongly to form a semicircular notch (Fig. 1); 
lateral carinae of basal area parallel ..... 

I. pedipullus, n. sp. 

2. Third to sixth tergites with dense punctures 

SOS 51S Use Lone Sah aE I. sanxiaensis Wang 

— Third to sixth tergites with dense and fine 
(immense INNES » ocaecccacccooooeueoer 3 

3. Palpi white; coxae mostly brown to darkish red 
brown; basal area and areola separated by a 
strong carina; first and second tergites without 
transverse lines, or only median portion at hind 
margin of second tergite with fine transverse 
lines; ovipositor in Fig. 3c ............ 


I. sapporensis Uchida 

— Palpi brownish black; coxae red; basal area and 
areola not separated by a distinct carina; first 
and second tergites with dense and fine trans- 
verse lines; ovipositor in Fig. 3b 
REE Giichics a CeOROR Cec onc Caen See I. rusticus (Geoffroy) 


aay 
Lo 
Nn 


ACKNOWLEDGMENTS 


I am deeply grateful to Dr. K. Sch6nitzer, 
Zoologische Staatssammlung Mitinchen, 
Germany, Dr. Jacek Hilszczanski, Depart- 
ment of Forest Protection, Forest Research 
Institute, Warsaw, Poland, and Dr. Rikio 
Matsumoto, Osaka Museum of Natural His- 
tory, Osaka, Japan, for their kindness in of- 
fering valuable material and lending speci- 
mens. 


LITERATURE CITED 


Uchida, T. 1928. Dritter Beitrag zur Ichneumoniden- 
Fauna Japans. Journal of the Faculty of Agricul- 
ture, Hokkaido University 25: 1—115. 

Wang, S., J. Yao, and G. Wang. 1997. Hymenoptera: 
Ichneumonidae, pp. 1617-1646. /n Yang, X., ed. 
Insects of the Three Gorge Reservoir Area of Yan- 
tze River. Chongqing Publishing Company, 
Chongqing, China. 

Yu, D. S. and K. Horstmann. 1997. A catalogue of 
world Ichneumonidae (Hymenoptera). Memoirs of 
the American Entomological Institute 58, 1558 
Pp: 

Zhang, S. and Y. Zhao. 1996. The Geographical Dis- 
tribution of Agricultural and Forest Insects in Chi- 
na. China Agriculture Press, Beijing, 400 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 436-440 


A NEW SPECIES OF HARMSTONIA 
(DIPTERA: DOLICHOPODIDAE) FROM BOLIVIA 


HAROLD ROBINSON AND NORMAN E. WOODLEY 


(HR) Department of Botany, National Museum of Natural History, PO. Box 37012, 
Smithsonian Institution, Washington, DC 20013-7012, U.S.A.; (NEW) Systematic Ento- 
mology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, 
% National Museum of Natural History, Smithsonian Institution, PO. Box 37012, MRC 
168, Washington, DC 20013-7012, U.S.A. (e-mail: nwoodley @sel.barc.usda.gov). 


Abstract.—Harmstonia ichilo Robinson and Woodley, new species, is described from 
Ichilo Province, Santa Cruz Department, Bolivia and is the first record for the genus in 


South America. 


Key Words: 


The genus Harmstonia was described by 
Robinson (1964) on the basis of two species 
from the eastern United States. The genus 
has some of the key characters of Micro- 
morphus Mik, especially the mesoscutum 
with a flattened posterior slope and an ab- 
sence of acrostichal setulae. However, 
Harmstonia, with its large male genital cap- 
sule and diverging R,,; and M veins of the 
wing differs strikingly from Micromorphus, 
and the two genera are not now considered 
to be particularly closely related. Eleven ad- 
ditional species were described by Robin- 
son (1967), mostly from Mexico, with a 
species from Costa Rica and another from 
the Lesser Antilles. Four more species were 
added by Robinson (1975), two from Ja- 
maica and two from Panama. A species de- 
scribed in the genus by Botosaneanu and 
Vaillant (1973) from Cuba was transferred 
to Enlinia Aldrich by Robinson (1975) be- 
cause of the wing venation and the presence 
of acrostichals, in spite of some unusual 
features and its larger than usual size. Until 
now, Harmstonia had not been reported 
from the continent of South America, al- 
though it was expected to occur there. 


Harmstonia, Bolivia, South America 


In November 2003, on a field trip to Bo- 
livia, N. E. Woodley collected a species of 
Harmstonia in the lowlands of eastern Bo- 
livia in Santa Cruz Department. The series 
was mostly females, but one specimen was 
a male, which allows the following descrip- 
tion. 


Harmstonia ichilo Robinson and 
Woodley, new species 
(Figs. 1-7) 


Male.—Length 1 mm; wing 1.1 mm by 
0.4 mm. 

Front dark green with brownish pollen; 
face covered with brownish pollen, trian- 
gular, with eyes narrowly contiguous in 
lower part. Palpi and proboscis brownish. 
Antenna (Fig. 1) black; scape without bris- 
tles, pedicel short-annuliform, ringed with 
short bristles; first flagellomere broadly ob- 
long with slight lobe below arista, nearly 
truncate; arista with short pubescence. Or- 
bital setulae small, black. 

Thorax dark green, slightly metallic with 
brownish pollen, pollen denser on sides; 
mesoscutum short, strongly gibbous, dis- 
tinctly flattened on posterior slope (Fig. 4, 


VOLUME 107, NUMBER 2 


437 


Figs. 1-3. 


ESEM views of Harmstonia ichilo. 
broadened face of female. 3, Setae on face of female. 


as in female); setae dark; acrostichals lack- 
ing; 5 strong pairs of dorsocentrals; scutel- 
lum with 2 strong, widely separated bris- 
tles, without small setula on lateral margin. 

Coxae, femora, tibiae and most of basi- 
tarsi pale; fore tarsus from tip of second 
segment, and middle and hind legs from tip 
of basitarsi, dark; hind coxa with small ex- 


1, Antennae and narrowed face of male. 2, Antennae and 


ternal bristle. Femora without longer setae 
ventrally. Lengths of parts of legs as fol- 
lows: Fore leg: tibia, 0.32 mm; tarsomeres 
1-5, 0.12, 0.07, 0.06, 0.05, 0.06 mm; mid- 
dle leg: tibia 0.45 mm: tarsomeres 1I-—5, 
0.06, 0.10, 0.07, 0.05, 0.06 mm; hind leg: 
tibia, 0.5 mm; tarsomeres 1—5, 0.14, 0.2, 
0.1, 0.06, 0.06 mm. 


438 


Figs. 4—5. 
acrostichal setae. 5, Wing. 


Wing (Fig. 5, as in female) oval, hyaline, 
veins brownish, veins R,,;, M and CuA di- 
verging from base, essentially straight; 
crossvein dm-cu perpendicular to M, sube- 
qual to to slightly longer than last of section 
of CuA (Fig. 5 does not show this well be- 
cause the wing is at a slight angle, making 
dm-cu appear shorter than the last section 
of CuA). Calypter brown with black bris- 
tles; halter pale brown. 

Genital capsule large, extending well for- 
ward under the abdomen, blackish brown, 
with little pollen; cercus (Figs. 6, 7) pale 
brown, S-shaped, borne from dorso-apical 
corner upward along truncate apex of cap- 
sule, broadened cleaver-like distal part with 
irregular long bristles along margins, with- 
out inflated basal part; gonopods projecting 
forward, with stout terminal spur; aedeagal 
sheath stout, curving strongly to end near 
bases of cerci; small lamella-like surstylus 
exposed along ventral margin of gonopod. 

Female.—Similar to male but eyes not 
contiguous below, face with 2—4 small se- 
tae near middle (Figs. 2, 3), and first fla- 
gellomere of antenna slightly more truncate 
below arista (Fig. 2). Lengths of parts of 
legs as follows: Fore leg: tibia, 0.32 mm; 
tarsomeres 1—5, 0.12, 0.07, 0.05, 0.04, 0.06 


Harmstonia ichilo. 4, Thorax of female 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


citer Lire 


showing depressed area before scutellum and lack of 


mm; middle leg: tibia, 0.42 mm; tarsomeres 
1-5, 0.17, 0.1, 0.04, 0.05, 0.07 mm; hind 
leg: tibia, 0.475 mm; tarsomeres |—5, 0.125, 
OBIS, WI, OVO; O07 racron, 

Holotype.—Male: BOLIVIA: Santa Cruz 
Department, Ichilo Province, Hotel Flora y 
Fauna, 4-6 km SSE Buena Vista, 
17°29.95'S, 63°33.15'W, 400-500 m, 10 
Nov. 2003, N. E. Woodley, in the National 
Museum of Natural History, Smithsonian 
Institution (USNM). 

Paratypes.—8 Females (including allo- 
type), same locality, date, collector, habitat 
as holotype (USNM, MNKM). 

Etymology.—tThe species epithet, a noun 
in apposition, is based on the province in 
Bolivia where the type series was collected. 

Remarks.—Harmstonia ichilo lacks dis- 
tinct rows of dorsal bristles on the fore and 
hind tibiae, which would place the species 
in the typical group of the genus according 
to Robinson (1967). The shape of the cerci, 
without an inflated base, with a cleaver-like 
tip and without forked marginal setae re- 
sembles most closely the Mexican H. clav- 
icauda Robinson. The latter species, nev- 
ertheless, has a small genital capsule borne 
apically on the abdomen, smaller cerci, and 
a thinner aedeagal sheath. 


VOLUME 107, NUMBER 2 


439 


Figs. 6-7. Harmstonia ichilo, genital capsule and appendages of the male in slightly different views with 
preabdomen above and hind leg to left and below. 6, Capsule without back-scattering showing surface detail. 
7, Capsule with back-spattering showing clearer outline of parts. Abbreviation: c, cercus. 


The series collected by Woodley includes 
8 females and only a single male. The spec- 
imens were taken along a small, partly 
shaded stream with steep banks in second- 


ary forest. Some of the water formed still, 
shallow pools. The substrate was very fine, 
clean sand. The flies were not actually ob- 
served in situ, but were collected by general 


440 


sweeping at water edges along with Ephy- 
dridae and other Dolichopodidae. The pau- 
city of males collected apparently indicates 
that the sexes have slightly differing pref- 
erences in habitat, something already seen 
in other members of the genus. Robinson 
observed males of H. intricata Robinson in 
eastern North America which seemed to 
prefer drier parts of muddy ruts in a road, 
while females were found in places with 
some standing water. Males of H. pectini- 
cauda Robinson, from the same area were 
only in direct sunlight while females, were 
not as particular. 

The other Dolichopodidae collected at 
the site included single males of an appar- 
ently undescribed Sarcionus Aldrich and an 
apparently undescribed Discopygiella Rob- 
inson. The latter genus has not previously 
been reported from the South American 
continent, but the species is not described 
at this time because only a single specimen 
was obtained. 


ACKNOWLEDGMENTS 


The Philips XL30 ESEM scope used to 
produce the illustrations was operated by 
Scott Whitaker, manager of the National 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Museum of Natural History SEM Labora- 
tory. Marjorie Knowles, technician in Bot- 
any, is thanked for her work with the spec- 
imens and illustrations. NEW thanks the 
Entomology Department at the Museo de 
Historia Natural ““‘Noel Kempff Mercado” 
(MNKM), Santa Cruz, Bolivia for facilitat- 
ing collecting in Bolivia. We are grateful to 
Wayne Mathis (Smithsonian Institution) 
and Natalia Vandenberg and Allen Norr- 
bom (Systematic Entomology Laboratory, 
USDA) for reviewing the manuscript. 


LITERATURE CITED 


Botosaneanu, L. and E Vaillant. 1973. Quelques Dip- 
tera Dolichopodidae nouveaux de Cuba. Résultats 
des Expéditions biospéleologiques Cubano-Rou- 
maines a Cuba 1: 411—424. 

Robinson, H. 1964. A synopsis of the Dolichopodidae 
(Diptera) of the southeastern United States and ad- 
jacent regions. Miscellaneous Publications of the 
Entomological Society of America 4: 103-192, 
figures 1-139. 

. 1967. Revision of the genus Harmstonia (Dip- 

tera: Dolichopodidae). Proceedings of the United 

States National Museum 123(3515): 1-16. 

. 1975. Bredin—Archbold—Smithsonian Bio- 

logical Survey of Dominica. The family Dolicho- 

podidae with some related Antillean and Pana- 
manian species (Diptera). Smithsonian Contribu- 

tions to Zoology 185: i-1v, 1-141. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 441-459 


STUDIES ON THE DARKLING BEETLES 
(COLEOPTERA: TENEBRIONIDAE) KNOWN FROM 
GRAND BAHAMA ISLAND, WITH DESCRIPTIONS OF NEW SPECIES OF 
BRANCHUS AND ADELINA 


WARREN E. STEINER, JR. 


Department of Entomology, NHB-187, Smithsonian Institution, Washington, DC 20560, 
U.S.A. (e-mail: steinerw @si.edu) 


Abstract.—An annotated checklist of the darkling and comb-clawed beetles (Coleop- 
tera: Tenebrionidae) of Grand Bahama Island is given, with observations on habitats, 
biology, and biogeography. More than 55% are new records for the Bahamas and some 
others are new island records. Branchus saxatilis Steiner, new species, and Adelina 
maryjoae Steiner, new species, are described and illustrated. Nautes viridimicans (Horn 
1878), described from Tampa, Florida, is placed as a new synonym under Nautes azur- 
escens (Jacquelin du Val 1857), described from Cuba. Threats to some species via deg- 
radation of the native maritime scrub habitats are discussed. Of the 31 species now known 
from Grand Bahama, 25 also occur in Florida, and a subset of 18 of the Florida species 
also are known from Cuba. Five of the species are known or suspected to be adventive. 


Key Words: 


Bahamas, Casuarina, endemic insects, introduced species, island biogeog- 


raphy, new synonymy, threatened habitats 


Systematic studies on the darkling bee- 
tles (Coleoptera: Tenebrionidae) of the West 
Indies are proving to be of great value 
toward increased understanding of the pat- 
terns of island biogeography of the Carib- 
bean region. My interest in this group of 
insects and the distributions of adventive 
and endemic species on islands, after two 
visits to Grand Bahama Island, led to this 
study. Similar treatments of island faunas 
are planned for future research in the Ba- 
hamian region, in order to document the en- 
demicity and adventive elements in these 
islands, and assist in enrichment of the Ba- 
hamian National Insect Collection and of 
other institutions. 

While identifying the available speci- 
mens and reviewing the literature, it be- 
came apparent that very little is known of 
the Bahamian beetle fauna. Biodiversity 


surveys have been done on only a few is- 
lands, and systematic studies have focused 
primarily on Lepidoptera and Hymenoptera 
(Elliott 2003). Only 28 species of Tene- 
brionidae are documented as occurring in 
the Bahamas (Steiner in press) but in the 
short time spent collecting these beetles on 
Grand Bahama, 30 species were document- 
ed; most represent new distribution records 
for the Bahama Islands, and two are newly 
described in this paper. Observations on the 
biology (including excerpts from field 
notes) and known distribution of each spe- 
cies are given for the new material, and 
other records in the literature are noted. 
West End, Grand Bahama 
about 100 km east of the Florida Peninsula 


Island lies 
at latitude 26°N; the elongate island extends 
eastward approximately 140 km and has a 
total land area of 1,356 km/’. It is surround- 


442 


ed by other Bahamian islands and small 
cays to the north, east and south at varying 
distances and these islands have a great 
range in size. The nearest other island of 
comparable size is Abaco, arcing around 
and east of Grand Bahama’s eastern end 
and separated from it at one point by only 
about 20 km. Abaco, Grand Bahama, and 
their associated offshore cays are all part of 
the Little Bahama Bank (Shattuck and Mill- 
er 1905), which was a single large island 
during lower sea levels of the Pleistocene 
(Sealey 1994). For these reasons, Grand 
Bahama and Abaco would be expected to 
have similar insect faunas (Browne et al. 
1993). While closer to its neighboring lands 
during that period, there was no land bridge 
between the Florida peninsula nor the Great 
Bahama Bank (“‘Paleoprovidence’’). Of the 
two primary types of islands (Gillespie and 
Roderick 2002), Grand Bahama can be con- 
sidered a “‘Darwinian” island in that sense 
as well as a “‘fragment”’ island in the other, 
because it has separated from a larger land 
mass since the Pleistocene. 

The natural vegetation of the island 
interior is forested, with the dominant tree 
being Caribbean pine (Pinus caribaea Mo- 
relet) on a rugged karst terrain which is no- 
where elevated more than a few meters 
above sea level (highest point less than 21 
m). Thickets of Sabal palmetto (Walt.) 
Lodd. ex JA & JH Schultes form much of 
the understory, mixed with various shrubs; 
large hardwood species are uncommon, but 
this may be the result of the historical har- 
vest of wood. The northern coast of Grand 
Bahama is irregular, with mangrove inlets 
and cays. The southern coast has beach and 
dune formations of varying width, slope, 
and sand composition. The calcareous sand 
deposits of the western Bahamas provide a 
mosaic of differing substrate deposits and 
textures due to storm action and sea level 
fluctuation (Multer 1977, Sealey 1994) and 
this provides a dynamic mixture of micro- 
sites for colonization by geophilous beetles. 
With the combination of coastal soils and 
interior forest, the island offers habitats 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Gazetteer of label localities for Tenebrion- 
idae of Grand Bahama Island. 


Label Locality Lat./Long. 

Bootle Bay Village 26°39'N, 78°56’ W 
Eight Mile Rock 26°31'N, 78°47'W 
Freeport 26°30'N, 78°41'W 
Garden of the Groves 26°32'N, 78°30'W 
Holmes Rock Village 

(5 km NW) 26°36'N, 78°52'’W 
Lucaya (beach) 26°29'N, 78°37'W 
Lucaya (19 km ENE of) 26°34'N, 78°26’ W 
McLean’s Town 26°39'N, 77°59'W 
West End 26°42'N, 79°00'W 
Xanadu Beach 26°29'N, 78°42'W 


(Correll 1979) for several guilds of geo- 
philous, lignophilous and fungivorous 
Tenebrionidae. However, native maritime 
scrub of the southern strand has in most ar- 
eas been severely altered with the invasion 
of Casuarina equisetifolia L. and related 
species (Hammerton 2001) with resulting 
impacts on beetle ocurrences in such habi- 
tats noted (Steiner 1991, in press). 


METHODS 


Many of the specimens reported here 
were taken in June 1987 (8 days of col- 
lecting effort) on the south side of Freeport, 
at fluorescent lights around a building that 
was located next to forested land, and by 
operation of black lights on several 
evenings near this site. One Malaise trap 
and several yellow pan traps at forest edges 
were also used at this locality. Various mi- 
crohabitats were investigated with hand- 
collecting techniques, at this and other lo- 
calities as noted under each species record 
below. Specimen label localities and their 
coordinates are listed in Table 1. All labels 
from the June 1987 collection read 
“GRAND BAHAMA ISLAND” (first 
line), followed by the locality and date (giv- 
en under each species listing, below) and 
ending with (collectors) ““W. E. Steiner, M. 
J. & R. Molineaux.”? October 1995 (3 days 
of collecting effort) specimens are labeled 
similarly but have coordinates of the local- 
ity given, and end with “coll. W. E. Steiner 


VOLUME 107, NUMBER 2 


& J. M. Swearingen.” Label data on type 
material, and the few other specimens from 
other sources, are spelled out in full. Spec- 
imens are deposited in the collections of the 
National Museum of Natural History, 
Smithsonian Institution, Washington, DC 
(USNM), the Bahamian National Insect 
Collection, Nassau (BNIC), and other insti- 
tutions as noted in the specimen records. 

A search for additional material in mu- 
seum collections leading up to this study 
was made, but was largely unsuccessful (as 
discussed below). Literature records were 
also searched. 


RESULTS 


The checklist (Table 2) of known Tene- 
brionidae from Grand Bahama Island in- 
cludes 31 species representing 7 subfami- 
lies and 15 tribes as presently recognized 
(Aalbu et al. 2002). Annotations given 
below for each of the species follow this 
sequence. The majority of species are con- 
sidered to be indigenous to the island and 
the region, but three are identified as being 
introduced from outside the region and two 
are probably recent arrivals from adjacent 
land areas; six are evidently endemic to the 
Bahama Islands. Most are new island 
records and the first reported for the Ba- 
hamas; in many cases they represent a 
range extension from some adjacent area, 
principally Florida. Few Grand Bahama 
specimens and literature records were 
found, probably because of the historical 
lack of focus on this group in the archipel- 
ago (Steiner, in press), and the relatively 
little general work by entomologists on this 
island. A relatively low number of Ceram- 
bycidae are recorded (Browne et al. 1993) 
from Grand Bahama, larger than some 
islands with greater richness documented, 
illustrating the lack of comparable collect- 
ing there. Perhaps the largest known Grand 
Bahama insect collection is that of D. J. 
MacDonald, kept at the Rand Nature Cen- 
tre, Freeport (Elliott 2003), but a family- 
level inventory did not list any Tenebrion- 
idae (N. B. Elliott, personal communica- 


443 


tion). In addition, the BNIC, Nassau, cur- 
rently holds mostly Lepidoptera, 
Hymenoptera, Orthoptera, and aquatic in- 
sects. 


Epitragodes tomentosus (LeConte) 


“5 km. NW Holmes Rock Village, 24 June 
NOS aay): 


This unique female was found by Jenni- 
fer Molineaux “at top of dune, under a 
white shell” and other field notes describe 
the beach as “fairly narrow and sloped, 
with a steplike slip-face at dune edge; an 
elevated rolling dune area fairly broad, and 
stabilized with trailing grasses, [pomoea, 
and succulents. Behind this and down in 
slack area was an abrupt dense scrub of old 
Coccoloba; no Australian pine here yet.” 
Except for a recent listing for “Bahamas” 
(Peck and Thomas 1998) this beetle was not 
previously known outside of the continental 
USA (Freude 1968). This was an unex- 
pected Voccintence.. but) the) species as 
winged and does occur along similar beach- 
es, barrier islands and sandy areas along the 
southern Atlantic coast; larvae live in sandy 
soils (Hoffman et al. 2003; Steiner 1988, 
1995, unpublished data). Compared with 
individuals from Florida, the specimen is 
larger than average and more robust with 
larger, denser punctures and setae of the 
dorsal surface. Two subspecies are recog- 
nized and separated on such differences 
(Freude 1968) and it is possible that the 
Bahamian form represents a third, but more 
material will be needed to determine its 
distinctness. M. A. Ivie (personal commu- 
nication) has seen a specimen from the 
Bimini Islands labeled “‘South Bimini Isl., 
Bahamas, B.W.I., March 22-25 1958, A. 
M. Nadler,’’ determined by C. A. Triple- 
horn and in the collection of American 
Museum of Natural History. This probably 
is the source of the Bahamian record listed 
by Peck and Thomas (1998). 

Branchus saxatilis Steiner, new species 
(Figs. 1-3A—D) 

The occurrence of a Branchus on Grand 

Bahama was expected since members of the 


444 


Table 2. Checklist of Tenebrionidae known from 


Grand Bahama Island. Classification follows that of 


Aalbu et al. (2002). Abbreviations: A, adventive; B, 
new island record but previously known from the Ba- 
hamas; N, new record for the Bahamas; E, apparently 
endemic to the Bahamas; P, previously reported from 
the island; FL and CU, also known from Florida and/ 
or Cuba. Refer to the text on each species for more 
distributional data, specific information and discussion. 


Pimeliinae 
Epitragini 


|. Epitragodes tomentosus (LeConte)—N; FL 


Branchini 
2. Branchus saxatilis Steiner, n. sp.—N, E 
Diaperinae 
Phaleriini 
3. Phaleria picipes Say—B; FL+CU 
4. Phaleria punctipes LeConte—P; FL+CU 
5. Phaleria testacea Say—B; FL+CU 


Diaperini 
6. Adelina plana (Fabricius)—N; FL+CU 
7. Adelina bidens (Schaetfer)—B: FL+CU 
8. Adelina maryjoae Steiner, n. sp.—N, E(?) 
9. Diaperis maculata Olivier—B; FL+CU 
10. Platydema excavatum (Say)—N; FL+CU 
11. Platydema nigratum (Motschulsky)—N; FL+CU 
12. Platydema ruficorne (Sturn)—N; FL 
13. Platydema ruficorne (Sturn)—N; FL 
14. Neomida bicornis (Fabricius)—B; FL+CU 


Crypticini 
15. Gondwanocrypticus platensis (Fairmaire)—A, N; 
FL 
Hypophloeinae 
16. Corticeus thoracicus (Melsheimer)—N; FL 
Opatrinae 
Opatrini 
17. Blapstinus fortis LeConte—N; FL+CU 
18. Diastolinus bahamae Marcuzzi—P, E(?) 
Platynotini 
19. Alaetrinus pullus (Sahlberg)—N, A(?); FL+CU 
Leichenini 
20. Leichenum canaliculatum variegatum (Klug)—A, 
N; FL+CU 
Tenebrioninae 
Helopini 
21. Nautes azurescens Jacquelin du Val—N; 
FL+CU 
Triboliini 


22. Tharsus seditiosus LeConte—N; FL 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 2. Continued. 


Alphitobiini 
23. Alphitobius diaperinus Panzer—A, N; FL+CU 


Amarygmini 
24. Cymatohes tristis (Laporte)—N, FL+CU 
Alleculinae 

Alleculini 

25. Lobopoda nesiotica Campbell—P, E 

26. Hymenorus densus LeConte—P; FL 

27. Hymenorus bahamensis Campbell—P, E 

28. Hymenorus farri Campbell—N, A; FL+CU 

29. Hymenorus convexus Casey—P; FL+CU 

30. Hymenorus transversus Campbell—B, E 
Coelometopinae 

31. Glyptotus cribratus LeConte—P(?); FL 


genus occur in southern Florida (Triplehorn 
and Weems 1964) and on New Providence 
(LeConte 1866) and Cuba (Spilman 1961) 
and specimens from other Bahamian islands 
had been seen by me. After extensive 
search in the most likely habitats (Steiner 
1991), a single female specimen was found 
at Eight Mile Rock, with associated larvae, 
on the first visit in June 1987. The beetle 
was distinct from any of the neighboring 
forms, but more specimens were needed to 
watrant a description. This prompted the re- 
turn to the site in October 1995, and it 
proved successful. 

Description.—Holotype, male (Figs. 1— 
3): Body color very dark brown, outline 
oval, nearly continous except for small in- 
dentation between posterior angle of pron- 
otum and elytral humerus; form convex, 
about equally so dorsally and ventrally; 
body length 12 mm from anterior angle of 
pronotum to elytral apex, greatest width 7.2 
mm at basal % of elytra; dorsum dull, cov- 
ered with fine, decumbent, pointed and ta- 
pered, golden brown setae. 

Head less than % as wide as pronotum, 
widest at eyes and equally so at epistomal 
canthi; frons with a triangular concavity 
across middle. Antenna with segment 3 
slightly shorter than combined length of 
segments 4 and 5; segments 9, 10 and 11 


VOLUME 107, NUMBER 2 


Fig. 1. Branchus saxatilis, holotype 


3, dorsal habitus. Length, 12 mm. 


445 


446 


wider than long, forming a weak club. 
Mentum somewhat heart-shaped, without 
sharp angles; surface flat, punctate, setose. 
Gena slightly produced anteriorly, not ex- 
tending to half the length of mentum. 

Prothorax about twice as wide as long, 
broadest at basal %, sides evenly arcuate, 
anterior margin broadly and evenly emar- 
ginate, anterior and posterior angles pro- 
duced, the latter with rounded, flattened 
apices directed posteriorly; lateral margin 
very slightly explanate, without a polished 
bead; pronotum evenly convex, surface 
with shallow rounded punctures, some con- 
tiguous, each bearing a flattened scalelike 
seta extending just beyond posterior margin 
of puncture; punctures absent along a nar- 
row smooth midline and on very small, 
slightly raised areas on middle of each side 
of disk; ventral surfaces more sparsely 
punctate with setae slender, hairlike, and 
not as flattened or decumbent. 

Elytron widest near basal ¥;'"*, about 2.5 
times longer than wide; dorsal surface gen- 
erally smooth except for basal plicae (Fig. 
1) and poorly defined rows of large deep 
punctures, mostly confined to basal %; 
scalelike setae smaller and more decurved 
than those on pronotum, arising from very 
inconspicuous punctures, distributed evenly 
or in slightly more dense patches but with 
gaps around the large punctures (Fig. 2). 
Margin of pseudepipleuron moderately well 
defined but not sharp or beaded; scalelike 
setae more dense than on elytral disk. Epi- 
pleuron long and tapered to elytral apex, 
generally “4 as wide as pseudepipleuron 
but abruptly widened at base. 

Sternites somewhat polished, with seti- 
ferous punctures on most areas but smaller, 
not as dense, especially sparse in median 
areas of abdominal sternites 1—4 which are 
smoother, shining. Legs with scalelike setae 
especially on tibiae; more hairlike on fem- 
ora and relatively dense on flat ventral sur- 
faces of meso- and metatibiae; tibial spurs 
well developed. Protibia broad and truncate 
at apex, forming a tooth at outer angle; 
meso- and metatibiae expanded at apex; 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. 
elytral surface including suture. Length of setae, ca. 
0.11 mm. 


Branchus saxatilis, holotype 6, detail of 


metatibia arcuate. Tarsi with conspicuous 
golden setae on ventral surfaces, especially 
on pro- and mesotarsomeres |—4. 

Tegmen (Fig. 3A—D) with apical piece 
slightly longer than basal piece, flattened 
and truncate at apex, sinuate across apex 
with slight asymmetry between small, 
sharp, lateral projections. Median lobe with 
a slender arching rodlike apex, ending with 
a slightly bulbous tip. 

Female: Similar to male but larger, 13— 
14 mm long and more robust, inflated. Sec- 
ondary sexual differences include lack of 
the arcuate form of the hind tibia, much re- 
duced setose flat ventral surfaces on the 
meso- and metatibiae, and more convex ab- 
dominal sterna. 

Variation.—The largest of the 5 males is 
12.3 mm long, the smallest is 11.5 mm 
long. Some specimens have a soil encrus- 
tation that obscures some of the punctures 
and setae. 

Type data.—Holotype ¢, 4 d paratypes 
and | @ paratype: “GRAND BAHAMA 


VOLUME 107, NUMBER 2 


Fig. 3. 


447 


Branchus saxatilis, holotype d, genitalia. A, Tegmen, dorsal view. B, Tegmen and median lobe, 


lateral view. C, Apical piece, dorsal view. D, Apical half of tegmen and median lobe, ventrolateral view. Length 


of tegmen, 5.3 mm. 


ISEAND Echt yiMile, Rock, 26230N- 
78°47'W, 25 October1995, coll. W. E. 
Steiner & J. M. Swearingen;” | 2 paratype, 
Same data except “26 June 1987; W. E. 
Steiner, M. J. & R. Molineaux”’ and 1 3 
paratype, ““Abaco Cays, Allans Cay, May 
9, 1953/Van Voast-A.M.N.H. Bahama Isls. 
Exped., Coll. E. B. Hayden.” The holotype 
and 4 paratypes are deposited in USNM; 
two paratypes, in BNIC, and the Abaco 
paratype, in the American Museum of Nat- 
ural History, New York. 

Etymology.—The specific name comes 
from the Greek word “‘saxatilis”” meaning 
“of the rock,” in reference to the name of 
the type locality, Eight Mile Rock. The 
coastal strand of beach at the locality is 
characterized by a ledge of coral rock on 
which the habitat of these beetles exists. 

Diagnosis and relationships.—Branchus 
saxatilis belongs to the “‘floridanus group” 


(Steiner 1991) that includes the type spe- 
cies, B. floridanus LeConte (1866), illus- 
trated by Triplehorn and Weems (1964) 
from southern Florida, and B. woodi Le- 
Conte (1866) from New Providence, and 
several other species to be described. All of 
the members of this group have the same 
distinctive form of the male genitalia, with 
minor variations; the external characters are 
more useful for recognizing the species, in 
particular the details of the scale-like elytral 
setae. These are very short, sparse and in- 
conspicuous in B. floridanus, narrow, fine 
and evenly scattered in B. woodi; the latter 
are generally smaller beetles, and neither 
species has the large, deep, elytral punc- 
tures seen in B. saxatilis. Branchus saxatilis 
appears to be endemic to islands of the Lit- 
tle Bahama Bank, but is most closely relat- 
ed to its neighboring congeners in Florida 
and islands of the Great Bahama Bank. A 


448 


key to Branchus species will be provided 
in a future monograph of the group. 

Biology.—At the type locality, digging 
by hand in coarse sand among low shrubs 
led to the discovery of several large larvae, 
later confirmed as belonging to Branchus. 
This preceded the finding of the first adult 
beetle, as described by me in field notes on 
26 June 1987: “‘Found a small pull-off at 
beach front on a rocky ledge—no beach at 
the surf line here, but rugged coral rock + 
pounding waves; dune sand was on upper 
part of coral bedrock, however, + had a low 
shrub zone, with no Casuarina. The large 
black Blapstinus [= Diastolinus bahamae| 
common here at edge of shrub patches un- 
der drift boards + leaf litter; associated lar- 
vae in sand a few cm deep. Dug deeper in 
loose sand under a piece of scrap lumber + 
found a big larva—looked like Branchus 
but larger than any B. larvae from Flori- 
da—kept digging around this local area at 
edge + under low spreading shrubs, finding 
more larvae in different sizes but most as 
large as the 1‘; after about half an hour, 
excavated a single 2 adult Branchus, about 
4—5 cm in sand, under a single layer of 
dead leaves in shade of low shrub, in 
among sticks + roots. Sand cool + dry, 
loose, + mixed with dark organic parti- 
cles.”’ On the return visit to this site eight 
years later, October 1995, adult beetles 
were more easily found, not as deep in sand 
but under leaf litter that was wet from re- 
cent rains; only one half-grown larva was 
found at this time. Known larvae of several 
Branchus species are to be described in a 
future work. 

This site was unique in being free of 
shade from Casuarina trees, presumably 
because thé shallow sand on the rock ledge 
has prevented their establishment, but ad- 
jacent to this section of coast was a dense 
stand of old Casuarina, typical of most of 
the shoreline on the southern strand. 
Searches for Branchus specimens at all oth- 
er sites visited were not successful; the 
dune crest and scrub vegetation behind the 
beach front in all other sites was shaded out 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


by these trees, or had been altered or lost 
by construction of buildings and roads. The 
threats to native biota by the dominance of 
the introduced Casuarina spp. are severe 
(Hammerton 2001) and Branchus species 
may be the most threatened of maritime 
scrub insects because of their habitat re- 
strictions (Steiner 1991, in press). Because 
Branchus beetles were searched for at all 
coastal sites visited, but found on Grand 
Bahama only at the type locality, B. saxa- 
tilis may be of conservation concern. Its 
status on Abaco is unknown at present. 


Phaleria picipes Say 
“Bootle Bay Village, 24 June 1987” (7). 


These specimens were among the larger 
series of P. testacea listed below; the two 
species often co-occur. My field notes de- 
scribe *“‘a well developed step-like beach of 
more coarse coral sand”’ and beetles taken 
‘“‘in moist sand under seaweed drift mats 
recently washed in.” Species of Phaleria 
are restricted to coastal beach sands where 
they feed on drift debris; this species and 
its two congeners listed below are wide- 
spread in the Caribbean region and highly 
variable in size and coloration (Triplehorn 
1991, Triplehorn and Watrous 1979, Wa- 
trous and Triplehorn 1982). The above 
specimens range from a uniform dark 
brown to a yellowish tan color; elytra are 
sometimes more pale than the pronotum 
and head. Phaleria picipes is known from 
the other Bahamian islands of South Bimini 
and Rum Cay (Triplehorn and Watrous 
IST). 


Phaleria punctipes LeConte 


“5 km. NW Holmes Rock Village, 24 June 
1987” (3); “Lucaya (beach), 25 June 
1987” (4); “19 km. ENE Lucaya, 25 Oc- 
tober 1995” (1); ““McLean’s Town, 26 
October 1995” (1); ““Xanadu Beach, 21 
june 1987 CA) 2s une MO Sie): 


For a flightless species, P. punctipes is 
surprisingly widespread on many islands 
(see papers cited under P. picipes) includ- 


VOLUME 107, NUMBER 2 


ing small ones, e.g., Pigeon Cay near An- 
dros (Anderson 1996). Beetles and larvae 
are found under washed-up sea drift that is 
generally more dry and higher on the beach 
than that used by P. testacea and others, 
e.g., as noted for the Xanadu Beach series: 
“‘under dry drift piles on a higher plateau 
of sand above normal high tide, and we 
found a concentration of them—in dry loose 
sand but where sand was moist 3—4 cm be- 
low surface. Some were even up at edge of 
beach grass and stabilized zone, and were 
found next to the Blapstinus.” 


Phaleria testacea Say 


“Bootle Bay Village, 24 June 1987” (48): 
“Lucaya (beach), 25 June 1987” (13). 


These were at first misidentified as P. pi- 
cipes (in field notes) because the mixed col- 
oration of Bahamian specimens is more 
typical of that species. Grand Bahama spec- 
imens range from yellowish brown to me- 
dium brown and about 10 % are bicolored 
(elytra mostly brown with a lateral stripe 
and basal band of yellow). The series from 
Lucaya was found with P. punctipes “on 
and in sand under seaweed drift mats along 
recent high tide line, upper part of wide, 
coral-sand beach.” 


Adelina plana (Fabricius) 


“Eight Mile Rock, 26 June 1987 / Under 
bark of dead Casuarina” (11); “Freeport, 
2Arune 9872 4G) 26 June 19S): 
‘27 October 1987” (14); “19 km. ENE 
ucaya25) October 1995— (2): 


This beetle is widespread in the Neotrop- 
ical region and reported from Cuba (Ardoin 
1977) and Florida (Peck and Thomas 
1998). They are attracted to lights and I 
have found them associated with larvae un- 
der dry thin bark of dead standing or re- 
cently fallen wood, not rotten, in semi- 
shaded sites. Notes on the specimens of 27 
October read ‘“‘series of small Adelina + 2 
assoc. larvae under tight bark of cut logs, 
probably Acacia or related legume.” 


449 


Adelina bidens (Schaeffer) 


“19 km. ENE Lucaya, 25 October 1995” 
6): 


Specimens were found co-occurring with 
the two A. plana cited above “under bark 
of fallen Casuarina” in a coastal forest 
dominated by that tree; ““Casuarinas domi- 
nant but many dead, standing.’ The species 
is not as widespread as A. plana but was 
reported from South Bimini (Vaurie 1952) 
and is also recorded from Cuba and Florida 
(Ardoin 1977, Peck and Thomas 1998); Ar- 
doin (1977) also listed the Dominican Re- 
public and Guatemala. 


Adelina maryjoae Steiner, new species 
(Figs. 4A—G) 


This distinctive species is known only 
from the holotype. It is different from all 
other species in the genus in having a con- 
spicuous, long basal antennomere or scapus 
(Figs. 4A—C). Ardoin (1977) recognized 18 
species from the Americas and: two others 
have been described since then (Triplehorn 
and Ivie 1983, Doyen 1984). 

Description.—Holotype, male (Figs. 4A— 
F): Length (from clypeal margin to elytral 
apex) 6 mm.; width (across middle of 
elytra) 2.4 mm.; greatest thickness (at 
metasternite) 0.7 mm. Body form very flat, 
elongate, parallel sided; surfaces smooth, 
shining; color reddish brown. Head finely 
punctate, nearly twice as wide as long, wid- 
est at laterally expanded epistomal canthus; 
anterior margin with an undulate appear- 
ance; area of frons between eyes slightly 
concave, then anteriorly with an abrupt de- 
clivity, with three shallow concavities, one 
median and others lateral to it. Labrum very 
small, setose; membrane at base not visible. 
Antenna long, reaching beyond basal 4" of 
elytron if folded back along body; scapus 
enlarged, clavate, nearly as long as head, 
setose on anterior side; pedicellus small, 
rounded; antennomeres 3 and 4 clavate, 
about twice as long as wide, with setae 
around widest area near apex; antennom- 
eres 5—10 of similar form, about 3 times as 


450 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


F 


Fig. 4. Adelina maryjoae, holotype ¢. A, Dorsal habitus. B, View of frons. C, Dorsal view of pronotum 
and head. D, Front leg, right, posterior view. E, Tegmen, dorsal view. EK Tegmen, lateral view. Length of beetle, 
6 mm; tegmen, 1.4 mm. 


VOLUME 107, NUMBER 2 


long as wide, antennomere 11 shorter than 
preceding, narrowed to a pointed apex. Pro- 
thorax broad, disk flattened, finely punctate 
laterally, less so medially; basolateral fo- 
veae conspicuous; anterior margin widely, 
evenly and deeply emarginate; basal margin 
nearly straight; lateral margin sinuate, finely 
beaded, widening rapidly from acute pos- 
terior corner to broadly rounded, inflated 
anterior corner. Scutellum flat, impunctate, 
with shape of a rounded-off triangle. Ely- 
tron about 3 times as long as wide; strial 
punctures small, regularly spaced, separated 
by about the diameter of a puncture; inter- 
vals with very fine scattered punctures; 
edge of lateral declivity of elytron begin- 
ning at the seventh stria, sharp basally, 
more rounded toward apex. Legs smooth, 
setae inconspicuous; front tibia straight, 
mostly parallel sided to apex, with a cren- 
ulate outer margin (Fig. 4D); front tibial 
spurs short, thick, larger curved; tarsomeres 
unmodified; basal first tarsomere only 
slightly thickened, without a ventral pro- 
cess. Tegmen (Fig. 4E) with basal piece 
more than twice as long as apical; base with 
asymmetric apex; apical piece narrowed 
and flattened from base to broadly rounded 
apex which is deflexed in lateral view (Fig. 
AP). 

Type data.—Holotype 3, ““GRAND BA- 
HAMA ISLAND, Freeport, 23 June 1987; 
W. E. Steiner, M. J. & R. Molineaux/At 
light on building; Caribbean pine and pal- 
metto scrub.”’ The holotype is deposited in 
USNM. 

Etymology.—The specific name “mary- 
joae” is derived from the names Mary + 
Jo + the Latin genitive possessive feminine 
ending ’’-ae.”’ I take pleasure in naming this 
species for Mary Jo Molineaux, entomolog- 
ical specialist with the Smithsonian Insti- 
tution; her invitation to accompany her 
family on the 1987 trip to Grand Bahama 
led to the discovery of this species and most 
of the records reported in this study. 

Diagnosis and relationships.— Adelina 
maryjoae belongs to the “premier groupe” 
of species recognized by Ardoin (1977) be- 


451 


cause of the position of the elytral declivity, 
but the large size and darker color of this 
beetle is more characteristic of members of 
the “‘second groupe.”” The lack of the 
ventral tooth on the first basal tarsomere is 
also atypical for the genus, as is the form 
of the front tibia, which is usually widening 
gradually in width from base to apex, and 
bearing a row of more distinct teeth along 
the outer margin. The relative lengths of the 
antennomeres offer a unique suite of char- 
acters; the enlarged basal segment is very 
unusual in Coleoptera, and the equal size 
and shape of segments 3 and 4 is unusual 
in Adelina species, in which segment 3 is 
generally longer and more slender than seg- 
ment 4. Because the species has a number 
of unique apomorphies, its sister relation- 
ships may be difficult to place, but identi- 
fication is easy. The female (unknown at 
present), however, may be found to have 
these characters less developed, as in other 
species of the genus. 


Diaperis maculata Olivier 


““McLean’s Town, 26 October 1995/Under 
bark of rotting trunk of Pinus caribaea 
with polypore fungi” (1). 


Previously recorded from Nassau (Triple- 
horn 1965), this species is widespread from 
North America to Panama and reported 
from a number of the larger Caribbean is- 
lands. This specimen was found in “pine 
flatwoods” on a fallen tree “‘at a roadside 
clearing edge—one trunk elevated from 
ground by root mass (wind thrown) and 
trunk fractured at one point—bark mostly 
tight except at this spot, where a soft fungal 
white mass of polypore was starting to 
emerge from under split bark—peeled bark 
back here and got a single teneral Diaperis 
maculata.” 


Platydema excavatum (Say) 
‘Freeport, 23 June 1987/At black light in 
Caribbean pine and palmetto scrub” (1); 
“On pine pulpwood, Grand Bahama Is- 
land/X-5—59, J. M. Henderson, Colr/ 


452 
Jacksonville, Florida—161, lot 59-— 
Mayers (1))). 


One of the most widespread species of 
the genus, it occurs from southern Canada 
to northern South America (Triplehorn 
1965); the small cosmopolitan bracket fun- 
gus Schizophyllum commune (L.) Fr. is the 
principal host (Leschen 1990). 


Platydema nigratum (Motschulsky) 


mETeepOnte22 JUNC IOS 7 aa) ee October 
1995 / Under bark of rotting trunk of P7- 
nus caribaea with polypore fungi” (1). 


This species is known from the southern 
U.S.A. (and is relatively common in Flori- 
da) to Costa Rica; the only Antillean re- 
cords are from Cuba (Triplehorn 1965). The 
specimens taken in June 1987 were “‘on 
woody polypore brackets under and on 
sides of logs.”” The large cut logs were 
dumped along a roadside in the native pine 
forest and appeared to be from some orna- 
mental, exotic species, possibly a Ficus. 
The unidentified fungus also was the host 
of Neomida bicornis (see below). The Oc- 
tober 1995 specimen was under the same 
piece of bark on a charred pine stump with 
the specimen of P. ruficorne, reported be- 
low. 


Platydema micans Zimmerman 


“Freeport, 21 June 1987” (3), “22 June” 
Ci), “24! oe Cb), “26 lune" CG). 


This is another Middle American species 
with distribution records that include the 
Greater Antilles and southeastern United 
States (Triplehorn 1965). It is known to in- 
habit leaf litter on the ground, unlike other 
Platydema species (Leschen 1990, Steiner 
1995). Notes from 21 June read ‘‘Under 
leaf litter on sandy loose soil under roadside 
shrubs, took a series of Gondwanocrypti- 
cus, + a few Platydema micans + Hyme- 
norus sp. This describes the typical 
microhabitat of this beetle. The other single 
specimens were taken at black light at for- 
est edge sites. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Platydema ruficorne (Sturm) 


“Freeport, 26 October 1995/Under bark of 
rotting trunk of Pinus caribaea with 
polypore fungi’ (1). 


This is the most common Platydema in 
eastern North America including the Flori- 
da Keys (Triplehorn 1965) but it has not 
been previously reported outside of this 
area. It has been associated with a number 
of polypore fungi (Leschen 1990). The sin- 
gle male was associated with the similar P. 
nigratum, reported above. 


Neomida bicornis (Fabricius) 


“Freeport, 22 June 1987” (9), “27 October 
1995” (8). 


Among the most abundant tenebrionids 
in eastern North America, N. bicornis 
shows considerable regional color variation 
over its range, from Canada to the Antilles 
(Cuba, Jamaica), Bermuda, and the Baha- 
mas (Andros), with larger and more bright- 
ly metallic specimens in the southern parts 
(Triplehorn 1965). Various polypore fungi 
are recorded as hosts (Leschen 1990). The 
June 1987 specimens were found in 
association with Platydema nigratum as 
described above; the series from October 
1995 was found “in whitish leathery poly- 
pores” on fallen rotting trunks of Pinus 
caribaea in forest interior. All of the spec- 
imens are of the West Indian color form 
(Triplehorn 1965), with a bright red pron- 
otum and metallic blue elytra. 


Gondwanocrypticus platensis (Fairmaire) 


“Freeport, 20 June 1987” (1), “21 June” 
C13), 24> June? Cy), “2 Jume? Cl) "26 
June” (1) “27 June” (1); ““Garden of the 
Groves, 8 km. E Lucaya, 23 June 1987” 
(9); ““West End, 24 June 1987” (2). 


This species has been recently separated 
from the North American G. obsoletus 
(Say) and recognized as a South American 
species adventive to many areas including 
the southeastern United States (Steiner 
1996, unpublished data). It is often com- 


VOLUME 107, NUMBER 2 


mon in open disturbed roadside areas with 
sandy soils, sparse turf, and leaf litter. The 
series taken 21 June 1987 is the Gondwan- 
ocrypticus mentioned in notes for Platyde- 
ma micans above; other singletons from 
Freeport were taken at lights except that on 
27 June, found running on a sunlit paved 
road after rain. The series from 23 June is 
noted “found some Blapstinus fortis + 
more Gondwanocrypticus under leaf litter, 
on sandy soil at edges of turf + gardens” 
and the specimens from West End were also 
associated with the same Blapstinus “‘under 
coconut husks, conch shells + plant debris— 
open weedy roadside.’ More information 
on the distribution and habits of this species 
will be given in a study of North American 
Crypticini now in preparation. 


Corticeus thoracicus (Melsheimer) 


misteepOntee2 OM UNE N9S 7 cab): 


No notes on the collection of this speci- 
men were made, but it probably was among 
the numerous small beetles taken at black 
light along a roadside gap in pine forest. 
This is the first record of this widespread 
species outside continental North America 
but it is reported from Key West, Florida, 
and in association with Pinus elliottii (Tri- 
plehorn 1990), a pine very similar to P. 
caribaea. 


Blapstinus fortis LeConte 


“Freeport, 24 June 1987” (2), “26 June”’ 
(1); ““Garden of the Groves, 8 km. E Lu- 
caya, 23 June 1987” (5); ““West End, 24 
June 1987” (4). 


This is a common species occurring from 
Florida to Texas and Central America, also 
Cuba and “‘Bahamas”’ (Peck and Thomas 
1998). The only reported specific island 
record that I am aware of is for New Prov- 
idence (Marcuzzi 1962). Its occurrence 
would not be unexpected on any of the larg- 
er islands. It can be abundant in open 
disturbed areas with sandy soils (see notes 
given for Gondwanocrypticus above) and 
the distribution appears to be expanding 


from its natural range via human activities. 
The specimens from Freeport were taken at 
black light; Blapstinus fortis has fully de- 
veloped flight wings. 


Diastolinus bahamae Marcuzzi 


“Eight Mile Rock, 26 June 1987” (25), 
“25, October 1995” (14); “5S km. NW 
Holmes Rock Village, 24 June 1987” (7): 
“19 km. ENE Lucaya, 25 October 1995” 
(1); “‘Pelican Point, 26 October 1995” 
(5); ““Xanadu Beach, 21 June 1987” (12), 
27 October 19951): 


This species was described from Gold 
Rock Creek, Grand Bahama (Marcuzzi 
1965) and in the same study, two other 
closely related forms were named from oth- 
er Bahamian islands. They belong to a 
group of species for which taxonomic re- 
vision is badly needed. Some specialists 
would include these in the large genus 
Blapstinus, while others consider the flight- 
less species (including this and others with 
fused elytra) as distinct, but there is consid- 
erable variation in the development of flight 
wings among other species currently placed 
in Diastolinus and Blapstinus. The Florida 
species Blapstinus alutaceus Casey and B. 
dispar Casey belong to the same close spe- 
cies group with Bahamian Diastolinus spe- 
cies. A reassessment of placement of all cir- 
cum-Caribbean forms is obviously neces- 
sary, but will require comparative studies of 
many taxa in the ““Blapstinus complex.” At 
this time I have chosen to retain the specific 
nomenclature. In the 1987 field notes, how- 
ever, I used Blapstinus for this taxon; it is 
the species mentioned in association with 
Phaleria punctipes, and figured 
(Steiner in press, fig. 8) as an example of a 
group in need of study. On Grand Bahama, 
the beetles are all from coastal sites, none 
from inland habitats. They were found most 
commonly at the upper edge of sand beach- 


above, 


es, on the sand surface but under leaf litter 
or drift debris, around the first spreading 
plants or shrubs of the dry but vegetated 


454 


zone and often at the edges of bare sand 
gaps among plants. 


Alaetrinus pullus (Sahlberg) 


‘Freeport, 23 June 1987” (1), “24 June” 
(1), both with second label ““Yellow pan 
trap in Caribbean pine and palmetto 
scrub.” 


Alaetrinus Iwan (1995), a subgenus of 
Opatrinus, was recently elevated to genus 
level (Iwan 2002). The two _ specimens 
above were studied and identified in the 
course of the review (Iwan 1995) of the ge- 
nus Opatrinus but were omitted in the list 
of specimens examined and the distribution 
map. No other Bahamian specimens are 
known; the distribution (Iwan 1995) in- 
cludes Bermuda, the Florida Keys and An- 
tillean islands from Cuba to St. Croix, and 
also the Yucatan region. Other extralimital 
occurrences were discussed as probable in- 
troductions and he believes that the species 
range is expanding due to anthropogenic 
activity. This may be the case on Grand Ba- 
hama, since the two specimens were taken 
in the same pan trap “‘at edge of open scrub 
and trash at side of parking lot.” 


Leichenum canaliculatum variegatum 
(Klug) 


‘Freeport, 26 June 1987” (1). 


This is an adventive species, presumably 
from Madagascar and established in the 
southeastern United States (Spilman 1959) 
as well as Cuba and Guadeloupe (Marcuzzi 
1984); it occurs on sandy soils and is now 
widespread in Florida (Peck and Thomas 
1998). The single specimen was found at 
black light at forest edge and “probably 
crawled up on to edge of ground sheet from 
dry sandy roadside bank.” Spilman (1959) 
mentioned “‘many occurrences at light” 
based on label data; beetles are fully 
winged. I have found the beetles (at other 
localities, including Madagascar) under leaf 
litter in open sandy coastal sites and reared 
a pupa (Steiner 1995) from larvae associ- 
ated with adults from pure dune sand. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Nautes azurescens (Jacquelin du Val) 


Helops viridimicans Horn 1878: 57. new 
synonymy. 

Freeport, 20 9une 1987 Gi) 2 unex 
@O); “22 juine* (W))5 "23 Jume (©), 24! 
June: (2), 25) June.s (8), Zo) dunes 
(5). 


All of the specimens were taken at lights 
with the exception of one dated 25 June, 
noted as being found dead in a swimming 
pool. Helopines are generally forest-dwell- 
ing beetles with immature stages in the soil 
(Steiner 1995, 1999) but the habits of this 
species are unknown. In making this iden- 
tification, I have compared these specimens 
with identified material from Cuba and also 
with Florida specimens identified as Nautes 
viridimicans (Horn). They appear to be con- 
specific. The form of the aedeagus does not 
differ significantly among males from the 
three lands. Cuban specimens are generally 
larger and metallic greenish, Florida speci- 
mens are more blue-green in color but less 
brightly metallic, and the Bahamian ones 
are intermediate in color and much less 
brightly shining. Bahamian specimens 
average smaller in size, but considerable 
variation and overlap in size is seen among 
all forms. I have not examined type speci- 
mens, but with confidence can place Helops 
viridimicans (Horn 1878), described from 
Tampa, Florida, in synonymy under Nautes 
azurescens (Jacquelin du Val 1857), de- 
scribed from Cuba (new synonymy). Both 
were described in the genus Helops and 
perhaps should have remained there; the 
genera of Helopini are in need of redefini- 
tion. Numerous related species, some flight- 
less, occur on islands throughout the Carib- 
bean region and most are undescribed 
(Steiner 2005). 


Tharsus seditiosus LeConte 


“lskeeororni, 2 Yuin Is (2), “Zs Jae / 
At black light in Caribbean pine and pal- 
metto scrub” (1), ““26 June” (1), “27 Oc- 
tober 1995 / Under bark of rotting trunk 


VOLUME 107, NUMBER 2 


of Pinus caribaea with polypore fungi” 


(5). 


Previously known only from the south- 
eastern United States including southern 
Florida (Peck and Thomas 1998), this is the 
first record of the beetle outside of that re- 
gion. It is the first record of the genus for 
the West Indies. The beetle is not common 
in collections; many U.S. records are from 
areas dominated by pine forest, as are the 
specimens above. One of the June 21 spec- 
imens was from a dead trunk of Pinus car- 
ibaea, “under bark slightly damp beneath, 
about 3 dm. above ground.’ The October 
1995 notes record a “‘series of Tharsus un- 
der bark on sides of fallen logs, with 1 as- 
soc. larva; bark moist beneath, more or less 
loose with a layer of frass under it.” 


Alphitobius diaperinus Panzer 


wEKecpone ZO Mune 98/0 ()) 3) 21 Junes 
CI), 2Ab Uomve< (CD) AX) dives: (yy 


All of these specimens were taken at 
lights, but details of the biology of this 
species are well studied and it has pest sta- 
tus as well as possible value in biological 
control (Despins et al. 1988). A nearly cos- 
mopolitan beetle of African origin, it is 
known from the United States and a number 
of West Indian Islands (Marcuzzi 1984). 


Cymatothes tristis (Laporte) 


mee pOn, 22. UNC mIION ai) 27 October 
1995/Under bark of rotting trunk of Pi- 
nus caribaea with polypore fungi” (1); 
“19 km. ENE Lucaya, 25 October 1995/ 
Under bark of rotting trunk of Pinus car- 
ibaea with polypore fungi” (5). 


Species in this genus were formerly 
placed in the genus Pyanisia LaPorte (Spil- 
man 1973). This species appears to be nat- 
urally distributed among West Indian 
Islands that have forest habitats, as well as 
the southern United States to Panama (Mar- 
cuzzi 1984; Peck and Thomas 1998; Spil- 
man 1969). The two specimens from June 
1987 were found after dark on the same 


455 


logs, probably Ficus sp., noted under Pla- 
tydema nigratum above. For the series tak- 
en 25 October 1995, I noted “‘Cleared areas 
with cut or fallen pines, some charred. Un- 
der loose bark got several adult and larval 
Cymatothes, some associated with bright 
orange fungal tissue’’ and the 27 October 
specimen was found under bark with the 
Tharsus specimens noted above. 


Lobopoda nesiotica Campbell 


Freeport, 20) June 19877 G); “21 Junev 
CI) 422) Tone? 4@) 23 ume cn (Ol) seer 
June’”’ (5), ““25 June”’ (5), “‘26 June”’ (6); 
“20-27 June 1987 / Malaise trap in Ca- 
ribbean pine and palmetto scrub” (11). 


Other than the Malaise-trapped series, all 
specimens were taken at lights. This species 
is among several that are apparently endem- 
ic to the Bahamas; it was described (Camp- 
bell 1971) from specimens from Andros 
(type locality), Grand Bahama (Freeport), 
and Great Abaco. 


Hymenorus bahamensis Campbell 


 Breeport, ZO Mune N98 7- G)): 


A variable species with a complex of 
island forms (Campbell 1971), this species 
is known only from the Bahamas and Cuba, 
and occurs on a number of the islands in- 
cluding one record from West End, Grand 
Bahama (Campbell 1971). 


Hymenorus convexus Casey 


“Freeport, 26 June 1987” (1); ““Xanadu 


Beach, 23 June 1987” (2). 


Campbell (1971) stated that this species 
“probably occurs on every island” of the 
Bahamas and also Cuba and southern 
Florida. He reported Grand Bahama records 
from Eight Mile Rock and West End. Spec- 
imens were taken at lights; the Xanadu 
Beach specimens came to a black light fac- 
ing inland (from the beach) over open dis- 
turbed growth of low scrub. 


456 


Hymenorus densus LeConte 


“Freeport, 20° June 1987” (1), 24 June™ 
(11), “25 June” (@), 26 June” (2). 


These specimens were taken at lights. 
Campbell (1971) reported this species as 
common along the U.S. coast from North 
Carolina to the Florida Keys, and gave a 
number of records from the Bahamas, in- 
cluding Grand Bahama (Pine Ridge). It is 
also listed from Mexico (Peck and Thomas 
1998) but is not known from other islands 
of the West Indies. 


Hymenorus farri Campbell 


‘Freeport, 20 June 1987” (4), “21 June” 
@), 24) lume Cy ~2o Jume (i). 


This would have represented a consid- 
erable range extension, since the species 
was described from Jamaica, Cuba and the 
Cayman Islands (Campbell 1971), but it is 
now known from southern Florida, Puerto 
Rico, Virgin Islands, and other Bahamian 
islands (Steiner 2004). Either the range was 
much wider than originally detected, or the 
species 1s expanding its range in recent 
decades. The latter scenario seems more 
likely, since it is often common, and recent 
faunistic works (Campbell 1971, Peck and 
Thomas 1998) did not detect it. Also, it 
does occur in open, disturbed habitats and 
is associated in the same microsites with the 
adventive Gondwanocrypticus, e.g., the 
specimens from 21 June, and collections 
from the other localities mentioned above. 


Hymenorus transversus Campbell 


“respon, 20) Jue Is7~ CY; “Zi Jumae™ 
(1), “24 June” (2); “20-27 June 1987 / 
Malaise trap in Caribbean pine and pal- 
metto scrub” (3). 


This is a new island record for this Ba- 
hamian species; Campbell (1971) described 
it from the nearby Bimini Islands and others 
in the northern part of the Bahamas. It is 
the smallest of the Hymenorus on Grand 
Bahama. Information on specific larval mi- 
crohabitats for this and the other alleculines 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


on the island can only be inferred from data 
on related members of the group (Steiner 
1995). 


Glyptotus cribratus LeConte 


No specimens examined. 


The species was listed by Leng and 
Mutchler (1914) from “‘Bahama” and [ 
provisionally interpret this to mean Grand 
Bahama. Peck and Thomas (1998) gave 
widespread Florida records and also listed 
*“Bahamas”’ based on that record. The orig- 
inal source of the record is unknown. It 
could have been collected by H. EK Wick- 
ham, but Grand Bahama was apparently not 
visited by him (Wickham 1895). Its occur- 
rence on Grand Bahama would not be un- 
expected. In South Carolina, beetles have 
been taken in hollow trees and under bark 
(Kirk 1969) and I have observed similar 
habits for the species in several U.S. local- 
ities, and reared the larvae from dead 
branches on hardwood trees. A larva iden- 
tified as that of G. cribratus by St. George 
(1924) came from Big Pine Key, Florida 
where adults were noted to occur “in large 
numbers.” The species is widespread along 
the southeastern U.S. coastal plain from 
Texas to Virginia (Hoffman et al. 2003). 


BIOGEOGRAPHY 


Of the 31 species (Table 2) now known 
from Grand Bahama, 25 also occur in Flor- 
ida, including the 5 species known or 
suspected to be adventive. A subset of 18 
of those 25 species is also reported from 
Cuba. Colonization from these two land 
masses (which were closer to each other 
during the Pleistocene), and possibly via 
other Bahamian islands, is the likely source 
of these faunal elements. Many of these are 
winged species and widespread in Middle 
America. Only three of the species are 
flightless, and two of these (Branchus sax- 
atilis and Diastolinus bahamae) are appatr- 
ently endemic to the Littlhe Bahama Bank. 
The diversity of tenebrionids on Grand Ba- 
hama can be attributed to the relatively var- 


VOLUME 107, NUMBER 2 


ied and complex habitats on the island, in 
spite of its low elevation, from the sea drift 
and zones of maritime scrub to the pine for- 
est interior. This offers breeding sites for 
the geophilous forms as well as the rotten- 
wood and fungus-feeding specialists, and 
would facilitate colonizations by natural or 
anthropogenic means. Proximity to the 
mainland and other islands is probably the 
reason that a low percentage of endemic 
species occurs there. In addition, the geo- 
logically young Bahamas, having been 
completely inundated during the Pliocene 
(Browne et al. 1993), would be more open 
to colonization via over-water dispersal and 
less likely to have provided isolation for 
speciation events (Peck 1989). Some of the 
alleculine species, all winged, are apparent- 
ly endemic to the Bahamas but not neces- 
sarily restricted to single islands or banks, 
and one species, Hymenorus convexus, 1s 
thought to have colonized southern Florida 
from the Bahamas (Campbell 1971). 

With further collection and field work, 
additional species of Tenebrionidae are ex- 
pected to be documented for Grand Bahama 
in the future, especially the cosmopolitan 
stored-product pests and those winged spe- 
cies that have a wide natural distribution in 
the Caribbean region. A comparable survey 
of Abaco and surrounding cays 1s desirable. 
The other islands of the Bahamian banks 
await similar surveys. Inter-island faunal re- 
lationships can be better described with 
equivalent specialized sampling efforts on 
each island that build collections of such 
information-rich taxa. Flightless forms such 
as Branchus and Trientoma spp. and mem- 
bers of the Blapstinus-Diastolinus complex 
will be of the greatest interest and value in 
studies of biogeography. 


ACKNOWLEDGMENTS 


For assistance and company in fieldwork, 
I thank Mary Jo and Richard Molineaux 
and family, and my wife, Jil M. Swearin- 
gen. Current research and fieldwork in the 
Bahamas has been facilitated and encour- 
aged by Eric Carey and Tanya Ferguson, 


457 


Department of Agriculture, Nassau. Nancy 
B. Elliott, Siena College, Loudonville, NY, 
and Michael A. Ivie, Montana State 
University, Bozeman, MT, provided infor- 
mation on a number of collection holdings. 
Lucrecia Rodriguez, Systematic Entomolo- 
gy Laboratory, USDA, assisted with imag- 
ing specimens. For reviews of the manu- 
script and helpful comments, I thank Mi- 
chael A. Ivie and one anonymous reviewer. 


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Anderson, R. A. 1996. Beetles of Pigeon Cay, Bahama 
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Ardoin, P. 1977. Contribution a l'étude des espéces 
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. 1961. A few tenebrionids new to Cuba. The 

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VOLUME 107, NUMBER 2 


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PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 460-462 


NOTE 


Physopleurella floridana Blatchley, 1925, a Synonym of Physopleurella mundula 
(White, 1877) (Hemiptera: Heteroptera: Cimicoidea: Anthocoridae) 


The genus Physopleurella Reuter (1884) 
was erected to contain Cardiastethus mun- 
dulus White, 1877, described from the Ha- 
waiian Islands with no specific locality. 
Physopleurella now contains 13 species, in- 
cluding P. floridana Blatchley, 1925 (Ford 
1979). The type species, P. mundula 
(White), is known to occur on the Hawaiian 
Islands of Hawaii, Kauai, Maui, Molokai, 
and Oahu (Zimmerman 1948; Nishida 
1994, 1997, 2002). Zimmerman (1948) in- 
cluded an illustration of the adult of P. 
mundula and Hiura (1959) published on P. 
armata Poppius from Japan and included 
illustrations of the adult and other anatom- 
ical details. Herring (1967) reported this 
species from nine widely separated islands 
in Micronesia. Ford (1979) reported twelve 
other species of Physopleurella from Africa 
(Camaroon, Ivory Coast, Madagascar, Re- 
union, and Zaire), Australia, Fiji, Japan, 
New Guinea, Seychelles, Sri Lanka, and 
Taiwan, and P. floridana Blatchley from 
southern Florida and possibly from Central 
America. Carayon (1972) included Physo- 
pleurella in the Anthocoridae subfamily 
Lyctocorinae, tribe Cardiastethini, together 
with such genera as Amphiareus Distant, 
Brachysteles Mulsant and Rey, Buchanan- 
iola Reuter, Cardiastethus Fieber, Dufour- 
iellus Kirkaldy, and Tiare Herring. Today, 
this assemblage of taxa, and others, are 
placed in the tribe Dufouriellini (Ford 1979, 
Schuh and Slater 1995). 

Blatchley (1925) described Physopleu- 
rella floridana from Florida and added Pan- 
ama as another locality in 1926. Henry re- 
peated these records in 1988. The type of 
Physopleurella floridana Blatchley was 
made available through the kindness of A. 
Provonsha, Collection Manager of the Pur- 
due Entomological Research Collection. 


The type specimen is lacking the head, pro- 
thorax, front legs, mid- and hind legs on the 
left side, part of the left forewing and there 
has been some damage to the tip of the ab- 
domen. The right side of the specimen is 
imbedded in glue on the point. The speci- 
men has been examined previously as evi- 
denced by a second hole in the locality la- 
bel. The label reads as follows: Dunedin, 
Fla., Apr. 3, 1923. Provonsha indicated that 
the paratype was no longer in the collection 
of Purdue University (June 2004). A review 
of the species description in Blatchley’s 
publication clearly indicates characters of 
Physopleurella: short beak, faint preapical 
transverse impression on pronotum, ostiolar 
canal curved backward (and continued in a 
forward direction as a narrow carina [see 
Herring 1965, fig. 18]), front femur en- 
larged with inner surface with a row of 
spinelike setae and another row of setae, the 
anterior tibia curved to fit against the femur, 
and the rugulose mesopleura. Blatchley’s 
species description mentioned a faint fus- 
cous band across the apical portion of the 
clavus and corium, legs yellow brown, 
punctures fine and irregular on the clavus, 
corium, embolium, with a small seta in each 
puncture, and size 2.8—3 mm. I have com- 
pared the remains of the type with recently 
collected specimens of Physopleurella 
mundula (White) from Oahu and other Ha- 
waiian Islands (courtesy of the Bishop Mu- 
seum, Honolulu), including a specimen I 
collected on Oahu, April, 2004) and find 
these two species to be identical. Physo- 
pleurella floridana Blatchley, 1925, is here 
considered a synonym of Physopleurella 
mundula (White, 1877) (new synonomy). 
A label has been placed on the type speci- 
men of P. floridana that indicates the cur- 
rent status of Blatchley’s species. 


VOLUME 107, NUMBER 2 


Specimens of Physopleurella floridana 
Blatchley and P. mundula (White) were re- 
ceived through the efforts of T.J. Henry. 
The three specimens had been identified by 
the late H.M. Harris (no date of identifica- 
tion cited), and were in the H.M. Harris 
Collection donated to the National Museum 
of Natural History, Smithsonian Institution 
(USNM) in 1977. The collection data fol- 
lows: 1 2 Mexico intercept/Phila. Pa.//June 
8, 1933/A.G. Wells//on banana leaf//flori- 
dana/H.M. Harris Coll. 1977; 1 2 Hondu- 
ras on bananas//intercept N.Orleans 2-17- 
36//floridana, H.M.Harris Coll.; 1 6 (ab- 
domen missing) Canal Zone; Barro Colo- 
rado, 18-vii-1924. N.Banks//Physopleurella 
floridana Det. H.M. Harris (no date)//flori- 
dana H.M.Harris Coll. 1977. The speci- 
mens of P. mundula (White) were identified 
by R.C. Froeschner in 1961 and had the 
following collection data: | d Honolulu, 
Feb. 10-43//Ti leaf, hula skirt, Hawaii No. 
842//ot No. 43-3889.//Physopleurella mun- 
dula (White), det. Froeschner ’61; 1 2 Bar- 
ber Point, Oahu, Hawaii, II-10-65//ex Ficus 
retusa//C.J.Davis Collector, ¢ Wahiawa, 
Oahu 7-58//light trap, E.J.Ford, Jr; 1 5 (?) 
(abdomen missing) Honolulu, T.-H. I-23-43/ 
Hawau 821//Ti leaf hula skirt//lot H43- 
2001. As indicated above, Physopleurella 
floridana is considered to be a synonym of 
P. mundula, accordingly, the three speci- 
mens cited above have been identified as 
such and a label Physopleurella mundula 
(White) placed on each specimen. Note that 
the records of P. mundula from Mexico, 
Honduras, and the Canal Zone constitute 
new country records of P. mundula, and the 
Florida specimens of P. floridana (now P. 
mundula) also represent a new record (Flor- 
ida) and add yet another non-indigenous 
species of Anthocoridae to the fauna of 
America north of Mexico.) The specimens 
from Oahu, Hawaiian Islands, will be in- 
cluded in a manuscript now in preparation 
on the Lasiochilidae, Lyctocoridae, and An- 
thocoridae of the Hawaiian Islands. Nishida 
(1997, 2002) reported P. mundula from the 
islands of Hawaii, Kauai, Maui, Molokai, 


461 


and Oahu. Elsewhere, this species has been 
reported from Guam (Usinger 1946), the 
Marshall Islands (Usinger 1951), and Fiji 
(Gross 1954). 

Most records on habitats for this species 
stated that specimens were beaten from 
dead leaves of palmetto (Blatchley 1926); 
beating dead leaves and branches (Usinger 
1946, 1951); banana leaves (USNM speci- 
mens), ti leaves of hula skirt(!); Ficus re- 
tusa (USNM specimens). Zimmerman 
(1948) reported a collection that included 
sugar cane, roofs of houses, dead cane 
leaves (preying on psocids and small leaf- 
hoppers) where it was most common where 
psocids occurred. The specimen I collected 
on Oahu was taken beating dead leaves. 
These habits and habitats are common to 
the habits of many Dufouriellini (e.g., Lat- 
tin 1999a, b). 

Acknowledgments.—My thanks to A. 
Provonsha, Collection Manager, Purdue En- 
tomological Research Collection; T. J. Hen- 
ry, U.S.D.A. Systematic Entomology Lab- 
oratory, Washington, D.C.; A. Ramsdale, 
Collection Manager, Bishop Museum, Hon- 
olulu; and to L. Parks, Oregon State Uni- 
versity, for careful attention to the manu- 
script, and to a careful reviewer. 


LITERATURE CITED 


Blatchley, W. S. 1925. Some additional new species of 
Heteroptera from the southern United States, with 
characterization of a new genus. Entomological 
News 36: 45-52. 

. 1926. Heteroptera or True Bugs of Eastern 

North America, with Especial Reference to the 


Faunas of Indiana and Florida. Nature Publishing 
Company, Indianapolis, 1,116 pp. 

Carayon, J. 1956. Trois especes afraicanes de Physo- 
pleurella (Hemipt. Bulletin du 
Muséum National d’ Histoire Naturelle (2) 28(1): 
102-110. 

. 1972. Cautéres systématiques et classification 


Anthocoridae). 


des Anthocoridae (Hemipt.). Annales de la Société 
Entomologique de France (N.S.). 8: 309-349. 
Ford, L. J. 
the Cimicoidea (Insecta: Hemiptera). Thesis, M.S. 
Degree, University of Connecticut, vii + 138 pp. 
Gross, G. EF 1954. A revision of the flower bugs (Het- 


1979. The phylogeny and biogeography of 


eroptera: Anthocoridae) of the Australian and ad- 


462 


jacent Pacific region. Part I. Records of the South 
Australian Museum 10: 129-164. 

Henry, T. J. 1988. Family Anthocoridae Fieber, 1837, 
pp. 12-28. Jn Henry, T. J. and R. C. Froeschner, 
eds. Catalog of the Heteroptera, or True Bugs, of 
Canada and the Continental United States. Brill, 
Leiden, 959 pp. 

Herring, J. L. 1965. Keys to genera of Anthocoridae 
of America north of Mexico, with description of 
a new genus (Hemiptera: Heteroptera). Florida 
Entomologist 59: 143-150. 

. 1967. Insects of Micronesia Heteroptera: An- 
thocoridae. Insects of Micronesia, Vol. 7, Number 
8: 390-414. 

Hiura, I. 1959. Contribution to the knowledge of An- 
thocoridae from Japan and its adjacent territories 
(Hemiptera: Heteroptera). 1. Bulletin of the Osaka 
Museum of Natural History, No. 11: 1-15. 

Lattin, J. D. 1999a. Dead leaf clusters as habitats for 
adult Calliodis temnostethoides and Cardiastethus 
luridellus and other anthocorids (Hemiptera: Het- 
eroptera: Anthocoridae). Great Lakes Entomolo- 
gist 32: 32-38. 

. 1999b. Bionomics of the Anthocoridae. An- 
nual Review of Entomology 44: 207-301. 

Nishida, G. M., ed. 1994. Hawaiian terrestrial arthro- 
pod checklist. Second edition. Hawaii Biological 
Survey, Contribution No. 94-04. Bishop Museum 
Technical Report No. 4, 287 pp. 

. 1997. Hawaiian terrestrial arthropod checklist. 

Third edition. Hawaii Biological Survey, Bishop 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Museum. Bishop Museum Technical Report No. 

12, 263 pp. 

. 2002. Hawaiian terrestrial arthropod checklist. 
Fourth edition. Hawaii Biological Survey, Bishop 
Museum. Bishop Museum Technical Report No. 
Wp Ba)\\ J0)9) 

Reuter, O. M. 1884. Monographic Anthocoridarum or- 
bis terrestris. Acta Societatis Scientiarum Fenni- 
cae 14: 555-758. Separate, Helsingfors, 1884, 1— 
204. 

Schuh, R. T. and J. A. Slater. 1995. True Bugs of the 
World (Hemiptera: Heteroptera). Comstock Pub- 
lishing Association, Ithaca, New York, 336 pp. 

Usinger, R. L. 1946. Hemiptera, pp. 11-103. Jn Insects 
of Guam. Bernice P. Bishop Museum. Bulletin 
189, 237 pp. 

1951. Heteroptera of the Marshall Islands. 
Proceedings, Hawaiian Entomological Society 
14(2): 315-321. 

White, F B. 1877. Description of new species of het- 
eropterous Hemiptera collected in the Hawaiian 
Islands by the Rev. T. Blackburn. Annals and 
Magazine of Natural History, Series 4, 10: 110— 
114. 

Zimmerman, E. C. 1948. Insects of Hawaii, Vol. 3 Het- 
eroptera. University of Hawaii Press, Honolulu, 
255 pp. 


John D. Lattin, Department of Botany 
and Plant Pathology, Oregon State Univer- 
sity, Corvallis, OR 97331-2902, U.S.A. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 463-465 


NOTE 


Larva and Possible Food Plant of Ancyloneura varipes (Cameron) 
(Hymenoptera: Pergidae) in Papua New Guinea 


The six known species of Ancyloneura 
Cameron (Pergidae: Euryinae) are endemic 
to New Guinea (Smith 1978). Nothing is 
known of the food plants, habits, or larvae 
of any species of the genus. During a study 
of caterpillars (Lepidoptera) feeding on the 
alien plant Spathodea campanulata (L.) 
Kunth. (Bignoniaceae) in Wamangu, Papua 
New Guinea, by DB, an adult of Ancylo- 
neura varipes (Cameron) was reared from 
larvae discovered on the foliage. Ancylo- 
neura varipes was described from *‘Aru”’ 
by Cameron (1877), and additional locali- 
ties in Irian Jaya and Papua New Guinea 
were given by Benson (1958) and Smith 
(1980). Although sketchy, this is the first 
information on the habits and larvae of An- 
cyloneura. 

Larvae were collected at Wamangu in 
East Sepik Province (143°49’E, 3°49'S). 
This area was sampled for Lepidoptera 
from 15 September to 2 October, 2003, cov- 
ering over 4,500 square meters of foliage 
area. Searches were made on saplings and 
lower branches of larger trees. Background 
of the project and sampling and rearing pro- 
cedures were provided by Miller et al. 
(2003). Larvae of A. varipes were not found 
on any of the other 90 woody plant species 
studied (Miller et al. 2003). 

Four pergid larvae were collected in the 
14 sampling days. The first two larvae were 
found together on the foliage of a sapling 
growing in the undergrowth of the second- 
ary forest area. They were about 60—80 cm 


above ground level. There were no signs of 


feeding or feeding damage on the leaf. The 
caterpillar-like features such as its black 10- 
segmented body and sclerotized head cap- 
sule indicated similar feeding habits as lep- 
idopteran caterpillars, but they did not feed 
on fresh leaves given to them. Rearing in 


the laboratory resulted in one larva reared 
to adult. This was from a larva that pupated 
five days after the first day of collection. 
The final larval instar took almost three 
days to complete pupation. The adult 
emerged seven days after pupation. The 
reared adult and the larva were preserved 
for identification. The other two larvae col- 
lected likewise did not show signs of feed- 
ing in the laboratory. The larvae may have 
been in their final instars when collected be- 
cause they soon pupated; however, no 
adults emerged. 

Although collections were from S$. cam- 
panulata and pupation and subsequent rear- 
ing to adult for one of the four larvae were 
successful in the laboratory, no larvae 
showed signs of feeding on this plant. Spa- 
thodea campanulata possibly is not the 
food plant. The larvae may have completed 
feeding and dropped from other plants in 
search of a pupation site and were only in- 
cidental on S. campanulata. Also, larvae 
may feed only on dead or dry leaves as do 
some other species of Euryinae in Austra- 
lia, thus not accepting fresh foliage. Moore 
(1957) recorded Polyclonus atratus (Kirby) 
from Australia feeding on dead or dying 
leaves of Eucalyptus and Angophora still 
attached to branches, and Tillyard (1926) 
mentioned this habit for a species of Di- 
phamorphos Rohwer. Other Euryinae in 
Australia are known to feed on dead or dry- 
ing leaves or leaves in leaf litter (Naumann 
1991) and one on ferns (Naumann and Bal- 
ciunas 1997), and most are found on or near 
the ground. 

The larva illustrated (Figs. 1, 2) may be 
a prepupa (last instar or non-feeding stage) 
when collected in the field. It is about 6.5 
mm long and entirely dark brown with the 
clypeus white, area below the eyes and an- 


464 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-2. 


tenna reddish brown, and prolegs pale 
brown. The antenna appears four segment- 
ed, with two small oval segments adjacent 
to one larger circular segment, and a small 
node within the circular segment. The head 
is covered with short spatulate setae, with 
simple stiff setae on the clypeus, mouth- 
parts, the lower area of the frons, and 
around the antennae. Each thoracic leg has 
a tarsal claw with a large fleshy lobe adja- 
cent to the claw (as in many Argidae). 
There are deep folds between the segments 
and the surpedal lobes are expanded later- 


Ancyloneura varipes, larva. 1, Lateral view. 2, Dorsal view. 


ally. The abdominal segments appear to 
have three annulets with the third larger 
than the first two; the tenth tergum is 
notched at the center in dorsal view. Prolegs 
are present on abdominal segments 2-8, 
with those on segment 2 smaller than the 
others. The thorax and abdomen are coyv- 
ered with short, spatulate setae, similar to 
those on the head; simple stiff setae are 
present on the thoracic legs. 

The larva is quite different from known 
Australian euryine larvae. Larvae of Clar- 
issa tasbates Naumann from Tasmania, 


VOLUME 107, NUMBER 2 


which are apparently ground dwelling and 
feed on dead Ranunculus leaves (Naumann 
1997), and larvae of Warra froggatti (Roh- 
wer), which feeds on emergent foliage of 
an aquatic fern (Naumann and Balciunas 
1997), both have four-annulate abdominal 
segments, have simple setae, apparently 
lack the large fleshy lobe adjacent to the 
tarsal claws, and do not have such deep 
folds and laterally expanded surpedal lobes 
on the body. 

The larva illustrated and reared adult are 
deposited in the National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, DC. Cathy Apgar, Systematic 
Entomology Laboratory, USDA, took the 
photographs. DB’s research in Papua New 
Guinea was supported by National Science 
Foundation grant DEB 0211591 to George 
Weiblen, Scott Miller, Vojtech Novotny, 
and Yves Basset, as well as the Czech 
Academy of Sciences and Darwin Initiative 
grant 162/10/030. 


LITERATURE CITED 


Benson, R. B. 1958. On some sawflies (Hymenoptera 
Symphyta) from New Guinea. Proceedings of the 
Entomological Society of London (B) 27: 15-18. 

Miller, S. E., V. Novotny, and Y. Basset. 2003. Studies 
on New Guinea moths. |. Introduction (Lepidop- 
tera). Proceedings of the Entomological Society of 
Washington 105: 1035-1043. 

Moore, K. M. 1957. Observations on some Australian 
forest insects. Proceedings of the Royal Zoologi- 
cal Society of New South Wales 1955—1956: 74— 
81. 


465 


Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, 
sawflies), pp. 916-1000. Jn CSIRO, The Insects 
of Australia, A Textbook for Students and Re- 
search Workers, Vol. 2. 
Press, pp. 543-1137. 

. 1997. A remarkable, new Australian sawfly 
with brachypterous, nocturnal or crepuscular fe- 
males (Hymenoptera: Symphyta: Pergidae). Jour- 
nal of Natural History 31: 1335-1345. 

Naumann, I, D. and J. K. Balciunas. 1997. A sawfly 
larva feeding on aquatic fern (Hymenoptera: Sym- 
phyta: Pergidae). Australian Entomologist 24: 39— 
47. 

Smith, D. R. 1978. Suborder Symphyta (Xyelidae, Par- 
achexyelidae, Parapamphiliidae, Xyelydidae, Kar- 
atavitidae, Gigasiricidae, Sepulcidae, Pseudosiri- 
cidae, Anaxyelidae, Siricidae, Xiphydriidae, Pa- 
roryssidae, Xylotomidae, Blasticotomidae, Pergi- 
dae). Jn van der Vecht, J. and R. D. Shenefelt, eds. 
Hymenopterorum Catalogus, pars 14, 193 pp. 

1980. Pergidae (Hymenoptera) from New 
Guinea and Australia in the Bishop Museum. Pa- 
cific Insects 22: 329-346. 

Tillyard, R. J. 1926. The Insects of Australia and New 
Zealand. Angus & Robertson, Ltd., Sydney, xv + 
560 pp. 


Melbourne University 


Darren Bito and David R. Smith. (DB) 
University of Papua New Guinea and New 
Guinea Binatang Research Center, P.O. 
Box 604, Madang, Papua New Guinea (e- 
mail: binatangi@datec.com.pg); (DRS) 
Systematic Entomology Laboratory, PSI, 
Agricultural Research Service, U.S. De- 
partment of Agriculture, c/o National Mu- 
seum of Natural History, Smithsonian In- 
stitution, P.O. Box 37012, MRC 168, Wash- 
ington, DC, 20013-7012, U.S.A. (e-mail: 
dsmith@ sel.barc.usda.gov) 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 466-468 


NOTE 


Dufouriellus ater (Puton), Macrotrachelia nigronitens (Stal), and Xylocoris (Arrostelus) 
flavipes (Reuter) (Hemiptera: Heteroptera: Cimicoidea: Anthocoridae): First Records 
from the Hawaiian Islands 


A recent shipment of unidentified Lasi- 
ochilidae, Lyctocoridae and Anthocoridae 
from the Bishop Museum, Honolulu, in- 
cluded specimens of three species not pre- 
viously reported from the Hawaiian Islands. 
Currently, I am reviewing these families 
found on the Hawaiian Islands. (Note: Fol- 
lowing Schuh and Stys (1991) and Schuh 
and Slater (1995), three families are now 
recognized: Lasiochilidae, Lyctocoridae, 
and Anthocoridae.) The species reported 
here are placed in the Anthocoridae. 

Dufouriellus ater was described from Eu- 
rope and Syria by Dufour (1833) (as Xylo- 
coris ater). Subsequently, this species has 
been reported from Europe, North Africa, 
Middle East, and Asia Minor (Péricart 
1972). It was first reported from North 
America by Van Duzee (1916) and subse- 
quently reported from B.C., Calif., Id., Ky., 
N.C., N.Y. and Ont. in Canada and the 
United States (Henry 1988) and Ore. (J.D. 
L., unpublished). Zheng and Bu (1990) re- 
ported it from China. Lattin (1999, 2000) 
reviewed this species and its habits that in- 
cluded occurrence under bark where it at- 
tacks bark beetles and elsewhere as a pred- 
ator of stored product pests. 

Two specimens of Dufouriellus ater, a 
male and a Sth instar nymph, were found 
in the material from the Bishop Museum. 
The label information reads: Hawaii, Ha- 
wauian I., Saddle Road at Pohakuloa, 1829 
m., 10-1-1971//Plasithymus borings//in 
Chenopodium//W.C. Gagné Collector, Bish- 
op Museum 1971: 366. This species is like- 
ly to be found under bark at higher eleva- 
tions for this is a temperate species and 
could be taken from stored products as it is 
a known predator of stored product pests. 

Macrotrachelia nigronitens (Stal) was 


described from Brazil (1860) (as Anthocoris 
nigronitens). Later, it was reported from 
Panama by Champion (1900). Specimens of 
this species were deliberately introduced 
into Oahu in 1965 from Cuernavaca, Mo- 
relos, Mexico, collected by Krauss and re- 
ported by Davis and Krauss (1966) (as Ma- 
crotrachelia thripiformis Champion). Six 
specimens (3 males and 3 females) of this 
original series were in the unidentified col- 
lections of the Bishop Museum and a large 
label stated “Part of series collected and re- 
leased.” These specimens were identified 
by me as Macrotrachelia nigronitens (Stal) 
rather than M. thripiformis utilizing the key 
of Champion and the illustrations of this 
species (Plate 19, figs. 18, 19, 19a, b) in the 
same publication. These specimens agreed 
in every way with the description and il- 
lustrations on M. nigronitens, not M. thri- 
piformis Champion. This is a correction of 
the information that appeared in several 
publications following the introduction 
(Davis for Chong 1967; Nishida 1994, 
1997, 2002). No additional specimens of 
Machrotrachelia nigronitens have been 
seen since its original release. Nishida re- 
ported its occurrence on Kauai, Oahu, Mo- 
lokai, Maui, and Hawaii (as M. thripifor- 
mis), but it appears that these island records 
actually apply to another Anthocoridae, 
very likely Montandoniola moraguesi (Pu- 
ton), introduced from the Philippine Islands 
in 1964 (Davis and Krauss 1966) into Kau- 
ai and Oahu and quickly spreading to Maui, 
Molokai, and Hawaii (Funasaki 1966). 
Montandoniola moraguesi was not included 
in the checklists of Nishida (1994, 1997). 
Xylocoris (Arrostelus) flavipes was de- 
scribed by Reuter from Lithuania in 1875 
(as Piezostethus flavipes). Péricart (1972) 


VOLUME 107, NUMBER 2 


published a review of this species and in- 
cluded illustrations of the brachypterous 
and macropterous adults. It was first re- 
ported from the United States by Bibby 
(1961) although specimens were dated well 
before that date (see Henry 1988). Péricart 
(1996) cited the occurrence of this species 
in North Africa, Turkey, China, Saudi Ara- 
bia, and Yemen, and North America, South 
America, India and Indonesia. A citation of 
particular interest is that of Orian (1956) 
who reported X. flavipes from the island of 
Mauritius. Henry (1988) reported this spe- 
cies from Ariz., D.C., Ga., Ks., Md., and 
Tex. Arbogast et al. (1971) reviewed the ac- 
tivities of this bug as a predator of stored 
grain pests. Awadalla and Tawfik (1972) 
published on the biology of this species in 
Egypt, followed by LaCato and Davis 
(1973) on the interactions of the bug with 
various species of stored grain pests in the 
United States. Press (1989) studied the 
compatibility of X. flavipes with a parasitic 
hymenopteran in the suppression of the al- 
mond moth. Two brachypterous specimens 
of Xylocoris flavipes were found in the col- 
lections of the Bishop Museum, one bra- 
chypterous male from Kalmuk, Oahu, 10- 
28-20, Coll. O.H.S./with O. oryzae L. 
[(now Sitophilus oryzae (L.)], in sorghum 
seed. The other brachypterous male came 
from Honolulu, T.H., 11-19-35/R.A. Smith/ 
ex organic fertilizer. While no more recent- 
ly collected specimens have been seen, this 
is a very small bug (1.7 mm) and easily 
overlooked when examining lots of organic 
materials. 

Thus, three additional non-indigenous 
species of Anthocoridae are added to the 
reported fauna, substantially increasing the 
number of non-native species known from 
the Hawaiian Islands (8 native, 14 non-na- 
tive). This is a remarkably high percentage 
of non-native species in the Hawaiian Is- 
lands, 64%, well above the number cited for 
all insects of the Hawaiian Islands by Nish- 
ida. Careful collecting at higher elevations 
on the islands will likely add additional spe- 
cies for the fauna—both native and non-na- 


467 


tive. These are small insects, and for that 
reason, they are likely to be overlooked by 
the average collector. It is appropriate to 
mention the important collections of Ha- 
waiian Hemiptera: Heteroptera made by the 
late Wayne C. Gagné. He added many in- 
teresting specimens of Heteroptera, includ- 
ing Anthocoridae, to the collections of the 
Bishop Museum. 

Acknowledgments.—My thanks go to A. 
Ramsdale, Collection Manager, Bishop Mu- 
seum, for the loan of specimens and to L. 
Parks, Oregon State University, Corvallis, 
for manuscript preparation, and to a careful 
reviewer. 


LITERATURE CITED 


Arbogast, R. T., M. M. Carthon, J. R. Roberts, Jr. 1971. 
Developmental stages of Xylocoris flavipes (He- 
miptera: Anthocoridae), a predator of stored prod- 
uct insects. Annals of the Entomological Society 
of America 64: 1131-1134. 

Awadallah, K. T. and M. E S. Tawfik. 1972. The bi- 
ology of Xylocoris (= Piezostethus) flavipes 
(Reut.) (Hemiptera: Anthocoridae). Bulletin de la 
Société Entomologique d’Egypte 56: 177-189. 

Bibby, R. E 1961. Notes on miscellaneous insects of 
Arizona. Journal of Economic Entomology 54: 
324-333. 

Champion, G. C. 1900. Insecta. Rhynchota. Hemip- 
tera: Heteroptera, Vol. Il. /n Biologia Centrali— 
Americana, London, xvi + 416 pp. 

Davis, C. J. and N. L. N. Krauss. 1966. Recent intro- 
ductions for biological control in Hawaii. XI. Pro- 
ceedings, Hawaiian Entomological Society 19(2): 
201—207. 

Davis, C. J. for M. Chong. 1967. Macrotrachelia thri- 
piformis Champion. Proceedings, Hawaiian En- 
tomological Society 19(2): 379. 

Dufour, L. 1833. Mémoire sur les genres Xy/locoris, 
Leptopus et Velia. Annals de la Société Entomo- 
logique de France 2: 104-118. 

Funasaki, G. Y. 1966. Studies on the life cycle and 
propagation technique of Montanandoniola mor- 
aguesi (Puton) (Heteroptera: Anthocoridae). Pro- 
ceedings, Hawaiian Entomological Society 19: 
209-211. 

Henry, T. J. 1988. Family Anthocoridae Fieber, 1837, 
pp. 12-28. /n Henry, T. J. and R. C. Froeschner, 
eds. 1988. Catalog of the Heteroptera, or True 
Bugs, of Canada and the Continental United 
States. E.J. Brill, Leiden, 958 pp. 

Lattin, J. D. 1999. Bionomics of the Anthocoridae. An- 
nual Review of Entomology 44: 207-231. 

—. 2000. Minute pirate bugs (Anthocoridae), pp. 


468 


607—637. In Schaefer, C. W. and A. R. Panizzi, 
eds. Heteroptera of Economic Importance. CRC 
Pres, Baton Raton, 828 pp. 

LeCato, G. L. and R. Davis. 1973. Preferences of the 
predator Xylocoris flavipes (Hemiptera: Anthocor- 
idae) for species and instars of stored-product in- 
sects. The Florida Entomologist 56(1): 57-59. 

Nishida, G. M., ed. 1994. Hawaiian terrestrial arthro- 
pod checklist. Second edition. Bishop Museum 
Technical Report No. 4, 287 pp. 

. 1997. Hawaiian terrestrial arthropod checklist. 

Third edition. Hawaii Biological Survey Bishop 

Museum. Bishop Museum Technical Report No. 

12, 263 pp. 

. 2002. Hawaiian terrestrial arthropod checklist. 
Fourth edition. Hawaii Biological Survey Bishop 
Museum. Bishop Museum Technical Report No. 
22, iv + 310 pp. 

Orian, A. J. E. 1956. Hemiptera (Heteroptera and Ho- 
moptera excluding Sternorhyncha) of Mauritius. 
Annals and Magazine of Natural History Series 
12, 9: 641-654. 

Péricart, J. 1972. Hémipteres Anthocoridae, Cimicidae 
et Microphysidae de lOuest—Paléarctique. In 
Faune de l’Europe et du Bassin Méditerranéen 7: 
i-iv, 1-404. Masson, Paris. 

. 1996. Family Anthocoridae Fieber, 1836, pp. 

108-140. In Aukema, B. and C. Rieger, eds. Cat- 

alogue of the Heteroptera of the Palearctic Region, 

Vol. 2. Cimicomorpha I. The Netherlands Ento- 

mological Society, The Netherlands, 359 pp. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Press, J. W. 1989. Compatibility of Xylocoris flavipes 
(Hemiptera: Anthocoridae) and Venturia canes- 
cens (Hymenoptera: Ichneumonidae) for suppres- 
sion of the almond moth Cadra cautella (Lepi- 
doptera: Pyralidae). Journal of Entomological Sci- 
ence 24: 156-160. 

Reuter, O. M. 1875. Genera Cimicidarum Europe. Bi- 
hang till Konglia Svenska Vetenskapsak-Ade- 
miens Handlingar 3: 3—66. 

Schuh, R. T. and J. A. Slater. 1995. True bugs of the 
world (Hemiptera: Heteroptera). Comstock Pub- 
lishing Association, Ithaca, New York, 336 pp. 

Schuh, R. T. and P. Stys. 1991. Phylogenetic analysis 
of cimicomorphan family relationships (Heterop- 
tera). Journal of the New York Entomological So- 
ciety 99: 298-350. 

Stal, C. 1860. Bidrag till Rio Janeiro—trakens Hem- 
ipter—Fauna. Kongliga Svenska Vetenskapsak- 
Ademiens Handlingar, Stockholm, II, No. 7: 1-84. 

Van Duzee, E. P. 1916. Checklist of the Hemiptera 
(excepting the Aphididae, Aleyrodidae and Coc- 
cidae) of America North of Mexico. New York 
Entomological Society, New York, xi + 111 pp. 

Zheng, L. Y. and W. J. Bu. 1990. A list of Anthocor- 
idae from China. Contributions from the Tianjin 
Natural History Museum 7: 23-27 (in Chinese). 


John D. Lattin, Department of Botany 
and Plant Pathology, Oregon State Univer- 
sity, Corvallis, OR 97331-2902, U.S.A. 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 469-470 


NOTE 


A New Non-Destructive DNA Extraction and Specimen Clearing Technique 
for Aphids (Hemiptera) 


Slide mounting soft bodied insects such 
as aphids is a time-consuming task that is 
often regarded as art and alchemy more 
than science. Many techniques have been 
developed to prepare specimens (Essig 
1948, Wilkey 1962, Martin 1983). The typ- 
ical use of potassium hydroxide (KOH) or 
sodium hydroxide (NaOH) complicates the 
process as one tries to strike a balance be- 
tween adequate clearing and rendering the 
specimen too fragile to handle. A technique 
developed by Hille Ris Lambers (1950) em- 
ploys chloral phenol in an effort to mitigate 
the risks to the specimen when using KOH. 
However, one key ingredient of chloral phe- 
nol, chloral hydrate, is a controlled sub- 
stance and is dangerous and unpleasant to 
use. I modified a standard DNA extraction 
technique that not only clears aphid speci- 
mens easily and with few complications, 
but also extracts the aphid’s DNA non-de- 
structively, allowing for specimen vouch- 
ering of genetic sources. 

Specimens of some taxa must be de- 
stroyed in order to extract their DNA. In 
fact, some organisms are known only by 
their DNA sequences (Pace 1997). How- 
ever, there are benefits in preserving the ac- 
tual specimen from which the DNA is ex- 
tracted. Various techniques have been de- 
veloped to extract DNA from insect speci- 
mens relatively nondestructively (Phillips 
and Simon 1995, Johnson et al. 2001, 
Starks and Peters 2002). Keeping a voucher 
specimen from which the DNA is taken is 
especially useful in the case of solitary 
aphids, when multiple clonal individuals 
from a single colony are not available. 

I adapted the DNA extraction protocol 
from Favret and Voegtlin (2004) to pre- 
serve the aphid cuticle intact for mounting 
to a microscope slide. The extraction solu- 


tion consists of 500 41 STE buffer (pH 7.5), 
25 wl of 10 mg/ml proteinase K, and 75 pl 
of 10% SDS (Hillis et al. 1996: 342-343). 
An incision along the length of the ventral 
aspect of the aphid abdomen is made under 
a microscope with a bent minuten pin 
mounted on the end of a thin dowel. The 
incision allows entry of the clearing solu- 
tion into the aphid body. The entire aphid 
is placed in a microvial containing the ex- 
traction solution and the vial placed in a 
55°C water bath overnight. The next day 
the aphid is removed from the buffer using 
sterilized forceps, DNA purification contin- 
ues as per the normal protocol (Hillis et al. 
1996, Favret and Voegtlin 2004), and the 
cuticle is dehydrated and mounted to a mi- 
croscope slide. The following steps are nec- 
essary to prepare the specimen for slide 
mounting in Canada balsam: 10 minutes in 
70% ethanol, 10 minutes in 95% ethanol, 2 
minutes in glacial acetic acid, 10 minutes 
in a 1:1 mixture of glacial acetic acid and 
terpineol, and 10 minutes in pure terpineol. 
After the specimen is positioned on the 
Slide in balsam, a cover slip is applied and 
the slide is cured on a slide warmer or in 
drying oven at 50°C for a week or more. 
For DNA extraction, each microvial must 
contain only one specimen, but several can 
be cleared simultaneously in a single mi- 
crovial if the DNA is not to be saved. The 
extraction solution clears the aphid similar- 
ly to conventional clearing techniques, yet 
leaves the aphid cuticle supple and strong. 
On rare occasions, large or heavily pig- 
mented aphids need additional clearing; for 
these, | remove the embryos from the body 
and repeat the clearing cycle. Aphids 
DNA extraction buffer and 
mounted on indistinguishable 
from those prepared by other means such 


cleared in 


slides are 


470 


as the Hille Ris Lambers (1950) method. 
This technique should be adaptable to most 
DNA extraction protocols such as the Qia- 
gen kit used by Johnson et al. (2001). They 
left lice specimens in the extraction buffer 
for 56 h, but I found overnight to be suf- 
ficient for most aphids, and most specimens 
yielded large, visible DNA pellets. 

The clearing technique presented here is 
not only easy to use and forgiving, but also 
allows for non-destructive DNA extraction 
and specimen vouchering. This latter advan- 
tage has positive ramifications for insect tax- 
onomy, and has allowed me to confirm the 
identity of Cinara atlantica (Wilson) in Bra- 
zil, which is not morphologically distin- 
guishable from C. ponderosae (Williams): 
CO-1 DNA sequence from a Brazilian spec- 
imen was identical to that of C. atlantica 
from Florida (GenBank accession number 
AY 300225), differed by one base from that 
of C. atlantica from South Carolina 
(AY300198), and differed from C. ponder- 
osae by 15 bases (AY300194). Likewise, I 
also have identified a Cinara nymph on an 
imported host, Pinus nigra Arnold in Ne- 
braska. Nymphs are difficult or impossible 
to identify morphologically and host-based 
keys are for European Cinara species only 
(Blackman and Eastop 1994). However, CO- 
1 DNA sequence data (AY300229) matched 
those found previously for C. arizonica 
(Wilson) (AY300222 and AY300230). This 
is the first record of C. arizonica from P. 
nigra and also the first record of this aphid 
species in Nebraska. Both the Brazilian and 
Nebraskan aphids whose DNA was extract- 
ed using the protocol described above are 
intact and deposited in the insect collection 
at the Illinois Natural History Survey, 
Champaign, IL: catalog numbers 18,292 and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


16,756 in the insect collection database at 
http://www.inhs.uiuc.edu. 


LITERATURE CITED 


Blackman, R. L. and V. E Eastop. 1994. Aphids on 
the World’s Trees: An Identification and Infor- 
mation Guide. CAB International, Wallingford, 
UK. 

Favret, C. and D. J. Voegtlin. 2004. Speciation by host- 
switching in pinyon Cinara (Insecta: Hemiptera: 
Aphididae). Molecular Phylogenetics and Evolu- 
tion 32: 139-151. 

Essig, E. O. 1948. Mounting aphids and other small 
insects on microscope slides. Pan-Pacific Ento- 
mologist 24: 9-22. 

Hille Ris Lambers, D. 1950. On mounting aphids and 
other soft-skinned insects. Entomologische Beri- 
chten (Amsterdam) 298: 55—58. 

Hillis, D. M., B. K. Mable, A. Larson, S. K. Davis, 
and E. A. Zimmer. 1996. Nucleic acids IV: se- 
quencing and cloning. Jn Hillis, D. M., C. Moritz., 
and B. K. Mable, eds. Molecular Systematics. Sin- 
nauer Associates, Sunderland, MA. 

Johnson, K. P., R. J. Adams, and D. H. Clayton. 2001. 
Molecular systematics of Goniodidae (Insecta: 
Phthiraptera). Journal of Parasitology 87: 862— 
869. 

Martin, J. H. 1983. The identification of common 
aphid pests of tropical agriculture. Tropical Pest 
Management 29: 395-411. 

Pace, N. R. 1997. A molecular view of microbial di- 
versity and the biosphere. Science 276: 734-740. 

Phillips, A. J. and C. Simon. 1995. Simple, efficient, 
and nondestructive DNA extraction protocol for 
arthropods. Annals of the Entomological! Society 
of America 88: 281—283. 

Starks, P. T. and J. M. Peters. 2002. Semi-nondestruc- 
tive genetic sampling from live eusocial wasps, 
Polistes dominulus and Polistes fuscatus. Insectes 
Sociaux 49: 20-22. 

Wilkey, R. E 1962. A simplified technique for clear- 
ing, staining and permanently mounting small ar- 
thropods. Annals of the Entomological Society of 
America 55: 606. 


Colin Favret, J//linois Natural History 
Survey, Center for Biodiversity, 607 E. Pea- 
body Dr., Champaign, IL 61820, U.S.A. (e- 
mail: crf@uiuc.edu) 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 471-473 


NOTE 


Fleas of the genus Ceratophyllus (Siphonaptera: Ceratophyllidae) 
in the southeastern United States 


Species of Ceratophyllus Curtis are rare 
in collections in the southeastern USA. Pre- 
vious records of this genus in the south- 
eastern USA include Ceratophyllus (Cera- 
tophyllus) idius Jordan and Rothschild from 
nests of the purple martin, Progne subis 
(Linnaeus), and Ceratophyllus (Monopsyl- 
lus) vison Baker from the nest of the red 
squirrel, Tamiasciurus hudsonicus (Erxle- 
ben), in Tennessee (e.g., Benton 1980, Dur- 
den and Kollars 1997). Collections were 
made to determine if this genus is present 
in Georgia, North Carolina, and South Car- 
olina. We report for the first time Cerato- 
phyllus (Ceratophyllus) celsus Jordan in 
Georgia and South Carolina, Ceratophyllus 
(Ceratophyllus) gallinae (Schrank) in North 
Carolina, and C. vison in North Carolina. 
Our new records increase the total number 
of fleas recorded from Georgia to 20, North 
Carolina to 19 (Benton 1980), and South 
Carolina to 26 (Durden et al. 1999). Echid- 
nophaga _ gallinacea (Westwood) (Pulici- 
dae) has been the only other bird flea re- 
ported from the states we investigated 
(Benton 1980). Due to the proximity of 
Georgia, North Carolina, and South Caro- 
lina to Appalachian records of C. idius in 
Tennessee, we suspect that this flea is also 
present in the mountains of these states. 
Further inspection of appropriate hosts, 
along with their respective nests and bur- 
rows, likely will produce new distribution 
records for species of Ceratophyllus in this 
region. Knowledge concerning the distri- 
butions of flea species will allow for vigi- 
lant monitoring of fleas as pests and pos- 
sible vectors of zoonotic agents. Adult fleas 
were identified using Benton (1983). 
Voucher specimens of C. celsus have been 
deposited in the Clemson University Ar- 
thropod Collection and the National Mu- 


seum of Natural History, Smithsonian In- 
stitution, and C. gallinae and C. vison in 
the Great Smoky Mountains National Park 
Museum (GSMNP),. 


Ceratophyllus (Ceratophyllus) celsus 
Jordan, 1926 


This species is an ectoparasite of cliff 
swallows, Petrochelidon pyrrhonota (Viel- 
lot), is found throughout much of North 
America. Except for a Virginia record (Eck- 
erlin et al. 2003) there have been no other 
confirmed records from the southeastern 
USA of this species (i.e., states east of the 
Mississippi and south of Indiana through 
Delaware). In the eastern United States, C. 
celsus has been recorded from Michigan, 
New York, and Vermont (Osgood 1964, 
Benton 1980). In eastern Canada, C. celsus 
has been recorded from New Brunswick, 
Ontario, and Québec (Holland 1985). 
Whether C. celsus is a vector or reservoir 
of zoonotic agents of disease remains un- 
known; however, it is a suspected vector of 
an avian-associated Trypanosoma sp. (Ho- 
pla and Loye 1983). 

New records.—20 @, 13 6, 3 pupae, 
west face of bridge over Seneca River, 
(34.6535°N, 82.8518°W), Oconee Co., SC, 
coll. Will K. Reeves (WKR), James A. Ko- 
recki (JAK), and Mark P. Nelder (MPN), 
12.VII.2004, ex: abandoned nest of P. pyr- 
rhonota; 15 2, 10 6, bridge going over 
Lake Keowee (34.8115°N, 82.9225°W), 
Pickens Co., SC, coll. JAK, 12.VII.2004, 
ex: abandoned nest of P. pyrrhonota; 3 pu- 
pae, 1-285 bridge over Chattahoochee Riv- 
er, (33.9016°N, 84.4420°W), Cobb Co., GA, 
WKR, 6.VIII.2004, abandoned 
nest of P. pyrrhonota. 


coll. ex: 


472 


Ceratophyllus (Ceratophyllus) gallinae 
(Schrank, 1803) 


Known as the European chicken flea, C. 
gallinae is a Holarctic ectoparasite of birds 
(mostly passerines) and mammals (mostly 
rodents). Ceratophyllus gallinae likely was 
introduced into eastern North America from 
Europe on domestic poultry (Lewis and 
Galloway 2001). In the eastern United 
States, C. gallinae has been recorded from 
Connecticut, Delaware, Maine, Massachu- 
setts, Michigan, New Hampshire, New Jer- 
sey, New York, Vermont, and West Virginia 
(Fox 1940, Osgood 1964, Shaw and Hovey 
1954, Benton 1980, Eckerlin and Painter 
2000). In eastern Canada, C. gallinae has 
been reported from New Brunswick, New- 
foundland, Nova Scotia, Ontario, Prince 
Edward Island, and Québec (Holland 
1985). Ceratophyllus gallinae is a biting 
pest of domestic poultry in Europe and the 
northeastern USA (Fox 1940, Shaw and 
Hovey 1954) and also bites and infests peo- 
ple working in chicken coops (Haas and 
Wilson 1973). 

New records.—2 2, 1 6, Purchase Creek 
(GSMNP), (35.5851°N, 83.0626°W), Hay- 
wood Co., NC, coll. WKR, 23.VII.2002, 
ex: abandoned blue bird nest inhabited by 
deer mice, Peromyscus maniculatus Walker, 
accession no. L-3002. 


Ceratophyllus (Monopsyllus) vison 
Baker, 1904 


This species is considered a specific ec- 
toparasite of the red squirrel, 7. hudsonicus, 
but has been reported from other rodents 
and mustelid carnivores (Fox 1940, Durden 
and Kollars 1997). In the eastern United 
States, C. vison has been recorded from 
Maine, Massachusetts, Michigan, New 
Hampshire, New York, Tennessee, West 
Virginia, and Vermont (Fox 1940, Osgood 
1964, Benton 1980). In eastern Canada, C. 
vison has been reported from Labrador 
(Newfoundland), New Brunswick, Nova 
Scotia, Ontario, and Québec, (e.g., Traub et 
al. 1983, Holland 1985). The potential of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


C. vison as a vector of zoonotic agents 1s 
unknown. 

New records.—l1 2, Clingman’s Dome 
(GSMNP), (35.5588°N, 83.4949°W), Sevier 
Co., TN, coll. E. Pivorun, 5.V1I.2004, ex. T. 
hudsonicus, accession no. L-2956; 3 2, 1 
36, Mt. LeConte (GSMNP), (35.6549°N, 
83.4408°W), Sevier Co., TN, coll. K. Wal- 
ters, 14.VI.2004, ex. 7. hudsonicus, acces- 
sion no. L-3048; 1 @, Oconaluftee 
(GSMNP), (35.5171°N, 83.3068°W), Swain 
Co., NC, coll. WKR, 5.VI.2004, ex. Tamias 
striatus (Linnaeus), accession no. L-2932. 

We thank Robert E. Lewis (Ames, IA) 
for confirming our identifications of C. cel- 
sus and C. gallinae and Peter H. Adler and 
Bill Wills (Clemson University) for review- 
ing an earlier version of the manuscript. 
This is technical contribution number 5049 
of the South Carolina Agriculture and For- 
estry Research System, Clemson Universi- 


ty. 
LITERATURE CITED 


Benton, A. H. 1980. An Atlas of the Fleas of the East- 
ern United States. Marginal Media, Fredonia, New 
York, 177 pp. 

. 1983. An illustrated key to the fleas of the 
eastern United States. Bioguide No. 3, Marginal 
Media, Fredonia, New York, 34 pp. 

Durden, L. A. and T. M. Kollars, Jr. 1997. The fleas 
(Siphonaptera) of Tennessee. Journal of Vector 
Ecology 22: 13-22. 

Durden, L. A., W. Wills, and K. L. Clark. 1999. The 
fleas (Siphonaptera) of South Carolina with an as- 
sessment of their vectorial importance. Journal of 
Vector Ecology 24: 171-181. 

Eckerlin, R. P. and H. E Painter. 2000. New records of 
fleas (Siphonaptera) from eastern West Virginia. 
Proceedings of the Entomological Society of 
Washington 102: 969-973. 

Eckerlin, R. P, H. E Painter, and R. B. Clapp. 2003. 
New flea and cimicid records from birds in Vir- 
ginia. Banisteria 22: 53-56. 

Fox, I. 1940. Fleas of the eastern United States. Iowa 
State College Press, Ames, Iowa, 191 pp. 

Haas, G. E. and N. Wilson. 1973. Siphonaptera of Wis- 
consin. Proceedings of the Entomological Society 
of Washington 73: 302-314. 

Hopla, C. E. and J. E. Loye. 1983. The ectoparasites 
and microorganisms associated with cliff swal- 
lows in west-central Oklahoma. I. Ticks and fleas. 
Bulletin of the Society of Vector Ecology 8: 111— 
121. 


VOLUME 107, NUMBER 2 


Holland, G. P. 1985. The fleas of Canada, Alaska, and 
Greenland (Siphonaptera). Memoirs of the Ento- 
mological Society of Canada 130: 1-631. 

Lewis, R. E. and T. D. Galloway. 2001. A taxonomic 
review of the Ceratophyllus Curtis, 1832 of North 
America (Siphonaptera: Ceratophyllidae: Cerato- 
phyllinae). Journal of Vector Ecology 26: 119— 
161. 

Osgood, FE L. 1964. Fleas of Vermont. Journal of the 
New York Entomological Society 72: 29-33. 
Shaw, E R. and C. L. Hovey. 1954. An infestation of 
the European hen flea in Maine. Journal of Eco- 

nomic Entomology 47: 942-943. 

Traub, R., M. Rothschild, and J. EF Haddow. 1983. The 
Rothschild collection of fleas: the Ceratophylli- 
dae. University Press, Cambridge, 288 pp. 


473 


Mark P. Nelder, Will K. Reeves, James 
A. Korecki, and Lance A. Durden. (MPN, 
JAK) Department of Entomology, Soils, and 
Plant Sciences, Clemson University, 114 
Long Hall, Box 340315, Clemson, SC 
29634, U.S.A.; (WKR) Centers for Disease 
Control and Prevention, Viral and Rickett- 
sial Zoonoses Branch, Mailstop G-13, 1600 
Clifton Rd. NE, Atlanta, GA 30333, U.S.A.; 
(LAD) Department of Biology, Georgia 
Southern University, P.O. Box 8042, States- 
boro, GA 30460-8042, U.S.A. 
mnelder@clemson.edu) 


(e-mail: 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 474-475 


NOTE 


Validation of Potamocloeon macafertiorum Lugo-Ortiz (Ephemeroptera: Baetidae) 


The genus. Potamocloeon Gillies (1990) 
(Ephemeroptera: Baetidae) was described 
for the species Cloeon dentatum Kimmins, 
which had been based on adults from Ugan- 
da (Kimmins 1956). Gillies (1988) associ- 
ated distinctive larvae with adults from 
Guinea that he attributed to C. dentatum. A 
second species, P. macafertiorum Lugo-Or- 
tiz, was described based on a larva from 
South Africa (Lugo-Ortiz and McCafferty 
1996). Gattolliat (2003) described two un- 
named Potamocloeon species from Mada- 
gascar—one from a larva and the other 
from a male adult. 

Gattolliat (2003) considered P. macafer- 
tiorum to be synonymous with P. dentatum. 
Gattolliat’s hypothesis of the equivalency of 
P. macafertiorum and P. dentatum was not 
based on variability that he observed in 
specimens of the two species in question. 
Rather, he cited variability that had been 
documented for species in some other gen- 
era. In particular, he noted the sexually di- 
morphic presence or absence of hindwings 
in Cheleocloeon Wuillot and Gillies and the 
variation in the length of maxillary palp 
segment 2 of certain Cloeodes Traver spe- 
cies. 

We examined the specimens on which 
Gillies (1988) based his larval description 
of P. dentatum. These specimens match the 
illustrations and descriptions he provided 
(Gillies 1988: figs. 14-30). Comparison of 
this associated larval material to the type 
specimen of P. macafertiorum indicates 
that the two species are distinctive. Addi- 
tional material of P. macafertiorum con- 
firmed this observation. The presence or ab- 
sence of hindwingpads (regardless of gen- 
der) and the shape and relative length of 
maxillary palp segments appear to be con- 
stant within individual Potamocloeon spe- 
cies. 


Based on comparative study, we must re- 
instate P. macafertiorum as a valid species. 
Potamocloeon dentatum larvae have no 
hindwingpads, segment 2 of the maxillary 
palp 0.55x the length of segment 1, and 
little to no speckling or coloration on terga 
or sterna. In contrast, P. macafertiorum l\ar- 
vae have conspicuous hindwingpads, seg- 
ment 2 of the maxillarly palp 0.85 the 
length of segment 1, some median and sub- 
median speckling on the terga, and lateral 
longitudinal dashes on the sterna. 

Material examined.—P. dentatum: GAM- 
BIA: R. Gambia, Fatoko, 19-II-1993, 
““Y 127-8,” one larva [The Natural History 
Museum, London, England (BMNH)]. 
GUINEA: R. Niandam, Sassambaya, on 
sand, 13-II-1986, ‘“‘X265,’’ one larva 
[BMNH]. IVORY COAST: R. Bouafleé, 
Maraoié (spelling?), VI-1976, JM Elouard, 
‘“W625-6,” three larvae [BMNH]. P. ma- 
cafertiorum: SOUTH AFRICA: Mooi R., 
above Rosetta, 29°18'10”S, 29°57'50”E, 
stones out of current, IIJ-1995, C Dickens, 
““MOI27AE,” seven larvae [Albany Muse- 
um, Grahamstown, South Africa (AMGS)]; 
Mooi R., Hornet Corner, 28°56'45’S, 
30°22'33”E, stones in current, III-1995, C 
Dickens, ““MOI22AP, MOI23AD,”’ nine lar- 
vae [AMGS]; Mpumalanga Prov., Kruger 
NP, Sabie R., NE Corner of Old Rhino 
Camp, 24-X-1990, WP&N McCafferty, one 
larva (P. macafertiorum holotype; mouth- 
parts, right foreleg, paraproct on slide SA37) 
[Purdue University Entomological Research 
Collection, West Lafayette, Indiana, USA 
(ER) 

Acknowledgments.—Helen Barber- 
James (Grahamstown, South Africa) and 
David Goodger (London, England) loaned 
some of the material examined. 


LITERATURE CITED 


Gattolliat, J.-L. 2003. The genera Demoulina Gillies 
and Potamocloeon Gillies (Ephemeroptera: Bae- 
tidae) in Madagascar. Zootaxa 184: 1-18. 


VOLUME 107, NUMBER 2 


Gillies, M. T. 1988. Descriptions of the nymphs of 
some Afrotropical Baetidae (Ephemeroptera), I. 
Cloeon Leach and Rhithrocloeon Gillies. Aquatic 
Insects 10: 49-59. 

. 1990. A new genus for the Afrotropical may- 
fly, Cloeon dentatum Kimmins (Ephem., Baeti- 
dae). Entomologist’s Monthly Magazine 126: 
207-208. 

Kimmins, D. E. 1956. New species of Ephemeroptera 
from Uganda. Bulletin of the British Museum 
(Natural History) Entomology 4: 71-87. 


475 


Lugo-Ortiz, C. R. and W. P. McCafferty. 1996. The 
Bugilliesia complex of African Baetidae (Ephem- 
eroptera). Transactions of the American Entomo- 
logical Society 122: 175-197. 


Luke M. Jacobus and W. P. McCafferty, 
Department of Entomology, Purdue Uni- 
versity, West Lafayette, IN 47907, U.S.A. 
(e-mail: luke-jacobus @ entm.purdue.edu) 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 476-477 


NOTE 


New State Record of Culex coronator Dyar and Knab (Diptera: Culicidae) 
From Mississippi 


Culex (Culex) coronator Dyar and Knab, 
is one of six species in the Culex coronator 
complex as defined by Bram (1967). This 
complex is very widely distributed between 
Argentina and the southern United States, 
but is poorly understood and differentiated 
primarily by male genitalia characters. 
Bram (1967) determined that Cx. coronator 
is the only member of the complex to occur 
in the United States. The currently recog- 
nized distribution for Cx. coronator in the 
United States is listed as Arizona, New 
Mexico and Texas (Darsie and Ward 1981). 
This paper reports the first collections of 
Cx. coronator from Mississippi, which rep- 
resents a considerable eastward and north- 
ern extension of the previous distribution. 

New collection records.—Culex corona- 
tor Dyar and Knab: Copiah Co., Mississip- 
pi, Copiah County Wildlife Management 
Area, 22 September 2004, CDC light trap 
baited with CO,, Collector: Jerome God- 
dard, 1 2; Copiah Co. Wildlife Manage- 
ment Area, 1 October 2004, CDC light trap 
baited with CO,, Collector: Wendy Varna- 
do, 1 2; Copiah Co., Mississippi, 7 Decem- 
ber 2004, CDC light trap baited with CO,, 
Collector: Wendy Varnado, 2 2. 

All specimens were collected as a result 
of a statewide survey of mosquitoes of Mis- 
sissippi funded by a West Nile virus grant 
from the Centers for Disease Control and 
Prevention. Collections were made by 
hanging CDC light traps baited with dry ice 
(CO,) in late afternoon in selected sites and 
retrieving the traps first thing the next day. 
Mosquitoes were killed by freezing, glued 
to pinned paper points, labeled with collec- 
tion information, and identified using Car- 
penter and LaCasse (1955), Darsie and 
Ward (1981), and Clark-Gil and Darsie 
(1983). The September—October specimens 


of Cx. coronator were collected at Copiah 
County Wildlife Management Area 
(CCWMA), an undisturbed area in central 
Mississippi containing numerous semi-per- 
manent pools fed by springs as well as nu- 
merous rain-filled woodland pools. The 
CCWMA is composed of open fields, vast 
pine forests, and creek bottoms containing 
mature hardwoods (predominantly oak and 
hickory). The December specimens of Cx. 
coronator were collected in a wooded area 
approximately six km east of the CCWMA 
and within thirty meters of a paved county 
road. 

After collecting Cx. coronator, extensive 
efforts were made to find the breeding 
site(s). Roadside ditches, swales, artificial 
containers, and woodland pools were sam- 
pled in a three square mile area by dipping 
and looking for larvae. All specimen iden- 
tifications, adults and larvae, were con- 
firmed by the third author. Specimens have 
been deposited in the Mississippi Entomo- 
logical Museum, Mississippi State Univer- 
sity, Starkville, MS, the Public Health Pest 
Management mosquito collection, Winston- 
Salem, NC, and the Smithsonian Institution 
mosquito collection, Washington, DC. 

The presence of Culex coronator in Mis- 
sissippi was unexpected in light of the gen- 
erally tropical nature of this species. Con- 
firmation that these collections represent an 
established breeding population of Cx. co- 
ronator was confirmed by finding hundreds 
of larvae in three widely separated areas 
near the original adult collection site. Lar- 
vae were found in a small spring in the 
woods, as well as in two separate roadside 
ditches/swales. 

The initial U.S. collections of Cx. coron- 
ator occurred in Texas (Randolph and 
O’Neill 1944, Rueger and Druce 1950, 


VOLUME 107, NUMBER 2 


Eads et al. 1951). These Texas records were 
extended by Hill et al. (1958), who record- 
ed Cx. coronator from Bexar, Bowie, Cam- 
eron, Dimmit, Hidalgo, Liveoak, Milam, 
Starr, Webb, and Willacy counties. In 1953, 
Richards et al. (1956) found this species in 
Cochise and Pima counties, Arizona, and 
Wolff et al. (1975) found it in two sites in 
Dona Ana County, New Mexico. However, 
there is another old record that should now 
be considered. Hill et al. (1958) reported 
one male and two females of this species 
collected in 1953 in Vernon Parish (Ft. 
Polk), Louisiana. This record was not in- 
cluded in the most recently published dis- 
tributions for this species (Darsie and Ward 
1981) because Carpenter (1970) suggested 
the Louisiana record should be deleted until 
it could be confirmed. Although finding Cx. 
coronator in Mississippi does not validate 
the Hill et al. (1958) record for Louisiana, 
it certainly suggests that those identifica- 
tions of Cx. coronator from Louisiana were 
correct. 


LITERATURE CITED 


Bram, R. A. 1967. Classification of Culex subgenus 
Culex in the New World (Diptera: Culicidae). Pro- 
ceedings of the Entomological Society of Wash- 
ington 120: 1—222. 

Carpenter, S. J. 1970. Review of recent literature on 
mosquitoes of North America. Supplement I. Cal- 
ifornia Vector Views 17: 39-65. 

Carpenter, S. J. and W. J. LaCasse. 1955. Mosquitoes 
of North America (North of Mexico). University 


477 


of California Press, Berkeley and Los Angeles, 
CA. 

Clark-Gil, S. and R. FE Darsie, Jr. 1983. The mosqui- 
toes of Guatemala. Mosquito Systematics 15: 
151-284. 

Darsie, R. F, Jr. and R. A. Ward. 1981. Identification 
and geographical distribution of the mosquitoes of 
North America, north of Mexico. Mosquito Sys- 
tematics Supplement |: 1—313. 

Eads, R. B., G. C. Menzies, and L. J. Ogden. 1951. 
Distribution records of west Texas mosquitoes. 
Mosquito News 11: 41—47. 

Hill, S. O., B. J. Smittle, and E M. Philips. 1958. Dis- 
tribution of mosquitoes in the fourth U.S. Army 
area. Entomology Division, Fourth U.S. Army 
Medical Laboratory, Fort Sam Houston, TX, 155 
PPp- 

Randolph, N. M. and K. O’Neill. 1944. The mosqui- 
toes of Texas. Bulletin of the Texas State Health 
Department, 100 pp. 

Richards, C. S., L. T. Nielsen, and D. M. Rees. 1956. 
Mosquito records from the great basin and drain- 
age of the lower Colorado River. Mosquito News 
16: 10-17. 

Rueger, M. E. and S. Druce. 1950. New mosquito dis- 
tribution records from Texas. Mosquito News 10: 
60-63. 

Wolff, T. A., L. T. Nielsen, and S. O. Hayes. 1975. A 
current list and bibliography of the mosquitoes of 
New Mexico. Mosquito Systematics 7: 13-17. 


Wendy C. Varnado, Jerome Goddard, and 
Bruce A. Harrison. (WCV, JG) Mississippi 
Department of Health, P.O. Box 1700, 
Jackson, MS 39215, U.S.A. (e-mail: 
Jgoddard@ msdh.state.ms.us); (BAH) North 
Carolina Department of Environment and 
Natural Resources, 585 Waughton Street, 
Winston-Salem, NC 27107, U.S.A. (e-mail: 
bruce.harrison@ncmail.net) 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, p. 478-479 


REPORTS OF OFFICERS 


EDITOR 


Volume 106 of the Proceedings included 
four issues with a total of 948 pages. Nine- 
ty-four regular papers, 16 notes, 3 book re- 
views, | obituary, minutes of Society meet- 
ings, reports of officers, instructions for au- 
thors, and the table of contents for volume 
106 were published. About 117 manuscripts 
(regular papers and notes) were submitted 
for consideration for publication from No- 
vember 1, 2003 to October 31, 2004. 

Memoir Number 25, ““A Catalog of the 
Cecidomylidae (Diptera) of the World” by 
Raymond J. Gagné, was published in Feb- 
ruary 2004. This 408 page catalog is avail- 
able from the Society for $50.00. 


I extend thanks to Ray Gagné, book re- 
view editor, for his excellent work in ob- 
taining book reviews for the Proceedings, 
and to Ray, Tom Henry, and Wayne Mathis 
of the Publications Committee for their en- 
couragement and support. I am especially 
grateful to the many reviewers for their 
time-consuming efforts and constructive re- 
views of manuscripts. Their contributions 
are essential to help increase the quality of 
papers published in the Proceedings. 


Respectfully submitted, 
David R. Smith, Editor 


‘TREASURER 


SUMMARY FINANCIAL STATEMENT FOR 2004 


Special 
General Publication Total 
Fund Fund Assets 
Assets: November 1, 2003 $53,567.44 $136,748.31 $190,315.75 
Total Receipts for 2004 $75,197.37 $6,330.56 $81,527.93 
Total Disbursements for 2004 $82.960.00 $22,832.87 $105,792.87 
Assets: October 31, 2004 $45,804.81 $120,246.00 $166,050.81 
Net Changes in Funds $7,762.63 ($16,502.31) ($24,264.94) 


Audited by the Auditing Committee, November 30, 2004 consisting of Michael W. Gates, 
Chairman, John W. Brown, and Allen L. Norrbom. Presented to the membership at the 


meeting of December 2, 2004. 


Respectfully submitted, 
Michael G. Pogue, Treasurer 


VOLUME 107, NUMBER 2 479 


MEMBERSHIP SECRETARY 


In 2004, the Society received applications for membership from 13 people: 


Stephen M. Bahr II Robert R. Kula 
Julieta Brambila Don A. Russo 
Onder Calmasur Steven J. Seybold 
Sylvio G. Codella, Jr. Donald C. Weber 
Jaime Estes Lijie Zhang 
Jeffrey Y. Honda Yalin Zhang 


Akito Y. Kawahara 


Each applicant was sent a letter of welcome to the Society and his/her name was read 
at a regular monthly meeting. The number of applications decreased 43% from 2003. 
Other letters from the Membership Secretary included six letters to guest speakers, 13 
letters to special fund contributors, and one reply to a request for information. 


Respectfully submitted, 
Holly B. William, 
Membership Secretary 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, p. 480-482 


SOCIETY MEETINGS 


1,084th Regular Meeting—July 8, 2004 


The 1,084th meeting of the Entomologi- 
cal Society of Washington consisted of the 
Annual Banquet at the Uniformed Services 
University of Health Sciences in Bethesda, 
Maryland. A cash bar opened at 6:00 p.m. 
and the formal banquet followed at 7:00 
p.m. Approximately 90 members and guests 
were in attendance. At 8:00 p.m. the speak- 
er, Keith Wilmott of The Natural History 
Museum, London, presented his talk, “A 
Day in the Life of a Tropical Butterfly.” 
This consisted of a quick overview of the 
diversity of form and life history among 
Ecuadorian butterflies, all magnificently il- 
lustrated with color photos of them in their 
habitat. The meeting adjourned at about 
9:30 p.m. 


Respectfully submitted, 
Stuart H. McKamey 
Recording Secretary 


1,085th Regular Meeting— 
October 7, 2004 


The 1,085th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby 
room of the National Museum of Natural 
History, Washington, D.C., by President 
Eric Grissell, at 7:10 p.m. The meeting was 
attended by 16 members and 7 guests. The 
minutes of the 1,083rd and 1,084th meet- 
ings were approved as read. 

There were no new applicants for mem- 
bership. One new member and 4 visitors 
were introduced. 

In miscellaneous business, President 
Grissell mentioned the need for a nominat- 
ing committee for next year’s officers— 
there were no takers. Thomas Henry later 
accepted the position. 

For exhibits, Dave Furth had eight new 
books, which dealt with the diversity, tax- 
onomy, and biogeography of Mexican ar- 
thropods, catalogues of Bulgarian bupres- 


tids and world epilachnine lady bird beetles, 
a revision of the silkmoth genus Samia, an 
illustrated catalogue of the carabid beetle 
genus Carabus, diversity and biology of 
spiders and other arachnids, and two coffee- 
table books, “Insects Revealed: Monsters 
or Marvels?” by J. Tonnancour, and “For 
Love of Insects” by T. Eisner. 

After being regaled with the adventures 
of dead dragonflies while the speaker cir- 
cumnavigated around endless roadblocks, 
the audience was finally treated to the pre- 
sentation ““A Different Olympiad: Moun- 
tain Hopping in Northern Greece From Mt. 
Gramos & Varnous Across to Mt. Rhodo- 
pi’ by Fred Paras of the Baltimore City 
Community College and perennial presi- 
dent of the Maryland Entomological Soci- 
ety. The slide show consisted of a travel- 
ogue rich with habitat, landscape, geogra- 
phy, plants, butterflies, and even the history 
and culture of Greece. From the rocky, dry 
habitats in the south to the lush forests of 
the north near Bulgaria and the overgrazed, 
battle scarred fields bordering Albania, ly- 
caenids dominate the butterfly fauna, fol- 
lowed by satyrids and nymphalids. The pro- 
tected species Parnassius apollo was found 
to be locally very abundant near Bulgaria 
at a location not reported in these minutes. 
Colorful neuropterans—diurnal owlflies 
and nemopterids—were encountered in 
Florina, and were included in the specimen 
drawers exhibited at the end of the meeting, 
at 9:15. 

Refreshments were provided by the So- 
ciety. 


Respectfully submitted, 
Stuart H. McKamey 
Recording Secretary 


1,086th Regular Meeting— 
November 4, 2004 


The 1,086th regular meeting of the En- 
tomological Society of Washington (ESW) 


VOLUME 107, NUMBER 2 


was called to order in the Cathy Kerby 
room of the National Museum of Natural 
History, Washington, D.C., by President 
Eric Grissell, at 7:05 p.m. The meeting was 
attended by 18 members and 10 guests. The 
minutes of the 1,085th meeting were ap- 
proved with modification. 

There were 6 new applicants for mem- 
bership from August through November: 
Julieta Brambila, Stephan Bahr, Akito Ka- 
wahara, Jaime Estes, Steven Seybold, and 
Jeffrey Honda. No new members were pres- 
ent. One visitor was introduced. 

In miscellaneous business, President 
Grissell announced the Nominating Com- 
mittee and encouraged nominations for all 
positions. A breath of relief swept the room 
as Grissell noted that nominations already 
had been accepted for all positions. Voting 
takes place in December. 

For exhibits, Edd Barrows mentioned 
that Virginia Trail Guides are still available 
and had a copy for perusal. Warren Steiner 
had a new publication about a tenebrionid 
associated with Affa ant garbage dumps, 
and a vial with two recently collected “‘fish 
lice,” which are huge, nasty-looking iso- 
pods. Dave Furth had three new books and 
two other items: “‘Proceedings of the XI In- 
ternational Symposium on Insect-Plant Re- 
lationships” by J. K. Nielson, C. Kjaer, and 
L. M. Schoonhoven (eds.); ““A Guide to the 
Winged Aphids of Costa Rica’ by D. Voeg- 
tlin and C. Rivera (eds.); ““Lucanidae of 
Thailand” by A. Pinratana and J.-M. Maes; 
an insect specimen catalogue from Insects 
International (formerly Combined Scientific 
Supply); and a poster on Vanishing Polli- 
nators, noting that they were still available. 

Dave Furth introduced the evening’s 
speaker, Dr. John S. LaPolla, presently a 
post-doctoral fellow at the Smithsonian In- 
stitution. His presentation, entitled “The 
Rise of Ranching in the Ants & the Evo- 
lution of Trophophoresy,’”’ consisted of a 
summary of his dissertation research. Two 
well known habits of ants include fungi- 
culture (plant-based farming) and tropho- 
biosis (feeding on honey-dew exudates of 


48] 


scales, aphids, treehoppers, lycaenids, or 
mealybugs). Dr. LaPolla then focused on 
obligate trophobiosis and coined the word 
trophophoresy—take out food, insect-style. 
This is epitomized by his study organism, 
the ant genus Acropyga, tiny formicines 
that are entirely subterranean except for 
mating swarms and that farm a specific 
tribe of mealybugs. The close relationship 
among Acropyga and mealybugs was high- 
lighted with a picture of a queen ant car- 
rying a mealybug—both preserved in 10— 
20 million year old Miocene amber from 
Hispaniola. Although the mostly pantropi- 
cal genus has only 37 described species, Dr. 
LaPolla recently collected three new Acro- 
Pyga species (and two new mealybug spe- 
cies) in one square meter of soil in Guyana, 
so there are undoubtedly many, many more 
remaining to be described. Dr. LaPolla 
found that in Acropyga, male morphology 
was a good source for phylogenetic fea- 
tures, which is unusual among ants. 

The meeting was adjourned at 8:10. Re- 
freshments were provided by the Society. 


Respectfully submitted, 
Stuart H. McKamey 
Recording Secretary 


1,087th Regular Meeting— 
December 2, 2004 


The 1,087th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby 
room of the National Museum of Natural 
History, Washington, D.C., by President 
Eric Grissell, at 7:04 p.m. The meeting was 
attended by 17 members and 9 guests. The 
minutes of the 1,086th meeting were ap- 
proved as read. 

There were no new applicants 
bership or new members present. Two vis- 


for mem- 


itors were introduced. 

In miscellaneous business, reports from 
officers were given. The slate of officer 
candidates for 2005 was approved unani- 
mously. 

For exhibits, Terry Nuhn discussed his 


482 


work with microhymenopterans and 
brought literature and synoptic drawers, in- 
cluding the recently described New Zealand 
family Maamingidae. Dave Furth had one 
new book for display: ““Artrépodos de Cha- 
mela” by A. N. Garcia Aldrete and R. Ay- 
ala Barajas (eds.). 

Dave Furth introduced Dr. Sean Brady, a 
postdoctoral fellow of the Smithsonian In- 
stitution, who gave a presentation on “Un- 
covering the Evolutionary History of 
Ants.’ This consisted of a phylogenetic as- 
sessment of two case studies: “dinosaur 
ants,’ supposedly the most primitive, and 
army ants, whose similarities across three 
subfamilies and two continents is generally 
thought to be convergence. Dinosaur ants 
lack foraging cooperation or trail phero- 
mones, and queens revert to foraging if 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


needed. Army ants all have obligate group 
foraging, nomadism, and wingless queens 
alternating between a sturdy running form 
and an immobile egg-factory. Using data 
from molecules and morphology, Dr. Brady 
debunked both dogmas. The “‘dinosaur 
ants’’ seem closely related to 40—45 million 
year old fossils from Argentina so their 
Australian distribution appears to be relic- 
tual, and the army ants formed a monophy- 
letic group, probably attaining their distri- 
butions with the break-up of South America 
and Africa about 100 million years ago. 
The meeting was adjourned at 8:30. Re- 
freshments were provided by the Society. 


Respectfully submitted, 
Stuart H. McKamey 
Recording Secretary 


PROC. ENTOMOL. SOC. WASH. 
107(2), 2005, pp. 483-484 


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A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E. 
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Grissell and Michael E. Schauff. 87 pp. 1997 


Revision of the Oriental Species of Aphthona Chevrolat (Coleoptera: Chrysomelidae), by Alexander S. 
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Revision of the Genus Anoplophora (Coleoptera: Cerambycidae), by Steven W. Lingafelter and E. 
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No.5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 __. 
No.6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi 


Takahasi. 230 pp. 1969 


No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato- 
pogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 


No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 
S773] NAS Toe RE SRE hk RE ee NE RE aR CO EN OEE. E,Mae Ace ame aL 


No. 14. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174 
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SMITHSONIAN NSTITUTION LIBRARIES 


wu QU 


3 9088 01140 9398 
CONTENTS 


(Continued from front cover) 

ROBBINS, ROBERT K. and MARCELO DUARTE—Phylogenetic analysis of Cyanophrys 
Clench, a synopsis of its species, and the potentially threatened C. bertha (Jones) 
(Lycaenidae: Theclinae? Himmaeuann) (sac iie eign as oleae aerate” elec sq ial elola ee irbale sles thn) ate ne etait ca etal 

ROBINSON, HAROLD AND NORMAN E. WOODLEY—A new species of Harmstonia 
(Diptera: Dolichopodidae) fromyB olivia te is 2 M20 este eels iourciatate’s nip sie sixieletsirielghi inlet a) eke once aie 


SAINI, MALKIAT S. and DAVID R. SMITH—Revision of the southeastern Asian sawfly genus 
Busarbia. Cameron (i ymenoptera: slenthmedimid ac) yeni. sh leey. sia pialeyn areal leeenelaye are reer 


SCHAWAROCH, VALERIE, DAVID GRIMALDI, and ANGELA V. KLAUS—Focusing on mor- 
phology: Applications and implications of confocal laser scanning microscopy (Diptera: 
€ampichoetidae; CamillidaeDrosophilidae). Jcgae els ss. Vere eee> sntiee a> ghee ne ae eerie aie 


SHENG, MAO-LING—The genus Jschnoceros Grayenhorst (Hymenoptera: Ichneumonidae) 
TOIVA @) orien Ee Re ROR Be ae SE BAL et on iE Dina Ron Sale Ae AG RUDDERS ComcetGs aE Guu mac. Adomda dcr 


SMITH, DAVID R.—Two new fern-feeding sawflies of the genus Aneugmenus Hartig 
(Hymenoptera: Tenthredinidac) from South America, i.) oc mg. eres +n ieee eee 


STEINER, WARREN E., JR.—Studies on the darkling beetles (Coleoptera: Tenebrionidae) 
known from Grand Bahama Island, with descriptions of new species of Branchus 
PVG LO Ke (ed Roe Neat Ook ae aR t! “Ac ) vais Men A eat Png ee ied Codi a iat ee ae ae UNI BS LL eae Siciy ob 


TOGASHI, ICHIJI—A new species of the Eriocampa ovata group (Hymenoptera: 
Tenthredinidae) feeding on Alnus matsumurae Call. (Dicotyledoneae: Fagaceae) from Japan 


TOGASHI, ICHIJI—Description of a new species of the genus Apethymus Benson (Hymenoptera: 
Tenthredinidae) feeding on Quercus acutissima Carruthers (Fagaceae) in Japan.............. 


WHEELER, A. G., JR.—Blissus minutus (Blatchley) and Toonglasa umbrata (Distant): Seldom- 
collected native chinch bugs (Hemiptera: Lygaeoidea: Blissidae) as colonists of the African 
bunchegrass Eragrostis curvula, and their association with other grasses in the southern 
Wife GyS tates eres eh ee Sate UA Nc AE Ns SS EMRRR AR Peach oa is a aT nce ens Saco Maree 


YOUNG, DANIEL K.—Taxonomic notes on South American Pogonoceromorphus Pic 
(Coleoptera), including transfer from Pyrochroidae (Pyrochroinae) to Anthicidae 
(Bina fore OVUNOVEIS) here der rs OVUM ye CUBA AH aaa Nh iad ASS raD eC SR ASW S 2 bbe ocagi coor mds 


ZHANG, YALIN and MIN HUANG—Two new leafhopper genera, Direnaia and Xaniona 
(Hemiptera: Cicadellidae: Typhlocybinae: Typhlocybini) from China ........................ 
NOTES 


BITO, DARREN and DAVID R. SMITH—Larva and possible food plant of Ancyloneura varipes 
(Cameron) (Hymenoptera: Pergidae) in Papua New Guinea ......................-...--2-2--- 


FAVRET, COLIN—A new non-destructive DNA extraction and specimen clearing technique for 
aphids) @Hemipteral) 27x te eae ccpua ciety tits oats aecelcihe hata c Si Se Ren Sta thay aol eas ate me eae 


JACOBUS, LUKE M. and W. P. McCAFFERTY—vValidation of Potamocloeon macafertiorum 
bugo: Ortiz, (Ephemeropteram Bacttdae) ie ice! 5 toe to ese anya ieee sie tictec* oie metre aes a cee reeeare eye 


LATTIN, JOHN D.—Physopleurella floridana Blatchley, 1925, a synonym of Physopleurella 
mundula (White, 1877) (Hemiptera: Heteroptera: Cimicoidea: Anthocoridae) ................ 


LATTIN, JOHN D.—Dufouriellus ater (Puton), Macrotrachelia nigronitens (Stal), and Xyvlocoris 
- (Arrostelus) flavipes (Reuter) (Hemiptera: Heteroptera: Cimicoidea: Anthocoridae): First 
records mom: the; Haw alae Slam Si se) Saes cere ee ee eee yeas eee ae ene ae nce egies na 


NELDER, MARK P, WILL K. REEVES, JAMES A. KORECKI, and LANCE A. DURDEN— 
Fleas of the genus Ceratophyllus (Siphonaptera: Ceratophyllidae) in the southeastern 
Winited ‘Statesie a yuapio tei eat oa RE aN PN ede Ae gee (5 he pits LUSENe ca Dia UG ARN eFC aC 


VARNADO, WENDY C., JEROME GODDARD, and BRUCE A. HARRISON—New state record 
for Culex coronator Dyar and Knab (Diptera: Culicidae) from Mississippi ................-- 


MISCELLANEOUS 
Reports: off OPACers) ie A IG RU CTS he ace) ee UO td 1 SANS ate cpa ete lee 
SOGIStyA MISS tini gs enh ete la sia Ska PN AR) yl Maoh lear Une ye) RP eat saa aU PUR Pay eee eter fer 


Instructions Tor A UOT i ay eles ee I ire Phe Sa SARS OEM 2 pid SR ee ete pannel ata a RNP ae 


474 


460 


466 


471 


VOL. 107 JULY 2005 NO. 3 
QO i (ISSN 0013-8797) 


U bl CATRSON AAD 


—t PROCEEDINGS 1132 


of the LIBRARIES 


ENTOMOLOGICAL SOCIETY 
of WASHINGTON 


PUBLISHED 
QUARTERLY 


CONTENTS 


BARBA-ALVAREZ, R. and J. BUENO-SORIA—New species of the genus Polycentropus Curtis 
(inchoptera-/Lolycentropodidae) trom, Mexico iy eae tne lon vos sd eivinladeene cliche alee rue aoe 663 


BARROWS, EDWARD M., ANNE M. McINTYRE, and OLIVER S. FLINT, JR.—Alderfly 
(Neuroptera: Sialidae) flight periods, sex ratios, and habitat use in a Virginia freshwater tidal 


MALSHMOW MOLeSL FANGMNEIT/ECOLOMES: (che katie tee ses Put Mode de Old plac ety re toe 693 
BROOKS, SCOTT E. and TERRY A. WHEELER—Ethiromyia, a new genus of Holarctic 
Bolichapadmac (DipterasDolichopodidaeyi as. 22.0. sak vas ce a tidve ad Wrestle see ue eae 489 
CARPENTER, TERRY L.—Notes on the life of Dr. Clara Southmayd Ludlow, Ph.D., 
MedicabeniM@mOlorish (USS) ODA ie Vane sliee es Uy agen ont JAM bia rains, Wao tka a me 657 
ESPINASA, LUIS—A new genus of the subfamily Cubacubaninae (Zygentoma: Nicoletiidae) 
HLOMAY CLACIUZAMVLC SI COR He mie t WRAY. ni. /s sen ene SE No bek Aaa. Serna peste, KB wan Deve Be 510 


GAMEZ-VIRUES, SAGRARIO and ASTRID EBEN—Predatory behavior of Repipta flavicans 
Stal (Hemiptera: Reduviidae), a natural enemy of Diabroticina (Coleoptera: Chrysomelidae) 642 


HARRISON, T. L.—A new species of Douglasiidae (Lepidoptera) from the eastern Nearctic .... 596 


HELLENTHAL, RONALD A. and ROGER D. PRICE—Two new species of Myrsidea Waterston 
(Phthiraptera: Amblycera: Menoponidae) from the jewel-babblers (Passeriformes: Eupetidae) 
HiSAM ChAT LC Tet Hower ste Meal meal meina Ue Ti ek MS Sis te URN eS a Co eth SS, 485 


HELLENTHAL, RONALD A., ROGER D. PRICE, and JASON D. WECKSTEIN—The genus 
Ramphasticola Carriker (Phthiraptera: Amblycera: Menoponidae) from the toucans 
(Piciformes: Ramphastidae), with description of a new species ..............0 0c cece eee eee ees 565 


HESPENHEIDE, HENRY A.—Weevils of the genera Archocopturus Heller and Zygopsella 
Champion: Sibling species and mimetic homoplasy (Coleoptera: Curculionidae: 
BACARRA Pt Acces Niche, AAA Ia head wae ey ale ns 2 IR Sele ea Mes in oP aoe meniedgern yale.n alee s Ube we 671 

HUANG, YIAU-MIN—Cornetius, a new subgenus of Aedes, and a redescription of Aedes 
KGcOetins Kaos ANGOrnen (Ointera:: CUNCldde) s)sccmaict . Saraddre vkr Shar be ew emus oo Mb ek du eemidins 517 


JENNINGS, JOHN T. and DAVID R. SMITH—The taxonomic placement of several New World 
and Oriental gasteruptiid wasps (Hymenoptera: Gasteruptiidae) .....................062-0000- 686 


(Continued on hack cover) 


THE 


~ ENTOMOLOGICAL SOCIETY 
OF WASHINGTON 


OFFICERS FOR 2005 


JASON P. W. HALL, President Jon A. Lewis, Custodian 
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PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 485-488 


TWO NEW SPECIES OF MYRSIDEA WATERSTON 
(PHTHIRAPTERA: AMBLYCERA: MENOPONIDAE) FROM THE 
JEWEL-BABBLERS (PASSERIFORMES: EUPETIDAE) FROM 
NEW GUINEA 


RONALD A. HELLENTHAL AND ROGER D. PRICE 


(RAH) Department of Biological Sciences, P.O. Box 369, University of Notre Dame, 
Notre Dame, IN 46556-0369. U.S.A. (e-mail: ronald.a.hellenthal. 1 @nd.edu); (RDP) 4202 
Stanard Circle, Fort Smith, AR 72903-1906, U.S.A. (e-mail: rpricelice@ aol.com) 


Abstract.—Two new species of Myrsidea are described and illustrated: M. castanon- 
otae from the type host Ptilorrhoa castanonota, the chestnut-backed jewel-babbler, and 
M. leucostictae from the type host Ptilorrhoa leucosticta, the spotted jewel-babbler. These 
represent the first species of this chewing louse genus described from the Eupetidae. 


Key Words: 
tidae 


In a continuing survey of the species of 
the chewing louse genus Myrsidea Water- 
ston from the Passeriformes, we here de- 
scribe and illustrate two new species from 
hosts within the Eupetidae, the jewel-bab- 
blers. A summary of the features of the 
Myrsidea is given in Clay (1966) and Hel- 
lenthal and Price (2003); these will not be 
repeated here but the principal features will 
be incorporated into the species descrip- 
tions. As the Myrsidea of various passerine 
families have been reviewed, it has become 
apparent that the Myrsidea from each host 
family are restricted to that family. Because 
this louse genus contains well over 200 spe- 
cific names (see Price et al. 2003), and 
probably infests all passerines it is an un- 
realistically large task to review the entire 
genus at one time. Thus we continue to fol- 
low the only practical course available, that 
of reviewing the lice by host family. 

In the following descriptions, all mea- 
surements are in millimeters. Abbreviations 
are TW, temple width; HL, head length; 
PW, prothorax width; MW, metathorax 


chewing lice, Myrsidea, Phthiraptera, Menoponidae, jewel-babblers, Eupe- 


width; AWIV, abdomen width at segment 
IV; TL, total length; ANW, female anus 
width; and GL, male genitalia length. The 
host nomenclature below order follows that 
of Dickinson (2003). The holotypes and 
paratypes of both new species are in the K. 
C. Emerson Museum, Oklahoma State Uni- 
versity, Stillwater. The specific name for 
each new species is derived from the spe- 
cles name of type host. 


Myrsidea castanonotae Hellenthal and 
Price, new species 
(Figs. 1-3) 


Type host.—Ptilorrhoa castanonota 
(Salvadori), the chestnut-backed jewel-bab- 
bler. 

Male.—As in Fig. 1. Anterior margin of 
head evenly rounded, without preocular 
notch or slit; outer occipital seta much 
shorter than inner; without ventral spinous 
process; gula with 5, less often 4, setae; hy- 
popharyngeal sclerites well developed. 
Pronotum with 6 long setae at posterior 
margin, 3 short setae at each lateral corner, 


486 


% 


Figs. 1-3. 
rax and abdomen. 


and no dorsal setae; mesonotum with single 
pair of minute setae posterior to postnotum, 
prosternal plate well developed, with single 
pair of minute anterior setae; metanotal 
posterior margin and metasternal plate each 
with 6—8 setae. Femur III with ventral setal 
brush. Abdomen with undivided tergites; 
without anterior tergal or pleural setae. Ter- 
gal setae, including postspiracular setae: I, 
16-20; II-III, 20-22; IV—VII, 14—20; VIII, 
10—12. Postspiracular setae very long on I— 
II, IV, and VII-VIII, shorter on III and V— 
VI. Sternite I small, without setae; sternite 
II enlarged, with aster of 6—7 prominent 
heavy setae at each ljateroposterior corner. 
Sternal setae: II, 15—20 anterior, 17—18 mar- 
ginal setae in addition to those in asters; III, 
37-43; IV-V, 52-62; VI, 49-58; VII, 33— 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


WS 


ABER sesn ewe 


Myrsidea castanonotae. 1, Male dorsoventral. 2, Male genitalia. 3, Female dorsoventral metatho- 


43; VIII, 17-25. Genitalia (Fig. 2) with 
slender straight parameres, lightly spiculate 
sac, and elongate sclerite as shown. Dimen- 
sions: TW, 0.43—0.46; HL, 0.31—0.32; PW, 
0.28-0.31; MW, 0.41—0.44; AWIV, 0.50— 
0.51; TL, 1.39—1.46; GL, 0.41-0.46. 
Female.—Head and thorax much as for 
male. Metathorax and abdomen as in Fig. 
3. Metanotum with convex posterior mar- 
gin. Tergite I with extensive slender medi- 
oposterior projection, II—VII medially nar- 
rowed and displaced as shown, VIII nor- 
mal. Tergal setae (including postspiracular 
setae): I-III, 19—28; IV, 9-10; V, 8-9; VI— 
VIII, 8. Postspiracular setae as for male. 
Sternal setae: I, 0; Il, 4—6 short mostly la- 
teroanterior setae and 16—19 marginal setae 
in addition to 7—8 setae in each aster; III, 


VOLUME 107, NUMBER 3 
4 
iy 


ARS a= 


V 


wes if 
(Sy 


Y 


RTT at 
te 


487 


Figs. 4-6. Myrsidea leucostictae. 4, Male dorsoventral metathorax and abdomen. 5, Male genitalia. 6, Fe- 


male dorsoventral metathorax and abdomen. 


35-48; IV—V, 43-68; VI, 33-43; VU, 11- 
17. Subgenital plate with 13—15 marginal, 
14—17 anterior setae. Anus oval, without in- 
ner setae, and with 34—42 setae in dorsal 
fringe, 36—45 in ventral fringe. Dimensions: 
TW, 0.50-—0.51; HL, 0.33-—0.35; PW, 0.32; 
MW, 0.54—0.58; AWIV, 0.68—0.73; TL, 
1.73-1.77; ANW, 0.23-0.25. 

Type material.—Holotype 2, ex P. cas- 
tanonota, New Guinea: West Sepik, Mt. So- 
moro, 23 May 1975, 104794; 1 paratype 6, 
same as holotype; | d, 1 2 paratypes, same 
as holotype except 20 May 1975, 104754; 
1 3, 1 2 paratypes, same except Madang 
Dist., Wanuma, 13 Mar. 1974, 104203. 

Remarks.—The female of this new spe- 
cies is readily recognized by the shape of 
its abdominal tergites and the associated 
chaetotaxy, the male by its genitalia with 
the slender straight parameres and genital 
sac sclerite as in Fig. 2. 


Myrsidea leucostictae Hellenthal and 
Price, new species 
(Figs. 4—6) 


Type host.—Ptilorrhoa leucosticta (P. L. 
Sclater), the spotted jewel-babbler. 

Male.—Head and thorax as in Fig. 1. Ab- 
domen as in Fig. 4, differing from M. cas- 
tanonotae as follows. Metasternal plate 
with 8—9 setae. Fewer tergal setae: I, 12— 
16; II-III, 15-18; IV—VII, 14—16; VIII, 8- 
9. Postspiracular setae shorter on I and lon- 
ger on III. Sternal setae: Il, 26—27 anterior 
setae, 14—15 marginal in addition to 7 in 
each lateroposterior aster; III, 21—26; [V—V, 
60—67; VI, 56-60. Genitalia (Fig. 5) with 
short outwardly curved parameres and sac 
sclerite as shown. Dimensions: AWIV, 
0.56—0.57. 

Female.—Head and thorax much as for 
male. Metathorax and abdomen as in Fig. 


488 


6. Metanotum with slightly convex poste- 
rior margin. Tergites II-VI with small me- 
dioposterior convexity, I and VII—VIII 
straight. Tergal setae (including postspira- 
cular setae): I, 15—16; I-III, 20—26; [V— 
VII, 16—20; VIII, 12—14. Postspiracular se- 
tae as for male. Sternal setae: I, 0; II, 42— 
48 anterior and 16—18 marginal setae in ad- 
dition to 7—8 setae in each aster; III, 46— 
49; IV-V, 90-98; VI, 81-91; VII, 42—56. 
Subgenital plate with 15—18 marginal, 34— 
38 anterior setae. Anus oval, without inner 
setae, and with 46—56 setae in dorsal fringe, 
46—49 in ventral fringe. Dimensions: TW, 
0.49-0.51; HL, 0.34—0.35; PW, 0.32-0.33; 
MW, 0.49-0.52; AWIV, 0.76—0.80; TL, 
1.78-1.87; ANW, 0.25-—0.27. 

Type material.—Holotype 2, ex P. leu- 
costicta, New Guinea: Morobe Dist., Moi- 
mo, 15 Dec. 1969, BBM-98129; | paratype 
3d, same as holotype; | 6, 3 ¢ paratypes, 
same except Wau, Kainde Rd., 12 Aug. 
1969; BBM-97632. 

Remarks.—The female is recognizable 
from that of M. castanonotae by its lack of 
modified abdominal tergites and its much 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


larger number of setae on all abdominal 
sternites and on tergites [V—VIII. The male 
is separable by the different shape of the 
genitalic parameres and by having more an- 
terior setae on sternite II and fewer setae on 
sternite III. 


LITERATURE CITED 


Clay, T. 1966. Contributions towards a revision of 
Myrsidea Waterston. I. (Menoponidae: Mallopha- 
ga). Bulletin of the British Museum (Natural His- 
tory) Entomology 17: 327-395. 

Dickinson, E. C., ed. 2003. The Howard and Moore 
Complete Checklist of the Birds of the World. 3™ 
edition. Princeton Univ. Press, Princeton, New 
Jersey. 1,039 pp. 

Hellenthal, R. A. and R. D. Price. 2003. The genus 
Myrsidea Waterston (Phthiraptera: Menoponidae) 
from bulbuls (Passeriformes: Pycnonotidae), with 
descriptions of 16 new species. Zootaxa 354: 1— 
20. 

Price, R. D., R. A. Hellenthal, and R. L. Palma. 2003. 
World checklist of chewing lice with host asso- 
ciations and keys to families and genera, pp. 1— 
448 in Price, R. D., R. A. Hellenthal, R. L. Palma, 
K. P. Johnson, and D. H. Clayton. The Chewing 
Lice: World Checklist and Biological Overview. 
Illinois Natural History Survey Special Publica- 
tion 24. x + 501 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 489-500 


ETHIROMYIA, A NEW GENUS OF HCLARCTIC DOLICHOPODINAE 
(DIPTERA: DOLICHOPODIDAE) 


Scott E. BROOKS AND TERRY A. WHEELER 


(SEB) Diptera Unit, Invertebrate Biodiversity, Agriculture and Agri-Food Canada, K.W. 
Neatby Building, 960 Carling Ave., Ottawa, Ontario, KIA OC6, Canada (email: 
brookss @agr.gc.ca); (TAW) Department of Natural Resource Sciences, McGill University 
(Macdonald Campus), Ste-Anne-de-Bellevue, Quebec, H9X 3V9, Canada (email: 
wheeler @nrs.mcgill.ca) 


Abstract.—Ethiromyia Brooks, new genus, is described, and distinguished from other 
Dolichopodinae by the following features: wing vein M straight and subparallel to R,.;; 
notum of thorax metallic greenish black or bluish black, with violet reflections; thoracic 
pleuron with a cluster of fine hairs in front of the posterior spiracle; foretibia of males 
with an elongate apicoventral seta; cercus of males whitish with black border, large, 
rounded, with long, fine marginal setae; dorsal surstylus with preapical dorsal notch and 
keel-like projection; female terminalia with inner medial pair of spines on tergite 10. The 
genus comprises three species: Ethiromyia purpurata (Van Duzee), new combination 
(eastern Nearctic); Ethiromyia violacea (Van Duzee), new combination (eastern Nearc- 
tic); and Ethiromyia chalybea (Wiedemann), new combination (Europe). All species are 


redescribed and a key is provided. 


Key Words: 


Gymnopternus Loew is a diverse (more 
than 100 species) and widespread genus of 
dolichopodine flies found in the Nearctic, 
Palearctic, and Oriental regions. Dolicho- 
podid workers in the Old and New World 
have differed in their views of the taxonom- 
ic rank of Gymnopternus. Although Loew 
(1857) originally assigned Gymnopternus 
generic rank, subsequent Old World work- 
ers treated it as a synonym (Becker 1917— 
1918, Negrobov 1991) or subgenus (Pollet 
1990, Wei 1997, Chandler 1998, Yang and 
Grootaert 1999, Yang and Saigusa 1999) of 
Hercostomus Loew. Pollet (2004) recently 
restored Gymnopternus to generic rank and 
transferred the Palearctic species from the 
subgenus Hercostomus (Gymnopternus) to 
Gymnopternus. In the Nearctic, Gymnopter- 
nus has long been recognized as a genus 


Dolichopodidae, Dolichopodinae, Ethiromyia, new genus, Holarctic 


(e.g., Curran 1933, 1934; Robinson 1964; 
Foote et al. 1965; Robinson and Vockeroth 
1981; Pollet et al. 2004) and that classifi- 
cation was supported by a phylogenetic 
analysis (Brooks 2005) that placed Gym- 
nopternus not with Hercostomus, but in- 
stead closely related to Dolichopus Latreille 
(including species formerly placed in Licht- 
wardtia Enderlein), based on the synapo- 
morphic possession of a cluster of fine hairs 
on the thoracic pleura in front of the pos- 
terior spiracle. 

Gymnopternus is a morphologically uni- 
form genus, with the exception of an enig- 
matic species group that includes Gymnop- 
ternus chalybeus (Wiedemann), Gymnop- 
ternus purpuratus (Van Duzee), and Gym- 
nopternus violaceus (Van Duzee) (herein 
referred to as the chalybeus group). In his 


490 


revision of the European species of the sub- 
genus Hercostomus (Gymnopternus), Pollet 
(1990) noted several differences between 
G. chalybeus and its European congeners, 
but ultimately left that species in the sub- 
genus Hercostomus. Recently, Pollet (2004) 
further noted a number of differences be- 
tween the chalybeus group and Gymnopter- 
nus, based on a study of over thirty species 
from the Nearctic and Palaearctic Regions. 
In the same paper, he hypothesized a sister 
group relationship between the chalybeus 
group and the remaining Gymnopternus 
based on subparallel wing veins R,,; and 
M. In contrast, Brooks (2005) considered 
that character state plesiomorphic, and 
found that the chalybeus group 1s the sister 
group to Dolichopus based on the posses- 
sion of a distinctive, dorsally notched dorsal 
surstylar lobe of the male genitalia (Figs. 4, 
5, 9,10) and a pair of inner, medial spines 
on tergite 10 of the female terminalia (Figs. 
7, 8). The monophyly of Gymmnopternus, 
exclusive of the chalybeus group, is sup- 
ported by the possession of elongate pro- 
jections on the base of the ejaculatory apo- 
deme and a broad, lobate postgonite 
(Brooks 2005). 

Dolichopus is considered to be mono- 
phyletic based on the synapomorphic pos- 
session of one or more strong dorsal setae 
on the hind basitarsus, a distinctive S- 
shaped bend in wing vein M, and a T- 
shaped ejaculatory apodeme (Brooks 2005). 
The monophyly of the chalybeus group is 
supported by the possession of an elongate 
apicoventral seta on the male foretibia, and 
the distinctive male cercus, characterized by 
long, fine marginal setae (Pollet 2004, 
Brooks 2005). Despite the fact that both of 
these features are homoplasious within the 
Dolichopodinae and have arisen in some 
species of Dolichopus and Hercostomus, 
they are part of a suite of congruent nested 
synapomorphies that support the monophy- 
ly of the chalybeus group. In this paper the 
new genus Ethiromyia is established for the 
chalybeus group, and its three included spe- 
cies are redescribed. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


MATERIALS AND METHODS 


This study is based on material housed 
in the Canadian National Collection of In- 
sects, Ottawa, Ontario, Canada (CNC), Ly- 
man Entomological Museum, McGill Uni- 
versity, Ste-Anne-de-Bellevue, Quebec, 
Canada (LEM), Museum fiir Naturkunde 
der Humboldt Universitat zu Berlin, Berlin, 
Germany (ZMHB) and Naturhistorisches 
Museum Wien, Wien, Austria (NMW). 
Morphological terminology mainly follows 
McAlpine (1981); terms for male genitalia 
follow Cumming et al. (1995) and Sinclair 
(2000). Body length is measured from the 
base of the antenna to the tip of the abdo- 
men. Wing length is measured from the hu- 
meral crossvein to the wing apex. Relative 
lengths of each tarsomere are representative 
ratios expressed using the following for- 
mula: t,/t,/t,/t,/t;, where t, is the basitarsus. 

Male and female terminalia were mac- 
erated in either 10% KOH, which was heat- 
ed on a hot plate for about 10 minutes, or 
in 85% lactic acid, heated in a microwave 
oven. Each microwave heating interval 
comprised 30 seconds and was followed by 
a 1—2 minute cooling period during which 
macerated muscle tissue was removed with 
a fine probe. 

Figures showing male genitalia in lateral 
view are oriented as they appear on the in- 
tact specimen (rotated 180° and lateroflexed 
to the right), with the morphologically ven- 
tral surface up, dorsal surface down, ante- 
rior end facing right and posterior end fac- 
ing left. Figures showing the male genitalia 
in ventral view are correspondingly orient- 
ed with the anterior end facing right and 
posterior end facing left. The following ab- 
breviations are used in the figures: apv lobe: 
apicoventral epandrial lobe; bv lobe: basi- 
ventral epandrial lobe; bv seta: basiventral 
epandrial seta; cerc: cercus; dsur: dorsal 
lobe of surstylus; ejap: ejaculatory apode- 
me; epand: epandrium; hy: hypandrium; 
hyap: hypandrial apodeme; pgon: postgon- 
ite; ph: phallus; S: sternite; T: tergite; vsur: 
ventral lobe of surstylus. Abbreviations 


VOLUME 107, NUMBER 3 


used in the text include T: abdominal ter- 
gite, and S: abdominal sternite. 


Ethiromyia Brooks, new genus 


Type species: Hercostomus purpuratus Van 
Duzee 1925: 185 [Nearctic], by present 
designation. 


Recognition.—Species of Ethiromyia can 
be recognized by the following combina- 
tion of characters: wing vein M straight and 
subparallel to R,,;; notum of thorax metal- 
lic greenish black or bluish black, with vi- 
olet reflections; thoracic pleuron with a 
cluster of fine hairs in front of posterior spi- 
racle; male foretibia with an elongate api- 
coventral seta; male cercus whitish with 
black border, large, rounded, with long, fine 
marginal setae; dorsal surstylus with preap- 
ical dorsal notch and keel-like projection; 
female terminalia with inner medial pair of 
spines on T10. Species of Ethiromyia lack 
a distinct anterodorsal row of strong setae 
on the foretibia (present in most Gymnop- 
ternus), and also possess 1—3 strong poster- 
oventral setae on the foretibia (absent in 
most Gymnopternus). In addition, species 
of Ethiromyia lack the dorsal setae on the 
hind basitarsus that are found in species of 
Dolichopus. 

Description.—Head: Vertex not excavat- 
ed, 1 pair of strong divergent ocellar setae, 
1 pair of strong vertical setae, stronger than 
postverticals. Frons about 2—2.8 wider 
than high, sides weakly convergent anteri- 
orly. Face broad in male, sides slightly con- 
vergent below or subparallel, broader in fe- 
male with sides subparallel. Clypeus slight- 
ly produced to strongly bulging, especially 
in female, lower margin straight or slightly 
emarginate, ending well above lower eye 
margin. Palp ovoid, with weak setae on dis- 
tal half of outer surface and a distinct apical 
seta. Proboscis large and projecting or mod- 
erate in size. Antenna inserted above mid- 
dle of head; scape subconical, dorsally se- 
tose, with well-developed acute medioven- 
tral process; pedicel short, with nipplelike 
medial condyle; first flagellomere subtrian- 


49] 


gular to ovoid, about as long as wide; arista 
dorsal, 2-segmented, second segment weak- 
ly to strongly pubescent. Postocular setae 
uniseriate, lowermost seta sometimes stron- 
ger. One pair of postvertical setae, subequal 
to distinctly stronger than uppermost pair of 
postoculars. 

Thorax: Notum metallic greenish black 
or bluish black with violet reflections. Ac- 
rostichals biserial; 6 dorsocentrals, fifth pair 
distinctly offset medially; postpronotum 
with | strong medioclinate seta and 2-3 
weaker outer setae; 1 strong outer posthu- 
meral, | weak inner posthumeral; 2 noto- 
pleurals; 1 presutural; 1 sutural; 2 supra- 
alars; 1 postalar. Upper and lower part of 
propleuron with fine hairs; lower part of 
propleuron with | strong prothoracic seta; 
pleural surface in front of posterior spiracle 
with a cluster or row of fine hairs; metepis- 
ternum with a cluster of several fine hairs. 
Scutellum with 1 strong inner seta and | 
small outer seta on lateral margin, dorsum 
with sparse hairs, posterior margin with 
sparse short hairs or long dense hairs. 
Legs: Pulvilli normal. Foreleg: Tibia with 
2—4 anterodorsal setae, 2 dorsals, 1—3 pos- 
teroventrals, 2 apicals, | long, fine apicov- 
entral in male, without distinct, regular, an- 
terodorsal row of strong setae. Midleg: Fe- 
mur with | anterior preapical seta; tibia 
with 3—5 anterodorsals, 2 dorsals, 1—3 an- 
teroventrals, 5 apicals. Hind leg: Coxa with 
strong lateral seta near or slightly below 
middle; femur with | anterodorsal preapical 
seta; tibia with 3—6 anterodorsals, 1 preap- 
ical dorsal, 3—6 posterodorsals, 3—6 ven- 
trals, 2 apicals, apex with weak to indistinct 
ridgelike process posterodorsally in male; 
basitarsus subequal to or slightly shorter 
than second tarsomere, without dorsal setae, 
with distinct basiventral seta, male with 
hooklike process posterobasally. 

Wing (Figs. 1-3): Brownish to grey. 
Male costa with or without pterostigma 
near insertion of R,; R,,, relatively straight 
to weakly convex; R,,; straight with pos- 
terior curve in distal section; distal section 
of M beyond crossvein dm-cu with barely 


492 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


3 


1mm 


Figs. 1-3. 


discernable sinuous bend before middle, 
straight, or with slight convex curve in dis- 
tal section similar to that of R,,;, ending 
near wing apex; R,,; and M subparallel; 
crossvein dm-cu subequal to or shorter than 
distal section of CuA,. 

Abdomen: Subconical. T1l—5 setose. 
Male: T6 bare; S2 unmodified; S3 unmod- 
ified or emarginate and mainly membranous 
posteromedially; S4 strongly emarginate or 
divided, membranous medially; S5 mainly 
to entirely membranous; S6 mainly mem- 
branous, sclerotized along anterior margin; 
segment 7 bare, forming well-developed 
peduncle; S8 subquadrate to subtriangular, 
setose. Hypopygium (Figs. 4—6, 9-10) 


Wings. 1, Ethiromyia purpurata, male. 2 


, E. purpurata, female. 3, E. violacea, male. 


large. Epandrium subtriangular in lateral 
view, about 1.5—2x longer than high, fo- 
ramen lateral, well-separated from base of 
cerci; basiventral epandrial !obe weakly de- 
veloped, basiventral epandrial seta present; 
apicoventral epandrial lobe well-developed, 
subquadrate, rounded or flared apically, 
with | lateral and 2 apical setae. Surstylus 
2-lobed. Ventral lobe more or less digiti- 
form, with or without dorsal hump, with 
weak dorsal to dorsomedial preapical pro- 
jection, apex with short, stout seta. Dorsal 
lobe larger than ventral lobe, with 1-2 
strong dorsomedial setae and 1 preapical 
lateral seta, dorsal surface notched preapi- 
cally with distinct to weakly developed 


VOLUME 107, NUMBER 3 


keellike projection across notch bearing a 
short seta (Fig. 5). Postgonite with anter- 
oventral portion weakly sclerotized, nearly 
membranous and bifurcate anteriorly; pos- 
terodorsal portion vestigial (Figs. 9-10), or 
well-developed and digitiform (Fig. 5). 
Proctiger brushes absent. Cercus (Figs. 4, 
9-10) large, round to ovoid, pale with dark 
margin; apical and lateral margin jagged, 
sometimes with well-developed digitiform 
projections (Figs. 4, 9); lateral and/or apical 
margin with very long, fine setae. Hypan- 
drium elongate and slender, troughlike, free 
laterally with membranous connection to 
epandrium basally; hypandrial arms con- 
nected to hypandrium; hypandrial apodeme 
well-developed, with knob-like apex. 
Sperm pump cylindrical; ejaculatory apo- 
deme rodlike; basal sclerite of sperm pump 
well-developed, thick and heavily sclero- 
tized, broadly V-shaped in dorsal view. 
Phallus elongate and slender, apical portion 
with weak rounded projection (Figs. 5, 10), 
or finely serrate (Fig. 9). Female (Figs. 7— 
8): T6, T7, S6 and S7 undivided; T8 and 
S8 divided medially, tergite and sternite 
fused anterolaterally. Furca narrow and 
weakly sclerotized or absent. T10 divided 
medially into hemitergites each bearing 4— 
5 spines along outer margin and a single 
inner medial spine (Fig. 7), spines pointed 
to blunt apically. Upper lobe of cercus with 
short apical seta. 

Etymology.—The generic name is de- 
rived from the Greek etheria (hair) in ref- 
erence to the long hairs on the male cercus, 
and the Greek myia (fly). The gender is 
feminine. 

Remarks.—Ethiromyia was referred to as 
‘“New Genus A” in Brooks (2005). 


KEY TO SPECIES OF ETHIROMYIA 


1. Mid- and hind tibiae yellow with dark spots at 
insertion points of setae; midtibia with | strong 
ventral seta at distal third; male with wing mar- 
gin incised between veins M and CuA, (Fig. 
3); male cercus with weakly developed digiti- 
form projections on apicodorsal margin (Fig. 
10) (eastern Nearctic) ... violacea (Van Duzee) 

— Mid- and hind tibiae without dark spots at in- 


493 


sertion points of setae, hind tibia yellow or 
brown; midtibia lacking strong ventral seta at 
distal third; male with wing evenly convex be- 
tween M and CuA, (Fig. 1); male cercus with 
well-developed digitiform projections on api- 
codorsal margin (Figs. 4,9) ............. 2 
. Palp blackish brown; antenna entirely black; 
forecoxa dark, forefemur usually brown dor- 
sally. Male: Hind leg with long, fine posterior 
hairs on distal half of femur and basal part of 
tibia; hind tibia with four closely spaced, flat- 
tened posterodorsal setae on basal half; wing 
with pterostigma near insertion of R, (Fig. 1); 
T2 and T3 velvety black laterally; cercus with 
scythe-shaped apicoventral seta on first elon- 
gate digitiform projection, marginal setae about 
as long as width of cercus (Fig. 4) (eastern Ne- 
ALCO) iuspekieerst es 6 = Sao purpurata (Van Duzee) 
— Palp yellow, dark basally; antenna with scape 
and pedicel pale ventrally; forecoxa mainly 
pale, infuscated basally on outer side, forefe- 
mur entirely yellow. Male: Hind leg lacking 
fine posterior hairs on femur and tibia; hind 
tibia without four flattened posterodorsal setae 
on basal half; wing without pterostigma; T2 
and T3 not velvety black laterally; cercus with 
spatulate apicoventral seta on first elongate 
digitiform projection, marginal hairs distinctly 
longer than width of cercus (Fig. 9) (Europe) 
chalybea (Wiedemann) 


iw) 


Ethiromyia chalybea (Wiedemann), 
new combination 
(Fig. 9) 


Dolichopus chalybeus Wiedemann 1817: 
V2. 

Dolichopus cinereomaculatus Roser 1840: 
5) OF 

Gymnopternus chalybeus (Wiedemann): 
Loew 1857: 21; Pollet 2004: 546. 

Hercostomus (Gymnopternus) chalybeus 


(Wiedemann): Lundbeck 1912: 189; 
Chandler 1998: 90. 
Hercostomus chalybeus (Wiedemann): 


Becker 1917: 212: Pollet 1990: 361. 
Gymnopternus cinereomaculatus (Roser): 
Pollet 2004: 546. 


Male.—Body length = 3.6—4.4 mm, 
wing length = 3.8—4.6 mm. Head: Frons 
metallic violet bronze or violet blackish, 
lower and lateral margins often metallic 
bluish green. Face and clypeus silvery-grey 
pollinose; face about 0.2 as wide as head; 


494 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


pane Se od 
ph eee 
bv lobe 


apv lobe 


epand 


bv lobe 


Figs. 4-8. Ethiromyia purpurata. 4, Male genitalia, left lateral view (external). 5, Male genitalia, left lateral 
view (internal). 6, Male genitalia, ventral view (postgonite, surstylus and cercus not shown). 7, Female genitalia, 
dorsal view. 8, Female genitalia, left lateral view. 


VOLUME 107, NUMBER 3 


495 


a ar 
bv lobe 


apv lobe 


epand 


10 


Figs. 9-10. Ethiromyia chalybea and E. violacea. 9, E. chalybea, male genitalia, left lateral view. 10, E. 


violacea, male genitalia, left lateral view. 


clypeus strongly bulging. Palp mainly yel- 
low, brownish black near base, with black 
hair. Proboscis large and usually projecting. 
Scape and pedicel yellow ventrally, dark 
brown dorsally; first flagellomere blackish 
brown, apex weakly pointed or rounded; 
arista dark brown, strongly pubescent. Post- 
ocular setae black. Ocellar tubercle with 
several hairs medially. Postvertical seta 
slightly stronger than upper postocular seta. 

Thorax: Notum metallic greenish black 
or bluish black, with violet reflections. 
Pleuron dark metallic greenish blue and 


bronze with whitish pollinosity. Scutellum 
black with metallic violet, green and blue 
reflections, with long, dense hairs on pos- 
terior margin. Notum and scutellum paler 
black bronze in older specimens. 

Legs: Forecoxa, femora, fore- and mid- 
tibiae mainly yellow, mid- and hind coxae 
concolorous with thoracic pleuron, hind tib- 
ia more or less brownish, darker towards 
apex. Foreleg: Coxa darkened basally on 
outer side; tibia with 1—2 strong anterodor- 
sal setae in basal part, 1—2 weaker distal 
anterodorsals, 2 dorsals, 1—2 posteroven- 


496 


trals, distal seta often weaker or absent, 2 
apicals in addition to long, fine apicoven- 
tral; basitarsus mainly yellow with brown 
apex, tarsomeres 2—5 brown, tarsomere ra- 
tio: 4.0/1.7/1.3/1.0/1.0. Midleg: Tibia with 
3 anterodorsal setae, 2 dorsals, 2 anterov- 
entrals, 5 apicals; basitarsus mainly yellow 
with brown apex, tarsomeres 2—5 brown, 
tarsomere ratio: 4.0/2.0/1.5/1.1/1.0. Hind 
leg: Coxa often yellow apically; femur 
brown apicodorsally; tibia with 4—6 anter- 
odorsal setae, 1 preapical dorsal, 4—6 pos- 
terodorsals, 3—5 ventrals, 2 apicals, and 
dense, well-developed hairs posteriorly; tar- 
sus blackish brown, tarsomere ratio: 2.9/ 
S\N SYM P21 (0). 

Wing: Brown, darker on anterior half; 
pterostigma absent; margin evenly convex 
between M and CuA,; calypter with black 
setae; halter pale yellow. 

Abdomen: T1—5 dark bronze dorsally, 
metallic greenish blue laterally with whitish 
pollinosity, with weak violet reflections; S5 
with eversible membranous sac; T6, seg- 
ment 7 and S8 dark brown. Hypopygium 
(Fig. 9): Epandrium dark brown, abruptly 
narrowed in distal half, ventral margin (in- 
cluding basiventral and apicoventral epan- 
drial lobes) pale yellow; basiventral epan- 
drial lobe very weakly developed; apicov- 
entral epandrial lobe flared apically. Sursty- 
lus pale yellow; ventral lobe slender, 
without dorsal hump; dorsal lobe with 2 
strong, dark dorsomedial setae. Postgonite: 
posterodorsal portion vestigial. Cercus 
ovoid with well-developed digitiform pro- 
jections, first elongate digitiform projection 
with apicoventral spatulate seta, marginal 
setae very long. Hypandrium amber, lack- 
ing dorsal process, tubular basally, open 
along right side exposing phallus. Phallus 
with a pair of serrate longitudinal bands 
apically. 

Female.—Body length = 4.0—4.7 mm, 
wing length = 4.1—4.6 mm. Similar to male 
except as follows: Head: Face and clypeus 
broader; face about 0.3 as wide as head, 
slightly darker; clypeus weakly pollinose, 
bronze, especially on upper part; proboscis 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


larger. Legs: Foretibia lacking long, fine 
apicoventral; posterior hairs on hind tibia 
not as strongly developed. 

Type material examined.—Syntypes: 2 
eunle 2) (ZMEB: Nor 29311); i Gee 
(NMW), GERMANY: Kiel. 

Other material examined.—BELGIUM: 
Oost-Vlaanderen: 16 5, 20 2, Heurne, Het 
Dal Nature Reserve, white water traps, 
1997, M. Pollet & P Grootaert (LEM); 
AUSTRIA: Carinthia: 1 6, nr Drobollach 
am Faaker See, reedbed, 3.vii.1992, C.E. 
Dyte (LEM); 1 @, same data except nr 
Faaker See, woodland (LEM). 

Distribution.—Great Britain, Ireland, 
France, Belgium, The Netherlands, Den- 
mark, Germany, Switzerland, Italy, Austria, 
Czech Republic, Slovakia, Poland, Hunga- 
ry, Romania, former Yugoslavia, Sweden, 
Norway, Finland, Belarus, Ukraine, and 
European Russia (Negrobov 1991, Pollet, 
unpublished data). 

Ecology.—Ethiromyia chalybea prefers 
reedmarshes and humid woodlands (carrs) 
with an undergrowth of sedges, where it 1s 
often found in large numbers (see Pollet 
1992, Pollet et al. 1992). 


Ethiromyia purpurata (Van Duzee), 
new combination 
(Figs. 1-2, 4-8) 


Hercostomus purpuratus Van Duzee 1925: 
185. 

Gymnopternus purpuratus (Van Duzee): 
Foote et al. 1965: 499; Pollet et al. 2004: 
41. 


Male.—Body length = 4.2—5.0 mm, 
wing length = 3.7—4.3 mm. Head: Frons 
bronze to blackish bronze, with violet me- 
tallic reflections, lower and lateral margins 
usually metallic bluish green. Face and 
clypeus silvery-grey pollinose, sometimes 
with weak blue-green reflections; face 
about 0.3 X as wide as head; clypeus strong- 
ly bulging. Palp blackish brown with black 
hairs. Proboscis large and projecting. An- 
tenna black; first flagellomere rounded or 
weakly pointed apically; arista with short 


VOLUME 107, NUMBER 3 


pubescence. Postocular setae black. Ocellar 
tubercle with several hairs medially. Post- 
vertical seta stronger than upper postocular 
seta. 

Thorax: Notum metallic greenish black 
with violet, blue and bronze reflections. 
Pleuron dark metallic greenish blue with 
whitish pollinosity. Scutellum blackish with 
metallic violet, green and blue reflections; 
with fine, short hairs on posterior margin. 

Legs: Coxae blackish brown with metal- 
lic green tinge, concolorous with thoracic 
pleuron; femora and tibiae mainly yellow. 
Foreleg: Femur usually brown dorsally; tib- 
ia with 1—3 anterodorsal setae, distal antero- 
dorsals sometimes weaker, 2 dorsals, 1-3 
posteroventrals, 2 apicals in addition to 
long, fine apicoventral; basal two-thirds of 
basitarsus yellow, distal third black, tarso- 
meres 2—5 black, tarsomere ratio: 3.5/1.6/ 
1.2/1.0/1.0. Midleg: Tibia with 3—5 antero- 
dorsal setae, 2 dorsals, 1—3 anteroventrals, 
5 apicals; basal two-thirds of basitarsus yel- 
low, distal third black, tarsomeres 2—5 
black, tarsomere ratio: 4.3/2.2/1.5/1.1/1.0. 
Hind leg: Distal half of femur and basal 
part of tibia with long, fine hairs posteri- 
orly, hairs about as long as width of femur; 
tibia with 4—5 anterodorsal setae (distal 1— 
2 setae sometimes dorsal), 1 preapical dor- 
sal, 4 close set, flattened posterodorsals on 
basal half, 3—6 weaker ventrals, 2 apicals, 
apex of tibia brown posteriorly; tarsus 
blackish brown, tarsomere ratio: 2.8/3.1/ 
2.0/1.2/1.0. 

Wing (Fig. 1): Evenly dark brown; ptero- 
stigma present near insertion of R,; wing 
margin evenly convex between M _ and 
CuA,; calypter with black setae; halter pale 
yellow. 

Abdomen: T1—5 dark metallic greenish, 
lateral part of T2 and T3 velvety black with 
fine hairs, T4 also with fine hairs laterally; 
T6, segment 7 and S8 dark brown. Hypo- 
pygium (Figs. 4, 5, 6): Epandrium mainly 
dark brown, ventral margin (including bas- 
iventral and apicoventral epandrial lobes) 
pale yellow, left basiventral epandrial lobe 
larger than right lobe (Fig. 6); apicoventral 


497 


epandrial lobe subquadrate. Surstylus pale 
yellow; ventral lobe with dorsal hump; dor- 
sal lobe with 1 strong, dark dorsomedial 
seta. Postgonite with posterodorsal portion 
pale, digitiform, bent ventrally. Cercus 
round with well-developed digitiform pro- 
jections, first elongate digitiform projection 
with apicoventral scythe-shaped seta. Hy- 
pandrium amber with dorsal process near 
middle, tubular basally, open along right 
side exposing phallus. Phallus with weak 
rounded process in apical portion. 

Female.—Body length = 4.3-5.5 mm, 
wing length = 3.9—4.5 mm. Similar to male 
except as follows: Head: Face and clypeus 
darker, broader; face about 0.35 as wide 
as head, clypeus very strongly bulging. Pro- 
boscis slightly larger. Legs: Foretibia lack- 
ing long, fine apicoventral; hind femur and 
basal part of hind tibia lacking long, fine 
hairs posteriorly; hind tibia with three nor- 
mally developed posterodorsals on basal 
half. Wing: Pterostigma absent. Abdomen: 
Dark metallic green with violet, blue and 
bronze reflections, T2 and T3 without lat- 
eral velvety black patch, T2—4 without fine 
hairs laterally. 

Type material examined.—Holotype 
3d: CANADA: Manitoba: Stockton, 
29.vii.1924, N. Criddle (CNC, No. 1413). 
Allotype ¢: same data as holotype. 

Other material examined.—CANADA: 
Ontario: 1 6, Swastika, 7.vii.1987, J.R. 
Vockeroth (CNC); Quebec: 1 6d, Beach- 
grove (45°37'N, 76°8’W), 24.vi.1988, J.R. 
Vockeroth (CNC); Lac St-Francois Natural 
Wildlife Area: 18 36, 6 2, Marais Fraser, 
45°02.37'N, 74°27.73'W, Carex meadow, 
pan trap, 03.vi—11.vi.1999, EK Beaulieu 
(LEM); 7 6, 3 2, same data except 26.v— 
03.vi.1999 (LEM); 1 2, same except sweep 
net, 28.v.1999 (LEM); 4 3, 3 2, same ex- 
cept sweep net, 05.vi.1999 (LEM); 8 6, 4 
2, same except 1|1.vi—19.vi.1999 (LEM); 3 
6, 5 ¢, same except 45°02.40’N, 
74°28.03'W (LEM); 2 &, same except 
26.v—03.vi.1999 (LEM); 3 6, 6 2, same 
except 11.vi-19.vi.1999 (LEM); 5 6, 12 2, 
NW of Aménagement Therrien, close to 


498 


ruisseau Therrien, 45°00.39'N, 74°30.99'W, 
Carex meadow, pan trap, 03.vi—11.vi.1999, 
F Beaulieu (LEM); 2 6, 3 2, same except 
11.vi.—19.vi.1999 (LEM); 1 6, same except 
19.vi.—26.vi.1999 (LEM); 8 3, 3 2, same 
except sweep net, 05.vi.1999 (LEM); 1 6, 
same except 45°00.17'N, 74°30.63'W 
(LEM); 8 3, 1 2, same except pan trap, 
03.vi-11.vi.1999 (LEM); 1 3, same except 
Pievi=l9 vie 1999) (EE MI 2G le same 
except 19.vi—26.v1.1999 (LEM). 
Distribution.—Manitoba, Michigan, 
northern Ontario and southwestern Quebec. 
Remarks.—Beaulieu and Wheeler (2001) 
collected large numbers of this species (as 
‘*Gymnopternus n. sp. 1”) in lakeside sedge 
meadows in southwestern Quebec. 


Ethiromyia violacea (Van Duzee), 
new combination 
(Figs. 3, 10) 


Proarchus violaceus Van Duzee 1921: 123 

Hercostomus (Proarchus) violaceus (Van 
Duzee): Leonard 1928: 782. 

Hercostomus violaceus (Van Duzee): Stey- 
skal 1959: 5. 

Gymnopternus violaceus (Van Duzee): 
Robinson 1964: 158; Foote et al. 1965: 
500; Pollet et al. 2004: 41. 


Male.—Body length = 3.6—4.6 mm, 
wing length = 3.7—4.4 mm. Head: Frons 
bronze to blackish, with violet and bluish 
green metallic reflections. Face and clypeus 
silvery grey pollinose; face about 0.2 as 
wide as head; clypeus weakly or strongly 
bulging. Palp mainly yellow, brown basally, 
with black hairs. Proboscis medium sized. 
Scape and pedicel mainly yellow, brown 
dorsally; first flagellomere yellow basally, 
distal portion blackish-brown, apex round- 
ed or weakly pointed; arista blackish 
brown, strongly pubescent. Postocular setae 
black. Ocellar tubercle with several hairs 
medially. Postvertical seta stronger than or 
subequal to upper postocular seta. 

Thorax: Notum metallic greenish black 
to greenish bronze, with violet and bluish 
reflections. Pleuron dark metallic greenish 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


grey to greenish bronze, with whitish pol- 
linosity. Scutellum dark bronze or blackish, 
with violet, green, and blue reflections; with 
fine, short hairs on posterior margin. 

Legs: Fore- and hind coxae mainly yel- 
low, brown at base on outer side, midcoxa 
more or less concolorous with thoracic 
pleuron; femora and tibiae yellow, mid- and 
hind tibiae with dark spots at insertion 
points of setae. Foreleg: Tibia with 2—4 an- 
terodorsal setae, distal 2 anterodorsals usu- 
ally weaker if developed, 2 dorsals, 1—3 
posteroventrals, 2 apicals in addition to 
long, fine apicoventral; basitarsus mainly 
yellow with brown apex, tarsomeres 2—5 
brown, tarsomere ratio: 4.1/1.7/1.3/1.0/1.0. 
Midleg: Femur with long, fine hairs basov- 
entrally, hairs slightly shorter than width of 
femur; tibia with 3—4 anterodorsal setae, 2 
dorsals, 2—3 anteroventrals, | ventral at dis- 
tal third, 5 apicals; basitarsus mainly yellow 
with brown apex, tarsomeres 2—5 brown, 
tarsomere ratio: 4.0/2.0/1.5/1.1/1.0. Hind 
leg: Femur with well-developed setae dor- 
sally; tibia with 3—4 anterodorsal setae, 3— 
4 posterodorsals, 3—4 ventrals, 1 preapical 
dorsal, 2 apicals; basitarsus mainly yellow 
with brown apex, tarsomeres 2—5 brown, 
tarsomere ratio: 2.6/3.2/2.0/1.3/1.0. 

Wing (Fig. 3): Grey; pterostigma absent; 
wing margin incised between M and CuA,; 
calypter with black setae; halter pale yel- 
low. 

Abdomen: T1-—5 dark metallic green with 
greyish pollen laterally, T1—3 metallic black 
or bronze dorsomedially; T6, segment 7 
dark brown; S8 brown or metallic greenish 
brown. Hypopygium (Fig. 10): Epandrium 
dark brown, sometimes dark metallic green 
basally, ventral margin (including basiven- 
tral and apicoventral epandrial lobes) pale 
yellow amber; basiventral epandrial lobe 
very weakly developed; apicoventral epan- 
drial lobe rounded apically. Surstylus pale 
yellow; ventral lobe slender, without dorsal 
hump; dorsal lobe with 2 strong, dark dor- 
somedial setae. Postgonite: posterodorsal 
portion vestigial. Cercus ovoid with jagged 
margin, lacking well developed digitiform 


VOLUME 107, NUMBER 3 


projections, marginal setae very long. Hy- 
pandrium pale amber, apex enlarged. Phal- 
lus with weak rounded preapical projection. 

Female.—Body length = 4.2 mm, wing 
length = 3.8—4.4 mm. Similar to male ex- 
cept as follows: Head: Face and clypeus 
broader; face about 0.3 as wide as head; 
clypeus strongly bulging. Proboscis slightly 
larger. Legs: Foretibia lacking long, fine 
apicoventral. Wing: Margin not distinctly 
incised between M and CuA,. 

Type material.—Holotype ¢: USA: New 
York: Erie County, Dayton, 5.vii.1920, 
M.C. Van Duzee (California Academy of 
Sciences, No. 3467) (not examined). 

Material examined.—CANADA: Ontar- 
io: 1 d, Ottawa, 8.viii.1993, J.R. Vockeroth 
(CNC); Quebec: 2 6d, Rigaud, Chemin 
de la Mairie, Parc Lévy Macdonald, 
6.vili.2000, sweep net, S.E. Brooks (LEM); 
1 2, Old Chelsea, 24.vi.1956, J.R. Vock- 
eroth (CNC); USA: North Carolina: 1 6, 
Highlands, 3,800’, 7.vi.1957, W.R.M. Ma- 
son (CNC); 1 3d, same except 10.vi.1957, 
J.R. Vockeroth (CNC); 1 6, same except 
LOW I95S7 (ONG) i 2. seine Gwoge 
20.vi.1957 (CNC); Tennessee: | 6, Knox- 
ville, Univ. Farm, 20.v.1957, J.R. Vocker- 
oth (CNC); 1 3, Knoxville Co., 26.v.1957 
(CNC); 1 2, same except 30.v.1957 (CNC). 

Distribution.—Ontario, Michigan, New 
York, Quebec to Massachusetts, south to 
Ohio, Virginia, Tennessee, North Carolina, 
and South Carolina (Pollet et al. 2004). 


ACKNOWLEDGMENTS 


Thanks to Frédéric Beaulieu whose field- 
work in Quebec sedge meadows provided 
many specimens of FE. purpurata and to 
Marc Pollet who first recognized those 
specimens as E. purpurata. Thanks to Jeff 
Cumming (CNC), Joachim Ziegler 
(ZMHB), Peter Sehnal (NMW), Marc Pol- 
let, and Peter Dyte for the loan of types and 
other specimens. We also thank Richard 
Hurley, Marc Pollet and Harold Robinson 
for reviewing an earlier version of the man- 
uscript. Magnolia Press is acknowledged 
for the use of figures 4—10 previously pub- 


499 


lished in Zootaxa 857 (figs. 11 and 12). 
Funding was provided by an NSERC Dis- 
covery Grant to T.A. Wheeler. 


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PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 501-509 


PHYLOGENETIC RELATIONSHIPS AMONG THE SPECIES OF PANTHIADES 
HUBNER (LYCAENIDAE: THECLINAE: EUMAEINI) 


ROBERT K. ROBBINS 


Department of Entomology, P.O. Box 37012, NHB Stop 127, Smithsonian Institution, 
Washington, DC 20013-7012, U.S.A. (e-mail: RobbinsR @ SI.edu) 


Abstract.—A species level phylogenetic analysis of Panthiades Hiibner was performed 
using twelve characters of wing pattern, androconia, and male and female genitalia. The 
purposes were to determine whether Panthiades is monophyletic without Cycnus Hiibner 
and to provide a cladogram for a project on the evolution of “false head” wing patterns. 
Parsimony analysis with all characters unordered yielded two trees. One was the strict 
consensus of the two trees, and the other was the only most parsimonious tree when one 
of the multi-state characters was ordered. Panthiades is characterized by five hypothesized 
synapomorphies. If Cycnus is recognized, Panthiades is not monophyletic on either of 
the most parsimonious cladograms. The “‘classic’’ false head wing patterns in Panthiades 


appear to have evolved once. 


Key Words: 


Nicolay’s (1976) taxonomic treatment of 
Panthiades Hiibner and Cycnus Hiibner has 
been stable with minor exceptions. Robbins 
(2004a, b) changed two specific epithets for 
nomenclatural reasons and synonymized 
the monotypic Cycnus with Panthiades, 
stating that Panthiades was probably not 
monophyletic without Cycnus. Consistent 
with this synonymy, Nicolay (1976:3) had 
noted that “‘the entry of the ductus semin- 
alis on the ventral-lateral side of the corpus 
bursae”’ is shared by Panthiades and Cyc- 
nus, but not by other close relatives. 

Panthiades is of biological interest be- 
cause it contains two species, P. bathildis 
(Reakirt) and P. phaleros (L.), that have 
wing patterns (Fig. 10) traditionally asso- 
ciated with the “‘false head”? hypothesis of 
predator avoidance (Robbins 1980). These 
wing patterns show a significantly greater 
incidence of unsuccessful predator attacks 
directed to the “‘false head”’ than other ly- 
caenid wing patterns (Robbins 1981). It is 


hairstreaks, false head hypothesis, Cycnus 


unclear whether the classic “‘false head”’ 
wing pattern evolved once or twice in the 
Panthiades/Cycnus lineage, especially since 
Nicolay (1976) placed one in Cycnus and 
one in Panthiades. 

In this paper, I infer phylogenetic rela- 
tions among the eight species of Panthiades 
for the purposes of assessing the monophy- 
ly of Panthiades without Cycnus and of 
providing a cladogram for ongoing studies 
of the evolution of wing patterns associated 
with the “‘false head”’ hypothesis. 


MATERIALS AND METHODS 


Coded characters were derived from a 
comparison of adult morphology using 
1,009 pinned specimens of Panthiades in 
the National Museum of Natural History, 
Smithsonian Institution, Washington, DC, 
USA, plus numerous specimens borrowed 
from other museums. In addition, 41 geni- 
talic dissections of both sexes of the eight 
Panthiades species were examined. Pan- 


502 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Character matrix for Panthiades. The outgroups are the type species of Parrhasius, Thepytus, and 


Porthecla. Characters and their states are detailed in the text. 


Characters 

Species | 2 3 4 5 6 7 8 9 10 11 12 
Panthiades bitias 0) 0) | 0) | | 3 2 | 0) 0) 0) 
Panthiades hebraeus 0) 10) | 0) | | 3 2; | 0 O 0) 
Panthiades aeolus 0) 0 0) ? 0) | | 0) | 0) 0) O 
Panthiades boreas 2 0) | | 0) 0 3 2 | 0) 0) 0) 
Panthiades ochus 0) 0 | | 0) 0) 3 2 | 0 0) O 
Panthiades paphlagon 0) 0) | | 0) 0) 3 2 ] 0) O 0) 
Panthiades bathildis Z l l 0) 0) | 2 | 0) 0) 0 
Panthiades phaleros 2 | | 0) O | 2 | ] ] 0) 0) 
Parrhasius polibetes | 0) 0) | ? | 0) 7 0) 2 | | 
Thepytus epytus 1 0) 0) 0) 0) | O 0) ] D, 1 1 
Porthecla porthura | 0) O O 2B | 0) 0) O 2 1 l 


thiades genitalia have been illustrated (Nic- 
olay 1976) except for the female of P. bo- 
reas (Felder and Felder), which is figured 
in this paper. Genitalic terms follow those 
in Klots (1970), as illustrated in Robbins 
and Nicolay (2002). Androconial terminol- 
ogy follows Robbins (1991), and wing vein 
names follow Nicolay (1971, 1977). 

The terminal taxa are the eight species 
that have been placed in Panthiades or Cyc- 
nus (Nicolay 1976, Robbins 2004b). They 
are: P. bitias (Cramer, 1777), P. hebraeus 
(Hewitson, 1867), P. aeolus (Fabricius, 
1775), P. boreas (C. Felder and R. Felder, 
1865), P. ochus (Godman and Salvin, 
1887), P. paphlagon (C. Felder and R. 
Felder, 1865), P. bathildis (C. Felder and 
R. Felder, 1865), and P. phaleros (Linnae- 
us, 1767). These taxa can be identified us- 
ing the key in Nicolay (1976). The adults 
were illustrated in D’ Abrera (1995). 

One outgroup for the analysis is Par- 
rhasius polibetes (Cramer), the type species 
of Parrhasius Hiibner, a genus that has been 
considered to be congeneric with Panthia- 
des (Clench 1961) or its closest relative 
(Nicolay 1976). To test the robustness of 
the root, I also used as outgroups the type 
species of Thepytus Robbins, T. epytus 
(Godman and Salvin), and of Porthecla 
Robbins, P. porthura (H.H. Druce). Among 
the genera of the Panthiades Section (Rob- 
bins 2004a,b), their male genitalia, espe- 


cially the valvae, are phenetically most sim- 
ilar to those of Panthiades. 

The character state for each Panthiades 
and outgroup species is listed in the char- 
acter matrix (Table 1). I used the implicit 
enumeration option of Hennig86 software 
to derive a most parsimonious cladogram. 
A strict consensus tree was determined. To 
test the assumption of equally weighted 
characters, successive weighting was per- 
formed (Farris 1969) and a consensus of the 
resulting trees was determined. All charac- 
ters were unordered, except for multi-state 
character 7. Because it forms a morphocline 
(cf., Pogue and Mickevich 1990), it was an- 
alyzed ordered and unordered. Mapping of 
characters on trees was done with Winclada 
software (Nixon 2002) with the fast opti- 
mization option. Jackknife support was de- 
termined in Winclada using Nona (1000 
replications with mult*10, memory 1000 
trees). 


MORPHOLOGY AND CODED CHARACTERS 


Ventral wing pattern.—The ventral wing 
pattern in Panthiades is highly variable. 
However, determining homology among the 
different wing pattern elements is difficult 
because these elements are not recognizable 
in some species. Further, I am interested in 
using the resulting cladogram to examine 
wing pattern evolution within Panthiades. 
For these reasons, the only character coded 


VOLUME 107, NUMBER 3 


has one state in Panthiades and a second 
state in all other Panthiades Section genera, 
so it does not affect the inferred phyloge- 
netic relations within the genus. 

Character 1: Ventral hindwing postme- 
dian line segment in cell Sc+R1-Rs (0) co- 
linear with remainder of postmedian line, 
(1) basally displaced. In three Panthiades 
species, the postmedian line is not recog- 
nizable, so these species were coded with a 
question mark. 

Dorsal wing pattern.—Character 2: Dor- 
sal wings of female with (0) shining blue- 
green iridescence, (1) a varying amount of 
dull, “chalky” blue scales. The second 
character state is restricted in Panthiades to 
P. bathildis and P. phaleros, where it varies 
within each species from no blue to a dull 
blue sheen that is distinguishable from the 
shining iridescent blue-green of the other 
species. 

Androconia.—Androconia in Panthiades 
are restricted to the dorsal forewing and are 
composed of three parts. The first is a black 
(rarely gray or tan) scent pad in the discal 
cell surrounded by a conspicuous ring of 
scales (Figs. 1-2, also fig. 122 in Eliot 
1973), which are usually gray in color. 
These androconia and the surrounding ring 
of scales are tightly attached to the wing 
membrane. They occur in all Panthiades 
except P. aeolus (Fig. 3), but nowhere else 
in the Eumaeini. 

The second part is a gray to dark char- 
coal colored scent pad that is universal in 
the Panthiades Section. It is distal of the 
ring of scales (Figs. 1—2, 4) and is, in turn, 
composed of two parts. The first is very 
roughly oval and usually covers the upper 
and middle disco-cellular veins and parts of 
the wings basal and distal of these veins 
(Fig. 2). There is sometimes a second part 
at the base of vein M3 (most conspicuous 
in P. aeolus, Fig. 3, arrow C), but its pres- 
ence is intraspecifically variable in some 
species, such as P. bitias, for which reason 
it is not coded. 

The third part is a patch of black scales, 
usually on the distal half of the wings (Fig. 


503 


4). It is unclear whether these scales form 
a scent patch, but I tentatively treat them as 
androconia because they are restricted to 
males. They occur in three species of Pan- 
thiades, one Thepytus, and two Parrhasius 
(Nicolay 1979). The size and shape of the 
black scales often varies geographically, at 
least in Panthiades and Parrhasius (Nico- 
lay 1976, 1979). There is a patch of dark 
brown scales in P. aeolus between the scent 
pads on the disco-cellular veins and at the 
base of vein M3 (Fig. 3, arrow B) that could 
possibly be homologous with the black 
scales, for which reason I code the second 
character below with a question mark for P. 
aeolus. 

Character 3: Scent pad in the discal cell 
surrounded by a ring of scales that are tight- 
ly attached to the wing membrane (Q) ab- 
sent (Fig. 3), (1) present (Fig. 4). 

Character 4: Black scent patch distal of 
the discal cell (0) absent, (1) present (Fig. 
4, letter B). 

Male genitalia——As noted by Nicolay 
(1976), the genitalia of Panthiades are more 
interspecifically variable than those of 
many other eumaeine genera. Five charac- 
ters are coded. 

Character 5: Number of cornuti in penis 
(O) 1, (1) 2. Cornuti are usually easily 
scored in eumaeines, including Panthiades. 
However, outgroup genera Parrhasius and 
Porthecla are problematic in that the folded 
vesica within the penis has patches of vary- 
ing sclerotization, making it difficult to de- 
termine what is and is not a cornutus. De- 
spite this problem in Parrhasius and Por- 
thecla, for which reason they are coded 
with question marks, the two states within 
Panthiades are clear. 

Character 6: Gnathos tips (QO) flared, (1) 
not flared. Modification of the gnathos is 
unusual in the Panthiades Section, and the 
first state was illustrated by Nicolay (1976). 
The gnathos of P. aeolus have a laminate 
carina (sensu Field 1967) at the elbow, but 
it is the only species in the Panthiades Sec- 
tion with such a clearly developed carina 


504 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-4. Dorsal forewing androconia. 1, Panthiades phaleros, arrow A points to ring of scales around one 
scent pad (Character 3), arrow B to second scent pad distal of ring of scales. 2, P. phaleros wing slide showing 
scent pads in relation to wing veins, arrows A and B as in previous figure. 3, P. aeolus, arrow A points to scent 
pad covering the upper disco-cellular veins, arrow B points to dark scales that may be androconia, arrow C to 
scent pad at base of vein M3. 4, P. ochus, letter B is in middle of dark scales that may be androconia (Char- 
acter 4). 


on the elbow, for which reason it was not 
coded. 

Character 7: Ventral surface of valvae 
fused anteriorly (0) O-30% of length, (1) 
30-50% of length, (2) 50-70% of length, 
(3) 70—100% of length. Nicolay (1976) first 
reported the fused valvae in Panthiades 
(Figs. 5—7). 

Character 8: Shape of valva in ventral 


aspect (OQ) roughly triangular, but more than 
twice as long as wide (Fig. 5), (1) duplex 
with a well-developed internal ridge de- 
marcating the two parts (Fig. 6), (2) roughly 
an equilateral triangle (Fig. 7). Because the 
valva of outgroup P. polibetes is so differ- 
ent in shape from that in Panthiades (cf. fig. 
2 in Nicolay 1979), it is coded with a ques- 
tion mark. 


VOLUME 107, NUMBER 3 


Figs. 5-7. 


505 


| eh ; | —_ 
Ww { } ) 5 J 
\ WSS i Tiff 
2 cae fi Y 
eS ae 
\..\ y) 
\\ \ / 
\ \\ i 
\ fy 


Male genitalia saccus and valvae in ventral aspect (anterior is down, digitized from Nicolay 


1976). 5, Panthiades aeolus, each valva is longer than wide. 6, P. phaleros, each valva is duplex, separated by 
a well-developed internal ridge (arrow). 7, P. boreas, each valva is roughly an equilateral triangle. 


Character 9: Ventral of the notch where 
the labides meet, the length of the presumed 
remnant uncus is (0) <0.05mm, (1) 
~0O.l1mm. So far as I am aware, the second 
character state only occurs in Panthiades 
and Thepytus. 


Female genitalia— Character 10: Signa 
(O) skillet-shaped (Fig. 8), (1) rectangular 
and narrow (fig. 22 in Nicolay 1976), (2) 
with a single central spine (fig. 3 in Nicolay 
1979). The skillet-shaped signa (Fig. 8, ter- 
minology from Nicolay 1976) occurs only 


Fig. 8. 
Scale | mm. 


Female genitalia (bursa copulatrix) of Panthiades boreas in lateral (left) and ventral (right) aspects. 


506 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


enytus 
polibetes 
porthura 


(74) [ _ horeas 


9 


Fig. 9. One of two most parsimonious cladograms (character 7 unordered, 17 steps, CI = 0.94, RI = 0.96) 
for Panthiades species. Thepytus epytus, Parrhasius polibetes, and Porthecla porthura are the outgroups. Char- 
acter numbers are placed above nodes and character state numbers below nodes. Open circles represent reversal 
or convergence of the character state at that node. Jackknife support is noted in brackets above each node. This 
cladogram is also the consensus of the two most parsimonious trees. 


VOLUME 107, NUMBER 3 507 


9 


10 


Fig. 10. Most parsimonious cladogram (character 7 ordered, 17 steps, CI = 0.94, RI = 0.96) for Panthiades 
species. Thepytus epytus, Parrhasius polibetes, and Porthecla porthura are the outgroups. Character numbers 
are placed above nodes and character state numbers below nodes. Open circles represent reversal or convergence 
of the character state at that node. This cladogram is also one of the most parsimonious trees when character 7 
is unordered. The ventral wing pattern of each species is placed on the cladogram. The classic “false head” 
wing patterns in P. bathildis and P. phaleros appear to have evolved once in their immediate common ancestor. 


508 


in Panthiades, and the rectangular and nar- 
row signa occurs in P. phaleros (fig. 23 in 
Nicolay 1976). 

Character 11: Lamella postvaginalis in 
ventral aspect (0) sclerotized and fan- 
shaped (Fig. 8), (1) not fan-shaped (Fig. 3 
in Nicolay 1979). The first state occurs in 
Panthiades and not the other genera of the 
Panthiades Section. 

Character 12: Origin of ductus semin- 
alis (0) on the left ventral side of the corpus 
bursae (Fig. 8), (1) on the dorsal side of the 
corpus bursae. In the Panthiades Section, 
the origin is usually on the left side, but in 
Parrhasius, it is very close to the center. 


PHYLOGENETIC ANALYSES AND RESULTS 


I analyzed the coded data (Table 1) using 
the Hennig86 “‘ie*’’ option, which searches 
exhaustively for the most parsimonious 
cladograms. The analysis with character 7 
unordered yielded two equally parsimoni- 
ous 17-step trees with a consistency index 
of 0.94 and retention index of 0.96. The 
first (Fig. 9) is also the strict consensus tree. 
The second (Fig. 10) is the only most par- 
simonious tree when character 7 is treated 
as ordered. Successive weighting did not 
change the cladogram topology. Jackknife 
support values are reported for the consen- 
sus tree (Fig. 9) and were slightly lower 
than those for the other most parsimonious 
tree with character 7 ordered. In both of the 
most parsimonious trees, Panthiades is par- 
titioned into four monophyletic groups; P. 
aeolus, a lineage of P. phaleros and P. 
bathildis, a lineage of P. bitias and P. he- 
braeus, and a lineage of P. ochus, P. paph- 
lagon, and P. boreas. 


DISCUSSION 


Synapomorphies for Panthiades (Figs. 
9-10) are (1) ventral hindwing postmedian 
line segment in cell Sc+RI1-Rs co-linear 
with remainder of postmedian line, (2) ven- 
tral surface of valvae fused at their anterior 
base more than 30% of their length, (3) 
skillet shaped signa, (4) lamella postvagin- 
alis in ventral aspect sclerotized and fan- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


shaped, and (5) ductus seminalis arises on 
the left ventral side of the corpus bursae. 
Except for the first, these characters had 
been explicitly noted by Nicolay (1976). If 
P. phaleros is moved from Panthiades to 
Cycnus, then Panthiades is not monophy- 
letic on either of the most parsimonious 
trees (Figs. 9, 10), whether character as 
ordered or unordered. These results support 
the classification in Robbins (2004b). 

Panthiades bathildis and P. phaleros 
have wing patterns that have traditionally 
been associated with the “‘false head” hy- 
pothesis (Robbins 1980, 1981). The phy- 
logenetic results suggest that this wing pat- 
tern evolved in the ancestor of the two spe- 
cies (Fig. 10). In Nicolay’s (1976) classifi- 
cation, this result was not evident. 


ACKNOWLEDGMENTS 


I am grateful to Stan Nicolay for more 
than I can ever reasonably acknowledge, es- 
pecially sharing for three decades his broad 
knowledge of the Neotropics and its eu- 
maeine fauna. Specifically for this paper, he 
allowed me to scan digitally his genitalia 
drawings. For drawing the female genitalia 
of P. boreas, 1 thank Vichai Malikul. For 
making suggestions on the manuscript, I 
thank Marcelo Duarte, Gerardo Lamas, Tia- 
go Quental, John Shuey, and Andy Warren. 


LITERATURE CITED 


Clench, H. K. 1961. Tribe Theclini, pp. 177—220. In 
Ehrlich, P R. and A. H. Ehrlich, How to Know 
the Butterflies, Brown Company, Dubuque, Iowa. 

D’ Abrera, B.L. 1995. Butterflies of the Neotropical re- 
gion. Part VII. Lycaenidae. Hill House, Black 
Rock, pp. I-xi + 1098-1270. 

Eliot, J. N. 1973. The higher classification of the Ly- 
caenidae (Lepidoptera): A tentative arrangement. 
Bulletin of the British Museum (Natural History) 
Entomology 28: 371-505. 

Farris, J. S. 1969. A successive approximations ap- 
proach to character weighting. Systematic Zoolo- 
gy 18: 374-385. 

Field, W. D. 1967. Preliminary revision of butterflies 
of the genus Calycopis Scudder (Lycaenidae: The- 
clinae). Proceedings of the United States National 
Museum 119 (No. 3552), 48 pp. 

Klots, A.B. 1970. Lepidoptera, pp. 115-130. In Tuxen, 


VOLUME 107, NUMBER 3 


S. L. ed., Taxonomist’s Glossary of Genitalia in 
Insects. Munksgaard, Copenhagen. 

Nicolay, S. S. 1971. A new genus of hairstreak from 
Central and South America (Lycaenidae, Thecli- 
nae). Journal of the Lepidopterists’ Society 25, 
Supplement 1, 39 pp. 

. 1976. A review of the Htibnerian genera Pan- 

thiades and Cycnus (Lycaenidae: Eumaeini). Bul- 

letin of the Allyn Museum 35, 30 pp. 

. 1977. Studies in the genera of American hair- 

streaks. 4. A new genus of hairstreak from Central 

and South America (Lycaenidae: Eumaeini). Bul- 
letin of the Allyn Museum 44, 24 pp. 

. Studies in the genera of American hairstreaks. 
5. A review of the Hubnerian genus Parrhasius 
and description of a new genus Michaelus (Ly- 
caenidae: Eumaeini). Bulletin of the Allyn Mu- 
seum 56, 52 pp. 

Nixon, K. C. 2002. WinClada version 1.00.08. Pub- 
lished by the author, Ithaca, NY. 

Pogue, M. G. and M. FE Mickevich. 1990. Character 
definitions and character state delineation: the béte 
noire of phylogenetic inference. Cladistics 6: 319— 
361. 

Robbins, R. K. 1980. The lycaenid “‘false head’’ hy- 
pothesis: Historical review and quantitative anal- 


509 


ysis. Journal of the Lepidopterists’ Society 34: 

194-208. 

. 1981. The ‘‘false head”’ hypothesis: Predation 

and wing pattern variation of lycaenid butterflies. 

American Naturalist 118: 770-775. 

1991. Evolution, comparative morphology, 

and identification of the eumaeine butterfly genus 

Rekoa Kaye (Lycaenidae: Theclinae). Smithsoni- 

an Contributions to Zoology No. 498, 64 pp. 

. 2004a.Introduction to the checklist of Euma- 

eini (Lycaenidae), pp. xxiv—xxx. Jn Lamas, G. ed. 

Checklist: Part 4A. Hesperioidea—Papilionoidea. 

In Heppner, J. B. ed. Atlas of Neotropical Lepi- 

doptera, Vol. 5A. Association for Tropical Lepi- 

doptera, Scientific Publishers, Gainesville, xxxvi 

+ 439 pp. 

. 2004b. Lycaenidae. Theclinae. Tribe Euma- 
eini, pp. 118-137. In Lamas, G. ed. Checklist: 
Part 4A. Hesperioidea—Papilionoidea. Jn He- 
ppner, J. B. ed. Atlas of Neotropical Lepidoptera, 
Vol. 5A. Association for Tropical Lepidoptera, 
Scientific Publishers, Gainesville, xxxvi + 439 pp. 

Robbins, R. K. and S. S. Nicolay. 2002. An overview 
of Strymon Hiibner (Lycaenidae: Theclinae: Eu- 
maeini). Journal of the Lepidopterists’ Society 55: 
85-100. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 510-516 


A NEW GENUS OF THE SUBFAMILY CUBACUBANINAE 
(ZYGENTOMA: NICOLETIIDAE) FROM VERACRUZ, MEXICO 


Luis ESPINASA 


Natural Sciences Department, Shenandoah University, 1460 University Drive, 
Winchester, VA 22601, U.S.A. (e-mail: lespinas @su.edu) 


Abstract.—Acanthonima veracruzi, n. gen., n. sp., is described and separated from 
other species of the subfamily Cubacubaninae. It was collected from under rocks by the 
side of the Actopan River in Veracruz, Mexico. 


Key Words: 


While reviewing the Zygentoma collec- 
tion of the American Museum of Natural 
History, eight nicoletiids were found in a 
vial with ethanol labeled as *“‘Puente Acto- 
pan, 5 km SE Actopan, Veracruz, Mexico. 
25 Dec. 1976. J. Reddell, A. Grubbs.’’ One 
individual was an undescribed species of 
Squamigera Espinasa (1999a), another in- 
dividual belonged to the Atelurinae, and the 
six remaining individuals belonged to the 
subfamily Cubacubaninae, but these could 
not be assigned to any previously described 
genus. Dissections of a male and a female 
of this last group were made with aid of a 
stereo microscope and mounted in fixed 
preparations with Hoyer’s solution. The 
four remaining samples were left in a vial 
with ethanol. [Illustrations were made with 
aid of a camera lucida attached to a com- 
pound microscope. Here I describe and 
name a new genus and species. Types are 
deposited in the Zygentoma collection of 
the American Museum of Natural History, 
ING, INLYG 


Acanthonima Espinasa, new genus 


Diagnosis.—A member of the subfamily 
Cubacubaninae without scales. Cerci and 
median filament of mature males with 
spines. 


Thysanura, Zygentoma, Nicoletiidae, Cubacubaninae, Actopan 


Description.—Pedicellus of adult male 
with unicellular glands. Mouthparts not 
specialized. Mandible strongly sclerotized 
apically with usual teeth. Galea apically 
with sensory pegs. Lacinia heavily sclero- 
tized distally. First process of lacinia pec- 
tinate. Labium without prominent lateral 
lobes. 

Tarsi with four articles. Praetarsi with 
three simple claws. Median claw glabrous, 
slender and smaller than lateral claws. 
Urosterna II—VII subdivided into two cox- 
ites and one sternite. Urosterna VIII and IX 
of male entire. Median portion of sternites 
with | + 1 sublateral macrochaetae at hind 
borders, as well as | + 1 macrochoetae near 
suture at about middle of segment. Coxites 
on segments II-IX with stylets. Eversible 
vesicles on segments II-VI, pseudovesicles 
on VII. Urosterna III and IV of adult males 
apparently without modifications. Urosterna 
VII with a wide and not too deep posterior 
emargination. Tergum X with several sub- 
equal macrochaetae on posterior angles. 

Point of insertion of parameres apparent- 
ly shghtly deep and with modified setae on 
internal face of coxal processes. Parameres 
normal for subfamily; specialized setae on 
apex, but otherwise not subdivided nor 
somewhat constricted apically. Stylets IX 


VOLUME 107, NUMBER 3 


AWA 


% PN 


se hi area 


thee 
i i 
Se 
= 
— 
A 
= 
. sn 
VE ~S 
LEZ en 
VEZE fil a \ 
= Se 
= Vhs g 
= ah vet 
= RA 
H 


Wwe 
Lem \ We 
Mall Le 


a 


~~ / 
| Se 


Se 

Fig. 1. Acanthonima veracruzi, male holotype. Microchaetae partially shown. A, Body. B, Head and anten- 
nae. C, Pedicellus. D, Setae on border of insertion of antennae. E, Labial palp and labium. K Maxilla. G, Apex 
of maxilla. H, Mandible. I, Thoracic terga. 


512 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


apparently without spines. Opening of penis 
longitudinal. Cercus of male with sensory 
pegs, some may even be arranged in two 
rows. Appendix dorsalis also with sensory 
pegs. Female with a subgenital plate. 

Type species.—Acanthonima veracruzi, 
NSP: 

Etymology.—Akantha = spine, nima = 
filament (in nominative feminine singular). 
In reference to the spines or pegs along the 
appendix dorsalis, also known as middle 
terminal filament. 

Remarks.—Acanthonima belongs to the 
Cubacubaninae (Mendes 1988), character- 
ized by subdivided abdominal sterna II-VI 
and fused coxites of abdominal segments 
VIET and IX. Acanthonima is distinguished 
from almost all genera of this subfamily by 
having sensory pegs on the appendix dor- 
salis. Appendix dorsalis pegs were previ- 
ously described only in nicoletiids of the 
subfamilies Coletiniinae and Subnicoleti- 
inae (Mendes 1988). The only other speci- 
mens with pegs on the appendix dorsalis 
within the Cubacubaninae belong to an un- 
described Squamigera species from a cave 
in Chiapas, Mexico, and are currently under 
study. The two species can easily be differ- 
entiated because the Squamigera new spe- 
cies has a longer and more subdivided ovi- 
positor and a prominent downward pointing 
robust spine in the pedicellus, which is ab- 
sent in Acanthonima veracruzi. 

The genus Acanthonima can further be 
differentiated from Texoreddellia Wygod- 
zinsky (1973) and Squamigera by the ab- 
sence of scales; from Allonicoletia Mendes 
(1992) by the presence of stylets on uro- 
sternite I]; from Neonicoletia Paclt (1979) 
by the aspect of the endopodium; from 
Prosthecina Silvestri (1933) by the absence 
of conspicuous lateral lobes bearing numer- 
ous glandular pores in the submentum; 
from most Anelpistina Silvestri (1905) by 
the absence of articulated submedian ap- 
pendages in urosternite [V of males; from 
the Anelpistina Espinasa (1999b) without 
the appendages and from Cubacubana Wy- 
godzinsky and Hollinger (1977) by the sen- 


sory pegs of the appendix dorsalis. Pegs in 
cerci in the last two genera are also ar- 
ranged in a single row, while, at least in this 
new species, they are arranged in two rows 
at times. 

The pedicellus of the adult male in Acan- 
thonima veracruzi is also unique for the 
subfamily in having a cluster of unicellular 
glands on a basal bulge on the outer border 
of the antennae (Figs. 1B, C). Regrettably, 
this character is variable along the post- 
embryonic development and is absent in ju- 
veniles. Also, pedicellus modifications are 
quite variable among the genera. Until more 
Acanthonima species are described, it is un- 
known if this character will be diagnostic 
for the genus or if it only represents species 
variation within the genus. 


Acanthonima veracruzi Espinasa, 
new species 
(Figs. 1A—I, 2A--K 3A—F) 


Type material—Puente Actopan, 5 km 
SE Actopan, Veracruz, Mexico. 25 Dec. 
1976. J. Reddell, A. Grubbs cols. Male ho- 
lotype, 3 male paratypes, | adult female and 
1 juvenile female paratypes. 

Description.—Maximum body length of 
samples 9.5 mm. Maximum conserved 
length of antenna and caudal appendages 
6.0 mm and 5.0 mm. Body proportions as 
in Fig. 1A. General color light yellow to 
white. Basal articles of antenna of female 
simple. Pedicellus of male equal in length 
to first article and with five clusters of uni- 
cellular glands. Four ventral clusters bor- 
dered with a not too conspicuous row of 
microchaetae forming a “U.” Last cluster 
on basal bulge on outer border (Figs. 1B, 
C). Head with macrochaetae and micro- 
chaetae as shown in Figs. 1B and D, very 
abundant on border of insertion antennae. 
Mouthpart appendages relatively short, la- 
bial palp as in Fig. 1E, apical article barely 
longer than wide and barely longer than 
next to last article. Penultimate article with 
bulge containing two macrochaetae. Labi- 
um and first article of labial palp with ma- 
crochaetae. Maxilla as shown in Fig. 1F 


VOLUME 107, NUMBER 3 S15) 


Fig. 2. Acanthonima veracruzi, male holotype. Microchaetae partially shown. A, Third leg. B, Second leg. 
C, Apex of tibia. D, Urosterna I-III. E, Urosterna VII-IX. F Penis and coxal processes. 


Last article % longer than penultimate. totaxy as in Fig. 1H, with many macro- 
Apex of galea with two conules, one longer _ chaetae, although only four bifid. 

than wide and other wider than long (Fig. Pro-, meso-, and metanota with approx- 
1G). Two teeth on lacinia. Mandible chae- imately 3 macrochaetae slightly inside of 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


4mm \ 
TA n 
PUSS > 1p 71 77 LO \)s Z i () \, 
Po) gee es RRS AN XS 
hie hi Aleeg \ \ 
NG Lg Li : WR Wa \ 
A THN a I 
{ RE 1a 
LA 
TRAN 3 iS 
Sy : 
he nec 
INT BT 
Mad Br 


ee 
ae 


S 
| 
y 
3 


AN 
| 
TTTTTIN 
\ 
ad 
ee 
ESBS 


= 
a 
] 


= 


So VALE, 
ae 


SS” 
x 


w 


1) 


Js 


\ 
Loy 
a 


4 
r 
meee 


cq 


aQ 
SW 


"7 


Fig. 3. Acanthonima veracruzi, A—D, Male holotype. E, Female paratype. EK Juvenile female paratype. Mi- 


crochaetae partially shown. A, Urotergum X. B, Caudal appendages. C, Sensory pegs on cercus. Notice the 


double row of spines. D, Sensory pegs on appendix dorsalis. E, Subgenital plate and ovipositor. Tip of ovipositor 
broken. EK Subgenital plate and ovipositor in immature female. 


VOLUME 107, NUMBER 3 


lateral borders, apart from several setae of 
varied sizes at edge of borders (Fig. 11). 
Legs of medium size, hind tibia approxi- 
mately 5x longer than wide and ¥, shorter 
than tarsus (Fig. 2A). Tibia of second leg 
somewhat stouter (4x longer than wide) 
with 4 macrochaetae (Fig. 2B). Claws of 
normal size. 

Abdominal sterna as in Figs. 2D, E. 
Urosternum III and IV without modifica- 
tions. Urosternum VIII of male very slight- 
ly emarginated, its projections far apart, 
rounded and small (Fig. 2E). Urosternum 
IX of male as in Fig. 2E. Point of insertion 
of parameres in urosternum IX slightly be- 
low level of base of the stylets of this seg- 
ment. Base of internal faces of coxal pro- 
cesses with longer and slightly sclerotized 
macrochaetae (Fig. 2F). Penis and paramer- 
es as in Figs. 2E, EK Parameres attaining al- 
most % length of stylets IX. 

Stylets IX larger than others, without 
sensory cones, and with two macrochaetae 
and an extra subapical pair. Terminal spine 
with small teeth. Urotergite X shallowly 
emarginate in both sexes, posterior angles 
with several macrochaetae and a few rela- 
tively strong setae (Fig. 3A). Length of in- 
ner macrochaetae about half distance be- 
tween them. 

Cercus of adult male straight, with a lon- 
ger than wide basal article, sometimes fol- 
lowed by one or two articles wider than 
long, then a very long article, followed by 
numerous short articles. Sensory pegs of 
subequal size on very long article and on 
one or two of short articles (Fig. 3B). Some 
pegs arranged in double rows (Fig. 3C). 
Appendix dorsalis with sensory pegs (Figs. 
3B, D). Female cercus and appendix dor- 
salis simple. 

Subgenital plate of female rounded (Figs. 
3E, F). Ovipositor in single adult female 
surpassing apex of stylets IX by slightly 
less than 3X length of stylets (Fig. 3E), but 
in this specimen ovipositor’s tip broken and 
its left side partially deformed. Gonapo- 
physes with slightly more than 21 articles 
(last few articles missing). 


SS) 


Etymology.—The name veracruzi (in 
genitive singular) refers to the state of Ve- 
racruz, Mexico, where the samples were 
collected. 

Remarks.—The male postembryonic de- 
velopment is mostly unknown because 
samples ranged from 6.8 mm to 9 mm. The 
only difference in the two individuals 
smaller than 7.3 mm from the bigger indi- 
viduals is that the bulge in the pedicellus is 
absent. The length of the parameres and 
sensory pegs on the caudal appendages re- 
main the same. As for the females, only two 
were available for study, one measuring 8 
mm and the other 5.2 mm. In the smaller 
female, the ovipositor barely surpasses the 
apex of the stylets IX (Fig. 3F). 

The limits of distribution are currently 
unknown for the species, although as a gen- 
erality, members of the subfamily Cubacu- 
baninae are restricted to the Neotropics and 
each species is known from a single local- 
ity. It is likely that the species is limited to 
the State of Veracruz. The only individuals 
found to date are those collected by Reddell 
and Grubbs in 1976 in Actopan, Veracruz, 
Mexico. An extensive search on 11/19/01 
in the type locality provided many nicole- 
tiid specimens, most of them Anelpistina, 
but no new specimens of Acanthonima ver- 
acruzi could be found. The reason for their 
absence, is currently unknown. 


ACKNOWLEDGMENTS 


I thank Randall T. Schuh, George Willett 
Curator and Chair of the Division of Inver- 
tebrate Zoology of the American Museum 
of Natural History, for kindly offering the 
material in the collection for study. Thanks 
to Graeme Smith and Luis EF Mendes for 
reviewing the manuscript, and to Oscar Do- 
rado and Dulce Arias, both directors of 
CEAMISH, Universidad Autonoma del Es- 
tado de Morelos (UAEM), for their support 
of the study. Work was done while on sab- 
batical leave from UAEM, in facilities of 
the American Museum of Natural History 
and Shenandoah University. 


516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


LITERATURE CITED 


Espinasa, L. 1999a. A new genus of the subfamily Cu- 
bacubaninae (Insecta: Zygentoma: Nicoletiidae) 
from a Mexican cave. Proceedings of the Biolog- 
ical Society of Washington 112(1): 52-58. 

. 1999b. Two new species of the genus Anel- 
pistina (Insecta: Zygentoma: Nicoletiidae) from 
Mexican caves, with redescription of the genus. 
Proceedings of the Biological Society of Wash- 
ington 112(1): 59-69. 

Mendes, L. FE 1988. Sur deux nouvelles Nicoletiidae 
(Zygentoma) cavernicoles de Gréce et de Turquie 
et remarques sur la systématique de la famille. Re- 
vue Suisse de Zoologie 95(3): 751-772. 

. 1992. Novos dados sobre os tisanuros (Micro- 
coryphia e Zygentoma) da América do Norte. 
Garcia de Orta Sér. Zool. 16(1—2): 171-193. 

Paclt, J. 1979. Neue Beitrage zur Kenntnis der Apter- 
ygoten-Sammlung des Zoologischen Instituts und 
Zoologischen Museums der Universitat Hamburg. 


VI. Weitere Doppel- und Bortenschwanze (Diplu- 
ra: Campodeida: Thysanura: Lepismatidae und 
Nicoletiidae). Entomologische Mitteilungen aus 
dem zoologischen Museum Hamburg 6(105): 
221-228. 

Silvestri, E 1905. Materiali per lo studio dei Tisanuri. 
VI. Tre nuove specie di Nicoletia appartenenti ad 
un nuovo sottogenero. Redia (Firenze) 2: 111— 
120. 

. 1933. Nuovo contributo alla conoscenza dei 
Tisanuri del Messico. Bolletino del Laboratorio di 
Zoologia general e agraria di Portici 27: 127-144. 

Wygodzinsky, P. 1973. Description of a new genus of 
cave Thysanuran from Texas (Nicoletiidae, Thy- 
sanura, Insecta). American Museum Novitates 
2518: 1-8. 

Wygodzinsky, P. and A. M. Hollinger. 1977. A study 
of Nicoletiidae from Cuba (Thysanura). Résultats 
des Expéditions Biospéleologiques Cubano-Rou- 
maines a Cuba 2: 317-324. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 517-529 


CORNETIUS, A NEW SUBGENUS OF AEDES, AND A REDESCRIPTION OF 
AEDES (CORNETIUS) COZI CORNET (DIPTERA: CULICIDAE) 


YTAU-MIN HUANG 


Department of Entomology, MRC 534, Smithsonian Institution, PO. Box 37012, 
Washington, DC 20013-7012, U.S.A. (e-mail: huang.yiau-min@nmnh.si.edu) 


Abstract.—Cornetius, a new subgenus of Aedes Meigen is characterized and diag- 
nosed. Aedes cozi Cornet is removed from the subgenus Stegomyia Theobald and placed 
in the new monotypic subgenus Cornetius on the basis of a critical study of all known 
stages. The adult of both sexes, the pupa, and the larva of Ae. (Cornetius) cozi are 
described and illustrated. Its affinity to other subgenera of the genus Aedes is discussed. 
Information on type data, distribution, bionomics, and medical importance, and a taxo- 


nomic discussion of this species are presented. 


Key Words: 
Senegal 


Cornet (1973: 175) described Aedes cozi 
from specimens collected in eastern Sene- 
gal (Kedougou) and placed it in the sub- 
genus Stegomyia Theobald. After critical 
study of both adults and the immature stag- 
es, it is apparent that Ae. cozi should not be 
in Stegomyia, as | noted previously (Huang 
2001, 2002). This paper formally deals with 
the suggested taxonomic change for Ae. 
cozi with description herein of Cornetius, a 
new monotypic subgenus of the genus Ae- 
des Meigen. The new subgenus is very dis- 
tinct from other subgenera of the genus Ae- 
des in all known stages. The adult of both 
sexes, the pupa, and the larva are here de- 
scribed and illustrated. Keys for the iden- 
tification of this species were published pre- 
viously (Huang 2001, 2002). Information 
on the type data, distribution, bionomics, 
medical importance, and a taxonomic dis- 
cussion of this species are presented. The 
suggested abbreviation for the subgenus 
Cornetius is Cor. 


MATERIALS AND METHODS 


This study is based on specimens in the 
Department of Entomology, National Mu- 


Cornetius, new subgenus, Aedes cozi, characteristics, systematics, Culicidae, 


seum of Natural History, Smithsonian In- 
stitution (USNM). Other specimens .were 
borrowed from individuals and institutions 
mentioned in the acknowledgments. 

The terminology follows Harbach and 
Knight (1980, 1982) with the exception of 
“tarsal claws,’ which is retained for “‘un- 
gues.” The wing venational terms follow 
those of Belkin (1962). An asterisk (*) fol- 
lowing the abbreviation used (M = male, F 
= female, P = pupa and L = larva) indi- 
cates that all or some portion of that sex or 
stage is illustrated. 


Genus Aedes Meigen 
Cornetius Huang, new subgenus 


Type species: Aedes cozi Cornet, 1973, by 
present designation 
Aedes (Stegomyia) of Cornet 1973: 175. 


Characteristics —The subgenus Corne- 
tius is characterized by the following com- 
bination of characters: Adult (both sexes): 
(1) Vertex with all broad, flat decumbent 
scales, erect forked scales numerous, not re- 
stricted to occiput; (2) maxillary palpus of 


518 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


male about as long as proboscis, 5-seg- 
mented, dark, with a white band at base of 
palpomeres 2—5, those on palpomeres 4,5 
incomplete ventrally; palpomeres 4,5 very 
short (total length of palpomeres 4 and 5 
shorter than palpomere 3), somewhat down- 
turned with a few short setae; palpomere 4 
slightly swollen, with a few short stiff setae 
at apex; palpomere 5 shorter than palpo- 
mere 4, somewhat narrowed toward tip, 
downturned; maxillary palpus of female 
about 0.25 length of proboscis, dark, with 
white band at base of palpomeres 2 and 3, 
palpomere 4 minute, with all dark scales; 
(3) pedicel with long patch of white broad, 
flat scales on mesal surface; (4) acrostichal 
setae absent; (5) paratergite with broad 
white scales; (6) postspiracular setae pre- 
sent; (7) lower prealar scale-patch absent; 
(8) subspiracular area with broad white 
scales; (9) postprocoxal membrane without 
scales; (10) lower mesepimeral setae pre- 
sent; (11) scutellum with all broad scales 
and with broad white scales on all lobes; 
(12) hindtarsus with a basal white band at 
least on tarsomeres 1—3. Male Genitalia; 
(13) Aedeagus strongly toothed; (14) clas- 
pette well developed, with numerous setae; 
(15) gonostylus elongate, rather flat, slight- 
ly expanded just beyond middle, narrow 
and curved toward tip, with a blunt, stout 
gonostylar claw apically on inner margin of 
slightly expanded subapical part; (16) par- 
aproct with a sternal arm; cercal setae ab- 
sent; (17) sternum IX with setae. Female 
genitalia: (18) Insula as long as or slightly 
longer than broad, with tuberculi, without 
setae; upper vaginal sclerite present; lower 
vaginal sclerite absent; (19) cerci short and 
broad; (20) 3 spermathecae, one slightly 
larger than other 2. Pupa: (21) Seta 6-CT 
single, long, stout, much longer and stouter 
than 7-CT; (22) seta 9-VI,VII single, long, 
stout, much longer and stouter than 9-I-V; 
(23) paddle margins with fringe of very fine 
hairlike spicules; apex notched; seta 1-P 
single, long. Larva: (24) Seta 4-C well de- 
veloped, single, simple, closer to 6-C than 
5-C, cephalad and mesad of 6-C; (25) seta 


————. 1.0mm 
Aedes (Cornetius) cozi Cornet 


1.0mm 4 


Aedes (Stegomyia) aegypti (Linnaeus) 


c 1.0mm 4 


Aedes (Diceromyia) furcifer (Edwards) 


Fig. 1. Maxillary palpus and proboscis of males, 
lateral view. A, Aedes (Cornetius) cozi. B, Aedes 
(Stegomyia) aegypti. C, Aedes (Diceromyia) furcifer. 


12-I absent; (26) seta 2-VIII distant from 1- 
VIII; (27) comb scales in a single row; (28) 
ventral brush (seta 4-X) with 5 pairs of se- 
tae on grid; (29) without precratal tufts; 
(30) seta 1-S single, long, inserted distal 
and dorsad of pecten spines. This combi- 
nation of characters distinguishes Cornetius 
from all other subgenera of Aedes. 
Systematics.—Aedes cozi 1s very distinc- 


VOLUME 107, NUMBER 3 


tive and has several unique features in both 
adults and immatures, as noted above, but 
it also shows a number of similarities with 
both Diceromyia Theobald and Stegomyia. 
The male genitalia, pupa, and larva of Ae. 
cozi are so strikingly different from all the 
known species in these two subgenera, as 
well as from other subgenera of Aedes, that 
I believe a distinct subgenus should be rec- 
ognized for this species. 

The subgenus Cornetius possesses some 
rather important basic characters in com- 
mon with the subgenera Aedimorphus 
Theobald, Albuginosus Reinert, Diceromyia 
and Stegomyia of the genus Aedes: male 
maxillary palpus 5-segmented, aedeagus 
with conspicuous teeth, claspette devel- 
oped, female insula longer than broad, lar- 
val seta 12-I not developed and pecten 
spines present. These shared characters in- 
dicate the affinity of Cornetius to these four 
subgenera. 

The adult male and female of Cornetius 
are very similar in some characters to those 
of Stegomyia, such as having the vertex 
with all broad, flat decumbent scales and 
the scutellum with all broad scales. It can 
easily be distinguished from those of Stego- 
myia by the vertex with erect forked scales 
numerous, not restricted to occiput. The 
male of Cornetius is very similar to that of 
Stegomyia in having the maxillary palpus 
of the male 5-segmented, dark, with a white 
band at the bases of palpomeres 2—5. It can 
be easily distinguished from Stegomyia by 
the very short palpomeres 4,5 (the total 
length of palpomeres 4 and 5 shorter than 
palpomere 3), somewhat downturned and 
with a few short setae; palpomere 4 slightly 
swollen and with a few short stiff setae at 
apex; palpomere 5 shorter than palpomere 
4, somewhat narrowed toward tip and 
downturned (see Fig. 1A). In Stegomyia, 
the maxillary palpus of the male has long 
palpomeres 4,5 (the total length of palpo- 
meres 4 and 5 longer than palpomere 3), 
subequal in length, slender, upturned, and 
with only a few short setae (see Fig. 1B). 

The male genitalia of Cornetius are very 


519 


similar to those of Stegomyia in having the 
aedeagus strongly toothed and the claspette 
well developed and bearing numerous se- 
tae. They can easily be distinguished from 
those of Stegomyia by the gonostylus, 
which is elongate, rather flat, slightly ex- 
panded just beyond middle, narrow and 
curved toward apex, with a blunt, stout 
gonostylar claw apically on inner margin of 
slightly expanded subapical portion (see 
Fig. 4C), and by the sternum IX with setae 
(Sceunions ©): 

The maxillary palpus of the Cornetius 
male is very similar to that of species in 
subgenus Diceromyia. In Diceromyia, the 
maxillary palpus of the male has very short, 
swollen and downturned palpomeres 4,5 
(the total length of palpomeres 4 and 5 
shorter than palpomere 3), with a few short 
setae; palpomere 4 swollen, with a few 
short stiff setae at apex; and palpomere 5 
much shorter than palpomere 4, or palpo- 
mere 5 minute (see Fig. 1C). 

The adults of Cornetius can be distin- 
guished from those of Diceromyia by hav- 
ing the pedicel with a long patch of white 
broad, flat scales on the mesal surface. The 
male genitalia of Cornetius have the para- 
proct with a sternal arm that is markedly 
different from all known species of Dicer- 
omyia (see Fig. 3A). In Diceromyia, the 
paraproct lacks a sternal arm. The paraproct 
with a sternal arm is shared with many, but 
not all, species of Stegomyia. The female 
genitalia of Cornetius have the insula rather 
short and broad, which is strikingly differ- 
ent from all the known species of Dicero- 
myia (see Fig. 6A). In Diceromyia, the in- 
sula is long and narrow (see Reinert 2000, 
figs. 17-19). 

The pupa of Cornetius has seta 6-CT sin- 
gle, long, stout, much longer and stouter 


than 7-CT, and seta 9-VI,VII, is single, 
long, stout, and much longer and _ stouter 
than 9-I-V, which are strikingly different 


from all the known species of Diceromyia 
(see Figs. 4A, B). In Diceromyia, seta 6-CT 
is single and short, much shorter than 7-CT, 
and seta 9-VI is small, single, and similar 


520 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Aedes (Cornetius) cozi Cornet 


Fig. 2. Aedes (Cornetius) cozi. A, Thorax (dorsal view). B, Fore-. mid- and hindlegs (anterior view) of 
female. C, Fore-, mid- and hindlegs (anterior view) of male. D, Tarsal claws (fore-, mid- and hindlegs) of male. 


VOLUME 107, NUMBER 3 


to 9-I-V (see Huang 1986, figs. 4A, B). The 
larva of Cornetius has seta 4-C single, sim- 
ple, and cephalomesad of 6-C, which is 
strikingly different from all the known spe- 
cies of Diceromyia (see Fig. 5A). The po- 
sition of seta 4-C cephalomesad of 6-C is 
shared with Stegomyia (Huang 1979, 2004). 
In Diceromyia, seta 4-C is well developed, 
and caudomesad of 6-C (see Huang 1986, 
fig. SA). 

Etymology.—The subgeneric name, Cor- 
netius (gender, masculine), honors Dr. 
Michel Cornet, Medical Entomologist, Ser- 
vices Scientifiques Centraux, Office de la 
Recherche Scientifique et Technique Outre- 
Mer (ORSTOM), and Chief, Medical En- 
tomology Laboratory, Institute Pasteur de 
Dakar, Senegal. I am grateful for his kind- 
ness in welcoming me to the Medical En- 
tomology Laboratory and allowing me to 
study the mosquito collections at his Lab- 
oratory in Dakar. The subgeneric patronym 
also recognizes his many contributions to 
our knowledge of the mosquito fauna of 
Africa. 

Distribution.—Presently known only 
from Senegal. 

Bionomics.—Females have been collect- 
ed biting humans in gallery forests in Ke- 
dougou, Senegal. 

Medical importance.—Unknown. 


Aedes (Cornetius) cozi Cornet 
(Figs. 1A, 2—6) 


Aedes (Stegomyia) cozi Cornet 1973: 175 
QM, F*, P¥, L¥). 


Female.—Head: Proboscis entirely dark- 
scaled, about as long as forefemur; maxil- 
lary palpus about 0.25 length of proboscis, 
dark, with a white band at bases of palpo- 
meres 2 and 3, palpomere 4 dark, minute; 
antennal pedicel covered with broad white 
scales on mesal surface; flagellomere 1 with 
small patch of white scales on mesal sur- 
face; clypeus bare; erect forked scales dark, 
not restricted to occiput; vertex with medial 
stripe of broad dark scales and distinct sub- 
medial stripe of broad white scales, with 


521 


broad dark scales on each side interrupted 
by lateral stripe of broad white scales, fol- 
lowed ventrally by patch of broad white 
scales. Thorax (Fig. 2A): Scutum with nar- 
row dark scales, some white scales on an- 
terior promontory and along anterior scutal 
margin, distinct submedian longitudinal 
white line of narrow scales from anterior 
margin to about the level of wing root, 
broad longitudinal line of narrow white 
scales on posterior dorsocentral area, short 
median white line of narrow scales reaching 
prescutellar area; prescutellar line of narrow 
white scales absent, with only a few white 
scales; patch of white scales on lateral mar- 
gin in front of wing root; acrostichal setae 
absent; dorsocentral setae present and well 
developed; scutellum with broad white 
scales on all lobes and a few broad dark 
scales at apex of midlobe; antepronotum 
with broad white scales; postpronotum with 
stripe of broad white scales and some broad 
dark scales dorsally; paratergite with broad 
white scales; prespiracular setae absent; 
postspiracular setae present; postspiracular 
area with few broad white scales; hypostig- 
mal area with small patch of broad white 
scales; patches of broad white scales on 
propleuron, subspiracular area, upper and 
lower areas of mesokatepisternum, and me- 
sepimeron; lower mesepimeron with | seta; 
metameron and mesopostnotum bare. Wing: 
With dark scales on all veins except for mi- 
nute basal spot of white scales on costa; 
upper calypter fringed with many hairlike 
setae; alula with a row of fringe scales and 
3 decumbent scales on dorsal surface; vein 
1A ending well beyond base of fork of vein 
Cu; cell R, about 1.5 length of R,,,. Halter: 
With dark scales. Legs (Fig. 2B): Coxae 
with patches of white scales; forefemur 
with a white line on anteroventral surface; 
mid- and hindfemora and fore-, mid- and 
hindtibiae with a medial white line on an- 
terior surface; foretarsomere | with basal 
0.15—0.18 white on dorsal surface; foretar- 
somere 2 with basal 0.31—0.40 white on 
dorsal surface; foretarsomeres 3—5 all dark; 
midtarsomere | with basal 0.16—0.20 white 


522 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


IX tergum 


+— 0.1mm—4 


IX sternum 


ELE +——_ 0.1 m m ———— 


claspette 


-——— 0.1mm ——— 


aedeagus 


Aedes (Cornetius) cozi Cornet 


Fig. 3. 


Aedes (Cornetius) cozi. A, Paraproct (dorsal aspect). B, Claspette (dorsal aspect). C, IX tergum 


(dorsal aspects). D, [IX sternum (dorsal aspect). E, Aedeagus (dorsal aspect). 


on dorsal surface; midtarsomere 2 with bas- 
al 0.40—0.50 white on dorsal surface; mid- 
tarsomere 3 with basal 0.32—0.33 white on 
dorsal surface; midtarsomeres 4, 5 all dark; 
hindtarsus with a basal white band on tar- 
someres |—3, ratio of length of white band 
on dorsal surface to total length of tarso- 
meres 0.21—0.27, 0.40—0.45 and 0.66—0.75 


respectively; hindtarsomere 4 all white ex- 
cept at apex on ventral surface; hindtarso- 
mere 5 all white; fore-, mid- and hindlegs 
with tarsal claws equal, simple. Abdomen: 
Tergum I with white scales on laterotergite; 
terga II-VII with basal sublateral white 
spots only; tergum VIII with basal white 
band; sterna II—-VII largely with white 


VOLUME 107, NUMBER 3 


scales; segment VIII largely retracted. Gen- 
italia (Fig. 6): Apical margin of sternum 
VIII with median notch and conspicuous 
rounded lateral lobes; insula as long as or 
slightly longer than wide, with minute spic- 
ules and 10—12 tuberculi on apical 0.4; up- 
per vaginal sclerite present; lower vaginal 
sclerite absent; apical margin of tergum [X 
with well-developed lateral lobe, with 4 or 
5 setae; apical margin of postgenital plate 
without small notch; cercus short and 
broad; 3 spermathecae, one slightly larger 
than other 2. 

Male.—Essentially as in female, differ- 
ing in following characters: Head (Fig. 1A): 
Maxillary palpus about as long as probos- 
cis, 5-segmented, predominantly dark, with 
white band at bases of palpomeres 2—5, 
those on palpomeres 4,5 incomplete ven- 
trally; palpomeres 4,5 very short, somewhat 
downturned and with only few short setae; 
palpomeres 4 and 5 together very short, 
shorter than palpomere 3, 0.40—0.44 length 
of palpomere 3; palpomere 4 slightly swol- 
len, with a few short stiff setae at apex; pal- 
pomere 5 shorter than palpomere 4, some- 
what narrow toward tip and downturned; 
antenna plumose, shorter than proboscis. 
Wing: Cell R, 1.2—1.5 length of R,,,. Legs 
(Figs. 2C, D): Midtarsomere 3 all dark: 
foreleg with tarsal claws unequal, larger 
one toothed, smaller one simple; midleg 
with tarsal claws unequal, both simple. Ab- 
domen: Tergum II with basolateral white 
spots; terga III-VH each with small basal 
sublateral white spots; terga I-VII all dark 
dorsally (Holotype specimen); sternum VIII 
with basolateral white spots. Genitalia 
(Figs. 3, 4C): Gonocoxite 2.2 times as long 
as wide (width measured 0.5 from base), 
scales restricted to dorsolateral, lateral and 
ventral surfaces, with dense scales on mesal 
margin of ventral surface, and setae scat- 
tered on dorsomesal surface; claspette long, 
reaching to 0.82 of gonocoxite, with row of 
short setae along mesal margin of apical 
part, numerous long, curved setae along lat- 
eral margin of apical part, 8 straight setae 
on lateral side of basal part, and 9 straight 


523 


setae on mesal side of basal part; gonosty- 
lus elongate, rather flat, about 0.86 length 
of gonocoxite, slightly expanded just be- 
yond middle, narrow and curved toward tip, 
with a blunt, stout gonostylar claw apically 
on inner margin of slightly expanded sub- 
apical part; aedeagus with strongly toothed 
lateral plates; paraproct with a sternal arm; 
cercal setae absent; apical margin of tergum 
IX slightly concave medially, with 7—9 se- 
tae on each side; sternum IX with 8—10 
larger setae in one row and 6 smaller setae 
in second row. 

Pupa (Figs. 4A, B).—Cephalothorax: 
Trumpet 3.7 as long as wide (width mea- 
sured 0.5 from base); setae 1—5-CT single, 
short; 6-CT single, long, stout, much longer 
and stouter than 7-CT; 7-CT single, short; 
8,9-CT single; 10-CT single, caudomesad 
of 11-CT; 11-CT single, long; 12-CT sin- 
gle. Abdomen: Seta 1-I well developed, 
usually with 6 branches (6—8); 1-II single; 
3-I1, 111 single, short; 1-III,[V single; 5-IV,V 
single, short, not reaching beyond posterior 
margin of following segment; 6-III-V- sin- 
gle, short, about as long as 9-III-V; 9-I-V 
small, single, simple; 9-VI,VII single, long, 
stout, much longer and stouter than 9-I-V; 
9-VIII usually with 6 branches (3-8), 
barbed. Paddle: Oval, 2.0 as long as wide; 
margins with fringe of very fine hair-like 
spicules; apex notched; seta 1-P single, 
long. 

Larva (Fig. 5).—Head: Antenna without 
spicules; seta 1-A inserted on distal 0.25 of 
shaft, 2-branched; seta 1-C well developed, 
single, long, stout and curved; 4-C well de- 
veloped, single, simple, closer to 6-C than 
5-C, cephalad and mesad of 6-C; 6-C sin- 
gle, cephalad and mesad of 5-C; 5-C single, 
long, longer than 6-C; 4-C and 6-C cepha- 
lad of antenna base; 7-C usually with 6 
branches (6—8); 8-C single; 9-C usually 
with 2 branches (1,2); 10-C usually with 2 
branches (2—4); 11-C usually with 7 
branches (6—11), barbed; 12-C usually with 
2 branches (1,2); 13-C usually single (1-3); 
14-C usually with 4 branches (3,4), barbed; 
15-C usually with 2 branches (2—4); men- 


524 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


1.0mm 


C 


ee 
gonostylus nee 


gonostylar claw 


Aedes (Cornetius) cozi Cornet 


/ 


Fig. 4. Aedes (Cornetius) cozi. A, Dorsolateral aspect of the cephalothorax of the male pupa. B, Dorsal and 
ventral aspects of the metathorax and abdomen of the male pupa. C, Tergal aspect of the male genitalia. 


VOLUME 107, NUMBER 3 525 


7 
en 
7° 
8 2 
1.0mm 
4 3 Vi 10 


é 
5 
g 


i 


Aedes (Cornetius) cozi Cornet 


Fig. 5. Aedes (Cornetius) cozi. A, Dorsal and ventral aspects of the head of the fourth instar larva. B, Dorsal 
and ventral aspects of the thorax and abdomen of the fourth instar larva. C, Lateral aspect of the terminal 
abdominal segments of the fourth instar larva. 


526 


tum with 7 or 8 teeth on each side of central 
tooth. Thorax: Seta 1-P with 4 branches, 
barbed; 2—4-P single; 5-P usually with 4 
branches (3,4), barbed; 6-P single, long, 
barbed; 7-P usually with 3 branches (3,4), 
barbed; 9-P single; 11-P single, small; 5,7- 
M single, long, barbed; 6-M usually with 5 
branches (5,6), barbed; 8-M usually with 5 
branches (4,5), barbed; 9-M usually with 8 
branches (7,8), barbed; 10,12-M_ single, 
long, and barbed; 11-M single, small; 7-T 
with 5 branches, barbed; 9-T usually with 
9 branches (8,9), barbed; 10,11-T similar to 
10,11-M; 12-T much reduced, single, sim- 
ple; basal spine of meso- and metapleural 
setae long, pointed apically. Abdomen: Seta 
6-I with 5 branches, barbed; 7-I double, 
barbed; 6-II usually with 3 branches (3,4), 
barbed; 7-II with 2 branches, barbed; 6-III- 
V with 3—5 branches, barbed; 6-VI single, 
barbed; 1-VII single, long, barbed; 2-VII 
double; 2-VIII distant from 1-VIII; 1-VIII 
usually single (1,2), long, barbed; 3-VIII 
usually with 4 branches (4—6), barbed; 5- 
VIII single, barbed; 2,4-VIII single; comb 
usually with 4 (3—5) scales in a row, each 
scale with small basal denticles; saddle in- 
complete, marginal spicules very small, in- 
conspicuous; seta 1-X with 2 branches; 2- 
X usually with 6 branches (5,6); 3-X with 
4 branches; 4-X with 5 pairs of setae on 
grid, each seta usually with 3 or 4 branches 
(2-5); 4e-X very small, with 2 branches; no 
precratal tufts; anal papillae sausage-like, 
dorsal pair longer than ventral pair. Siphon: 
About 1.8 as long as width at 0.5 from base, 
acus absent; usually with 3 (2—5) pecten 
spines; each spine usually with 1—3 basal 
denticles; seta 1-S single, long, inserted be- 
yond apical pecten spine and dorsad of pec- 
ten spines; seta 2-S single, short, stout, 
curved. 

Type data.—Aedes (Stegomyia) cozi Cor- 
net, holotype ¢ (GA 457, ex. 2 7), in [OR- 
STOM]; type locality: SENEGAL.—Sene- 
gal Oriental: Kedougou (12°33'N, 
12°11'W), 6 km N. Kedougou, Galerie, 28- 
VII-1972 (M. Cornet). Paratypes: 1 6, 1 
Cel (ex, 2 7) same datayas holotype 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


[ORSTOM]; 2 ¢ (ex. @ 7), with genitalia 
on slides (MEP Acc. 724, 82/5, 82/27), 1 
2 (ex. 2 7), with genitalia on slide (MEP 
Acc. 724, 82/6) and 1 L (MEP Acc. 724), 
same data as holotype [USNM]. 

Other material examined.—SENE- 
GAL.—Senegal Oriental: Kedougou 
(12°33'’N, 12°11’W), 6 km N. Kedougou, 
Galerie, 19-XI-1973, M. Cornet, (B.M., 
1974-145), 1 2 [BMNH]; same data except 
22-XI-1973, M. Cornet, (B.M., 1974-145), 
io eo cen! (MER Acc 7/ 19537773) 
[BMNH]; same data except 10 km N. Ke- 
dougou, Galerie, 19-XI-1973, M. Cornet, 
(BEMViE, 19742145) hale? = (MIER Nees) 
[BMNH]; 3 L, 4 P and 3 4" instar larvae 
on slides (MEP Acc. 724), same data as ho- 
lotype [ORSTOM]. Kedougou, ex. larva, 
IX-1983, J.P. Hervy, 4 6,4 d gen (SAMP 
Acc. 1083, 93/116, 93/117, 04/55, 04/56), 
3 2,3 2 gen (SAMP Acc. 1083, 93/119, 
04/57, 04/58) [USNM]; same data except 
WIEWIEIOS3 IP leanne 2 oO, 2 6 gen 
(SAMP Acc. 1083, 04/59, 04/60), 1 2 
(SEM), de 2 tgen (SAME Accs 108351937 
118) [USNM]. 

Distribution.—This species is known 
only from Kedougou, Senegal. 

Taxonomic discussion.—Aedes cozi, for- 
merly placed in the subgenus Stegomyia, 
differs significantly from all other Stego- 
myia species and should be excluded from 
that subgenus (see the discussion mentioned 
under the Systematics of Cornetius). 

The female genitalia of Ae. cozi possess 
characters that agree with Reinert’s (2000) 
diagnostic characters of the female genitalia 
of genus Aedes. 

The most important adult characters for 
determining the subgeneric position in the 
genus Aedes are those of the male genitalia. 
The male genitalia of Ae. cozi possess some 
rather basic characters in common, and sug- 
gest affinities with four subgenera of Aedes, 
namely Aedimorphus, Albuginosus, Dicer- 
omyia, and Stegomyia. However, Ae. cozi 
shares more important characters in both 
adult and immature stages with Stegomyia 


VOLUME 107, NUMBER 3 


tuberculus 


spermathecae 


-— 0.1mm—— 


Si) 


IX tergum 


Aedes (Cornetius) cozi Cornet 


Fig. 6. Aedes (Cornetius) cozi. A, Ventral aspect of the female genitalia; B, VIII sternum (dorsal aspect); 
C, VII tergum (dorsal aspect); D, Spermathecae; E, IX tergum (dorsal aspect). 


than with any other subgenus, suggesting 
the strongest affinities with that subgenus. 

Bionomics.—The holotype and paratypes 
of Aedes cozi were reared from eggs col- 
lected from a female (No. 7) biting humans 
in a gallery forest, 6 km N. of Kedougou, 
eastern Senegal. The females of this species 
were also taken biting/landing on humans 
in different gallery forests between 1700— 
2000 h at Kedougou, Senegal. 

Medical importance.—Unknown. 

Remarks.—Although Reinert et al. 
(2004) substantially revised the classifica- 


tion of the tribe Aedini and some of their 
conclusions are warranted, I do not fully 
accept their classification for two primary 
reasons. First, their results were based on a 
preliminary study. Many groups were only 
partially treated and large numbers of spe- 
cies remained unplaced in their classifica- 
tion. 

Second, the selection of exemplars in 
some cases did not represent the known 
complexity and diversity within a group. 
The subgenus Diceromyia, for example, is 
represented in their study by two exem- 


528 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


plars, furcifer and taylori. Both of these 
species, however, are in the same species 
complex and species group (Edwards’ 
Group A or African species), and the other 
species group (Group B), which includes 14 
Oriental species, is not represented and may 
not be monophyletic with the African spe- 
cies. Indeed, my study of Diceromyia 1n- 
dicates that this subgenus is presently a het- 
erogeneous assemblage of species that does 
not form a monophyletic group. 

Another subgenus for which the exem- 
plars were not representative is Stegomyia. 
Six species (aegypti, africanus, albopictus, 
desmotes, futunae, and scutellaris) were 
used as exemplars in their analysis. These 
six species, however, represent only four of 
the eight species groups then recognized 
(Belkin 1962; Bohart 1956; Huang 1972, 
1977, 1979, 1990, 1997). Moreover, I now 
recognize 11 species groups in the Afro- 
tropical Region alone (Huang 2004), and 
there are three more species groups in the 
Oriental and Oceanian regions. 

These two subgenera (Diceromyia, 28 
species, see Reinert et al. 2004: 357; and 
Stegomyia, 127 species including cozi Cor- 
net, see Reinert et al. 2004: 365-366) will 
undoubtedly be shown to be paraphyletic, 
if not polyphyletic, and the selection of ex- 
amplars used in their analysis does not re- 
flect this complexity and diversity. Thus, I 
feel that it is premature to raise these sub- 
genera to generic status, and I retain both 
Diceromyia and Stegomyia as subgenera in 
the present work. 


ACKNOWLEDGMENTS 


I express my sincere appreciation to Dr. 
Wayne N. Mathis, Department of Entomol- 
ogy, Smithsonian Institution, Drs. Richard 
C. Wilkerson and Leopoldo M. Rueda, Wal- 
ter Reed Biosystematics Unit (WRBU), and 
Dr. Maria Anice Mureb Sallum, University 
of Sao Paulo, Sao Paulo, Brazil/NRC Re- 
search Associate Fellow, for critically re- 
viewing this manuscript and for their valu- 
able comments. 

I am most grateful to Drs. Peter EK Mat- 


tingly (deceased), Graham B. White (for- 
merly of BMNH), and Miss Theresa M. 
Howard, Department of Entomology, The 
Natural History Museum [BMNH], Lon- 
don, England, for the loan of specimens; to 
Drs. M. Germain, A. Rickenbach, M. Cor- 
net (formerly of ORSTOM), and Dr. B. 
Geoffroy, Institut de Recherche pour le De- 
veloppement (IRD), Department Sante BP 
5045, 34032 Montpellier Cedex O1 (for- 
merly Services Scientifiques Centraux de 
V’O.R.S.T.0.M., Bondy) [ORSTOM], 
France, for the loan of the specimens used 
in this study. The Senegal material from Dr. 
M. Cornet and Mr. J. PB. Hervy (ORSTOM) 
and Medical Entomology Laboratory, Insti- 
tut Pasteur de Dakar, Senegal, are acknowl- 
edged with sincere appreciation. 

I also express my gratitude to Mr. Young 
T. Sohn, scientific illustrator, Department of 
Entomology, Smithsonian Institution, for 
preparing the illustrations. 


LITERATURE CITED 


Belkin, J. N. 1962. The Mosquitoes of the South Pa- 
cific (Diptera, Culicidae). University of California 
Press, Berkeley and Los Angeles. 2 vols., 608 and 
412 pp. 

Bohart, R. M. 1956. Insects of Micronesia (Diptera: 
Culicidae). Bernice P. Bishop Museum, Insects of 
Micronesia 12: 1—84. 

Cornet, M. 1973. Aedes (Stegomyia) cozi n. sp., une 
nouvelle espece de Culicidae au Senegal. Cahiers 
de Office de la Recherche Scientifique et Tech- 
nique Outre-Mer Serie Entomologie Medicale et 
Parasitologie 11: 175-180. 

Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ 
Glossary of Mosquito Anatomy. Plexus Publish- 
ing, Inc., Marlton, New Jersey, 415 pp. 

. 1982. Corrections and additions to Taxono- 
mists’ Glossary of Mosquito Anatomy. Mosquito 
Systematics (1981)13: 201-217. 

Huang, Y.-M. 1972. Contributions to the Mosquito 
Fauna of Southeast Asia. XIV. The subgenus 
Stegomyia of Aedes in Southeast Asia. 1. The scu- 
tellaris group of species. Contribution of the 
American Entomological Institute (Ann Arbor) 
9(1): 1-109. 

. 1977. Medical Entomology Studies. VII. The 

subgenus Stegomyia of Aedes in Southeast Asia. 

II. The edwardsi group of species. III. The w-al- 

bus group of species (Diptera: Culicidae). Contri- 

bution of the American Entomological Institute 

(Ann Arbor) 14(1): 1-111. 


VOLUME 107, NUMBER 3 529 


. 1979. Medical Entomology Studies. XI. The Afrotropical Region (Diptera: Culicidae). Pro- 
subgenus Stegomyia of Aedes in the Oriental Re- ceedings of the Entomological Society of Wash- 
gion with keys to the species (Diptera: Culicidae). ington 103: 1—S3. 

Contributions of the American Entomological In- . 2002. A pictorial key to the mosquito genera 
stitute (Ann Arbor) 15(6): 1—79. of the world, including subgenera of Aedes and 

. 1986. Notes on the Aedes (Diceromyia) fur- Ochlerotatus (Diptera: Culicidae). Center for In- 
cifer group, with a description of a new species sect Systematics, Korea, Insecta Koreana 19(1): 
(Diptera: Culicidae). Proceedings of the Entomo- 1-130. 
logical Society of Washington 88: 634-649. . 2004. The subgenus Stegomyia of Aedes in 

. 1990. The subgenus Stegomyia of Aedes in the Afrotropical Region with keys to the species 
the Afrotropical Region. I. The africanus group of (Diptera: Culicidae). Zootaxa 700, 120 pp. 
species (Diptera: Culicidae). Contribution of the Reinert, J. E 2000. Comparative anatomy of the female 
American Entomological Institute (Gainesville) genitalia of genera and subgenera in tribe Aedini 
26(1): 1-90. (Diptera: Culicidae). Part V. Genus Aedes Meigen. 

. 1997. The subgenus Stegomyia of Aedes in Contributions of the American Entomological In- 
the Afrotropical Region. Il. The dendrophilus stitute (Gainesville) 32(3): 1-102. 
group of species (Diptera: Culicidae). Contribu- Reinert, J. E, R. H. Harbach, and I. J. Kitching. 2004. 
tion of the American Entomological Institution Phylogeny and classification of Aedini (Diptera: 
(Gainesville) 29(4): 1-104. Culicidae), based on morphological characters of 

. 2001. A pictorial key for the identification of all life stages. The Linnean Society of London, 
the subfamilies of Culicidae, genera of Culici- Zoological Journal of the Linnean Society 142: 


nae, and subgenera of Aedes mosquitoes of the 289-368. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 530-535 


A NEW TRIGONALID WASP (HYMENOPTERA: TRIGONALIDAE) FROM 
EASTERN NORTH AMERICA 


DAvID R. SMITH AND IAN C. STOCKS 


(DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, % National Museum of Natural History, Smithsonian Insti- 
tution, PO. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. (e-mail: 
dsmith @sel.barc.usda.gov); (ICS) Department of Entomology, Soils, and Plant Sciences, 
Clemson University, Clemson, SC 29634-0315, U.S.A. (e-mail: istocks @clemson.edu) 


Abstract.—Orthogonalys bella, n. sp., is described from Great Smoky Mountains Na- 
tional Park, Tennessee. It is the second species of Orthogonalys Schulz and the fifth 
species of Nearctic Trigonalidae. Characters separating it from Orthogonalys pulchella 


(Cresson) are given. 


Key Words: 


Inasmuch as a new trigonalid has not 
been discovered in the Nearctic for 108 
years, one would assume that the fauna is 
completely known. Nothing is sure, how- 
ever, and surprisingly a new species was 
discovered during the All Taxa Biodiversity 
Inventory (ATBI) investigations of Great 
Smoky Mountains National Park (GRSM). 
One specimen, belonging to the genus Or- 
thogonalys Schulz as defined by Carmean 
and Kimsey (1998), is so unique that it can- 
not be confused with the only other North 
American species of the genus, O. pulchella 
(Cresson 1869). In thousands of Malaise 
trap samples from GRSM and elsewhere in 
eastern United States (e.g., Smith 1996), 
many thousands of O. pulchella have been 
captured, but no Orthogonalys so remark- 
ably different as the species described here. 
Only a single specimen is known, but an- 
other may not be found in a lifetime, and 
we consider it significant to document its 
existence. 

About 88 species in 16 genera of Trigon- 
alidae are known worldwide (Carmean and 
Kimsey 1998). Four genera, each with one 


hyperparasitoid, parasitoid, wasp, Trigonalyidae 


species, occur in the Nearctic Region. 
Townes (1956) was the last to revise the 
Nearctic fauna, Carlson (1979) cataloged 
the family, and Smith (1996) gave a key 
and reviewed the eastern North American 
species. Carmean (1995) reviewed and gave 
keys to the genera of Costa Rica, and Car- 
mean and Kimsey (1998) presented a com- 
prehensive phylogeny and review of the 
world fauna. Most trigonalids are hyperpar- 
asitoids of endoparasitic Ichneumonoidea 
and Tachinidae or parasitoids of Vespidae 
larvae. Completion of their life cycle de- 
pends on a sequence of chance circum- 
stances leaving one to wonder how they 
survive. In brief, numerous eggs are laid 
randomly on plant foliage; the eggs must 
be ingested by a phytophagous insect, usu- 
ally a caterpillar, which is already parasit- 
ized; the eggs hatch in the gut and the lar- 
vae enter the body of the caterpillar to 
search for a parasitoid on which the trigon- 
alid larva develops. There is no further de- 
velopment unless the caterpillar is parasit- 
ized by another parasitoid. In species that 
are parasitoids of Vespidae, a caterpillar in- 


VOLUME 107, NUMBER 3 


fected by a trigonalid must be taken as prey 
back to the vespid nest where the vespid 
larva serves as the final host. Reviews of 
biology are given by Townes (1956), Carl- 
son (1979), Weinstein and Austin (1991), 
Carmean (1991, 1995), Smith (1996) and 
Carmean and Kimsey (1998). 


Orthogonalys bella Smith and Stocks, 
new species 
(Figs. 1—5) 


Female.—Length, 5.5 mm; forewing 
length, 4.8 mm. Color: Antenna with seg- 
ments 1—6 reddish brown to orange, seg- 
ments 7—9 white, with only extreme apex 
of 9 blackish, segments 10 to apex black. 
Head black, with spot at middle of inner 
orbit, spot on lower inner orbit, clypeus, 
and mouthparts dark orange; palpi more 
whitish; apices of teeth of mandible black. 
Mesosoma black, with tegula whitish. Legs 
orange with coxae black, trochanters more 
whitish, and extreme apex of hind femur 
black. Metasoma black, with about anterior 
half dark orange and apical half black. 
Wings hyaline; veins and stigma black. 

Head: Antenna 24-segmented, length 
slightly less than 3X head width. Eyes 
small, round; in lateral view, far from hind 
margin of head; in front view, far apart and 
slightly diverging below, with lower inter- 
ocular distance 1.8 eye height and upper 
interocular distance 1.3 eye height. Clyp- 
eus 3X broader than its medial length. Sec- 
ond tooth of left mandible much longer and 
broader than apical and third teeth (man- 
dibles closed, symmetry not observed). Ma- 
lar space long, nearly 0.5 eye height. Dis- 
tance between toruli about half distance be- 
tween inner margin of eye and torulus. Ge- 
nal carina distinct, broad, about equal to 0.7 
diameter of an ocellus, ending at hyposto- 
mal carina. In dorsal view, head strongly 
narrowing behind eyes. Head uniformly 
shining, with small punctures separated by 
several puncture diameters, somewhat dens- 
er and closer together on frons, and very 
fine on malar area, with surface appearing 
dull. Mesosoma: Smooth, shining; prono- 


531 


tum laterally with several straight carinae; 
mesepisternum with broad transverse sulcus 
with carinae, surface above sulcus shiny but 
roughened, below sulcus smooth, shining 
and sparsely punctate; mesonotum smooth, 
with scattered punctures, transverse sulcus 
anterior to scutellum with short carinae, 
scutellum low and evenly convex, without 
central groove, with scattered punctures; 
metanotum shining, with few punctures an- 
teriorly; propodeum with areolate-rugose 
sculpturing, Propodeal foramen U-shaped. 
Forewing with submarginal cell 2 not pet- 
iolate. Posterior margin of hind coxa with 
sharp carina extending length of coxa, with 
dorsal part of carina sharply angulate; an- 
terior margin of hind coxa with less devel- 
oped carina; hind coxa rectangular in lateral 
view. Hind trochanter 3-segmented. Inner 
hind tibial spur shorter than apical width of 
hind tibia. Tarsal claws bifid, with both 
teeth subequal in length. Metasoma: 
Smooth and shining, with short fine carinae 
at extreme base of first segment. Sternal ar- 
mature absent. Ovipositor sheath broad, 
with length subequal to its height. 

Male.—Unknown. 

Holotype.—Female, from Great Smoky 
Mountains National Park, Tennessee, la- 
beled ““TN: Sevier Co., GRSM, ATBI Plot 
Brushy Mountain, Malaise trap MT 14, 
280000E, 3950599N, 3-18 June 2002, 
BMerritt, MT1420020618.” Provisionally 
deposited in the National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, DC, pending mutual resolution 
and agreement with the National Park Ser- 
vice regarding specimen deposition. 

Habitat.—Malaise trap #14 at Brushy 
Mountain (elevation 4,810’ [1,466 m]) was 
located in a Rhododendron heath bald. The 
bald habitat is a Kalmia latifolia L.—Rho- 
dodendron catawbiense Michx.—(Gaylus- 
sacia baccata (Wangenh.) K. Koch., Pieris 


floribunda Benth. & Hook., Vaccinium cor- 


ymbosum L.) |Ericaceae] shrubland alli- 
ance, as documented by NatureServe, a 
non-profit conservation organization work- 


532 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-5. 
view. 5, Apex of abdomen and sheath, lateral view. 


ing in partnership with GRSM to inventory 
plant communities. 

Etymology.—The name reflects the au- 
thors’ concept of the species—beautiful, 
from the Latin bellus. 

Diagnosis and discussion.—Orthogona- 
lys pulchella, the only other North Ameri- 
can species of the genus, occurs from 


Orthogonalys bella. 1, Lateral view. 2, Head, lateral view. 3, Head, dorsal view. 4, Head, front 


southeastern Canada through eastern United 
States to Mexico. Comparison of O. bella 
with O. pulchella is shown in Figs. 1—10. 
Numerous characteristics separate the two 
species. Orthogonalys pulchella is much 
more richly marked with yellow and or- 
ange; the antennal length is more than 3X 
the head width; antennal segments 9—13 


VOLUME 107, NUMBER 3 


533 


Figs. 6-10. Orthogonalys pulchella. 6, Lateral view. 7, Head, lateral view. 8, Head, dorsal view. 9, Head, 
front view. 10, Apex of abdomen and sheath, lateral view. 


and sometimes 14 are white; the genal ca- 


rina is narrower, less than half the diameter 


of an ocellus; the eyes are larger, more oval, 
closer together in front view, and closer to 
the hind margin of the head in lateral view; 
in front view, the inner margin of the eyes 
are subparallel, not diverging below; the 


apical three teeth of the left mandible are 
subequal in size; the clypeus is about 2X 
broader than its medial length; the malar 
space is shorter, about 0.2 the eye height; 
in dorsal view the head is straighter behind 
the eyes and is more rectangularly shaped; 
the mesoscutellum has a shallow, longitu- 


534 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


dinal groove; there is no sharp carina on the 
posterior margin of the hind coxa and the 
hind coxa is elongate oval in lateral view; 
the inner apical tibial spur is longer than the 
apical width of the hind tibia; the tarsal 
claws are bifid, with the inner tooth longer 
and broader than outer tooth: and the sheath 
is narrower, nearly twice as long as broad. 

Carmean and Kimsey (1998) listed 11 
world species of Orthogonalys, one from 
North America, one from South America, 
three from Japan, three from southeastern 
Asia, and three from Africa. The only South 
American species, O. boliviana Schulz 
1905 is almost indistinguishable from O. 
pulchella. The overall black coloration re- 
sembles some of the Asian species, which 
are mostly black, with few white or brown 
markings as opposed to the rich white and 
orange markings of species from the rest of 
the world (Carmean and Kimsey 1998). Or- 
thogonalys bella does not key to any of the 
Japanese species treated by Tsuneki (1991). 
It is closest to O. fukuiensis Tsuneki 1991, 
but in O. fukuiensis only the first two me- 
tasomal tergites are broadly brownish white 
and the thorax has no yellow or brown 
marks. Additionally, based on examination 
of O. hagoromensis Teranishi 1929 and 
based on Tsuneki (1991), the Japanese spe- 
cies have less developed carinae on the hind 
coxae, the second tooth of the left mandible 
is the same size as the first and third teeth, 
the malar space is narrow (as in O. pul- 
chella), and the tarsal claws have a large 
inner tooth. Orthogonalys bella, mostly 
characterized by its predominantly black 
color, does not agree with descriptions of 
the other species listed by Carmean and 
Kimsey (1998). 

The possible resemblance of O. bella to 
eastern Asian species rather than the North 
America species may be another example 
of biogeographical similarities between 
eastern United States and eastern Asia, 
though color is not a good indication of re- 
lationships. There are undoubtedly many 
examples, but it is reminiscent of a sawfly, 
Strongylogaster lata Smith and Naito 1995 


(Tenthredinidae), which is known from 
only a few specimens from the mid-Atlantic 
states. Strongylogaster lata, only discov- 
ered after 100 or more years of collecting 
in such a well-collected area, belongs to a 
group of species known only from eastern 
Asia and is very unlike the other North 
American species of the genus (Smith and 
Naito 1995). 

Besides the two species of Orthogonalys, 
the only other trigonalid collected in GRSM 
is Taeniogonalys gundlachi (Cresson 
1865), which occurs from eastern United 
States to Costa Rica. Lycogaster pullata 
Schuckard 1841, another eastern North 
American species, probably occurs there, 
but they are se!dom collected in Malaise 
traps (Smith 1996). No other trigonalids 
were collected in the traps at Brushy Moun- 
tain during the inventory of 2001 and 2002. 


ACKNOWLEDGMENTS 


Cathy Apgar, Systematic Entomology 
Laboratory, U.S. Department of Agriculture 
(SEL) took the Auto-Montage™ photos and 
arranged the plates. We appreciate the re- 
views of David Carmean, Simon Fraser 
University, Burnaby, BC, and M. G. Pogue 
and T. J. Henry, SEL, Washington, DC. ICS 
was supported with funding from the U:S. 
Geological Survey Biological Resources 
Discipline Natural Resources Protection 
Program grant 1434-OOHQAG0044, given 
to C. R. Parker (U.S. Geological Survey) 
and E. C. Bernard (University of Tennessee, 
Knoxville). Brian Merritt collected the 
specimen. The natural resources staff of 
Great Smoky Mountains National Park, 
Discover Life in America, and volunteers 
too numerous to name individually provid- 
ed financial and much logistical support to 
the ATBI initiative. 


LITERATURE CITED 


Carlson, R. W. 1979. Trigonalidae, pp. 1107-1198. In 
Krombein, K. V., P. D. Hurd, Jr, D. R. Smith and 
B. D. Burks, eds. Catalog of Hymenoptera in 
America North of Mexico, Vol. 1, Symphyta and 
Apocrita (Parasitica). Smithsonian Institution 
Press, Washington, DC., 1,198 pp. 


VOLUME 107, NUMBER 3 


Carmean, D. 1991. Biology of the Trigonalyidae (Hy- 
menoptera), with notes on the vespine parasitoid 
Bareogonalos canadensis. New Zealand Journal 
of Zoology 18: 209-214. 

. 1995. Trigonalyidae, pp. 187—192. In Hanson, 
P. E. and I. D. Gauld, eds. The Hymenoptera of 
Costa Rica. Oxford University Press, Oxford, 893 
pp. 

Carmean, D. and L. Kimsey. 1998. Phylogenetic re- 
vision of the parasitoid wasp family Trigonalidae 
(Hymenoptera). Systematic Entomology 23: 35— 
76. 

Smith, D. R. 1996. Trigonalyidae (Hymenoptera) in 
the eastern United States: Seasonal flight activity, 
distributions, hosts. Proceedings of the Entomo- 
logical Society of Washington 98: 109-118. 


535 


Smith, D. R. and T. Naito. 1995. A new species of 
Strongylogaster (Hymenoptera: Tenthredinidae) 
from North America. Entomological News 106: 
57-60. 

Townes, H. 1956. The Nearctic species of trigonalid 
wasps. Proceedings of the United States National 
Museum 106: 295-304. 

Tsuneki, K. 1991. Revision of the Trigonalidae of Ja- 
pan and her adjacent territories. Special Publica- 
tions of the Japan Hymenopterists Association No. 
37, 68 pp. 

Weinstein, P. and A. D. Austin. 1991. The host-rela- 
tionships of trigonalyid wasps (Hymenoptera: Tri- 
gonalyidae), with a review of their biology and 
catalogue to world species. Journal of Natural His- 
tory 18: 209-214. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 536-542 


MYSTAXIOPS: A NEW GENUS OF SMALL MINNOW MAYFLIES 
(EPHEMEROPTERA: BAETIDAE) FROM PAPUA NEW GUINEA 


W. P. MCCAFFERTY AND L. SUN 


Department of Entomology, Purdue University, West Lafayette, IN 47907, U.S.A. 


(e-mail: mecaffer @ purdue.edu) 


Abstract.—Newly discovered larvae of the mayfly family Baetidae in Papua New Guin- 
ea are described as Mystaxiops venatoris, new genus and new species. The new genus 
is shown to belong to a monophyletic, predatory subgroup of the Centroptiloides complex, 
which was previously known only from the Afrotropics. The labrum of the new genus is 
extremely produced into a friction pad, and other mouthpart characteristics also demon- 
strate adaptations for predation. Labial palps and hypopharynx are distinct from all other 
related genera. Functional morphology of mouthparts and historical biogeography are 


hypothesized. 
Key Words: 


An unusual predatory subgroup of the 
Afrotropical Centroptiloides complex of the 
mayfly family Baetidae has been known to 
include the genera Barnumus McCafferty 
and Lugo-Ortiz (southern Africa), Centrop- 
tiloides Demoulin (eastern and southern Af- 
rica), Guloptiloides Gattolliat and Sartori 
(Madagascar), Herbrossus McCafferty and 
Lugo-Ortiz (Madagascar), and Nesoptilo- 
ides Demoulin (Madagascar). The Centrop- 
tiloides complex and this particular sub- 
group were first recognized and defined by 
Lugo-Ortiz and McCafferty (1998). The def- 
inition of the complex was more recently 
modified somewhat by McCafferty (2002). 
All known species of the subgroup contain- 
ing the genera mentioned above are appar- 
ently predatory at least as mature larvae. 
Such larvae are all nearly 10 mm or more 
in body length (relatively large for lotic 
baetid species) and demonstrate certain spe- 
cialized characteristics in common that set 
them apart from other genera of the Cen- 
troptiloides complex. These characteristics 
include, for example, a short, broad and 


mayflies, Papua New Guinea, Baetidae, new genus 


thickened labrum; mandibles with more or 
less lateral convexity together with various 
degrees of setation in the basal half of the 
mandible; and heavily sclerotized and well- 
developed apical spines on the galealaci- 
niae. The presence of highly developed 
spines of the galealaciniae is a common 
adaptive trait associated with predatory lar- 
vae in Ephemeroptera. The relative sharp- 
ness and development of mandibular inci- 
sors and reduction of the mola or its mod- 
ification into a secondary incisor are also 
adaptive traits commonly associated with 
predation in Ephemeroptera, but they vary 
in degree of development among the genera 
of this predatory subgroup. Raptorial fore- 
legs may or may not be variously developed 
in predatory mayflies, and this is also the 
case among this predatory subgroup of the 
Centroptiloides complex. 

Recently we identified another member 
of this subgroup of predatory baetids from 
Papua New Guinea, which we describe 
herein. Significantly this discovery expands 
the known geographic range of the sub- 


VOLUME 107, NUMBER 3 


group and suggests a possible Gondwanan 
or East Gondwanan origin, with subsequent 
vicariance giving rise to the present-day, 
possibly relictual, distribution of the sub- 


group. 


Mystaxiops McCafferty and Sun, 
new genus 
(Figs. 1-10) 


Larva.—Head: Labrum (Figs. 1, 8—9) 
broadly V-shaped at distal margin, consid- 
erably broader than long, extremely thick- 
ened dorsoventrally as best seen in lateral 
(Fig. 1) or ventral view of hypognathous 
head; lateral lobes curving somewhat pos- 
teriorly; distal expansive area of labrum 
forming shallow, inverted U-shaped friction 
pad (Fig. 8) as viewed ventrally; friction 
pad covered with simple hairlike setae and 
more marginally with pinnate hairlike setae 
with very small setules (Fig. 9). Lingua of 
hypopharynx (Fig. 2) with pronounced, 
narrow, parallel-sided medioapical lobe. 
Angulate and planate mandibles (Figs. 3—4) 
somewhat narrow-elongate, with lateral 
margins convex and with elongate patch of 
moderately long setae in basal half of lat- 
eral margin, and with only somewhat re- 
duced mola and with one set of broadly 
based incisors (appearing fused for much of 
their length); tuft of setae on apical margin 
between prostheca and mola absent but area 
roughened with extremely small rudimen- 
tary armature detectable only at very high 
magnification; prostheca somewhat robust, 
more so on angulate mandible. Maxilla 
(Figs. 1, 5) with two-segmented maxillary 
palp extending beyond tip of galealacinia; 
inner medial subdistal hump of galealacinia 
with only single small seta. Segment 3 of 
labial palp (Fig. 6) broadly rounded and 
nearly as broad as long; segment 2 extreme- 
ly broadened medially from base, at mid- 
length nearly twice breadth of terminal seg- 
ment, and with distinct medioapical, thumb- 
like, subacute extension; glossae narrowing 
distally and considerably smaller than par- 
aglossae. Thorax: Pronotum (Fig. 10) with- 
out lateral flanges. Forefemur (Fig. 10) 


S/ 


broadened mostly in basal half, with 
smoothly convex ventral margin, and with 
only short, sparse setae dorsally. Claws 
with one strong row of denticles and one 
weaker row of denticles. Abdomen: Gills 
(Fig. 10) large and asymmetrically subov- 
ate; tracheation well developed. Caudal fil- 
aments with moderately well developed lat- 
eral swimming hairs; middle caudal fila- 
ment nearly as long as cerci. 

Adult.—Unknown. 

Type species.—Mystaxiops venatoris 
McCafferty and Sun. 

Etymology.—The generic nomen is from 
the Greek ‘‘mystax’’ (mustache) and 
““iops”” (minnow), a masculine name allud- 
ing to the profuse setation of the labrum 
and thus the mustachioed minnowlike may- 
fly. 

Discussion.— Mystaxiops larvae are dis- 
tinct from the Afrotropical genera of the 
predaceous subgroup of the Centroptiloides 
complex. The broadened labial palps with 
thumb and well-developed segment 3 (Figs. 
1, 6) are much different than the narrow, 
thumbless palps with a caplike or narrowed 
segment 3, typical of the other genera. The 
well-developed and peculiar median lobe of 
the hypopharynx (Fig. 2) is also apparently 
distinct, although the hypopharynx of Gul- 
optiloides remains undescribed. The friction 
disc is extremely well developed in Mys- 
taxiops (Figs. 1, 8-9), but is nearly as de- 
veloped in Nesoptiloides. The mandibular 
mola of Mystaxiops (Figs. 3—4) is some- 
what reduced as it is in Barnumus, Her- 
brossus, and Nesoptiloides, but it is not 
modified into a secondary incisor as it is in 
Centroptiloides and Guloptiloides. 

When the arrangement of the mouth- 
parts of Mystaxiops larvae are viewed ven- 
trally, a hypothesis regarding the predatory 
function of the suite of characters becomes 
possible. The large and dominant friction 
disc of the labrum could be used to hold 
prey in place, while at the same time the 
heavily sclerotized maxillary spines and 
mandibular incisors could be used to im- 
pale and cut the prey, and the intricate la- 


538 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


6 


Figs. 1-7. Mystaxiops venatoris, larva. 1, Head (lateral). 2, Hypopharynx. 3, Angulate mandible. 4, Planate 
mandible. 5, Maxilla. 6, Labium. 7, Paraproct. 


VOLUME 107, NUMBER 3 539 


— FOfim 


Figs. 8-9. Mystaxiops venatoris, larva. 8, Labral friction pad (ventrodistal view). 9, Friction pad setae. 


540 


bial palps used to manipulate the prey. The 
extraordinary, well-developed friction disc 
may compensate for forelegs that in Mys- 
taxiops show no adaptations for being rap- 
torial. Nesoptiloides, as another example 
within the subgroup, does demonstrate 
forelegs that are both appropriately con- 
toured and fitted with ventral marginal ar- 
mature to suggest a raptorial function (see 
fig. 87 in Lugo-Ortiz and McCafferty 
1998), in addition to having a cuplike (per- 
haps suctioning) labrum that could assist 
in stabilizing prey items. Such differences 
among the genera of the Centroptiloides 
subgroup of predatory genera could also 
be related to differences in prey preference 
and the various degrees to which they rely 
on carnivory during different phases of 
their larval development. 

The distribution of the predatory sub- 
group of the Centroptiloides complex now 
appears to be disjunct (part of the Afro- 
tropical Region and Austral-Asian subre- 
gion) and therefore could be interpreted as 
representing a fragmentary relictual pat- 
tern remaining from an old East Gond- 
wanan general distribution. These may- 
flies are not only large and distinctive, 
they are also relatively easy to collect 
from riffle areas of streams, as observed 
by WPM collecting Centroptiloides and 
Barnumus in South Africa. If they occur 
in India and Sri Lanka or other parts of 
the Oriental Region as might be expected 
from their known distribution [see e.g., 
biogeographical analysis of East Gond- 
wanan family Teloganodidae by Mc- 
Cafferty and Wang (1997) and Mc- 
Cafferty and Benstead (2002)], they 
would have probably been found and re- 
ported by now. 


Mystaxiops venatoris McCafferty and 
Sun, new species 
(Figs. 1-10) 


Late instar larva.—Body (Fig. 10) 
length 11.5 mm. Cercus length 7.0 mm. 
Median caudal filament length 5.0 mm. 
General coloration tan dorsally to cream 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ventrally, with brown margination. Head 
capsule (Fig. 10) with white epicranial su- 
ture and distinct white border medial and 
anterior to compound eyes. Segment | of 
maxillary palp (Fig. 5) with numerous 
short bristlelike setae laterally. Other 
mouthpart setation as shown in Figs. 2—6. 
Mandibular incisors and maxillary apical 
spines heavily sclerotized and more dark- 
ly pigmented than other mouthparts when 
viewing head ventrally. Pronotum some- 
times marked as shown in Fig. 10. Femora 
(Fig. 10) with ventral marginal and medial 
areas of anterior face distinctly patterned 
with brown blotch extending narrowly ba- 
soventrally, but not extending to dorsal 
margin or distal anterior surface. Abdom- 
inal terga essentially concolorous tan ex- 
cept for brown intersegmental margin- 
ation and sometimes faint patterning on 
posterior terga as shown in Fig. 10. Ab- 
dominal sterna unmarked. Gills (Fig. 10) 
profusely tracheated, with short, sparse, 
blunt, hairlike microsetae along distal and 
inner margins, and with some sparse, mi- 
nute spines at outer-distal margin. Para- 
procts (Fig. 7) with 33 or more marginal 
spines and numerous scattered scale bases 
on surface. 

Material examined.—Holotype: Late in- 
star larva, Papua New Guinea, Chimbu 
Province, Wara Sera Research Station, Cra- 
ter Mountain Conservation Area, VII-11- 
2001, Bradler, Jarvis, Svenson (deposited in 
the Purdue Entomological Research Collec- 
tion, West Lafayette, Indiana). Other mate- 
rial: Two middle instar larvae, some parts 
slide-mounted, same data and deposition as 
holotype. 

Etymology.—tThe specific epithet, vena- 
toris, is a Latin masculine noun meaning 
“the hunter.” 

Discussion.—Because this is the first 
species described for Mystaxiops, there is 
no basis for a diagnosis. The highly dis- 
tinctive patterning on the femora may prove 
to be useful for identification of the species, 
as might the high degree of spination of the 
paraprocts. It is possible that some charac- 


541 


VOLUME 107, NUMBER 3 


a 
= 
~ 
we 
a 


MT 
saereaee 


= 


Fig. 10. Mystaxiops venatoris larva, dorsal habitus. 
hump of the galealaciniae may vary among 


teristics given in the generic description 
may prove to be variable among different species. 
species if more species are found. For ex- No habitat data are available for the new 
ample, the number of setae on the inner species. 


542 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ACKNOWLEDGMENTS 


We thank H. Ogden (Provo, Utah) for 
providing materials from New Guinea for 
our study, and A. Provonsha (West Lafay- 
ette, Indiana) for rendering the artwork used 
herein. Scanning electron microscopy was 
conducted at the Life Science Microscopy 
Facility, Purdue University. 


LITERATURE CITED 


Lugo-Ortiz, C. R. and W. P. McCafferty. 1998. The 
Centroptiloides complex of Afrotropical small 


minnow mayflies (Ephemeroptera: Baetidae). An- 
nals of the Entomological Society of America 91: 
1-26. 

McCafferty, W. P. 2002. Gose’s African Ephemerop- 
tera (Baetidae, Heptageniidae). Entomological 
News 113: 294-302. 

McCafferty, W. P. and J. P. Benstead. 2002. Cladistic 
resolution and ecology of the Madagascar genus 
Manohyphella (Ephemeroptera: Teloganodidae). 
Annales de Limnologie 38: 41—52. 

McCafferty, W. P. and T. Q. Wang. 1997. Phylogenetic 
systematics of the family Teloganodidae (Ephem- 
eroptera: Pannota). Annals of the Cape Provincial 
Museums (Natural History) 19: 387—437. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 543-547 


CHARACTERISTICS OF THE LOYOLA NAVAS MALE 
(NEUROPTERA: CHRYSOPIDAE: APOCHRYSINAE) 


CATHERINE A. TAUBER, GILBERTO S. ALBUQUERQUE, AND MAURICE J. TAUBER 


(CAT, MJT) Department of Entomology, Comstock Hall, Cornell University, Ithaca, 
NY 14853-0901 (e-mail: mjt4@cornell.edu); (GSA) Laborat6rio de Protecdo de Plantas, 
CCTA, Universidade Estadual do Norte Fluminense, Campos dos Goytacazes, Rio de 
Janeiro, Brazil 28013-600 (e-mail: gsa@uenf.br) 


Abstract.—The only known male specimen of the rare New World genus Loyola Navas 
has genitalic characteristics that are generally consistent with those described for its sub- 
family, Apochrysinae. The genital characters are also consistent with the placement of the 
genus in the Joguina clade of Apochrysinae. Our specimen of Loyola croesus (Gerstaeck- 
er), one of three species in the genus, has several distinguishing genitalic features; whether 


these features typify the genus Loyola or only L. croesus, is unknown. 


Key Words: 


Apochrysinae is a small, largely tropical 
subfamily of strikingly handsome, but rare- 
ly encountered green lacewings. Adults 
have slender, graceful bodies and broad, 
delicate wings that are often elaborately 
marked or have densely reticulate venation. 
For some genera only one sex is known, 
and with one exception, Apochrysa (= Na- 
caura) matsumurae (Okamoto) (Tsukaguchi 
1995), the biological characteristics and im- 
mature stages of all members of the sub- 
family are unknown. 

In a recent phylogenetic study, Winterton 
and Brooks (2002) confirmed the monophy- 
ly of the subfamily and reduced the 13 pre- 
viously recognized genera to six. Of these, 
three occur in the New World, specifically, 
Central and South America. They are Do- 
menechus Navas (known only from the 
New World; two species—one each from 
Guatemala and Brazil), Joguina Navas (five 
species in total: three from the Orient, two 
from the New World—one each from Gua- 
temala and South America), and Loyola Na- 
vas (known only from three New World 


Chrysopidae, Loyola, Joguina clade, Apochrysinae, green lacewings 


species: two from Brazil, the third from 
Mexico, Nicaragua, Costa Rica, Panama, 
Venezuela, and Brazil; previously known 
only from females). Together with the Ori- 
ental genus, Nobilinus Navas, these three 
genera comprise the Joguina clade of 
Apochrysinae (Winterton and Brooks 
2002). 

The male terminalia of Apochrysinae are 
reduced and vary only in minor respects 
among taxa. Nevertheless, they provide 
some characters that are useful in identify- 
ing species and in helping to discern phy- 
logenetic relationships, both within Apoch- 
rysinae and between Apochrysinae and oth- 
er chrysopid taxa (Winterton and Brooks 
2002). Thus, the lack of information on 
males in the genus Loyola left a distinct gap 
in the knowledge of New World Chryso- 
pidae. 

During a recent field trip in the Mata At- 
lantica of the State of Rio de Janeiro, Bra- 
zil, one of us (GSA) collected a male of 
Loyola croesus (Gerstaecker). Our short 
note here describes the external and internal 


544 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


characteristics of the abdomen of this spec- 
imen. We hope to acquire additional males 
and females of this rare species and to rear 
and describe the immatures. 


MATERIALS AND METHODS 


The specimen, which was teneral when 
collected, was taken to the laboratory where 
it matured for ten days on a diet of auto- 
lyzed yeast, fructose, and honey (1:1:1 mix- 
ture by volume), photoperiod of LD 16:8, 
temperature of 24 + 1°C. It was then frozen 
and pinned; subsequently, the abdomen was 
removed (at segment 3), cleared in a warm 
solution of 10% KOH, rinsed, stained with 
Chlorozol black, and preserved in glycerin. 
The specimen is deposited in the insect col- 
lection of the Universidade Estadual do 
Norte Fluminense, Campos dos Goytacaz- 
es, RJ, Brazil. 


RESULTS AND DISCUSSION 
Loyola croesus (Gerstaecker) 


Collection record.—The specimen was 
taken in a small fragment of secondary for- 
est at Fazenda Carrapeta (22°10’S, 
41°52'W), Conceigaéo de Macabu, Rio de 
Janeiro State, 200 m altitude, November 5, 
2002, G. S. Albuquerque and Ederaldo A. 
Silva, collectors. Collection of the teneral 
specimen occurred on the wing, in the for- 
est on a rainy day. 

Description of male abdomen.—Abdo- 
men thin from segment | through 6, swol- 
len apically from segment 7 (Figs. 1A, B). 
Dorsum and sides green with tergites yel- 
low. Sternites 1 to 3 (S1—3) yellow; S4 yel- 
low anteriorly, brown posteriorly; S5 
brown; S6 to 8+9 yellow. Callus cerci yel- 
low. 

Segments 4 to terminus (cleared and 
stained): Integument (sclerites and inter- 
sclerotic membrane) with moderately dense 
covering of microsetae and more sparsely 
scattered, straight setae; without microtholi. 
Spiracles positioned mesally in interseg- 
mental membrane, not enlarged or elabo- 
rate, with shallow, bilobed atria. 


Tergites 3 through 6 (T3—6) well defined, 
but without dark margins, largely quadrate 
in lateral view. Anterior of T7 (lateral view) 
approximately 1.3 times depth posteriorly. 
T8 short; anterior margin (lateral view) 
deeper than posterior margin of T7. T9 very 
weakly sclerotized, probably not fused with 
ectoprocts; apodemes not distinguished. Ec- 
toprocts fused dorsally via a broad band, 
without dorsal invagination, rounded basal- 
ly and apically, very near each other pos- 
teroventrally. Callus cerci oval; ~35—40 
relatively short trichobothria, not extending 
much beyond margins of callus cerci. Ster- 
nites through S7 quadrate in ventral, lateral 
views; margins well defined, darkly stained; 
intersegmental membranes between S3, S4, 
and S5 very short; connections between S5, 
S6, S7 and S8 distinctly membranous. 
S8+9 fused, tapering and rounded distally, 
without lateral processes. 

Genitalia well sclerotized. Tignum, gon- 
apsis absent. Gonarcus (Fig. 1C) broadly 
arcuate, dorsoventrally flattened, bearing a 
pair of upturned entoprocesses laterally. 
Mediuncus [= arcessus of Winterton and 
Brooks (2000)] well sclerotized, broad and 
bulbous basally, tapering to two rounded 
lobes apically, apparently with a trough 
dorsally, eight pairs of short gonosetae api- 
cally. Hypandrium internum (Fig. 1D) ar- 
row-shaped, with pointed tip apically and 
raised, rounded keel mesally; comes small, 
curved upward. 

Abdominal characteristics.—In their cla- 
distic analysis of the Apochrysinae, Win- 
terton and Brooks (2002) used six charac- 
ters from the male abdomen. Here we code 
L. croesus for each and discuss its relation- 
ship with other genera in Apochrysinae (see 
Table 1). The numbered characters refer to 
Winterton and Brooks’ characters: 

30. Male Ectoprocts. (0) Separated dor- 
sally. (1) Fused dorsally. 

With the exception of the genus Joguina 
(= Joguina + Lainius), the ectoprocts of 
males in Apochrysinae species are typically 
joined. In L. croesus, they are connected via 
a broad, sclerotized band (1). 


VOLUME 107, NUMBER 3 545 


T8 T9-memb 


i 
\——— anus 
cia 


comes 


~ 


_~ T9-memb 


Fig. 1. Loyola croesus, male abdomen. A, Terminal segments, lateral view. B, Terminal segments, ventral 
view. C, Gonarcus: left, dorsal view; right, lateral view. D, Hypandrium internum: upper, dorsal view; lower, 
lateral view. cc, callus cerci; ect, ectoproct; ent, entoprocessus; gs, gonarcus; gse, gonosetae, la, lateral arms of 
gonarcus; mu, mediuncus (= arcessus of Winterton and Brooks 2002); S7, seventh sternite; S8+9, fused eighth 
and ninth sternites; sa, subanale; T7, seventh tergite; T8, eighth tergite; T9-memb, very weakly sclerotized or 
membraneous ninth tergite. 


546 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Male abdominal characters among exemplars of Apochrysinae (Chrysopidae). Coding follows Win- 
terton and Brooks (2002). Data for Loyola croesus are added to those published by Winterton and Brooks (2002) 
and Tjeder (1966) * for the other species. 


EEE SSS 
Winterton and Brooks Character No. 


Species 30 31 32 33 34 35 


See ee ee 


Apochrysa 


montrouziert (Girard) ] (0) 0 0 10) 0 

lutea (Walker) 1 0 0 0) 0 0) 

voeltzkowi (Weele) 1 0 0 0 0 0) 

matsumurae (Okamoto) 1 1 0 1 0) 1 

wagneri (Hd6lzel) 1 1 (0) (0) ? 1 

retivenosa (Winterton) 1 0) 0 (0) (0) 1 

*africana (Kimmins) 1 0/1 1 0) 0) 0) 

*/eptalea (Rambur) 1 0 0 0 1 1 
Domenechus 

mirificus (Gerstaecker) 1 1 (0) (0) 0 1 
Joguina 

nicobarica (Brauer) 0 1 0 0 0 1 

constellatus (Navas) 0 1 (0) 0 0) 0) 
Loyola 

beata (Walker) ? ? 2 2 2 o) 

croesus (Gerstaecker) 1 1 1 0 0) 1 
Nobilinus 

albardae insignitus (Navas) 1 0) 0) 0) 0 0) 
Nothancyla 

verreauxi (Navas) 1 0 1 0 1 0) 


EE 


31. Male Ectoprocts and T9. (0) Fused. Apochrysa, neither of which is in the Jo- 
(1) Separate. guina Clade. In Nothancyla (= Anapochry- 

In L. croseus, T9 is very weakly sclero- sa) verreauxi Navas the entoprocesses are 
tized and appears to be separate from the well defined and shaped like those in L. 
ectoprocts (1). However, the condition is croesus, but they are less prominent (Tsu- 
not entirely clear on our specimen. kaguchi 1995, fig. 106i; mentioned in Win- 

A character state of male ectoprocts sep- terton and Brooks 2002; not visible in 
arate from T9 is consistent with that in oth- sketch by New, 1980, p. 88). In Apochrysa 
er members of the Joguina clade. But, itis africana (Kimmins) the entoprocesses are 
noteworthy that the trait can vary among much smaller (Tjeder 1966, fig. 1861) than 
species within a genus [as in Apochrysa in L. croesus. 


(Winterton and Brooks 2002)], and we sus- 33. Gonarcus. (0) Arcuate. (1) Trans- 

pect that it may vary with maturation. Thus, _ verse. 

re-evaluation across all species within the Within the Chrysopidae, an arcuate gon- 

subfamily is warranted. arcus is plesiomorphic. Except for Apoch- 
32. Entoprocesses. (0) Absent. (1) Pres- rysa matsumurae (Okamoto) (Winterton 

ent. and Brooks 2002), the condition holds for 


Distinct, upturned entoprocesses are pre- all known Apochrysinae, now including 
sent on the L. croesus gonarcus (1). The Loyola (0). 
other Apochrysinae genera that are reported 34. Mediuncus (= Arcessus) Shape. (0) 
to have entoprocesses are Nothancyla and _ Short, broad. (1) Elongate, narrow. 


VOLUME 107, NUMBER 3 


As in all other studied species of Apoch- 
rysinae (except N. verreauxi), the mediun- 
cus of L. croesus is relatively short, broadly 
based and tapered apically (0). 

35. Gonosetae on Arcessus. (O) Absent. 
(1) Present. 

Eight pairs of gonosetae occur on the L. 
croesus arcessus. In this respect, L. croesus 
resembles Domenechus, several Apochrysa 
species [but not A. africana (Kimmins), A. 
lutea (Walker), A. montrouzieri (Girard) or 
A. voeltzkowi (Weele)], and Joguina nico- 
barica (Brauer) [but not Joguina constellus 
(Navas)] (Tjeder 1966, Tsukaguchi 1995, 
Winterton 1995, Winterton and Brooks 
2002). Nothancyla verreauxi is reported to 
have gonosetae, but is not coded for them 
(Winterton and Brooks 2002). 

Conclusion.—Our specimen of L. croe- 
sus exhibits most of the male abdominal 
and genital characteristics that Brooks and 
Barnard (1990) used to distinguish the sub- 
family Apochrysinae (Table 1). The main 
exception, one that was previously noted by 
Winterton and Brooks (2002) for another 
genus of Apochrysinae, is the presence of 
entoprocesses on the gonarcus. Also, all of 
the male character-states of L. croesus, ex- 
cept one (the presence of entoprocesses on 
the gonarcus), are consistent with Winterton 
and Brooks’ (2002) placement of the genus 
in the Joguina clade. 

Our specimen of L. croesus has several 
features that were previously undescribed 
for any Apochrysinae: a very weakly scler- 
otized ninth tergite (that probably is not 
fused with the ectoprocts); ectoprocts that 
are connected dorsally via a broad band; a 
broadly arcuate, well sclerotized gonarcus 


547 


bearing a pair of upturned entoprocesses; 
and a broadly based, tapering arcessus bear- 
ing gonosetae. Whether these unique fea- 
tures are typical of the genus Loyola is un- 
known. We hope that males from the other 
Loyola species will become available for 
comparison. 


ACKNOWLEDGMENTS 

This work was supported, in part, by the 
National Science Foundation Grants INT- 
9817231 and INT-9912449 (MJT, CAT, 
GSA), National Geographic Society (MJT, 
CAT, GSA), Conselho Nacional de Desen- 
volvimento Cientifico e Tecnoldgico 
(GSA), Regional Project W-1185 (MJT, 
CAT), Cornell University (MJT, CAT), and 
Universidade Estadual do Norte Flumi- 
nense (GSA). We thank Shaun L. Winterton 
for helpful comments on the manuscript. 


LITERATURE CITED 


Brooks, S. J. and P. C. Barnard, 1990. The green lace- 
wings of the world: A generic review (Neuroptera: 
Chrysopidae). Bulletin of the British Museum of 
Natural History (Entomology) 59: 117-286. - 

New, T. R. 1980. A revision of the Australian Chry- 
sopidae (Insecta: Neuroptera). Australian Journal 
of Zoology. Supplemental Series 77: 1—43. 

Tjeder, B. 1966. Neuroptera-Planipennia. South Afri- 
can Animal Life, Vol. 12: 228-534. Statens Na- 
turvetenskapliga Forskningsrad, Stockholm 23. 

Tsukaguchi, S. 1995. Chrysopidae of Japan (Insecta, 
Neuroptera). S. Tsukaguchi, Aioi-cho 6-14-102, 
Nishinomiya-shi, Hyogo, 662 Japan. 

Winterton, S. L. 1995. A new genus and species of 
Apochrysinae (Neuroptera: Chrysopidae) from 
Australia, with a checklist of Australian Chryso- 
pidae. Journal of the Australian Entomological 
Society 34: 139-145. 

Winterton, S. L. and S. J. Brooks. 2002. Phylogeny of 
the apochrysine green lacewings (Neuroptera: 
Chrysopidae: Apochrysinae). Annals of the En- 
tomological Society of America 95: 16—28. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 548-553 


CULEX MALARIAGER, N. SP. (DIPTERA: CULICIDAE) FROM DOMINICAN 
AMBER: THE FIRST FOSSIL MOSQUITO VECTOR OF PLASMODIUM 


GEORGE POINAR, JR. 


Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. (e-mail: 
poinarg @ science.oregonstate.edu) 


Abstract.—A new species of fossil mosquito, Culex malariager (Culicidae: Diptera), 
is described from Dominican amber. This species differs from extant members of the 
genus by the following combination of characters: 1) proboscis shorter than antennae; 2) 
wing scales both long and narrow and short and broad; 3) base of R vein (remigium) 
with two groups of dorsal setae; 4) clypeus as long as broad; 5) empodium on foretarsus; 
6) vein 2A with rows of erect, short setae, and 7) postgenital lobe covering base of cerci. 
Inside the body cavity of C. malariensis are various stages (oocysts, sporozoites, ookinete, 
and microgametocyte) of a Plasmodium malarial parasite. This is the first record of a 


fossil mosquito vector of a vertebrate pathogen. 


Key Words: 
modium 


During an ongoing investigation of fossil 
haematophagous insects, a species of Culex 
in Dominican amber was discovered to con- 
tain sporogonic stages of a Plasmodium 
within its body cavity. The description of 
the Plasmodium will be reported in a sep- 
arate paper (Poinar, in press). The present 
study describes this fossil mosquito as a 
new species in the genus Culex L. 

Although not abundant, fossil culicids 
are known from Tertiary amber (Poinar 
1992) and a single specimen has been de- 
scribed from Cretaceous amber, along with 
a critical synopsis of previously described 
fossil mosquitoes (Poinar et al. 2000). 
Whereas Culex is the most common genus 
of mosquitoes in Dominican amber (Poinar 
and Poinar 1999), the only described mos- 
quito from these deposits is Anopheles 
dominicanus Zavortink and Poinar, 2000. 
Dating of Dominican amber is still contro- 
versial with the latest proposed age of 20— 
15 mya based on foraminifera (Iturralde- 


Culex malariager n. sp., Culicidae, Dominican amber, malaria vector, Plas- 


Vincent and MacPhee 1996) and the earliest 
as 45—30 mya based on coccoliths (Cépek 
in Schlee 1999). What makes dating of the 
amber difficult is that it is secondarily de- 
posited in turbiditic sandstones of the Up- 
per Eocene to Lower Miocene Mamey 
Group (Draper et al. 1994). The plant spe- 
cies that formed the amber is a member of 
the legume family (Hymenaea protera 
Poinar 1991) and the original environment 
was similar to a present day moist tropical 
forest (Poinar and Poinar 1999). 


MATERIALS AND METHODS 


The piece of amber originated from 
mines in the northern area of the Dominican 
Republic, between the cities of Puerto Plata 
and Santiago. It is trapezoidal in outline, 
measuring 14 X 17 X 25 X 15 mm on the 
sides, 5 mm in thickness and weighs 2 
grams. Observations and photographs were 
made with a Nikon stereoscopic microscope 
SMZ-10 R and Nikon Optiphot™ at mag- 


VOLUME 107, NUMBER 3 


nifications up to 600. All measurements 
are in microns unless otherwise noted. 
The fossil female mosquito is essentially 
complete, but somewhat distorted with 
many scales and bristles detached (except 
for those on the wings). Only three legs are 
attached to the body (both forelegs and one 
midleg), however the others are adjacent to 
the specimen. The thorax and abdomen are 
partially cleared, thus making it possible to 
see internal structures. Since the fossil 
could not be identified with any extant spe- 
cies it is described as a new species below. 


Culex malariager Poinar, new species 
(Figs. 1-10) 


Description.—Size small, head bent 
down partly under thorax; length from 
pronotum to tip of abdomen 2.9 mm (how- 
ever head, thorax and abdomen measured 
separately, length 4.1 mm). 

Head: Length 0.59 mm, width 0.66 mm; 
vertex and occiput bearing 76 large aveoli 
(38 on each side; positions vary slightly 
from side to side); clypeus rounded in front, 
without scales or setae, length 136 pm, 
width 133 jm; eyes nearly contiguous with 
narrow interocular space; eye facets mea- 
suring approximately 18 x 21 wm; maxil- 
lary palpus with 3 palpomeres, no banding 
obvious, palpomeres 2 and 3 bear setae and 
few small scales, palpomere | and palpifer 
bare; lengths, palpomere 1 (with palpifer) 
95 wm; palpomere 2, 123 ym; palpomere 
3, 155 wm; proboscis length 1.53 mm, pro- 
boscis covered with mostly broad, but some 
narrow scales; labellum length 136 wm, la- 
bellum width 35 wm; distal and proximal 
sclerites of labellum apparently fused; lig- 
ula pointed; antenna length 1.90 mm; scape 
bare, length 32 ym; pedicel globose, with 
few setae; pedicel length 116 jm, pedicel 
width 123 pm; flagellomeres verticillate, 
however most setae absent; basal and ter- 
minal flagellomeres equal in length; lengths 
of flagellomeres: 1, 175 ym; 2, 140 wm; 3, 
168 wm; 4, 161 wm; 5, 161 pm; 6, 154 wm; 
7, 147m; 8, 154 wm; 9, 154 pm;10, 140 
tome el OSeum: 12% 105 poms 135 175° pm: 


549 


Thorax: Length 1.22 mm; antepronotum 
narrow, with 6 aveoli; scutal fossa (pres- 
cutum) with 6 aveoli; scutum with both se- 
tae and alveoli; scutellum (length 85 pm; 
width 372 jm) with 3 lobes bearing 10 av- 
eoli; no spiracular, post- spiracular, acros- 
tichal or postnotal setae; 3 prealar setae and 
4 aveoli on mesokatepisternum; 2 lower 
mesepimeral setae; wing length 2.7 mm, 
greatest width 0.7 mm; microtrichia pre- 
sent, posterior margin of wing with fringe 
of scales; wing banding not visible; cell R, 
longer than its stem (R ,,,); cell M almost 
twice as long as its stem; br and bm cells 
extend to middle of wing; anal vein termi- 
nates well past fork of CuA; plume scales 
on veins R, and R, narrow to slightly wid- 
ened; Sc and R contiguous for 360 ym from 
wing base; base of vein R with 2 groups of 
dorsal setae; group 1 with 7 setae 32-79 
zm long occurring 204 ym from wing base, 
group 2 with 8 setae 23-57 pm long oc- 
curring 45 wm from wing base; veins Rs 
and R ,,; without basal spurs; crossveins 
without scales; r-m distad to m-cu; alula 
(length 196 p.m) with outer row of scales; 
no setae on lower surface of wing at base 
of subcostal vein; scales on wing veins vary 
in length and breadth, scales on vein R, R 
5.3, R, and R,, distal portions of veins A , 
and CuA, long and narrow, scales on R,, R 
415, M,, M, and basal portions of vein A, 
and CuA, short and broad; other veins with 
intermediate scales; vein 2A without scales 
but bearing rows of short setae 7—10 .m in 
length (setae differ in structure and size 
from wing microtrichia); halter length 315 
wm; legs brown-scaled, no pattern or or- 
namentation visible; forecoxa with few al- 
veoli near apex; forefemur swollen slightly 
at base, length 1.38 mm; foretibia 1.8 mm; 
ventral surface of foretarsomere 1 with 
short spine, opposite surface with scales 
and longer spines; length of tarsomeres of 
right fore tarsus; 1, 1.5 mm; 2, 0.57 mm; 3, 
0.36 mm; 4, 0.157 mm; 5, 0.14 mm; claw 
length 50 ppm; pulvillus length 20 ppm; for- 
etarsus with slender empodium arising be- 
tween pulvilli, its filament short with a sin- 


550 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ee ee 


Figs. 1-4. Culex malariager in Dominican amber. 1, Anterolateral view. Bar = 0.9 mm. 2, Scales at the 
base of veins R2 and R3. Bar = 45 wm. 3, Labella with prementum (P), closely adjacent distal and proximal 
sclerites (S) and ligula (L). Bar = 45 pm. 4, Foretarsomere with simple claw (C), pulvillus (P) and empodium 
(arrow). Bar = 23 wm. Insert shows a portion of vein 2A bearing rows of short, erect setae. Bar = 30 pm. 


VOLUME 107, NUMBER 3 


Nn 
Nn 
— 


Figs. 5-10. Culex malariager in Dominican amber. 5, Right wing (ventral view) with venation. Bar = 210 
wm. 6, Dorsal surface of head with aveoli on occiput and vertex. Bar = 70 pm. 7, Cerci, postgenital lobe, and 


gonotreme. Bar = 49 pum. 8, Right flagellomeres. Bar 


10, Aveoli on trilobed scutellum. Bar = 114 pm. 


gle branch, basal sclerite not visible, em- 
podium not seen on mid- and hindtarsi, all 
pulvilli with fine setae on lower surface; 
midtarsomere 5 length 0.13 mm; claw 
length 46 pm; pulvillus length 17 pm; 
hindtarsomere 5 length 0.15 mm; claw 
length 23 pm; pulvillus length 18 pm; all 
claws paired, simple, equal. 

Abdomen: Length 2.3 mm; covered with 
mostly short, broad scales; tergum 8 with 


252 wpm. 9, Aveoli on antepronotum. Bar = 70 p.m. 


very numerous long setae and scales sur- 
rounding cerci; cercus length 50 zm; cercus 
width at base 26 jm; postgenital lobe large, 
covering bases of cerci. 

Ratios: Palpus to proboscis 0.24; pro- 
boscis to antenna 0.81; palpus to clypeus 
2.7; proboscis to forefemur 1.11; clypeus to 
pedicel 1.17. 

Material examined.—Holotype female in 
Dominican amber, deposited in the Poinar 


552 


amber collection maintained at Oregon 
State University (accession number D-7— 
6B). 

Type locality—La Toca amber mine in 
the Dominican Republic. 

Etymology.—‘“‘malariager”’ is based on 
the vector association of the fossil mosquito 
with Plasmodium. It is formed with the Lat- 
in suffix —ger meaning bear, carry or have. 

Diagnosis.—The following characters 
place C. malariager in the genus Culex: 1) 
short palps; 2) tip of abdomen blunt; 3) scu- 
tellum trilobed with each lobe bearing sep- 
arate tufts of setae; 4) no spiracular or 
postspiracular setae; 5) short, partly con- 
cealed cerci; 6) claws simple; 7) pulvilli 
present; 8) vein 1A long; 9) wing microtri- 
chia distinct; 10) eyes nearly contiguous; 
11) terga and sterna II—VII uniformly 
scaled; 12) cell R2 longer than vein R,,;; 
and 13) hairs on underside of costal vein 
absent. It is not possible to assign this spe- 
cies to an extant subgenus since these taxa 
are defined on the types of scales found at 
particular locations, scale and setal posi- 
tions on the thorax, and color of scales, ab- 
domen and pleural integument (Berlin and 
Belkin 1980, Casal and Garcia 1968, Har- 
bach and Peyton 1992), most of which can- 
not be determined in the fossil. 

The species can be distinguished from 
extant Culex by the following combination 
of characters: 1) proboscis shorter than an- 
tennae; 2) wing scales both long and narrow 
and short and broad; 3) base of vein R (re- 
migium) with two groups of dorsal setae; 
4) clypeus as long as broad; 5) empodium 
on foretarsus; 6) vein 2A with rows of 
erect, short setae; and 7) postgenital lobe 
covering base of cerci. Especially signifi- 
cant are items 3 and 6 since they do not 
appear to have been reported previously on 
any Culex spp. (Tom Zavortink, personal 
communication). 


DISCUSSION 


Two other Culex fossils are known from 
North America, C. damnatorum Scudder, 
1890 from Eocene Green River deposits in 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Wyoming and C. winchesteri Cockerell, 
1919 from Eocene deposits in Colorado. 
Neither of these is similar to C. malariager. 

There were two other hematophagous in- 
vertebrates in the same piece of amber, a 
triatomid bug and a female Ornithodorus 
dominicana Poinar, 1995. The presence of 
these three fossils suggests that the resin 
was produced in or close to a domicile of 
vertebrates, which probably served as hosts 
to these parasites. Since the stages of the 
malarial parasite inside C. malariager re- 
semble those of an extant avian Plasmodi- 
um species (Poinar, in press) and it is 
known that Culex transmit avian malaria 
(Riper et al. 1994), at least one of the ver- 
tebrate hosts of C. malariager was a bird. 
This is the first record of a fossil mosquito 
vector of a vertebrate pathogen. 


ACKNOWLEDGMENTS 


I thank Tom Zavortink for discussions on 
mosquito systematics and literature and 
reading a draft copy of this manuscript and 
Roberta Poinar for comments on the man- 
uscript. I also appreciate the help of two 
anonymous reviewers. 


LITERATURE CITED 


Berlin, O. G. W. and J. N. Belkin. 1980. Mosquito 
studies (Diptera, Culicidae). XXXVI. Subgenera 
Aedinus, Tinolestes and Anoedioporpa of Culex. 
Contributions of the American Entomological In- 
stitute (Ann Arbor) 17(2): 1-104. 

Casal, O. H. and M. Garcia. 1968. Culex (Allimanta), 
nuevo subgenero para Culex (Culex) tramazay- 
guesi Duret, 1954 (Diptera, Culicidae). Physis 
(Bueno Aires) 27: 329-335. 

Cockerell, T. D. A. 1919. The oldest mosquitoes. Na- 
ture 103: 44. 

Draper, G., P. Mann, and J. KF Lewis. 1994. Hispaniola, 
pp.129—150. In Donovan, S. and T. A. Jackson, 
eds. Caribbean Geology: An Introduction. The 
University of the West Indies Publishers’ Associ- 
ation, Kingston, Jamaica. 

Harbach, R. E. and E. L. Peyton. 1992. A new sub- 
genus of Culex in the Neotropical Region (Dip- 
tera: Culicidae). Mosquito Systematics 24: 242— 
252. 

Iturralde-Vincent, M. A. and R. D. E. MacPhee 1996. 
Age and Paleogeographic origin of Dominican 
amber. Science 273: 1850-1852. 

Poinar Jr., G. O. 1991. Hymenaea protera sp. n. (Leg- 


VOLUME 107, NUMBER 3 


uminoseae, Caesalpinioideae) from Dominican 

amber has African affinities. Experientia 47: 

1075-1082. 

1992. Life in Amber. Stanford University 

Press, Stanford, California, 350 pp. 

. 1995. First fossil soft ticks, Ornithodoros an- 

tiquus n. sp. (Acari: Argasidae) in Dominican am- 

ber, with evidence of their mammalian host. Ex- 

perientia 51: 384-387. 

. In press. Plasmodium dominicana n. sp., 
(Plasmodiidae: Haemospororida) from Tertiary 
Dominican amber. Systematic Parasitology. 

Poinar, Jr., G. O. and R. Poinar. 1999. The Amber For- 
est. Princeton University Press, Princeton, 239 pp. 

Poinar, Jr., G. O., T. J. Zavortink, T. Pike, and P. A. 
Johnston. 2000. Paleoculicis minutus (Diptera: 


553 


Culicidae) n. gen., n.sp., from Cretaceous Cana- 
dian amber, with a summary of described fossil 
mosquitoes. Acta Geologica Hispanica 35: 119— 
128. 

Riper III, C. van, C. T. Atkinson, and T. M. Seed. 1994. 
Plasmodia of Birds, pp. 73-140. Jn Kreier, J. P. 
ed. Parasitic Protozoa (2™ Edition), Vol. 7. Aca- 
demic Press, San Diego. 

Schlee, D. 1990. Das Bernstein-Kabinett. Stuttgarter 
Beitrager fur Naturkunde, Ser. C. Vol. 28: 100 pp. 

Scudder, S. H. 1890. The Tertiary insects of North 
America. Report of the United States Geological 
survey of the Territories 13: 1—734. 

Zavortink, T. J. and G. O. Poinar, Jr., 2000. Anopheles 
(Nyssorhynchus) dominicanus sp. n. (Diptera: Cu- 
licidae) from Dominican amber. Annals of the En- 
tomological Society of America 93: 1230-1235. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 554-564 


NONINDIGENOUS WOODBORING COLEOPTERA 
(CERAMBYCIDAE, CURCULIONIDAE: SCOLYTINAE) NEW TO OREGON 
AND WASHINGTON, 1999-2002: CONSEQUENCES OF THE 
INTRACONTINENTAL MOVEMENT OF RAW WOOD PRODUCTS AND 
SOLID WOOD PACKING MATERIALS 


J. R. LABONTE, A. D. MUDGE, AND K. J. R. JOHNSON 


Plant Division, Oregon Department of Agriculture, 635 Capitol Street, Salem, OR, 
97301-2532, U.S.A. (e-mail: jlabonte @ oda.state.or.us) 


Abstract.—Urban forests, port areas, mills and businesses known to have received or 
handled imported wood or wood products were surveyed for nonindigenous woodboring 
insects in Oregon and southernmost western Washington from 1999—2002, predominantly 
using Lindgren funnel traps, Intercept™ panel traps and/or Scots pine bait logs. Several 
other woodborer surveys or projects, using various traps and lures, also took place con- 
currently. Eight species of nonindigenous woodboring beetles new to Oregon, Washington, 
the western U.S., western North America, or North America are recorded for the first 
time: Phymatodes testaceus (L.), Tetropium castaneum L., Xylotrechus hircus (Gebler), 
and X. sagittatus sagittatus (Germar) (Cerambycidae), Monarthrum fasciatum (Say), Xy- 
losandrus crassiusculus (Motschulsky), X. germanus (Blandford), and an undetermined 
species of Xyleborus (Curculionidae: Scolytinae). Additional records are included for the 
following nonindigenous woodborers detected in 1997—1998 and reported in an earlier 
paper: Gnathotrichus materiarius (Fitch), Hylastes opacus Erichson, Xyleborinus alni (Ni- 
isima), Xyleborus californicus Wood, X. pfeili (Ratzeburg) (Scolytinae), and Xiphydria 
prolongata (Geoffroy) (Hymenoptera: Xiphydriidae). Seventy-five percent of the nonin- 
digenous woodborers treated in this and our earlier paper are known from both eastern 
and western North America. We believe these western records of five eastern indigenous 
species and eight extracontinental exotic species established in the East are evidence of 
the intracontinental movement of untreated domestic solid wood packing material and 
other raw woods as the probable pathways for these species to the West. 


Key Words: nonindigenous, woodborers, Cerambycidae, Scolytinae, Xiphydriidae 


Nonindigenous woodboring and wood- 
associated insects (NIWBI) introduced and 
established in North America and the Unit- 
ed States continue to be found at an alarm- 
ing rate (Atkinson et al. 1990, Hoebeke 


shrubs, quarantines of wood products and 
nursery stock, implementation of integrated 
pest management programs, and continuing 
survey, eradication, and research efforts 
(e.g., Haack et al. 1996, Haack et al. 1997, 


1994, Humble 2001, Mudge et al. 2001, 
Rabaglia and Valenti 2003). While the ef- 
fects of many of these species are unknown, 
several have already had significant eco- 
nomic impacts through damage to trees and 


Oliver and Mannion 2001, Poland et al. 
1998, McCullough and Roberts 2002). 
Such organisms threaten the health of North 
American forests, as well as other urban, 
rural, and natural environments (U.S. Con- 


VOLUME 107, NUMBER 3 


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gress, OTA 1993, Liebhold et al. 1995, 
Campbell and Schlarbaum 2002). 

As part of the USDA’s Cooperative Ag- 
ricultural Pest Survey (CAPS) program, the 
Oregon Department of Agriculture (ODA) 
has conducted surveys since 1997 at sites 
at risk for the introduction of NIWBI. The 
results of the first two years (1997 and 
1998) were reported in Mudge et al. (2001), 
where eight species of NIWBI were docu- 
mented from Oregon, Washington, the 
western United States, or North America 
for the first ttme. Surveys of high-risk sites 
continued in Oregon and southernmost 
western Washington from 1999 through 
2002. In addition, a research project was 
initiated in 2002 by the USDA Forest Ser- 
vice (USFS) in cooperation with ODA to 
develop more effective monitoring methods 
for ambrosia beetles (Curculionidae: Scol- 
ytinae). New state, regional, or continental 


MALHEUR 


HARNEY 


Survey sites for nonindigenous woodboring insects in Oregon 1997-2002. 


records from these projects reported here 
include eight species of nonindigenous 
woodboring Coleoptera (Cerambycidae and 
Scolytinae). Additional distributional re- 
cords for most of the species treated in 
Mudge et al. (2001) are included. Our belief 
that these records are evidence of the move- 
ment of nonindigenous woodboring insects 
throughout North America via the intracon- 
tinental movement of untreated solid wood 
packing material (SWPM) and raw wood 
products (RWP) is discussed. 

High-risk sites surveyed included ware- 
houses and businesses importing commod- 
ities with SWPM (this refers to all types of 
solid wood packing material, including 
crating, dunnage, pallets, etc.), mills im- 
porting RWP, wood recyclers, port and in- 
dustrial areas, and urban Of the 
sites surveyed, 57% were in the Portland 
metropolitan area (Clackamas, Multnomah, 


forests. 


556 


and Washington counties). However, addi- 
tional high-risk sites were surveyed in 14 
other counties in Oregon (Fig. 1), as well 
as Clark and Klickitat counties in southern 
Washington. 

Survey methods were essentially those of 
Mudge et al. (2001). Lindgren multiple- 
funnel traps (12-funnel), hereafter referred 
to simply as funnel traps, were the primary 
survey means. Three “‘standard traps”’ were 
placed per site, baited, respectively, with 
exotic Jps spp. lures (IPS: ipsdienol, methyl 
butenol, and cis-verbenol), woodborer lures 
(aP—EtOH: ultra-high release [UHR] a- 
pinene and UHR ethanol pouches), or UHR 
ethanol lures (EtOH) (Phero Tech, Inc., 
Delta, BC). In 2002, lures for specific target 
species were also employed at various sites, 
sometimes independently and sometimes in 
conjunction with the standard traps. Specif- 
ic lures were a five-component, pine shoot 
beetle (Tomicus piniperda (L.): Scolytinae) 
lure (PSB: a-pinene, a-pinene oxide, myr- 
tenol, nonanol, and trans-verbenol, IPM 
Tech, Inc., Portland, OR) and an experi- 
mental blend of spruce volatiles (SV: a- 
pinene, 3-carene, limonene, and turpinole- 
ne, Phero Tech) for Tetropium castaneum 
(E) (Cerambycidae). 

The standard traps were deployed from 
February through mid-October. Traps tar- 
geting specific taxa were deployed and 
maintained over different periods, based 
upon the target taxa life histories, 1.e., Feb- 
ruary through late May for pine shoot bee- 
tle and mid-April through mid-October for 
T. castaneum. All traps were monitored bi- 
weekly. 

Intercept™ panel traps (IPM Tech) (here- 
after referred to as panel traps) were used 
in 2000 in a survey for nonindigenous 
wood-associated insects that could vector 
exotic plant pathogens as well as at all sites 
trapped for 7. castaneum in 2002. The 
USFS and ODA cooperative ambrosia bee- 
tle monitoring project utilized funnel traps, 
panel traps, and Japanese beetle traps (Tre- 
ce, Inc., Salinas, CA) baited with aP-EtOH 
lures. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Bait logs of Scots pine (Pinus sylvestris 
L.), a preferred host of pine shoot beetle, 
were also placed at most trap sites through 
2001. In 2002, this method was replaced by 
the use of funnel traps baited with PSB 
lures. Similarly, bait logs of Sitka spruce, 
Picea sitchensis (Bong.), were used at all 
of the sites trapped for 7. castaneum in 
2001 and many of the sites trapped for this 
species in 2002. Spruce logs were the sole 
survey method at some 7. castaneum sites 
in 2002. Upon removal from the field, bait 
logs were placed in emergence tubes and 
monitored for target species emergence. 
Pine shoot beetle bait logs were removed 
from survey sites in late April and early 
May and monitored through at least July of 
the same year in which they were deployed. 
Bait logs for 7. castaneum were removed 
in August and monitored through August of 
the following year. Unless otherwise noted, 
all collection records refer to specimens 
trapped in funnel traps and are reported in 
the following format: state, county, city, site 
type (in parentheses), collection date, lure. 
The number of specimens collected, by lure 
type, are in parentheses. 

Target taxa were initially identified by 
LaBonte, based upon available literature 
and comparison with identified material in 
the insect collections of the ODA (ODAC), 
Salem, OR, or the Oregon State Arthropod 
Collection (OSAC), Oregon State Univer- 
sity, Corvallis, OR. Cerambycidae and 
Scolytinae identifications were also provid- 
ed or confirmed by several taxonomic au- 
thorities (see Acknowledgments). Unless 
otherwise stated, specimens collected in the 
Oregon surveys are housed in the ODAC. 


NEw RECORDS 


The following data represent, to the best 
of our knowledge, the first published re- 
cords of these species from the specified re- 
gion. 


VOLUME 107, NUMBER 3 


COLEOPTERA 
Cerambycidae 
Phymatodes testaceus (L.) 


Records.—Oregon: Clackamas Co., Port- 
land (SWPM importer), 12 VII 2000, EtOH 
(1); Columbia Co., Scappoose (wood re- 
cycler), | VII 2002, aP-EtOH (1); Mult- 
nomah Co., Portland (SWPM importer, 
wood recyclers), 6 VII 1998, 17 VI & 22 
VII 1999, 26 VI & 12 VII 2000, 12 & 26 
VI 2002, aP-EtOH (3), IPS (1), EtOH (6). 
Washington: Clark Co., Vancouver (port 
area), 17 VI 1999, aP-EtOH (1). 

Comments.—These data represent new 
western U.S. and state records for OR and 
WA. This species has been previously re- 
corded from Europe, North Africa, and 
eastern North America, west to MN and IA 
(Linsley 1964) and recently has been found 
in BC (Humble 2001). It is probably a non- 
indigenous species in North America 
(Bousquet 1991). Oaks are preferred hosts, 
but other hosts include coniferous and de- 
ciduous trees, e.g., hemlock, spruce, apple, 
beech, cherry, chestnut, hickory, and wil- 
low (Linsley 1964). Our surveys suggest P. 
testaceous is widespread throughout the 
Portland-Vancouver metropolitan area. The 
1998 specimen was not determined until af- 
ter publication of Mudge et al. (2001). 


Tetropium castaneum L. 


Records.—Oregon, Wasco Co., The 
Dalles (mill site), 5 VI & 5 VII 2000, aP- 
EtOH (2). 

Comments.—These are the first records 
of this species being trapped in North 
America and OR. This species has previ- 
ously been intercepted in dunnage and 
SWPM entering the United States (includ- 
ing OR) and Canada (Humble et al. 2002, 
USDA APHIS PPQ, Port Interception Net- 
work). It is a Palearctic species known from 
most of Europe, northeastern Asia, China, 
and Japan (Bense 1995; M. Rejzek, unpub- 
lished data). Hosts are conifers, primarily 
spruce, but also include fir, larch, and pine 
(Bense 1995, Rejzek and Rebl 1999). 


Sa)// 


Adults attack stressed, dying, or recently 
dead trees (Juutinen 1955, Bense 1995). 

Delimitation surveys were conducted for 
T. castaneum by ODA in 2001 and 2002. 
In 2001, funnel traps baited with aP-EtOH 
were deployed at 17 sites throughout The 
Dalles, centered upon the original trap site. 
In 2002, 46 sites were surveyed over a 
somewhat larger area, using panel traps 
baited with SV lure. Spruce bait logs were 
used in conjunction with the traps both 
years. Traps and logs were placed near po- 
tential hosts. A similar, but much smaller 
preliminary survey for 7. castaneum was 
also conducted in 2002 in the vicinity of a 
Port of Portland terminal where freshly 
dead adult 7. castaneum were found with 
recently fumigated raw SWPM originating 
from the Russian Far East. No 7. castaneum 
were trapped in 2001 and 2002, nor did any 
T. castaneum emerge from the spruce bait 
logs deployed in 2001 and 2002. 

It is interesting to note that no RWP of 
Palearctic origins were known to have been 
received at the mill in The Dalles for at 
least a year prior to the trapping of 7. cas- 
taneum. The mill’s records for 1999 and 
2000 indicated raw railroad ties from ID, 
MO, TX, WA, and Canada were received, 
raising the possibility of an undetected pop- 
ulation of 7. castaneum in one of those re- 
gions. 


Xylotrechus hircus (Gebler) 


Multnomah Co., 
1999, 


Records.—Oregon, 
Portland, (wood recycler), 22 VII 
EtOH (1): 

Comments.—This is the first record of 
this species from North America and OR. 
This Palearctic species is recorded from 
northern China, Japan, Korea, northern 
Mongolia, eastern Siberia, Kazakhstan, and 
the Sakhalin and Kurile Islands (Svacha 
and Danilevsky 1987; M. Rejzek, unpub- 
lished data). Larvae feed in the dead wood 
of birch (Svacha and Danilevsky 1987) and 
possibly other deciduous trees. This species 
has been rarely collected in its native haunts 
(M. Rejzek, personal communication). In 


558 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


2000, additional traps were placed in the 
vicinity of the original trap site and the site 
was trapped via the standard ODA proto- 
cols in 2001. No further specimens have 
been trapped. 


Xylotrechus sagittatus sagittatus (Germar) 


Records.—Oregon, Wasco Co., The 
Dalles (residential area), 25 IX 2001, aP- 
IBiOlal (il). 

Comments.—This is a new western 
North American and OR record. This beetle 
is native to eastern North America, from 
eastern Canada south to FL and west to NM 
(Linsley 1964). Hosts are conifers, includ- 
ing fir, pine, and spruce (Linsley 1964). 

The specimen was trapped in a 2001 T. 
castaneum delimitation trap. No additional 
specimens have been trapped, despite the 
numerous 7. castaneum delimitation traps 
in The Dalles in 2002. ‘ 


CURCULIONIDAE: SCOLYTINAE 
Monarthrum fasciatum (Say) 


Records.—Oregon, Wasco Co., The 
Dalles (mill site), 23 IV 1999 & 1 V 2000, 
aP-EtOH, (1; panel trap, 1). 

Comments.—These data represent the 
first western North American and OR re- 
cords for M. fasciatum. This species is a 
native of eastern North America, west to 
TX, WI, and ON (Wood and Bright 1992, 
Bright and Skidmore 1997). Typical hosts 
are deciduous hardwoods, but it has also 
been recorded from pine (Wood and Bright 
1992, Solomon 1995) and grape (Bright 
and Skidmore 1997). Despite further trap- 
ping in The Dalles, this species was not col- 
lected in 2001 or 2002. 


Xylosandrus crassiusculus (Motschulsky) 


Records.—Oregon, Wasco Co., The 
Dalles (mill site), all sampling periods from 
18 VIII-8 X 1999, 5 VI & 19 VI 2000, aP- 
Et@HiG3) EtOH Vda): 

Comments.—These data represent the 
first western North American records from 
traps outside of warehouses (X. crassius- 
culus has been intercepted in SWPM and 


trapped near SWPM in warehouses in Brit- 
ish Columbia—see Humble 2001) and the 
first OR records. This African and Asian 
species was first documented from SC in 
1974 and is also known from FL, GA, LA, 
MS, NC, TN, and TX (Solomon 1995, Ol- 
iver and Mannion 2001). While hosts in- 
clude over 200 woody angiosperms, this 
species shows some preference for sweet- 
gum (Solomon 1995). It is one of the two 
major ambrosia beetle species attacking 
chestnut in TN (Oliver and Mannion 2001). 
It has also been recorded as attacking at 
least one species of pine (Wood and Bright 
1992). Both healthy and stressed hosts, as 
well as freshly cut material, are attacked 
(Solomon 1995). Despite the placement of 
traps at prior positive sites, as well as nu- 
merous 7. castaneum delimitation traps in 
the general vicinity, this species was not 
collected in 2001 or 2002. 


Xylosandrus germanus (Blandford) 


Records.—Oregon, Multnomah Co., 
Portland (SWPM importer, urban forest, 
wood recycler), 4 VI & 22 VII 1999, 24 IV 
2000, 29 V & 17 VI 2002, aP-EtOH (4), 
EtOH (2); Washington Co., Hillsboro 
(SWPM importer), 27 ITV 2001, aP-EtOH 
(1). 

Comments.—These data represent new 
western U.S. and OR records. This Asian 
species has been introduced into Europe 
and eastern North America (Wood and 
Bright 1992), as weil as British Columbia 
(Bright and Skidmore 1997, Humble 2001). 
First documented from NY in 1932, it has 
since been recorded as far west as MI in the 
U.S. (Solomon 1995). Over 200 species of 
broadleafed and coniferous trees and shrubs 
are hosts, although angiosperms are pre- 
ferred (Solomon 1995). Along with X. cras- 
siusculus, this is one of the two major am- 
brosia beetles attacking chestnut in TN (OI- 
iver and Mannion 2001). As with the pre- 
ceding species, this species attacks vigorous 
as well as stressed hosts and recently cut 
material (Solomon 1995). 


VOLUME 107, NUMBER 3 


Xyleborus sp. undetermined 


Records.—Oregon, Washington Co., 
Hillsboro (SWPM importer), 28 VI 2000, 
aP-EtOH (1). 

Comments.—This specimen was sent to 
Lawrence Kirkendall and Stephen Wood for 
identification. Both concluded it was an ex- 
otic species with which they were unfamil- 
iar and ultimately decided that no species 
determination could be made at this time. 
However, they stated that it was apparently 
related to X. volvulus (L.), suggesting that 
it may have tropical or subtropical origins. 
Traps placed at the trap site in 2001 and 
2002 yielded no additional specimens. 


ADDITIONAL RECORDS 


The following data represent additional 
records for nonindigenous species previous- 
ly reported in Mudge et al. (2001). An ab- 
breviated format is used for the numerous 
records of Xyleborinus alni, Xyleborus cal- 
ifornicus, and X. pfeili. 


COLEOPTERA 
Curculionidae: Scolytinae 
Gnathotrichus materiarius (Fitch) 


Records.—Oregon, Wasco Co., The 
Dalles (mill site), all sampling periods from 
23 I-22 VII 1999, 31 V 2002, aP-EtOH 
(41), IPS (167), EtOH (1). 

Comments.—These specimens were 
trapped from the same locality where this 
species was first found in OR in 1998. No 
G. materiarius were collected when the 
same sites were trapped in 2000 and 2001, 
although a single specimen was trapped in 
2002. 


Hylastes opacus Erichson 


Records.—Oregon: Clackamas Co., Port- 
land (SWPM importers), 6 [V—9 V 2000, 
aP-EtOH (6), IPS (1); Marion Co., Salem 
(wood recycler), 1 V 2001, aP-EtOH (1); 
Multnomah Co., Portland (port vicinity, 
wood recyclers), 24 IV 2000, 29 III 2001, 
bait log (1), aP-EtOH (2; panel trap, 1), 
EtOH (1); Washington Co., Tualatin 


559 


(SWPM importer), 12 ITV 2001, aP-EtOH 
(DY: 

Comments.—These data include three 
new OR county records: Clackamas, Mari- 
on, and Washington. Initial OR detections 
of this species were from bait logs of Pinus 
sylvestris placed in early 1997. No speci- 
mens were found in bait logs or traps in 
19985 1999" onm2 002: 


Xyleborinus alni (Niisima) 


Records.—Oregon: Benton Co., Adair 
(wood recycler), 1 III 2002, EtOH (1); 
Clackamas Co., Portland (SWPM import- 
ers), 23 III & 6 IV 2000, aP-EtOH (2), IPS 
(1); Columbia Co., Scappoose (wood re- 
cycler) 25 IV 2000) aP-EtOHi(@); IPSi@): 
Linn Co., Sweet Home (mill site), 23 HI & 
9 IV 2001, EtOH (7); Marion Co., Salem 
(wood recycler), 26 III 2001, aP-EtOH (1), 
EtOH (1); Multnomah Co., Portland 
(SWPM importers, urban forests, ware- 
houses, wood recyclers), 27 II—-20 V, 20 IX 
& 16 X, 1999-2002, aP-EtOH (45), IPS 
(8), EtOH (109); Polk Co., Salem (nursery), 
10 V 2001, in the trunk of flowering cherry 
Prunus “Canada Red” (1); Washington 
Co., Portland and surrounding areas 
(SWPM importers, urban forest, wood re- 
cyclers), 13 I-20 V & 24 VII, 1999-2002, 
aP-EtOH (16), IPS (7), EtOH (17), PSB 
(8); Yamhill Co., Newberg (wood recycler 
vic.), 28 III 2002, PSB (1). Washington: 
Clark Co., Vancouver (port area), 24 Ill 
1999 & 6 IV 2000, IPS (1), EtOH (2). 

Comments.—These data include seven 
new OR county records (Benton, Clacka- 
mas, Linn, Marion, Polk, Washington, and 
Yamhill) and a new WA county record 
(Clark Co.). Formerly thought to be restrict- 
ed to a relatively modest area in Portland 
and vicinity, this species now appears to be 
widespread throughout much of the Wil- 
lamette Valley of northwestern OR. The 
specimen extracted from a nursery’s flow- 
ering cherry is the first record of X. alni 
from a species of Prunus in North America. 


5600 


Xyleborus californicus Wood 


Records.—Oregon: Benton Co., Adair 
(wood recycler), 4 VI 2002, EtOH (2); 
Clackamas Co., Portland (SWPM import- 
ers), 24 IV—12 VI 2000, 26 VIL & 23 VIII 
2000, aP-EtOH (3), IPS (3), EtOH (10); 
Columbia Co., Scappoose (wood recycler), 
8 IV—28 VI 2000, aP-EtOH (1), IPS (24), 
EtOH (24); Hood River Co., Hood River 
(rural forest), 31 V & 12 VII 2002, EtOH 
(5); Lane Co., Eugene (industrial area/urban 
forest), 30 V 2000, aP-EtOH (1); Jasper 
(mill site and vicinity), 21 IV—21 VII 1999, 
aP-EtOH (1), EtOH (30); Linn Co., Albany 
(SWPM importer), 25 V & 8 VI 1999, 
EtOH (6); Sweet Home (mill site), 20 IV 
2001, EtOH (1); Multnomah Co., Portland 
(customs warehouse, SWPM importers, 
wood recyclers), 11 IV—26 V & 9-28 VIII, 
1999-2002, aP-EtOH (1), IPS (4), EtOH 
(14), PSB (1); Washington Co., Portland 
and surrounding areas (SWPM importers, 
urban forest, wood recyclers), 15 [I-22 
VII, 1999-2002, aP-EtOH (16), IPS (5), 
EtOH @5) eSB Gd): 

Comments.—These data include six new 
OR county records: Benton, Clackamas, 
Columbia, Hood River, Linn, and Washing- 
ton. First recorded from OR based upon a 
specimen collected in Marion Co. in 1980 
(Wood 1982), this species now appears to 
be widely distributed in northwestern OR. 


Xyleborus pfeili (Ratzeburg) 


Records.—Oregon: Lane Co., Jasper 
(mill site and vicinity), 2 VI-7 X 1999, 30 
V 2000, aP-EtOH (panel trap, 1), EtOH 
(25); Linn Co., Sweet Home (mill site), 26 
VII 2001, EtOH (1); Marion Co., Salem 
(SWPM importer), 4 VI-28 IX 2001, aP- 
EtOH (4), IPS (2), EtOH (40); Multnomah 
Co., Portland (exotic woods importer, port 
vicinity, SWPM importers, urban forests, 
wood recycler), 26 V—3 VI, 23 VIII-19 Ix 
& 19 X 1999-2002, aP-EtOH (2), EtOH 
(5), SV (1); Washington Co., Tualatin 
(SWPM importer), 24 V 2001, EtOH (1). 
Washington, Clark Co., Vancouver (wood 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


recycler), 7 VI & 3 VUI-1 X 2001, aP- 
EtOH (1), EtOH (5). 

Comments.—These data include four 
new OR county records (Linn, Marion, 
Multnomah, and Washington) and a new 
WA state record. In an effort to delimit the 
extent of the populations of X. pfeili and X. 
xylographus (Say) detected in 1997 and 
1998, additional funnel traps were placed in 
1999 near the Jasper, Lane Co. (OR) sites 
where these species were originally detect- 
ed. Four additional traps, one at each car- 
dinal point, were placed approximately 2 
km from the original sites. Xyleborus pfeili 
was found in all four delimitation traps in 
the Jasper vicinity, indicating it is well es- 
tablished in that area and has perhaps been 
present for some time. The large numbers 
from the Portland metropolitan area (in- 
cluding the Newberg and the Vancouver 
sites), as well as the Salem records, also 
suggest that X. pfeili has been established 
in the northern Willamette Valley for a con- 
siderable period. 


Xyleborus xylographus (Say) 


Records.—Oregon: Lane Co., Goshen 
(mill site), 9 & 24 VI 1999, EtOH (8); Mar- 
ion Co., Salem (wood recycler), 6 VI 2001, 
aP-EtOH (1); Washington Co., Tualatin 
(SWPM importer), 24 V 2001, aP-EtOH 
(2). 

Comments.—These data include a new 
OR record for Washington County. In 1999, 
this species was only found at the original 
site, not in any of the delimitation traps 
placed in that vicinity subsequent to its de- 
tection in 1998, suggesting this population 
stemmed from a recent introduction. No 
specimens were trapped in 2000 or 2002. 


HYMENOPTERA 
Xiphydriidae 
Xiphydria prolongata (Geoffroy) 


Records.—Oregon, Multnomah Co., 
Portland (wood recyclers), 6-18 VIII 1999, 
2 VIII 2001, 27 VI 2002, aP-EtOH (3), IPS 
(2), EtOH (1). 


VOLUME 107, NUMBER 3 


Comments.—No specimens were trapped 
in 2000, despite trapping at the same sites 
where this species was found in 1999. Ef- 
forts to find infested host material in 2000 
were also unsuccessful. 


DISCUSSION 


Eight species of NIWBI trapped in 
1999-2002 at various high-risk sites in 
northwestern Oregon and southwestern 
Washington represent new Oregon, Wash- 
ington, Pacific Northwest, western U.S., or 
North American records. Most of the spe- 
cies of NIWBI found in 1997 and 1998 
(documented in Mudge et al. 2001) were 
trapped again in 1999—2002. There were no 
further records of Micromalthis debilis 
LeConte (Coleoptera: Micromalthidae) or 
Xyloterinus politus (Say) (Scolytinae). 
Trapping only one or a few specimens of a 
species, e.g., Tetropium castaneum, Xylo- 
trechus hircus, and Monarthrum fasciatum, 
may indicate possible interceptions rather 
than established populations. 

Twelve of the sixteen species (75%) of 
NIWBI treated in this paper and Mudge et 
al. (2001) are now known from both eastern 
and western North America. Gnathotrichus 
materiarius, M. debilis, Monarthrum fas- 
ciatum, X. politus, and Xylotrechus sagitat- 
tus sagittatus are indigenous to eastern 
North America (Linsley 1964, Wood 1982, 
Wood and Bright 1992, Phillips and Young 
2001). Hylastes opacus, Phymatodes testa- 
ceus, Xiphydria prolongata, Xyleborus cal- 
ifornicus, X. pfeili, Xylosandrus crassius- 
culus, and X. germanus are nonindigenous 
species from other continents that have 
been established in the East for varying pe- 
riods (Smith 1983, Hoebeke 1994, Solomon 
1995, Vandenberg et al. 2000). 

SWPM/RWP. from foreign (extraconti- 
nental) sources is subject to regulation, cer- 
tification, and inspection at ports of entry 
into the U.S., although only a small per- 
centage is actually inspected (USDA 2003). 
With few exceptions, there are virtually no 
restrictions on the transport of domestic 
SWPM/RWP across Canadian and _ state 


561 


borders. Consequently, potentially infested 
SWPM/RWP, which is not inspected or 
treated, travels through and into the West 
from eastern North America daily. It seems 
likely that at least some of the Oregon and 
Washington records for nonindigenous spe- 
cies previously known from eastern North 
America were the result of SWPM/RWP in- 
fested in the East. This was almost certainly 
the case for the records of eastern indige- 
nous species. 

The “‘sharing’”’ of NIWBI among North 
American regions has not been unidirec- 
tional. Three western scolytine species have 
recently been detected in the central and 
eastern United States: Dendroctonus pseu- 
dotsugae Hopkins in Minnesota (Dziuk 
2003) and HAylesinus californicus Swaine 
and H. criddlei (Swaine) in Maryland (Ra- 
baglia and Williams 2002). These records 
were most likely due to infested SWPM/ 
RWP received from the West. This may 
also have been the case for populations of 
several of the exotic NIWBI species re- 
cently detected in the East, e.g., Xyleberus 
californicus and X. pfeili (Vandenburg et al. 
2000). 

These data underscore the potential for 
and the risks associated with introducing 
NIWBI from one region of North America 
to another. The introduction of exotic forest 
pests established elsewhere in North Amer- 
ica (see below) or forest pests indigenous 
to other regions of the U.S. (e.g., southern 
pine beetle, Dendroctonus frontalis Zim- 
merman (Scolytinae)) into the West could, 
in some instances, profoundly affect west- 
ern urban environments, forests, and asso- 
ciated resources. The daunting list of NI- 
WBI species in eastern North America 
compounds these concerns. Recent NIWBI 
species established outside of western 
North America include the emerald ash bor- 
er, Agrilus planipennis Fairmaire (Bupres- 
tidae), the Asian longhorned beetle, Ano- 
plophora glabripennis (Motschulsky), the 
smaller Japanese cedar longhorned beetle, 
Callidiellum rufipenne (Motschulsky), and 
the brown spruce longhorned beetle, Tetro- 


562 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


pium fuscum (FE) (Cerambycidae), as well 
as myriad bark and ambrosia beetles (Scol- 
ytinae), e.g., the red-haired pine bark beetle, 
Hylurgus ligniperda (FE), the pine shoot 
beetle, 7. piniperda, Pityogenes bidentatus 
(Herbst), Xyleborus similis Ferrari, and Xy- 
losandrus mutilatus (Blandford) (Hoebeke 
1994, Hoebeke 1999, Hoebeke et al. in 
prep., Maier and Lemmon 2000, CFIA 
2002, McCullough and Roberts 2002, 
USDA Pest Alert—Hylurgus ligniperda), 
Schieffer and Bright 2004). Woodborers in- 
digenous to or established in the West could 
also have severe effects should they be in- 
troduced elsewhere in North America, in- 
cluding species that are benign or econom- 
ically insignificant in the West. An analo- 
gous example would be the current impacts 
of the hemlock woolly adelgid (Hemiptera: 
Sternorhyncha: Adelgidae: Adelges tsugae 
Annand) on eastern forests. 

NIWBI could be transported intraconti- 
nentally by infested nursery stock and 
household plants, infested SWPM or RWP 
(including firewood), or hitchhiking on ve- 
hicles or other transportation. Untreated 
hardwood SWPM is strongly implicated as 
one likely pathway since 79% of NIWBI 
species documented in this paper and in 
Mudge et al. (2001) are known predomi- 
nantly or exclusively from hardwoods. To 
the best of our knowledge, relatively little 
hardwood RWP are imported into Oregon 
and Washington from the East. In contrast, 
ODA staff have often observed substantial 
volumes of hardwood SWPM received in 
Oregon that clearly originated in the East. 
Further support for this possibility is pro- 
vided by the frequent interceptions of 
woodboring insects in foreign SWPM 
(USDA/APHIS/FS 2000, Haack 2001). Re- 
cords of NIWBI species with conifer hosts 
could stem from either SWPM or RWP. For 
instance, there is a much greater volume of 
eastern RWP versus SWPM received at one 
of the mill sites where several conifer-at- 
tacking NIWBI species have been detected. 
There is also evidence that nonwoodboring 
insects may be transported among North 


American regions via SWPM/RWP (e.g., 
Lattin and LaBonte 2002; ODA, unpub- 
lished data). 

The movement of unregulated, untreated 
SWPM/RWP within North America poses 
a major threat to the health of our forests 
and shrublands, nursery industry, and or- 
namental plantings. Combined with the on- 
going risks of the introduction of new 
woodboring insect pests from foreign 
sources, the potential for substantial eco- 
nomic and environmental disruption is 
great. The challenges associated with reg- 
ulation and treatment of domestic SWPM/ 
RWP are daunting. However, if the current 
situation persists, it is only a matter of time 
until a woodboring or wood-associated in- 
sect pest established in or indigenous to one 
North American region becomes a devas- 
tating new addition to another region. 


ACKNOWLEDGMENTS 


Robert Rabaglia, Maryland Department 
of Agriculture, Annapolis, MD, and Ste- 
phen L. Wood, Brigham Young University, 
Provo, UT graciously made or confirmed 
many of the Scolytinae determinations and 
provided information in support of these 
surveys. Steven W. Lingafelter, Systematic 
Entomology Laboratory, USDA, Washing- 
ton, D.C., determined the specimens of T. 
castaneum and X. hircus. 

E. Richard Hoebeke (Department of En- 
tomology, Cornell University, Ithaca, NY) 
and Leland M. Humble, Natural Resources 
Canada, Canadian Forest Service, Victoria, 
BC, provided advice and information which 
substantially aided in the development of 
this manuscript. Several colleagues and 
anonymous reviewers supplied helpful sug- 
gestions which greatly improved the man- 
uscript. 

These surveys could not have been ac- 
complished without the extensive support 
with survey implementation and logistics 
provided by the ODA staff, including 
Wayne Estabrook, Karl Puls, Terri Stafford, 
Kerri Schwarz, and Mike Savelich. Christy 
Brown and Jennifer Williams (ODA) sorted 


VOLUME 107, NUMBER 3 


and prepared many thousands of specimens. 
Their assistance, perseverence, and sharp 
eyes were greatly appreciated. Kerri 
Schwarz also prepared the map (Fig. 1). 
Phillip Coombs (ODA) also assisted with 
specimen sorting and preparation. Alan 
Kanaskie and David Overhulser (Oregon 
Department of Forestry) made it possible to 
obtain spruce bait logs from state forest 
lands. 

Funding for these surveys was provided 
by USDA APHIS PPQ, Western Region, 
Cooperative Agricultural Pest Survey Grant 
8584-0260-CA (1999-2002) and the 
Oregon general fund. 


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PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 565-571 


THE GENUS RAMPHASTICOLA CARRIKER 
(PHTHIRAPTERA: AMBLYCERA: MENOPONIDAE) FROM THE TOUCANS 
(PICIFORMES: RAMPHASTIDAE), WITH DESCRIPTION OF 
A NEW SPECIES 


RONALD A. HELLENTHAL, ROGER D. PRICE, AND JASON D. WECKSTEIN 


(RAH) Department of Biological Sciences, RO. Box 369, University of Notre Dame, 
Notre Dame, IN 46556-0369, U.S.A. (e-mail: ronald.a.hellenthal. 1 @nd.edu); (RDP) 4202 
Stanard Circle, Fort Smith, AR 72903-1906, U.S.A. (e-mail: rpricelice @aol.com); (JDW) 
Illinois Natural History Survey, 607 East Peabody Drive, Champaign, IL 61820-6970, 
U.S.A. (e-mail: jweckst @inhs.uiuc.edu) 


Abstract.—The diagnostic characteristics of the genus Ramphasticola Carriker are re- 
evaluated. They are of sufficient magnitude to justify the removal of Ramphasticola from 
synonymy with Myrsidea Waterston and merit its recognition as a distinct genus. The 
currently recognized three species of Ramphasticola are illustrated and redescribed. A 
new species, R. moylei, from the type host Ramphastos tucanus cuvieri Wagler in Peru, 


is described. 


Key Words: 
phastidae 


The chewing lice of the genus Myrsidea 
Waterston from toucans have not been the 
subject of systematic work for over 40 
years. There are more than 200 species of 
the genus recognized, with the majority of 
these on avian hosts within the Passerifor- 
mes. Three species are known from hum- 
mingbirds (Apodiformes: Trochilidae) (Dal- 
gleish and Price 2003) and 14 Myrsidea 
names currently are associated with species 
on toucans (Piciformes: Ramphastidae). Six 
of these originally were described in the ge- 
nus Ramphasticola Carriker and the re- 
maining eight were placed in what we now 
consider as typical Myrsidea (see Price et 
al. 2003). 

Hopkins and Clay (1952) did not recog- 
nize Ramphasticola as a distinct genus, in- 
cluding it within the Myrsidea. Without the 
benefit of having seen any material of Ram- 
phasticola and ignoring the opinion of Car- 


chewing lice, Ramphasticola, Phthiraptera, Menoponidae, toucans, Ram- 


riker (Carriker and Diaz-Ungria 1961), 
Price et al. (2003) followed Hopkins and 
Clay (1952) and maintained Ramphasticola 
as a junior synonym of Myrsidea. However, 
having now studied Carriker’s original type 
specimens and other material for all six 
Ramphasticola names, we concur with Car- 
riker that Ramphasticola merits generic sta- 
tus. It is our purpose here to give diagnostic 
details for the three species that we recog- 
nize to support the description of a new 
species. The redescriptions are accompa- 
nied by dorsoventral illustrations of female 
body shape, with details of the taxonomi- 
cally important thoracic area. Male recog- 
nition is quite tenuous. Although there are 
some useful characters for males, Ram- 
phasticola females are most easily and re- 
liably identified to species. 

In the following descriptions, all mea- 
surements are in millimeters. Abbreviations 


566 


are TW, temple width; HL, head length; 
PW, prothorax width; MW, metathorax 
width; ANW, female anus width; TL, total 
length; and GL, male genitalia length. Host 
classification below orders follows that of 
Dickinson (2003). The holotype and para- 
types of the new species are at the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. Abbreviations 
for the collectors of the lice are: MAC (M. 
A. Carriker, Jr), KE (K. Eckhardt), JH (J. 
Hill), DFL (D. E Lane), and JDW (J. D. 
Weckstein). 


Genus Ramphasticola Carriker 


Ramphasticola Carriker 1949:305. Type 
species: Ramphasticola hirsuta Carriker, 
by original designation. 


This genus was described by Carriker 
(1949) to accommodate the new species, R. 
hirsuta, from a toucan host. A second spe- 
cies, R. niethammeri, was described by Ei- 
chler (1954), but later was recognized as 
the subspecies R. hirsuta niethammeri by 
Carriker and Diaz-Ungria (1961) when the 
descriptions of the additional two species 
and two subspecies were given. Although 
there are some features in common with 
Myrsidea, a number of very important ones 
are different and justify the recognition of 
Ramphasticola as separate from Myrsidea. 
The principal features for separation of 
Ramphasticola are: (1) absence of an aster 
of strong setae on each lateroposterior cor- 
ner of an enlarged sternite II; (2) female 
with thoracic segments much enlarged and 
completely separated from each other; (3) 
female with a distinctive thoracic and/or ab- 
dominal chaetotaxy (see Figs. 1—4); and (4) 
three of four species with outer occipital se- 
tae long, of length equal to inner occipital 
setae. Consistent with both Carriker and 
Diaz-Ungria (1961) and Price et al. (2003), 
we recognize here three of the previously 
described species as valid and the remain- 
ing three as junior synonyms. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Ramphasticola hirsuta Carriker 


(Figs 1) 
Ramphasticola hirsuta Carriker 1949:305. 
Type host: Ramphastos swainsonit 
Gould. 


Ramphasticola niethammeri Eichler 1954: 
40. Type host: Ramphastos tucanus cu- 
viert Wagler. 

Ramphasticola hirsuta ambigua Carriker in 
Carriker and Diaz-Ungria 1961:19. Type 
host: Ramphastos ambiguus ambiguus 
Swainson. 

Ramphasticola hirsuta tucana Carriker in 
Carriker and Diaz-Ungria 1961:21. Type 
host: Ramphastos tucanus tucanus L. 


Female.—As in Fig. 1. Inner occipital se- 
tae long, outer minute. Pronotum with 8— 
13 long setae at posterior margin. Meson- 
otum large, evenly rounded posteriorly. 
Metanotum much wider than long, with 
sparse dorsal chaetotaxy distributed in 4 
patches; metasternal plate ventral to meson- 
otum, 0.30—0.34 wide, diamond-shaped, 
with 6—12 setae. Sternite II represented by 
50 or so short setae anterior to highly irreg- 
ular dense row of about 100 mostly long 
setae. Postspiracular setae very long on II- 
IV and VIII, short on V-VII. Anus with 35— 
41 ventral fringe setae, 31—33 dorsal. Sub- 
genital plate with 15—16 marginal setae. Di- 
mensions: TW, 0.49—0.53; HL, 0.35—0.36; 
PW, 0.36—0.40; MW, 0.62—0.68; ANW, 
0.21-0.27; TL, 1.62—1.76. 

Male.—Head as for female. Thorax and 
abdomen much as in Fig. 5. Pronotum with 
7—13 long marginal posterior setae; meson- 
otum and metanotum approximately equal 
in size, with latter having 10 marginal setae; 
metasternal plate with 7—9 setae. Abdomi- 
nal tergites and sternites without any un- 
usual enlargment. Tergites I-VI with 21—35 
setae; VII, 17—22; VIII, 9-12. Postspiracu- 
lar setae very long on I-IV and VI-VIII, 
shorter on V. Sternal setae: I, 0; II, 41-58; 
IlI-IV, 29—43; V—VII, 41-53; VIII, 25-35. 
Genitalia and genital sac sclerites as in Figs. 
6 and 7. Dimensions: TW, 0.45—0.48; HL, 


VOLUME 107, NUMBER 3 


Figs. 1-3. 


0.31—0.33; PW, 0.31—0.34; MW, 0.40—0.47; 
GL, 0.44-0.48; TL, 1.46—-1.51. 
Material Ex Ramphastos swainsonii, | 
2, 2 6 (including “Type’’ pair and para- 
type 6 of Ramphasticola hirsuta), Colom- 
bia: Santander N., Bella Vista (MAC-4643; 
1943): 4 2, 1 3, Colombia: Choc6, Rio 
Jurubidé (MAC-19873; 1951); 1 2, Pana- 
ma: La Laguna (200157; 1963). Ex R. t. 
cuvieri, 3 2, 1 36, Brazil: Para, ca 139 km 
SSW Santarem, W of Rio Tapajés, Alto Rio 
Arapiuns, 02°59.55’S, 55°50.37'W (JDW- 
458; 2000); 1 2, 2 6, Brazil: Mato Grosso, 
E bank Rio Teles Pires, 1.2 km up river 
from mouth Rio Cristalino, 09°38'22’S, 
5550250 We GD W=244-" 1999) 11 “21S: 
Brazil: Amazonas, Fazenda Toshiba, ca 8 
km NE Careiro, 03°47'S, 60°17'W (JDW- 
482; 2000); 2 2, 1 6, Peru: Loreto, 86 km 
SE Juanjui on E bank upper Rio Pauya, 
Ones 5100S 975.560 1 We (DEL=1252: 
2000); 1 2, 1 6, Peru: Loreto, 7 km SW 
Jeberos, 05°18'48"S, 76°16'32"W (KE-112; 
2001); 1 &, Peru: Huacomayo (MAC; 
1931); 1 6, Peru: Sapasoa (MAC; 1933); 1 


567 


Female dorsoventral thorax and abdomen. 1, Ramphasticola hirsuta. 2, R. aenigma. 3, R. mirabile. 


?, Bolivia: Huanay, Rio Bopi (MAC-9494; 
1934); 1 ¢, Venezuela: Sta. Elena, Gran Sa- 
bana (MAC; 1946). Ex R. t. tucanus, 2 @ 
paratypes of R. h. tucana, Venezuela: Cam- 
pamento Cecilia Magdalena, Rio Caura 
(2539; 1957); 1 3 paratype of R. h. tucana, 
Venezuela: Campamento Cecilia Magdale- 
na, Rio Caura (2540; 1957); 3 2, Guyana: 
Kartago Pt. (JH; 1984). Ex R. a. ambiguus, 
3 2 (holotype, paratypes of R. h. ambigua), 
Colombia: Belén, Dept. Huila (MAC- 
22009-1952): 

Remarks.—Ramphasticola hirsuta dif- 
fers from the other known members of this 
genus in having modestly modified female 
thoracic segments, minute outer occipital 
setae, and metasternal shape and chaetotaxy 
similar to toucan Myrsidea. The male gen- 
italia are similar to those of M. peruviana 
Eichler. The best differentiating feature is 
the possession by the female R. hirsuta of 
numerous long close-set setae across the 
anterior sternum (Fig. 1), this undoubtedly 
being responsible for the specific name of 
“hirsuta.” 


568 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Carriker included in his material a single 
collection of 2 males identified by him as 
R. hirsuta from Ramphastos vitellinus ci- 
treolaemus Gould taken in La Raya, Co- 
lombia. Given the absence of females, we 
prefer to defer recognition of this associa- 
tion until there is further confirmation. 


Ramphasticola aenigma Carriker 
(Fig. 2) 


Ramphasticola aenigma Carriker in Carri- 
ker and Diaz-Ungria 1961:24. Type host: 
Ramphastos tucanus tucanus L. 


Female.—As in Fig. 2. Both inner and 
outer occipital setae subequally long. Pron- 
otum with about 80—90 long close-set setae 
on posterior margin; prosternum with 4 se- 
tae. Mesonotum large, with posteriorly nar- 
rowed median process extending to abdo- 
men anterior margin. Metanotum bipartite, 
with each part oblong, encroaching into lat- 
eral margin of mesonotum, with sparse pe- 
ripheral chaetotaxy; metasternal plate ven- 
tral to mesonotum, large, 0.43—0.44 wide, 
hexagonal-shaped, with about 25-35 long 
setae. Probable sternite II represented by 
wide sclerite posterior to metasternum, with 
about 100 short heavy setae, these concen- 
trated on lateral portions of plate. Postspi- 
racular setae very long on I—II, short on II— 
VIII. Anus with 40—41 ventral fringe setae, 
24-28 dorsal. Subgenital plate with 17-18 
marginal setae. Dimensions: TW, 0.59— 
0.61; HL, 0.36—0.38; PW, 0.50—0.52; MW, 
0.71-0.79; ANW, 0.26—-0.28; TL, 1.88-— 
LS). 

Male.—Much as in Fig. 5. Pronotum 
with 17—21 long posterior marginal setae; 
prosternum with 4—5 setae. Metanotum 
with 13-17 marginal, 17—35 anterior setae; 
metasternal plate with 32—36 setae. Tergites 
I-IV with 23—27 setae; V, 17—20:; VI-VIL, 
14-17; VIII, 9-10. Postspiracular setae 
very long on II-IV and VI-VIII, shorter on 
I and V. Sternal setae: I, 0; II, 33-44; II— 
V, 40—48; VI, 37—42; VU, 30-31; VUI, 10— 
12. Genitalia and genital sac sclerites as in 
Figs. 6 and 7. Dimensions: TW, 0.47—0.51; 


HL, 0.31—0.34; PW, 0.36—-0.37; MW, 0.44— 
0.49; GL, 0.41—0.46; TL, 1.51-1.59. 

Material.—Ex R. t. tucanus, 1 °, 1 6 
paratypes of R. aenigma, Venezuela: Cam- 
pamento Cecilia Magdalena, Rio Caura 
(2540: 1957): 22, 4 6, Brazil? Paras Ba- 
zenda Morelandia, 8 km N Santa Barbara 
do Parad, 01°12'40"S, 48°14'47"W (JDW- 
284; 1999). 

Remarks.—The female of R. aenigma is 
readily separated from that of R. hirsuta by 
the differences in shapes of the thoracic no- 
tal sclerites and the ventral chaetotaxy (Fig. 
2 vs. Fig. 1). The male of R. aenigma is 
differentiated by having the outer occipital 
setae as long as the inner and the metaster- 
nal plate with many more setae. The spec- 
imens of the type series for both R. h. tu- 
cana and R. aenigma were taken from the 
same host individual in Venezuela. This oc- 
currence of two different species of the 
same genus on a single host is unusual. 
However, this pattern is also known for 
doves and their lice (Johnson et al. 2002). 
Both sexes of these Ramphasticola species 
are readily identifiable, minimizing the pos- 
sibility of incorrect association other than 
through host cross-contamination. 


Ramphasticola mirabile Carriker 
(Fig. 3) 


Ramphasticola mirabile Carriker in Carri- 
ker and Diaz-Ungria 1961:26. Type host: 
Ramphastos tucanus cuvieri Waglet. 


Female.—As in Fig. 3. Head as for R. 
daenigma. Pronotum disproportionately 
large, with 43—45 long posterior marginal 
setae, these more widely spaced than for R. 
aenigma. Mesonotum large, with posteri- 
orly narrowed median process broad and 
flattened at juncture with abdomen. Metan- 
otum bipartite, with each part relatively 
small, well separated, with chaetotaxy as 
shown; metasternal plate ventral to prono- 
tum, large, with 15—16 setae. Sternites not 
delineated, with only sparse short setae. 
Postspiracular setae very long on II and 
VII-VIII, much shorter on HI-VI. Anus 


VOLUME 107, NUMBER 3 


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Figs. 4-7. Ramphasticola moylei. 4, Female dorsoventral thorax and abdomen. 5, Entire male dorsoventral. 


6, Male genitalia. 7, Male genital sac sclerites. 


with 30—31 ventral fringe setae, 23—28 dor- 
sal. Subgenital plate with 18—19 marginal 
setae. Dimensions: TW, 0.59—0.60; HL, 
0.36; PW, 0.54; MW, 0.61—0.62; ANW, 
0.26—0.27; TL, 1.53-1.58. 

Male.—Much as in Fig. 5. Pronotum 
with 13-14 long posterior marginal setae. 
Metanotum with 12—16 marginal setae; me- 
tasternal plate with numerous setae, but 
mostly obscured. Tergites I-II with 18—20 
setae; II-VI, 20—29; VII, 18—23; VIII, 14— 
15. Postspiracular setae very long on II-IV 
and VI—VIII, shorter on I and V. Sternal 
setae: I-II, obscured; III, 36—38; IV—VI, 
44-52; VII, 32—40; VIII, 15-21. Genitalia 
and genital sac sclerites as in Figs. 6 and 7. 
Dimensions: TW, 0.47—0.49; HL, 0.31-— 
0.32; PW, 0.34—-0.36; MW, 0.42—0.44; GL, 
0.44—0.50; TL, 1.22—1.33. 

Material —Ex R. t. cuvieri, 3 2,3 ¢ (in- 
cluding holotype @, allotype d, 4 paratypes 


of R. mirabile), Bolivia: Chiniri, Rio Bopi 
(MAC-9752; 1934). 

Remarks.—As with the other species of 
the genus, the female of R. mirabile has dis- 
tinctively different shapes and chaetotaxy 
associated with the thoracic segments (Fig. 
3 vs. Figs. 1-2). The male of R. mirabile is 
separable from that of R. hirsuta on the ba- 
sis of its long outer occipital setae and its 
thoracic chaetotaxy, which are similar to R. 
aenigma. Males are tenuously separable 
from R. aenigma by having fewer pronotal 
and metanotal setae and more tergal and 
sternal setae on the posterior segments. 


Ramphasticola moylei Hellenthal, Price, 
and Weckstein, new species 
(Figs. 4—7) 


Type host.—Ramphastos tucanus cuvieri 
Wagler. 
Female.—As in Fig. 4; head as in Fig. 5. 


76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Outer occipital setae subequal in length to 
inner. Pronotum with 65—68 short to long 
setae on posterior margin; prosternal plate 
with 4 setae. Mesonotum large, tapered, 
with straight sides leading to point at junc- 
tion with anterior abdomen; metasternal 
plate ventral to mesonotum, with 18—24 se- 
tae. Metanotum essentially bipartite, with 
only narrow medial connecting bridge; sur- 
face of each side covered with short to long 
setae, including 2 very long setae. Only 
weak sclerite in position of sternite I ventral 
to mesonotum, with few widely spaced se- 
tae. Tergites I-II greatly reduced, hardly 
visible, but II with very long postspiracular 
seta; postspiracular setae short on III—VI, 
very long on VII-VIII. Total tergal setae: 
II-V, 14-20; VI-VH, 13-15; VIII, 10-12. 
Terminal segment as shown. Anus with 36— 
40 ventral fringe setae, 26—29 dorsal. Ster- 
nal setae: I-III, 10—15; IV, 17—22; V, 25-— 
33; VI-VII, 24-29; subgenital plate with 
lightly serrated posterior margin, 18—19 
marginal and 16—21 anterior setae. Dimen- 
sions: TW, 0.56—0.59; HL, 0.36—0.38; PW, 
0.46—-0.49; MW, 0.64—0.65; ANW, 0.26— 
O273 Wb, UoVARNI 7 

Male.—As in Fig. 5. Outer and inner oc- 
cipital setae subequally long. Pronotum 
with 14-15 long posterior marginal setae; 
prosternal plate with 3 setae. Mesonotum 
and metanotum of approximately equal 
lengths; metanotum with 18—23 marginal, 
20—25 anterior setae; metasternal plate with 
18-19 setae. Postspiracular setae very long 
on I-VIII. Tergal setae on I-VII, 20-28; 
VIII, 13-17. Sternal setae: I, 1—2; II, 34— 
42; Ill-V, 37-49; VI, 36-37; VII, 28-29; 
VI, 14-16. Genitalia as in Fig. 6, with 
genital sac sclerites as in Fig. 7. Dimen- 
sions: TW, 0.46—0.48; HL, 0.32—0.33; PW, 
0.33—0.35; MW, 0.44—0.47; GL, 0.44—0.45; 
TL, 1.45-1.49. 

Type material—Holotype @, ex R. t. cu- 
vieri, Peru: Loreto, ca 54 km NNW mouth 
Rio Morona on W bank, 04°16'51’S, 
77°14'16"W, 8 July 2000, KE-184; 3 2, 3 
3S paratypes, same data as holotype. 

Remarks.—The morphology of the fe- 


male thorax places this new species close 
to R. aenigma (Fig. 4 vs. Fig. 2), the ab- 
domen close to R. mirabile (Fig. 4 vs. Fig. 
3). The differentiating features are female 
R. moylei with: (1) a broad mesonotum with 
straight sides; (2) shorter metanotal scler- 
ites, but with denser chaetotaxy over their 
surface; and (3) without ventral dense short 
stout setae that are conspicuously present 
on R. aenigma. The male of R. moylei has 
very long postspiracular setae on V; it also 
has fewer metasternal setae and more tergal 
setae on V—VIII than does R. aenigma. The 
male genitalia and genital sac sclerites, 
though difficult to see, appear similar on all 
species. The similarity of these sac sclerites 
to those found for Myrsidea peruviana, 
whose host also includes R. ¢. cuvieri, 1s 
interesting and perhaps indicative of an un- 
derlying relationship. 

Etymology.—This species is named for 
Robert G. Moyle, American Museum of 
Natural History, in recognition of his ded- 
ication and interest in collecting lice. 


DISCUSSION 


Previous descriptions have found the 
louse Ramphasticola hirsuta on Ramphas- 
tos swainsonil, R. ambiguus, R. t. tucanus, 
and R. t. cuvieri. These host taxa make up 
a superspecies (Haffer 1974) and form a 
monophyletic clade in a phylogenetic re- 
construction of the Ramphastos toucans 
(Weckstein 2003). The distribution of R. 
hirsuta on these host taxa suggests some 
level of host specificity and perhaps cos- 
peciation at the level of host superspecies. 
More work, including molecular genetic 
analysis of R. hirsuta, may clarify whether 
populations of this widespread louse spe- 
cies represent host races or cryptic species. 

The remaining three Ramphasticola spe- 
cies—R. aenigma, R. mirabile, and R. moy- 
lei—are restricted to one host species, Ram- 
phastos tucanus, and are each known from 
only a relatively small geographic region. 
These three louse taxa share morphological 
similarities of female thoracic structure and 
form a group that is perhaps closely related 


VOLUME 107, NUMBER 3 


phylogenetically. The geographic distribu- 
tion of this group is comparable to that of 
R. hirsuta in the Amazon. In one case, R. 
hirsuta and R. aenigma both were collected 
from the same individual R. t. tucanus host 
in Venezuela (Carriker and Diaz-Ungria 
1961). Hosts at other localities within the 
Amazon also may carry multiple species of 
Ramphasticola. Further sampling is needed 
to establish geographic distribution, host as- 
sociations, and history of speciation within 
and between these louse taxa. 

As noted in the introduction, several au- 
thors (Hopkins and Clay 1952; Price et al. 
2003) did not recognize Ramphasticola as 
a genus distinct from Myrsidea. Our reas- 
sessment of the type specimens as well as 
new material supports the recognition of 
Ramphasticola as a valid genus. Ramphas- 
ticola shares several features with Myrsi- 
dea, including the general head shape and 
structure, the absence of a preocular slit or 
notch, temple setae 26 and 27 with alveoli 
well separated, the presence of only 2 me- 
dioanterior mesonotal setae, and similar 
male genitalic structure, all of which sug- 
gest that these genera share a close phylo- 
genetic relationship. 


ACKNOWLEDGMENTS 


We thank Nancy E. Adams, National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C., for the loan 
of critically important specimens, including 
Carriker types, so necessary for the com- 
pletion of this study. We also thank many 
generous bird collectors who helped to col- 
lect lice for this project: A. Aleixo, J. K. 
Anmentayke ‘eckhardt) D. EF Lane, J. P. 
O’Neill, T. Valqui, and C. C. Witt. This 
work was supported in part by NSF DEB- 
0104919 to JDW, NSF PEET DEB- 
0118794 to K. P. Johnson, the American 


571 


Museum of Natural History Chapman 
Fund, Sigma Xi, an American Ornitholo- 
gists’ Union Research Award, LSU Bird-a- 
thon, the T. Vinton Holmes endowment, the 
LSU Museum of Natural Science, and the 
LSU Department of Biological Sciences. 


LITERATURE CITED 


Carriker, M. A., Jr. 1949. Neotropical Mallophaga mis- 
cellany. V. New genera and species. Revista Bras- 
ileira de Biologia 9:297-313. 

Carriker, M. A., Jr. and C. Diaz-Ungria. 1961. New 
and little known Mallophaga from Venezuelan 
birds (Part I). Novedades Cientificas, Contribu- 
ciones Ocasionales del Museo de Historia Natural 
La Salle, No. 28:3—60. 

Dalgleish, R. C. and R. D. Price. 2003. Two new spe- 
cies of Myrsidea (Phthiraptera: Amblycera: Men- 
oponidae) from hummingbirds (Apodiformes: 
Trochilidae). Occasional Papers of the Western 
Foundation of Vertebrate Zoology, No. 6:1—9. 

Dickinson, E. C., ed. 2003. The Howard and Moore 
Complete Checklist of the Birds of the World. 3rd 
edition. Princeton University Press, Princeton, 
New Jersey, 1,039 pp. 

Eichler, W. 1954. Peruanische Mallophagen. Beitrage 
zur Fauna Perus 4:28—62. 

Haffer, J. 1974. Avian speciation in tropical South 
America. Publications of the Nuttall Ornithologi- 
cal Club No. 14:1—390. 

Hopkins, G. H. E. and T. Clay. 1952. A Check List of 
the Genera & Species of Mallophaga. British Mu- 
seum (Natural History), London, 362 pp. 

Johnson, K. P., B. L. Williams, D. M. Drown, R. J. 
Adams, and D. H. Clayton. 2002. The population 
genetics of host specificity: genetic differentiation 
in dove lice (Insecta: Phthiraptera). Molecular 
Ecology 11:25-38. 

Price, R. D., R. A. Hellenthal, and R. L. Palma. 2003. 
World checklist of chewing lice with host asso- 
ciations and keys to families and genera. pp. 1— 
448. In Price, R. D., R. A. Hellenthal, R. L. Pal- 
ma, K. P. Johnson, and D. H. Clayton. The Chew- 
ing Lice: World Checklist and Biological Over- 
view. Illinois Natural History Survey Special 
Publication 24, x + S5O1 pp. 

Weckstein, J. D. 2003. Systematics and cophylogenet- 
ics of toucans and their associated chewing lice. 
Ph.D. Dissertation, Louisiana State University, 
132 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 572-575 


FIRST RECORD OF CUTEREBRA ALMEIDAI (GUIMARAES AND CARRERA) 
FROM ARGENTINA, NEW HOST RECORDS FOR CUTEREBRA APICALIS 
GUERIN-MENEVILLE, AND LIST OF CUTEREBRA (DIPTERA: OESTRIDAE) 
IN THE COLLECTION OF THE INSTITUTO—FUNDACION MIGUEL LILLO, 
TUCUMAN, ARGENTINA 


C. MIGUEL PINTO AND GUILLERMO L. CLAPS 


(CMP) Escuela de Ciencias Biolégicas, Pontificia Universidad Catolica de Ecuador, Av. 
12 de Octubre y Roca, Apartado postal: 17-01-2184, Quito Ecuator (e-mail: cmpinto@ 
puce.edu.ec); (GLC). Instituto Superior de Entomologia ““Dr. Abraham Willink” (INSUE), 
Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucuman 
(UNT). Miguel Lillo 205 (4000), San Miguel de Tucumdan, Argentina (e-mail: 
guillermolclaps @csnat.unt.edu.ar) 


Abstract.—We review the collections of oestrid bot flies from the Instituto—Fundacion 
Miguel Lillo, Tucuman Argentina, with special emphasis on the genus Cuterebra Clark. 
The collection comprises seven species: Cuterebra almeidai (Guimaraes & Carrera), C. 
apicalis Guérin-Méneville, C. rufiventris Macquart, Cuterebra sp., Dermatobia hominis 
(Linnaeus Jr.), Gasterophilus nasalis (Linnaeus), and Oestrus ovis Linnaeus. We report 
the first record of C. almeidai for Argentina. We also include Oligoryzomys longicaudatus 
(Bennett) and Akodon caenosus Thomas (Rodentia: Muridae) as new host species of 
Cuterebra apicalis. 


Resumen.—Revisamos las colecciones de oestridos del Instituto—Fundacio6n Miguel 
Lillo, Tucuman Argentina, con especial énfasis en el género Cuterebra. La coleccién 
comprende siete especies: Cuterebra almeidai, C. apicalis, C. rufiventris, Cuterebra sp., 
Dermatobia hominis, Gasterophilus nasalis y Oestrus ovis. Reportamos el primer registro 
de C. almeidai para la Argentina. En base a trabajo de campo, también incluimos a 
Oligoryzomys longicaudatus y Akodon caenosus como nuevos hospederos de Cuterebra 
apicalis. 


Key Words: 


diptera, myiasis, rodents, taxonomy 


The family Oestridae is a group of my- 
iasis-causing flies comprising 25 valid 
world genera. There are two native Neo- 
tropical genera, Cuterebra Clark and Der- 
matobia Brauer, with Alouattamyia Town- 
send, Andinocuterebra Guimaraes, Meta- 
cuterebra Townsend, Pseudogametes Bi- 
schof, and Rogenhofera Brauer considered 
junior synonyms of Cuterebra (Pape 2001). 
Gasterophilus Leach, Hypoderma Latreille, 


and Oestrus Linneaus are introduced pests 
from Europe (Guimaraes and Papavero 
1999): 

Bot fly larvae of the Neotropical genus 
Cuterebra (Oestridae: Cuterebrinii) cause 
subdermal myiasis in more than 80 species 
of Neotropical mammals as well as a num- 
ber of introduced mammals. Sigmodontinae 
rodents (Rodentia: Muridae) and squirrels 
(Sciuridae) are among the most susceptible 


VOLUME 107, NUMBER 3 


(Catts 1982, Guimaraes and Papavero 
1999). 

The taxonomy of Cuterebra sensu lato is 
poorly understood, and the species in the 
genus have a large list of synonyms (Gui- 
maraes and Papavero 1999). The dispersed 
literature is a major problem as many of the 
descriptions were published during the 
nineteenth century in obscure journals of 
limited availability. Scant material is de- 
posited in scientific collections due to the 
difficulty in collecting adults and the rarity 
of rearing cuterebrid larvae to adulthood by 
mammalogists (Gardner 1988). 

There are relatively few reports of larvae 
of Cuterebra parasitizing rodents in Argen- 
tina (Guimaraes and Papavero 1999): Cu- 
terebra grandis (Guérin-Meneville) in Ako- 
don azarae (Fisher), Akodon molinae Con- 
treras, Oligorizomys flavescens (Water- 
house), Reithrodon physodes (Olfers), and 
Sciurus aestuans Linneus; Cuterebra api- 
calis Guérin-Méneville in Rattus norvegi- 
cus (Berkenhout); and Cuterebra sp. in 
Chinchilia lanigera (Molina). Guimaraes 
and Papavero (1999) provided a complete 
review on the taxonomy of the group, in- 
cluding a list of hosts and a bibliographic 
database. 


MATERIAL AND METHODS 


The Cuterebra collection of the Institu- 
to—Fundacion Miguel Lillo, Tucuman, Ar- 
gentina (IMLA), was identified using Gui- 
maraes and Papavero’s key (1999) and orig- 
inal descriptions. Terminology follows 
McAlpine (1981). Additionally, we collect- 
ed small mammals using Sherman traps 
(SFA model, H.B. Sherman, Tallahassee, 
FL) in Horco Molle, Tucuman Province, 
Argentina, in February of 2004. This area 
is a subtropical forest termed Yungas which 
is characterized by the presence of acid 
soils, year-round rainfall, and low temper- 
atures near O°C in the winter (Cabrera 
1976). Traps were baited with oatmeal. Ro- 
dents were euthanized with ether in order 
to collect ecto- and endoparasites. Rodents 
with bot fly maggots were dissected to ex- 


Sis} 


tract the larvae intact and allow pupation. 
Extracted larvae were placed in plastic jars 
with humid soil placed in the bottom. The 
jars were maintained at room temperature, 
with several drops of water added regularly 
to keep the soil humid (cf. Sabrosky 1986). 
Rodent hosts were identified following the 
guides by Barquez et al. (1991) and Diaz 
and Barquez (2002) and deposited in the 
Colecci6n Mamiferos Lillo de la Universi- 
dad Nacional de Tucuman (CML). 


RESULTS 


From the examination of the material 
housed at IMLA, and from the host speci- 
mens collected during fieldwork, we give a 
species list, report a new record of Cute- 
rebra for Argentina, and present new host 
records for C. apicalis. Additionally, we 
give taxonomic comments on the Cuterebra 
species to help clarify their identifications. 


Cuterebra almeidai 
(Guimaraes & Carrera) 


This is the first record of Cuterebra al- 
meidai in Argentina. The single specimen 
is characterized by its large size (body 
length 21.85 mm; wing length 18.47 mm), 
black body, abdomen wider than thorax, 
and tergites dark brown dorsally with yel- 
low hairs ventrally. Yellow hairs also occur 
on the sternites and sparsely on the pleura. 
Externally, this specimen resembles the 
darkly-colored species Cuterebra baeri 
Shannon & Greene and C. funebris (Aus- 
ten). Cuterebra almeidai differs from C. 
baeri by lacking yellowish veins in the 
wings and by the presence of yellow hairs 
on the pleura and abdomen. Differences in 
size are also evident. Cuterebra almeidai is 
large, whereas C. baeri is medium-sized 
(body length 18-20 mm; wing length 15 
mm). Guimaraes and Papavero (1999) sug- 
gested that C. almeidai might represent a 
color variation of C. funebris. We consider 
C. almeidai as a valid species based on the 
following characteristics: 1) presence of 
yellow hairs on the pleura and the abdomen 
in C. almeidai (black in C. funebris); 2) 


574 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


reddish-brown antennae (black in C. fune- 
bris); and 3) two distinctive longitudinal 
stripes on the prescutum (as noted in Gui- 
maraes and Carrera 1941) (three in C. fu- 
nebris). 

Specimen examined.—ARGENTINA. 
Salta: Parque Nacional Bariti, Rio Pesca- 
do—Porongal, 700 m, 30-X/7-XI-1978, 1 
adult, P. Fidalgo leg. 


Cuterebra apicalis Guérin-Méneville 


Five adult Cuterebra apicalis are in the 
IMLA. All specimens present typical char- 
acteristics of the species, including the 
shiny dark blue to dark brown tergites 1—3 
in dorsal view and yellowish hairs covering 
the thorax and fifth tergite (Leite and Wil- 
liams 1988b). 

During fieldwork in Horco Molle, we re- 
corded C. apicalis from two new host ro- 
dent species. We found a third instar larva 
in the left side of the rostrum in an adult 
individual of Akodon caenosus (CML 
06383, AA 156). The larva successfully pu- 
pated two days after it was removed from 
the host. Pupation lasted 26 days, unlike the 
range of 29—34 days reported by Leite and 
Williams (1988a). Based on the character- 
istics mentioned above, the adult that 
emerged clearly corresponds to C. apicalis. 
Moreover, we found a small third instar lar- 
va in the lower dorsum, just above the tail, 
in a juvenile of Oligoryzomys longicudatus 
(CML 06382, AA 158), but the larva died 
before pupation. Although the third instar 
larvae in Cuterebra are very similar under 
light microscopy (Leite and Williams 
1997), we consider this larva to be C. api- 
calis because all records from Horco Molle 
belong to this species. Furthermore, the en- 
tomological fauna at Horco Molle, the lo- 
cality where we collected the two parazited 
rodents, has been well documented for 
more than 30 years, and C. apicalis has 
been the only species of the genus recorded 
in the area. Some studies also indicate that 
all maggots found in rodents at a given lo- 
cality belong to the same species of Cute- 


rebra (see Getz 1970, Hunter et al. 1972, 
Wolf and Batzli 2001). 

Specimens examined.—ARGENTINA. 
Tucumadn: Aconquija, 6/10-XII-1950, 1 
adult, R. Golbach leg. Horco Molle, 22- 
XII-1965, 3-IV-1974 and 23-II-1975, 3 
adults, L. Stange leg. 25-IJ-2004, 1 third in- 
star larvae ex: Oligorizomys longicaudatus 
(CML 06382; AA 158), 25-II-2004 (1 third 
instar larvae), 27-IJ-2004 (pupa), 25-ILI- 
2004 (emerging adult), 1 adult and pupari- 
um, ex: Akodon caenosus (CML 06383; 
AA 156-R13), C. M. Pinto et al. leg. 


Cuterebra rufiventris Macquart 


We identified two specimens assignable 
to Cuterebra rufiventris based on the fol- 
lowing characteristics: whitish microtomen- 
tum on the face; yellow hairs on the pleura; 
rufous scutellum; and rufous abdomen. One 
of the specimens has the typical five dark 
stripes on the scutum; the scutum of the 
other fly is lightly damaged, and the stripes 
are not visible. The specimens cannot be 
confused with C. grandis because the arista 
is plumose on the upper side, whereas in C. 
grandis the arista is plumose on both sides 
(Guimaraes and Papavero, 1999). 

Specimens examined.—PERU. Hudnuco: 
Tingo Maria, Rio Huallaga, 700 m, 8-VIII- 
1947 (third instar larvae), 10-IX-1947 
(emerging adult), 1 adult and puparium, ex: 
“‘Larve unter Haut von Hausratte’’ (Rattus 
rattus?), W. K. Weyrauch leg. Tingo Maria, 
670 m, no date, 1 adult, W. K. Weyrauch 
leg. 


Cuterebra sp. 


One specimen from the collection of 
IMLA possesses characteristics that do not 
resemble any described species. This spec- 
imen resembles Cuterebra infulata Lutz by 
having a well-marked yellow hair stripe 
around the thorax. However, C. infulata 
Lutz has black hairs on the scutellum, while 
our specimen has rufous microtomentum 
covering the scutellum. This specimen may 
be a new taxon and will be treated else- 
where. 


VOLUME 107, NUMBER 3 


Specimen examined.—ARGENTINA. 
Salta: Rio Pescado, cerca de Oran, 24-II- 
1971, 1 adult, €. Porter leg. 


ACKNOWLEDGMENTS 


We express gratitude to the instructors 
and students of the course: ‘“‘Pardsitos de 
micromamiferos: Técnicas de estudio y 
diagnostico de los principales grupos. As- 
pectos de la Biologia y Ecologia,” orga- 
nized by Instituto Superior de Entomologia 
“Dr. Abraham Willink”’ of the Universidad 
Nacional de Tucuman (UNT), for their help 
and comments during this research. We 
thank Analia Autino for providing valuable 
information related to the rodents included 
in this work. [liana Alcocer, Donald Gettin- 
ger, Pablo Jarrin and Norman Woodley pro- 
vided literature resources to CMP. René 
Fonseca, Marcela Lareschi, Nathan Much- 
hala, Nelson Papavero, Norman Woodley, 
and two anonymous reviewers read earlier 
versions of the manuscript and made help- 
ful suggestions. This research was partially 
supported by small grants from the Labor- 
atorio de Investigacidn en Enfermedades 
Infecciosas (LIEI) and the Federacion de 
Estudiantes (FEUCE), both from the Pon- 
tificia Universidad Catdlica del Ecuador 
(PUCE), to CMP. This work is dedicated to 
the memory of René Fonseca (1976-2004). 


LITERATURE CITED 


Barquez, R. M., M. A. Mares, and R. A. Ojeda. 1991. 
Mammals of Tucuman/Mamiferos de Tucuman. 
Oklahoma Museum of Natural History, University 
of Oklahoma, Norman, 282 pp. 

Cabrera, A. L. 1976. Regiones fitogeograficas de la 
Argentina. Enciclopedia Argentina de Agricultura 
y Jardineria, Buenos Aires, 83 pp. 

Catts, E. P. 1982. Biology of new world bot flies: Cu- 
terebridae. Annual Review Entomology 27: 313— 
338. 


575 


Diaz, M. M. and R. M. Barquez. 2002. Los mamiferos 
de Jujuy, Argentina. Literature of Latin America. 
Buenos Aires, 326 pp. 

Gardner, A. L. 1988. Book review of: North American 
species of Cuterebra, the rabbit and rodent bot 
flies (Diptera: Cuterebridae), Sabrosky, C. W. 
Journal of Mammalogy 69(2): 436—437. 

Getz, L. L. 1970. Botfly infestations in Microtus penn- 
sylvanicus in southern Wisconsin. Journal of 
Mammalogy 84(1): 187-197. 

Guimaraes, J. H. and N. Papavero. 1999. Myiasis in 
man and animals in the Neotropical region. Bib- 
liographic database. FAPESP. Editora Pléiade. Sao 
Paulo, 308 pp. 

Guimaraes, L. R. and M. Carrera. 1941. Contribuicao 
ao conhecimento dos cuterebrideos do Brasil. Ar- 
quivos do Zoologia Estado do Sao Paulo 3(1): 1— 
12 + 3 plates. 

Hunter, D. M., R. M. E S. Sadleir, and J. M. Webster. 
1972. Studies on the ecology of cuterebrid para- 
sitism in deermice. Canadian Journal of Zoology 
50: 25-29. 

Leite A. C. R. and P. Williams. 1988a. The life cycle 
of Metacuterebra apicalis (Diptera: Cuterebridae). 
Memorias do Instituto Oswaldo Cruz 83(4): 485— 
491. 

. 1988b. Redescrigao de adultos de Metacuter- 

ebra apicalis (Diptera: Cuterebridae). Memorias 

do Instituto Oswaldo Cruz 83(4): 493-507. 

. 1997. External morphology of immature stag- 
es of Metacuterebra apicalis. Memorias do Insti- 
tuto Oswaldo Cruz 92(6): 775-783. 

McAlpine, J. K 1981. Chapter 2. Morphology and ter- 
minology—Adults, pp. 9-63. In McAlpine, J. FE, 
B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. 
Vockeroth, and D. M. Wood, eds. Manual of Ne- 
arctic Diptera, Vol. 1. Research Branch Agricul- 
ture Canada, Monograph 27. Ottawa, vi + 1—674 
PPp- 

Pape, T. 2001. Phylogeny of Oestridae (Insecta: Dip- 
tera). Systematic Entomology 26: 133-171. 

Sabrosky, C. W. 1986. North American species of Cu- 
terebra, the rabbit and rodent bot flies (Diptera: 
Cuterebridae). Thomas Say Foundation Mono- 
graphs. Entomological Society of America 11, vii, 
frontispiece + 240 pp. 

Wolf, M. and G. O. Batzli. 2001. Increased prevalence 
of bot flies (Cuterebra fontinella) on white-footed 
mice (Peromyscus leucopus) near forest edges. 
Canadian Journal of Zoology 79: 106—109. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 576-595 


REVISION OF THE NORTHERN SOUTH AMERICAN TIPHIID GENUS 
MERITHYNNUS KIMSEY, 1991 (HYMENOPTERA: TIPHIIDAE: THYNNINAE) 


LYNN S. KIMSEY 


Bohart Museum of Entomology, University of California, One Shields Ave., Davis 
95616, (e-mail: Iskimsey @ucdavis.edu) 


Abstract.—The South American genus Merithynnus is revised. Five new species from 
Colombia, M. bilobatus, M. intermedius, M. montanus, M. nasalis and M. paradoxus, 
are described. Of these species only paradoxus and intermedius are known from both 
sexes. The remaining species are based on males. Seven previously described species, 
three from Venezuela (osorioi, pecki, and turbulentus) and four from Colombia (acumi- 
natus, bogotanus, desertus, and mimulus) are redescribed, and a distribution map and key 
to the species are provided. Merithynnus desertus is known only from the female and 


acuminatus, bogotanus, mimulus, and turbulentus are known only from males. 


Key Words: 


For a number of years Michael Sharkey, 
University of Kentucky, in collaboration 
with the Alexander von Humboldt Biolog- 
ical Resources Research Institute, Bogota, 
has intensively collected Hymenoptera in 
parks across Colombia. This project has re- 
vealed a heretofore unknown fauna of wasp 
species, particularly in the tiphiid subfamily 
Thynninae. Only two genera of Thynninae 
are recorded from Colombia, Merithynnus 
and Aelurus (Rhagigasterini). All of the 
Colombian thynnines in the tribe Elaphrop- 
terini belong to the genus Merithynnus. 
This study increases the number of known 
Merithynnus species from seven to twelve. 
Oddly, none of the previously described 
species from Colombia were represented in 
the material collected by Sharkey and the 
Humboldt Institute. 

Prior to the study of Merithynnus in Co- 
lombia, the genus was known from fewer 
than 60 specimens, in most species just 
from the type series. Four of these, acumi- 
natus, bogotanus, desertus, and mimulus, 
were described from Bogota, Colombia. 


Merithynnus, Thynninae, Elaphropterini, Colombia 


The remainder of the previously described 
species, osorioi, pecki, and turbulentus, are 
from Venezuela. In the new Colombian ma- 
terial the type series of paradoxus alone is 
represented by more than 200 specimens. 
Although Merithynnus species are undoubt- 
edly parasitoids, like other members of the 
family, nothing is known of the natural his- 
tory of the genus. 


MATERIALS AND METHODS 


Specimens used in this study were bor- 
rowed from the following museums and in- 
stitutions. Type depositions are indicated in 
the text by the city name of the respective 
museum in capital letters: BERLIN—-Hum- 
boldt Museum, Humboldt-Universitat, Ber- 
lin, Germany (EF Koch); BOGOTA—Alex- 
ander von Humboldt Biological Resources 
Research Institute, Bogota, Colombia (D. 
Arias); BUENOS AIRES—Division Ento- 
mologia, Museo Argentino de Ciencias Na- 
turales, Buenos Aires, Argentina (M. J. Vi- 
ana); CAMBRIDGE—Museum of Compar- 
ative Zoology, Harvard University, Cam- 


VOLUME 107, NUMBER 3 


bridge, Massachusetts, USA (P. Perkins); 
DAVIS—Bohart Museum of Entomology, 
University of California, Davis, USA (S. L. 
Heydon); GAINESVILLE—Florida State 
Collection of Arthropods, Gainesville, USA 
(L. A. Stange); LONDON—The Natural 
History Museum, London, England (S. 
Lewis); OTTAWA—Hymenoptera Section, 
Biosystematics Institute, Agriculture Cana- 
da, Ottawa (L. Masner); OXFORD—Hope 
Entomological Collections, Oxford Univer- 
sity Museum, England (C. O’Toole). Pri- 
mary types were studied for all species. 

Modifications of the head, mesosoma and 
terminalia are most important to distinguish 
species. In females the shape of the protho- 
rax, propodeum and pygidium are most sig- 
nificant. Diagnostic features of males in- 
clude the configuration of different com- 
ponents of the face, body coloration, and 
the shape of the hypopygium, paramere and 
aedeagus. The terms terga and sterna are 
used to refer to segments of the metasoma. 
The slender, elongate, curved apex of the 
aedeagus is referred to below as the “apical 
strap.’’ Puncture diameter, given in the text 
as PD, is used to describe the density of 
punctation of the integument. 


Merithynnus Kimsey 


Merithynnus Kimsey 1991:74. Type spe- 
cies: Merithynnus pecki Kimsey 1991:75. 
Original designation. 


Male.—Mandible bidentate, usually with 
small angle or tooth subbasally on dorsal 
margin; pronotal transverse anterior ridge 
laterally notched; scutellum with trans- 
versely septate groove along anterior mar- 
gin; epipygium apically truncate to round- 
ed, with lateral carina; hypopygium notched 
submedially, ligulate or horse hoof-shaped 
apically; paramere elongate, bending ven- 
trally along longitudinal axis, with apical 
tooth or lobe bending toward midline vol- 
sella appressed against aedeagus, with short 
broadly rounded to slender, free apical lobe; 
aedeagal column short, often with large 


SiT/ 


ventral lobe or dorsal one, before apical 
strap, apical strap bending dorsally. 
Female.—Mandible slender, edentate; 
propleuron elongate and strongly convex 
ventrally; scutum and metanotum obscured; 
scutellum elevated and ovoid dorsally; me- 
tasomal tergum I strongly narrowed anieri- 
orly with short, often densely setose, ante- 
rior surface; tergum II with two strong 
transverse ridges, apical margin strongly 
upcurved; pygidium coarsely punctate, with 
lateral carina, usually with one or more api- 
cal lobes or teeth, tufts of setae laterally and 
One or more on the surface of the plate and 
on apical margin; sternum VI apical plate 
broadly ovoid or forming half circle. 
Distribution.—Species of Merithynnus 
appear to be restricted to higher elevations 
of the northern reaches of the Andes in Co- 
lombia and Venezuela, in the Cordillera 
Oriental and Cordillera Merida (Fig. 1). 
None of the specimens examined for this 
study were found below 1,000 m and the 
majority of specimens were from between 
2,000 m and 3,000 m in elevation. , 
Discussion.—The genus Merithynnus 
was described by Kimsey (1991) to include 
a variety of species, some of which were 
originally described in Scotaena and Ela- 
phroptera. Merithynnus is characterized by 
a number of features, some shared with oth- 
er members of the Tribe Elaphropterini, and 
some unique to the genus. Unique features 
include the configuration of the female py- 
gidium, with lateral carina, irregularly 
sculptured posterior surface, 1—4 apical 
lobes or teeth, and two or more tufts of se- 
tae, male scutellum with septate, transverse 
anterior groove, male hypopygium apically 
hoof-like, lateral margin with submedial 
notch or indentation, and paramere apically 
bending inward. Females are only known 
for six species: bogotanus, desertus, inter- 
medius, osorioi, pecki, and paradoxus. The 
female of desertus is undoubtedly synony- 
mous with one of the other species de- 
scribed by Turner (1910) based on males 
from Bogota, Colombia, but this cannot be 
resolved without collecting pairs in copula. 


578 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


EQUATOR 


[ct 


1 - acuminata 7 - montanus 

2 - bilobatus 8 -nasalis 

3 - bogotanus 9 - osorioi 

4 - desertus 10 - pecki 

5 - intermedius 1 - paradoxus 
6 - mimulus 12 - turbulentus 


piles | 


Fig. 1. Distribution map of Merithynnus species in Colombia and Venezuela. 


The genus is redescribed in light of phylo- 
genetic analyses of the genera of Thynninae 
currently underway. 


KEY TO THE SPECIES OF MERITHYNNUS 


Nl, Wineless Gems) occccoccacaecccecee DD 
= Wingeds(male) yo airs cy eerie Gitosoxcmoees etre 6 
2. Pronotal disk anteriorly trilobate (Fig. 32); 
pygidium with two subapical lobes or teeth 
between apical tufts of setae (Fig. 35); pro- 
podeum abruptly concave adjacent to metan- 
otum (FIs) 32) cee eG a aS SE 
LE oI paradoxus Kimsey, new species 
—  Pronotal disk bilobate, convex or flat anteri- 
orly (as in Figs. 24-25, 30, 37); pygidium 
emarginate or unilobate between apical tufts 
of setae (Figs. 29, 33-34); propodeum strong- 
ly convex adjacent to metanotum (Figs. 24, 
QD's SUD creations BEN Lire tee een Osea ere 3 
3. Propodeum dorsomedially dentate (Fig. 30); 
metasomal sternum VI apically forming half 
circled (Rie33) Nie eae iene osorioi Kimsey 
—  Propodeum dorsomedially rounded (Figs. 24— 
25, 37); metasomal sternum VI apically semi- 
circulan!(@sminyRicss295953536) meer 4 


Pronotal medial disk strongly excavate, al- 
most trough-like anteromedially, excavation 
planar with anterior collar (Fig. 24); pygidium 
with carina enclosed apical snout or plate 
(Fig. 33) .... intermedius Kimsey, new species 
Pronotum medial disk convex or flattened, 
strongly to somewhat elevated above anterior 
collar; pygidium without carina enclosed api- 
calisnoutfon plates eae eee 5 
Propodeum saddled in profile (Fig. 37); pron- 
otum with medial disk convex, strongly ele- 
vated above anterior collar (Fig. 37); pygidi- 

um apically quadrilobate (Fig. 36) ..... 

Be Correa eos moncle pecki Kimsey 
Propodeum convex in profile, not saddled 
(Fig. 25); pronotum nearly flat, medial disk 
only slightly elevated above anterior collar 
(Fig. 25); pygidium apically trilobite, with 
acute medial lobe (Fig. 29) .. desertus (Turner) 
Clypeus with medial lobe, tooth or projection 
(as in Figs. 3, 10); metasoma with extensive 
orange or reddish markings 
Clypeus without medial tooth or projection 
(as in Figs. 4, 11); metasoma without orange 
or reddish markings, except mimulus ..... 9 


VOLUME 107, NUMBER 3 


Fig. 2. 


Forecoxa with elongate ventral spine (Fig. 
13); clypeus apicomedial margin forming 
elongate, narrow, upcurved lobe (Fig. 10); an- 
tennal lobe sharply angulate apically (Fig. 10) 
tettishs ter eicteks paradoxus Kimsey, new species 
Forecoxa without ventral spine; clypeal api- 
comedial margin unilobate, trilobate, or 
broadly truncate (as in Figs. 3—9, 11—12); an- 
tennal lobe rounded 
Clypeus apical margin with small medial lobe 
and small obtuse sublateral lobes (Fig. 3) 
Bd! Goa CEN Rae me teres a acuminatus (Turner) 
Clypeus apicomedially slightly produced and 
flat medially, with small acute to obtuse sub- 
medial angles (Fig. 8) 
RAC SATESET er oda lanes. nasalis Kimsey, new species 
Metasoma and legs extensively reddish; man- 
dible broadest submedially (Fig. 6); clypeal 
apical projection with apex equal to distance 
between antennal sockets (Fig. 6) 
mimulus (Turner) 


Metasoma and legs without reddish colora- 
tion, black and yellow; mandible broadest 


579 


Side view of male and female Merithynnus paradoxus. 


subapically across subsidiary tooth (as in Fig. 
9); clypeal apical projection with apex broad- 
er than (as in Figs. 9, 11) or less than half 
distance between antennal sockets (Fig. 4) 10 


. Clypeus apicomedial projection with sides 


convex to lobate, apex narrowly bidentate 
(Fig. 4); forewing marginal cell more darkly 
stained, noticeably darker than rest of wing 
Siege Cota bilobatus Kimsey, new species 
Clypeus apicomedial projection with sides 
straight to concave, apex broadly bilobate, 
flat, bidentate or indented (as in Figs. 7, 9, 
11); forewing marginal cell at most slightly 
darker than rest of wing membrane 


. Clypeus convex medially, rounded apicome- 


dially; mandible broadest submedially; face 
with continuous yellow stripe along inner eye 
margin connecting behind ocelli, forming 
continuous stripe 
intermedius Kimsey, new species 


Clypeus flat medially, apicomedially angu- 
late, not rounded in front view; mandible 
broadest subbasally (as in Figs. 5, 9, 11); face 


580 


with discrete yellow stripe along inner eye 
margin not extending beyond ocelli 
12. Clypeus with apicomedial angles obtuse to 
90° (Figs. 7, 11); mandible slender, with small 
subapical tooth, broadest subbasally, without 
subbasal angle on inner margin (Figs. 7, 11) 


— Clypeus with apicomedial angles acute (Figs. 
4, 10, 13); mandible with large subapical 
tooth, broadest submedially, with small sub- 
basal angle on inner margin (Figs. 4, 10, 12) 

ay che SucB SESE ORES REINS oo CURSE ROMeM oes a. ante ara 14 

13. Clypeus flat in lateral view (Fig. 22); mandi- 
ble without subbasal angle (Fig. 11); aedea- 
gus with large ventral lobe before apical strap 
(Fig. 46) pecki Kimsey 

— Clypeus saddle-shaped in lateral view (Fig. 
18); mandible with obtuse subbasal angle 
(Fig. 7); aedeagus without ventral lobe before 
ANCAl Stays (ENS, SA) o6c0840000000000 

AS a Ae ee montanus Kimsey, new species 

14. Clypeus apicomedial margin shallowly emar- 
ginate, with acute lateral angles (Fig. 9); pro- 
podeum finely shagreened; frons without yel- 
low spot between ocelli...... osorioi Kimsey 

— Clypeus apicomedial margin deeply emargin- 
ate, with narrow, slightly up-turned lobes on 
either side (Fig. 12) or rounded (Fig. 5); pro- 
podeum highly polished; frons with yellow 
Spotubetweenkocellieyw rin ie Tcaen ener 15 

15. Clypeus apicomedial projection with lateral 
margin obtuse or rounded (Fig. 5); face with 
complete yellow stripe along inner eye mar- 
gin, extending to vertex behind ocelli; terga 
I-V with bright yellow stripe or spot; fore- 
wing marginal cell only slightly darker than 
rest of wing bogotanus Kimsey 

— Clypeus apicomedial projection with lateral 
margin straight (Fig. 12); face with two nar- 
rowly to well-separated yellowish spots along 
eye margin, dorsal spot barely extending to 
ocelli, not beyond; terga I-IV with small 
whitish lateral spot; forewing marginal cell 
markedly darker than rest of wing 

Edie i Rg ass tl gg <a ee Maar gti turbulentus (Turner) 


Merithynnus acuminatus (Turner) 
(Figs. 3, 14, 26, 38, 48) 


Elaphroptera acuminata Turner 1910:194. 
Syntype males: Colombia: Bogota (BER- 
LIN). 


Male.—Body length 14-15 mm. Head 
(Figs. 3, 14): Clypeus impunctate ventrally, 
dorsally punctures contiguous to 0.5 PD 
apart, with small medial tubercle, apical 
margin barely produced, with acute medial 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


lobe and obtuse submedial ones; area be- 
tween antennal socket and inner eye margin 
with tiny punctures, punctures 1—3 PD 
apart; frons with punctures larger than low- 
er face, nearly contiguous, somewhat elon- 
gate; vertex punctures smaller than frons, 
contiguous; hindocellus 3 hindocellar di- 
ameters from eye margin; flagellomere I 
2.5X as long as broad; flageliomere II 3.5 
as long as broad; flagellomere XI 6X as 
long as broad; mandible broadest medially, 
with long apical tooth, large subsidiary 
tooth and strong medial angle. Mesosoma: 
Legs unmodified; pronotal punctures tiny, 
2-3 PD apart; scutal punctures 2—4 PD 
apart medially, becoming denser laterally, 
0.5—1.0 PD apart; scutellar punctures 1—3 
PD apart; metanotum polished, punctures 
2-5 PD apart; propodeum densely, trans- 
versely shagreened, punctures obscure, 2—4 
PD apart; mesopleuron densely, finely sha- 
greened, punctures 1—2 PD apart. Metaso- 
ma: Finely shagreened, punctures tiny, 3— 
5 PD apart; epipygium lateral carina strong- 
ly produced; hypopygium ligulate, apically 
rounded, roughened dorsally (Fig. 26). 
Genital capsule (Figs. 38, 48): Paramere 
subtriangular, ventrally curved, tapering 
apically in lateral view, ventral surface 
strongly concave, broad, apically subtrun- 
cate, inner angle slightly hooked in dorsal 
view; aedeagus with large dorsal lobe. Col- 
or: Black, with yellow to red markings: 
clypeus ventrally yellow, antennal lobe yel- 
low, mandibles mostly yellow, inner eye 
margin with yellow stripe extending to ver- 
tex, connecting to stripe extending from 
posterior eye margin; pronotum with broad, 
transverse medial yellow stripe, large yel- 
low lateral spot; scutum with large, trilobate 
medial yellow spot, two small lateral spots 
and small yellow spot anteriorly on notau- 
lus; scutellum with large yellow medial and 
smaller lateral spot; metanotum yellow me- 
dially and laterally; propodeum with broad 
U-shaped sublateral yellow spots, narrowly 
separated medially, interrupted laterally in 
some specimens, with yellow dorsolateral 
spot; mesopleuron with large, J-shaped, yel- 


VOLUME 107, NUMBER 3 581 


582 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


low dorsal spot and smaller ventral one; fo- 
recoxa anteriorly yellow; mid and hindcox- 
ae yellow dorsally and laterally; fore- and 
midfemora yellow ventrally; tibiae, tarsi 
and hindfemur red to orange; metasomal 
tergum I yellow, with dark brown medial 
spot and subapical orange spots; segments 
I-IV primarily yellow, with orange basal 
markings and orange to dark brown apical 
markings; segment V blackish, with large, 
lateral, yellow spot; segment VI black, with 
lateral yellow tergal spot; epipygium dark 
brown, with basolateral yellow spot; wing 
membrane yellow-tinted, veins yellowish 
brown. Vestiture: Pale golden. 

Female.—Unknown. 

Material examined.—COLOMBIA: Bo- 
gota (the type series). 

Discussion.—Based on the coloration of 
the male and modifications of the male 
clypeus acuminatus is most similar to the 
new species nasalis and paradoxus, de- 
scribed below. All of these species have ex- 
tensive orange coloration on the legs and 
metasoma, and the clypeus has a medial 
tooth or projection. Merithynnus acumina- 
tus lacks the spined forecoxa and strongly 
elongate clypeus of paradoxus. It can be 
distinguished from nasalis by the clypeus 
having a single apicomedial lobe and ob- 
tuse sublateral ones. 


Merithynnus bilobatus Kimsey, 
new species 
(Figs. 4, 15, 27, 39, 49) 


Male.—Body length 10-14 mm. Head 
(Figs. 4, 15): Clypeus flat in side view, ap- 
icomedially elongate, with bilobate apex, 
side of elongation strongly convex, punc- 
tures sparse and 4—6 PD apart apically, be- 
coming denser, 0.5—1 PD apart, dorsally 
and laterally; area between eye margin and 
antennal socket with dense small punctures, 
contiguous to 0.5 PD apart; antennal lobes 
evenly rounded; frons punctures larger than 
on lower face, 1—2 PD apart; vertex sparse- 
ly punctate, punctures small, 1—4 PD apart; 
hindocellus 3 ocellar diameters from eye 
margin; mandible with long apical tooth, 


large submedial tooth and subbasal obtuse 
angle; flagellomere I 2.5 as long as broad; 
flagellomere If 3.5 as long as broad; fla- 
gellomere XI 6X as long as broad. Meso- 
soma: Pronotum weakly punctate, punc- 
tures minute, 3—8 PD apart; scutum pol- 
ished, punctures 1—4 PD apart; scutellar 
punctures 1—4 PD apart, laterally, nearly 
impunctate medially; metanotum impunc- 
tate; propodeum finely shagreened, nearly 
impunctate posteriorly; mesopleuron with 
dorsal margin of scrobal sulcus bulging, 
punctures 1—2 PD apart; grooves above pet- 
iolar socket faintly septate; legs unmodi- 
fied. Metasoma: Terga and sterna finely 
shagreened, punctures shallow, 1—4 PD 
apart; hypopygium broadly rounded apical- 
ly, dorsal surface transversely wrinkled 
(Fig. 27). Genital capsule (Figs. 39, 49): 
Paramere slender, elongate, tapering apical- 
ly in lateral view, broadened and slightly 
hooked in dorsal view, with submedially 
bulging inner margin; aedeagus with large 
dorsal lobe. Color: Black, with pale yel- 
lowish markings on lower two-thirds of 
clypeus, antennal lobe, base of mandible, 
face along eye margin with large spot ven- 
trad and small spot dorsad, vertex with 
transverse stripe, often interrupted subme- 
dially, gena with large spot along lower eye 
margin, mandible pale yellow medially; 
pronotum with transverse submedial stripe 
and large lateral spot; scutum with small 
medial spot and lateral stripe; scutellum 
with two medial and one anterolateral 
spots; metanotum yellow medially, with an- 
terolateral stripe; mesopieuron with com- 
ma-shaped dorsal spot, medial spot and 
posterior one; propodeum with lateral spot 
and submedial longitudinal stripes; coxae 
with dorsal spot, femora with ventral sur- 
face yellow; metasomal terga I-IV with lat- 
eral spot; wing membrane lightly brown- 
tinted, darkest in medial cell, veins dark 
brown, stigma yellowish brown medially. 
Vestiture: Long, sparse, silvery. 

Female.—unknown. 

Type material.—Holotype d: Caqueta 
Parque Nacional Natural, Cordillera de los 


VOLUME 107, NUMBER 3 583 


 \ 


14. acuminatus 15. bilobatus 


16. bogotanus 17. mimulus 19. nasalis 
18. montanus 


ee ee 
€ 
24. intermedius 


20. osorioi 
es 21. paradoxus r) >. 
22. pecki 23. turbulentus ( )siieitass 


UU ee vt 


25. desertus 
27. bilobatus 28. pecki 


amide 


31. osorioi 


26. acuminatus 30. osorioi 


32. paradoxus 


at 
Bea 


37. pecki 


33. intermedius 
34. osorioi 35. paradoxus 


Figs. 14-37. 14-23, Side view of male face, antenna removed. 24, 25, 30, 32, 37, Side view of female 
mesosoma. 26-28, Apex of hypopygium, dorsal view. 31, Apex of hypopygium, side view. 29, 33-36, Posterior 
view of female metasomal segment VI. 


584 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Pichachos, 2°44N 74°53W, 1,560 m, Mal- 
aise trap, E. Gonzalez, 1-7 Nov. 1997, No. 
662 (BOGOTA). Paratypes: 4 ¢, same data 
as holotype (BOGOTA, DAVIS). 

Etymology.—tThis species is named bi- 
lobatus in reference to the narrowly bilo- 
bate clypeus. 

Discussion.—The most distinctive fea- 
tures of male bilobatus are the black and 
pale yellow coloration, clypeus with the 
medial projection apicomedially narrowly 
bilobate and laterally convex to lobate, and 
the forewing marginal cell darker than the 
rest of the wing. The male clypeus appears 
flat in profile. 


Merithynnus bogotanus (Turner) 
(Figs. 5, 16, 27, 40, 50) 


Elaphroptera bogotana Turner 1910:196. 
Syntype males: Colombia: Bogota (LON- 
DON, BERLIN). 


Male.—Body length 13-15 mm. Head 
(Figs. 5, 16): Cypeus nearly impunctate, 
strongly produced apicomedially, apex 
sharply bidentate; area between antennal 
socket and eye margin with small punctures 
0.5—1.0 PD apart; frons and vertex with 
punctures 0.5 PD apart; hindocellus three 
hindocellar diameters from eye margin; fla- 
gellomere I 2.5—2.6X as long as broad; fla- 
gellomere II 3X as long as broad; flagel- 
lomere XI 5.0—5.3X as long as broad; man- 
dible medially and subbasally angulate, 
broadest submedially, with long acute api- 
cal tooth, and smaller subsidiary tooth. Me- 
sosoma: Pronotum polished, nearly im- 
punctate; scutum polished, punctures | PD 
apart; scutellum impunctate medially, punc- 
tures 1—2 PD apart laterally and anteriorly; 
metanotum impunctate; propodeum with 
small punctures 3—4 PD apart, nearly im- 
punctate above petiolar socket; mesopleu- 
ron polished, punctures 1—2 PD apart. Me- 
tasoma: Integument densely, finely sha- 
greened, punctures shallow, 1—4 PD apart; 
hypopygium apically broadly rounded, with 
dorsal surface irregularly wrinkled (Fig. 
27). Genital capsule (Figs. 40, 50): Para- 


mere slender, elongate and tapering apically 
in lateral view, broadened and _ slightly 
hooked apically in dorsal view; aedeagus 
without dorsal lobe, one-half or more as 
long as paramere. Color: Black, with yel- 
low markings; clypeus yellow, except api- 
comedial spot and around tentorial pits, in- 
ner eye margin with yellow stripe extending 
to hindocellus, antennal lobe yellow, man- 
dible yellow medially, posterior eye margin 
with yellow stripe along lower three- 
fourths, vertex with transverse yellow 
stripe; pronotum with transverse medial 
yellow stripe, interrupted medially, and 
large lateral yellow spot, scutum with pos- 
teromedial yellow spot and yellow lateral 
stripe, scutellum with two large medial and 
lateral yellow spots, metanotum yellow me- 
dially and laterally, propodeum with large 
U-shaped yellow sublateral spot; forecoxa 
ventrally yellow, mid and hindcoxae with 
dorsal and ventral yellow spot; femora with 
yellow ventral stripe, rest of legs black; me- 
tasomal tergum I with large, irregular U- 
shaped yellow spot, segments II-IV with 
large lateral tergal and sternal yellow spots, 
tergum V with large lateral yellow spot; 
wing membrane faintly brown, veins 
brown. Vestiture: Silvery. 

Female.—unknown. 

Material examined.—Colombia: Bogota 
(2 syntype males). 

Discussion.—Merithynnus bogotanus 
and turbulentus are similar in coloration 
and overall appearance in the males. Male 
bogotanus can be distinguished by the 
shape of the clypeus, and the more exten- 
sive yellow markings. The male clypeus has 
the apicomedial margin obtuse or rounded, 
and the face has a complete yellow stripe 
that extends along the inner eye margin 
from the clypeal margin to beyond the hin- 
docellus in bogotanus. In turbulentus the 
clypeus apicomedial margin is straight and 
the yellow stripe along the inner eye margin 
barely extends to the ocelli. In addition, the 
marginal cell in bogotanus is only slightly 
darker brown than the rest of the wing; 


VOLUME 107, NUMBER 3 585 


Caead 


: : . mimulus 
39. bilobatus 40. bogotaensis 42. montanus 


aa 


47. turbulentus 


38. acuminatus 


43. nasalis 44. osorioi 45. paradoxus 


48. acuminatus 49. bilobatus 50. bogotaensis 51. mimulus 


53. nasalis 54. osorioi 55. paradoxus 56. pecki 


57. turbulentus 


Figs. 38-57. 38-47, Male genital capsule; side view (left). 48-57, Dorsal view (right). 


586 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


whereas the marginal cell in turbulentus is 
conspicuously darkened. 


Merithynnus desertus (Turner) 
(Figs. 25, 29) 


Elaphroptera deserta Turner 1910:199. 
Syntype females; Colombia: Bogota 
(BERLIN). 


Male.—Unknown. 

Female.—Body length 7-8 mm. Head: 
Mandible long, slender, edentate; frons and 
vertex with punctures 0.5—1.0 PD apart, 
vertex with long seta from each puncture; 
scape ventrally with row of stout erect se- 
tae. Mesosoma (Fig. 25): Pronotum with 
anterior collar half as long as elevated disk, 
disk strongly depressed along notauli ante- 
riorly, posteriorly deeply sunken medially, 
integument finely shagreened, punctures |— 
2 PD apart; propleuron markedly convex 
ventrally; scutellum conspicuously elevated 
above pronotum, ovoid, with contiguous 
punctures; scutum and metanotum barely 
visible dorsally; propodeum dorsal surface 
about as long as scutellum, markedly decli- 
vous posteriorly, without long erect setae. 
Metasoma: Tergum I rounded anteriorly, 
without dense, erect, long setae, with trans- 
verse subapical row of pits; tergum II with 
transverse, broadly U-shaped, submedial 
carina, followed by large transverse sub- 
apical ridge, coarsely sculptured between, 
posterior margin flared; terga III—V trans- 
versely, finely wrinkled, punctures obsoles- 
cent, with subapical, broadly W-shaped 
transverse groove before flat, smooth apical 
rim; apical surface of tergum VI carina- 
edged, carina sharply angulate, with lateral 
brush of setae, apicomedially with acute 
tooth, with a short tuft of setae on either 
side, with medial longitudinal row of setae 
(Fig. 29). Color: Yellowish brown to dark 
brown. 

Material examined.—COLOMBIA: Bo- 
gota (only the holotype of deserta was 
seen). 

Discussion.—The male is unknown for 
deserta. It is most likely that the male is 


one of the three Merithynnus described by 
Turner (1910) from Bogota: acuminatus, 
bogotanus and mimulus. The collector and 
locality labels are identical among the type 
series of these species. The female of de- 
serta can be immediately distinguished 
from other female Merithynnus by the api- 
comedial tooth on the pygidium and the 
posteromedially depressed pronotum. 


Merithynnus intermedius Kimsey, 
new species 
(Figs. 24, 33) 


Male.—Body length 12 mm. Head: 
Clypeus convex medially, strongly pro- 
duced apicomedially, apex rounded, sides 
concave, nearly impunctate medially and 
apically, dorsally with dense, nearly contig- 
uous punctures; area between antennal 
socket and eye margin with small punctures 
0.5—1.0 PD apart; frons and vertex punc- 
tures 0.5—1.0 PD apart, punctures smaller 
on vertex; hindocellus 2.8—3.0 ocellar di- 
ameters from eye margin; flagellomere I 
2.5-2.6X as long as broad; flagellomere II 
3X as long as broad; flagellomere XI miss- 
ing; mandible with long acute apical tooth, 
medially with smaller subsidiary tooth, sub- 
basally angulate, broadest submedially. Me- 
sosoma: Pronotum polished, nearly im- 
punctate; scutum polished, punctures 1 PD 
apart; scutellum impunctate medially, lat- 
erally punctures 1—2 PD apart; metanotum 
nearly impunctate; propodeum finely sha- 
greened medially, with small punctures 2— 
3 PD apart, punctures smaller and 1—2 PD 
apart laterally; mesopleuron polished, punc- 
tures 2—4 PD apart. Metasoma: Integument 
densely, finely shagreened, punctures shal- 
low, 2—4 PD apart; hypopygium missing. 
Genital capsule: Missing. Color: Black, 
with yellow markings; clypeus yellow, ex- 
cept along dorsal margin, inner eye margins 
with yellow stripes meeting behind hindo- 
cellus, antennal lobe yellow, mandible yel- 
low medially, posterior eye margin with 
large U-shaped, yellow stripe along lower 
four-fifths; pronotum with transverse me- 
dial yellow stripe, interrupted medially, and 


VOLUME 107, NUMBER 3 


large lateral yellow spot, scutum with pos- 
teromedial yellow spot and yellow lateral 
stripe, scutellum with one large medial and 
one lateral yellow spot, metanotum yellow 
medially and laterally, propodeum with 
large U-shaped yellow sublateral spots; 
coxae dorsally yellow; femora with yellow 
ventral stripe, rest of legs black; metasomal 
tergum I with large, irregular U-shaped yel- 
low spot, tergum II with transverse medial 
yellow stripe; terga III—-V with lateral yel- 
low spot; sterna II-III each with small lat- 
eral yellow spot; wing membrane yellow- 
tinted, veins brown. Vestiture: Silvery. 
Female.—Body length 9 mm. Head: 
Mandible long, slender, edentate; frons with 
ovoid depression adjacent to upper inner 
eye margin; frons and vertex with punctures 
1—3 PD apart, frons with long seta from 
each puncture; scape ventrally with row of 
stout, erect setae. Mesosoma (Fig. 24): 
Pronotum flattened, with anterior collar 
more than half as long as elevated disk, disk 
strongly depressed, trough-like anterome- 
dially, floor of depression planar with an- 
terior collar, integument polished, with 
punctures 2—3 PD apart posteriorly, nearly 
impunctate anteriorly; propleuron strongly 
bulging ventrally and laterally; scutellum 
elevated above pronotum, ovoid, with 
punctures 1—3 PD apart;.scutum and metan- 
otum barely visible dorsally; propodeum 
dorsal surface about as long as scutellum, 
markedly concave posteriorly, conspicuous- 
ly angled laterally, with long, erect setae 
dorsally and laterally; forecoxa narrowly at- 
tached to prothorax (Fig. 24). Metasoma: 
Tergum I markedly narrowed anteriorly, 
with dense, erect, long setae, with trans- 
verse subapical row of pits; tergum II with 
transverse, broadly U-shaped, submedial 
carina, followed by large transverse sub- 
apical ridge, shagreened between, posterior 
margin flared; terga III—-V transversely, 
finely wrinkled, punctures obsolescent, with 
subapical, broadly W-shaped transverse 
groove before flat, smooth apical rim; api- 
cal surface of tergum VI with narrow cari- 
na-edged enclosure, dorsal carina indented 


587 


medially, with narrow, medial brush of se- 
tae, ventral edge with broad, short medial 
brush of setae (Fig. 33). Color: Yellowish 
brown to dark brown, facial depressions 
pale. 

Type material.—Holotype male and 
paratype female; COLOMBIA: Cundina- 
marca Prov., Monterredondo, Jan. 1959, 
1,420 m, Forester (DAVIS). 

Etymology.—The name is chosen to in- 
dicate the intermediate nature of structural 
features of the male between montanus and 
pecki. 

Discussion.—Merithynnus intermedius 
most closely resembles montanus and 
pecki, based in the male on the black and 
yellow coloration, the clypeus without a 
medial projection and apicomedially drawn 
out into an elongate lobe. It can be distin- 
guished from these two species by the api- 
cally rounded clypeal projection, clypeus 
convex in side view (not flat or concave), 
face with complete yellow stripe extending 
behind the hindocelli, and scutum and scu- 
tellum with a single large yellow spot (two 
on both structures in pecki, and two only 
on the scutelium in montanus). Females can 
be distinguished by the deeply medially im- 
pressed pronotum, strongly bulging pro- 
pleuron and narrowly attached forecoxa. 


Merithynnus mimulus (Turner) 
(Figs. 6, 17, 41, 51) 


Elaphroptera mimula Turner 1910:197. Ho- 
lotype male; Colombia: Bogota (BER- 
LIN). 


Male.—Body length 12-15 mm. Head 
(Figs. 6, 17): Clypeus concave medially, 
with elongate narrow apicomedial trunca- 
tion, truncation apex equal to or narrower 
than distance between antennal sockets, 
punctures |—3 PD apart medially, becoming 
denser along dorsal margin; vertex and 
frons with punctures contiguous to 0.5 PD 
apart, punctures smaller along lower eye 
margin than above antennal sockets; hin- 
docellus 2.5 ocellar diameters from eye 
margin; flagellomere I 2.5 as long as 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


broad; flagellomere II 3X as long as broad; 
mandible broadest submedially, with one 
large subapical tooth. Mesosoma: Prono- 
tum sparsely punctate, punctures 1—3 PD 
apart; scutal punctures larger than pronotal, 
0.5—1.0 PD apart; scutellar punctures 2—3 
PD apart; metanotum impunctate; propo- 
deum polished, with small punctures, nearly 
impunctate above petiolar socket, becoming 
more densely punctate laterally, punctures 
1—2 PD apart; mesopleuron polished, punc- 
tures small, 2-4 PD apart. Metasoma: Seg- 
ments finely, densely shagreened, punctures 
shallow, 1-2 PD apart; hypopygium apex 
rounded, dorsal surface finely and densely 
rugose. Genital capsule (Figs. 41, 51): Par- 
amere slender, elongate and tapering api- 
cally in lateral view, broadened and slightly 
hooked in dorsal view, with submedially 
bulging inner margin; aedeagus without 
dorsal lobe. Color: Head and mesosoma 
black, with yellow markings, metasoma 
red, with yellow and black markings, legs 
red; clypeus ventrally yellow, antennal lobe 
with large yellow spot, one large and one 
small yellow spot along inner eye margin, 
posterior eye margin with yellow stripe ba- 
sally, vertex with transverse yellow stripe 
interrupted medially, mandible yellow me- 
dially; pronotum with transverse medial 
yellow stripe and lateral yellow spot, me- 
sopleuron with one dorsal and two ventral 
yellow spots, scutum with yellow postero- 
medial spot and lateral stripe, scutellum 
with two medial and one lateral yellow 
spot, metanotum medially yellow, with lat- 
eral yellow stripe, propodeum with subla- 
teral U-shaped yellow, orange or red spot; 
metasomal tergum I blackish basally, with 
subapical transverse yellow stripe, inter- 
rupted medially in some specimens, seg- 
ments II-IV each with large lateral yellow 
spot, tergum V blackish basally and apical- 
ly, terga VI-VII and sterna V—VII dark 
brown to black; wing membrane yellow- 
tinted, veins yellowish brown to dark 
brown. Vestiture: Long, pale golden. 
Female.—Unknown. 


Material examined.~COLOMBIA: Bo- 
gota (only the type specimens were seen). 

Discussion.—Merithynnus mimulus 1s 
the only species in the male with red col- 
oration on the legs and metasoma that lacks 
a medial tooth or projection on the clypeus. 
All of the other Merithynnus species with 
red coloration have a medial clypeal tooth. 
In addition, the apex of the apicomedial ex- 
tension of the clypeus is narrow, as wide as 
the distance between the antennal sockets. 
In montanus, acuminatus, nasalis, osoriot 
and pecki the apex is broader than the in- 
terantennal distance. The shape of the par- 
amere and aedeagus resembles those of bo- 
gotanus. 


Merithynnus montanus Kimsey, 
new species 
(Figs. 7, 18, 42, 52) 


Male.—Body length 12-17 mm. Head 
(Figs. 7, 18): Clypeus slightly bulging dor- 
sally, concave medially, broadly subtrun- 
cate apicomedially, apical margin of trun- 
cation indented, punctures sparse and 4—6 
PD apart apically, becoming denser dorsally 
and laterally, 0.5—1 PD apart; area between 
eye margin and antennal socket with dense 
small punctures, 1 PD apart; antennal lobes 
evenly rounded; frons punctures larger than 
on lower face, contiguous to 0.5 PD apart; 
mandible with subapical tooth and broadly 
obtuse subbasal angle on inner margin; ver- 
tex with punctures 0.5—1.0 PD apart; hin- 
docellus 2.5 ocellar diameters from eye 
margin; flagellomere I 2.5 as long as 
broad; flagellomere If 3.5xX as long as 
broad; flagellomere XI 6X as long as broad. 
Mesosoma: Pronotum nearly impunctate; 
scutal punctures 1—2 PD apart; scutellum 
sparsely punctate, punctures 1—4 PD apart, 
nearly impunctate medially; metanotum im- 
punctate; propodeum polished, punctures 
2—4 PD apart, except impunctate above pet- 
iolar socket, grooves above petiolar socket 
conspicuously septate; mesopleuron with 
dorsal margin of scrobal sulcus bulging, 
punctures 1—2 PD apart, posteriorly finely 
shagreened; legs unmodified. Metasoma: 


VOLUME 107, NUMBER 3 


Terga and sterna finely shagreened, punc- 
tures shallow, 1—4 PD apart; hypopygium 
broadly rounded apically, dorsal surface 
transversely wrinkled. Genital capsule 
(Figs. 42, 52): Paramere slender, elongate, 
tapering apically in lateral view, broadened 
and slightly hooked in dorsal view, with 
submedially bulging inner margin; aedea- 
gus without dorsal lobe. Color: Body 
black, with yellow markings; clypeus yel- 
low except around tentorial pits, mandibles 
yellow medially, antennal lobes yellow; eye 
surrounded by yellow stripe, except poste- 
riorly near vertex, posterior stripe with 
branch extending across vertex, interrupted 
submedially; pronotum with transverse me- 
dial yellow stripe and large lateral yellow 
spot, scutum with yellow medial spot and 
lateral stripe; scutellum with two medial 
and one lateral yellow spot; metanotum yel- 
low medially and laterally, mesopleuron 
with three large yellow spots, propodeum 
with large comma-shaped sublateral stripe 
and lateral spot, coxae with anterior yellow 
spot; femora with ventral yellow stripe, me- 
tasomal terga I—V and sterna I-IV each 
with irregular lateral yellow spot; wing 
membrane brown-tinted, veins dark brown. 
Vestiture: Sparse and silvery. 

Female.—Unknown. 

Type material—Holotype 6, COLOM- 
BIA: Boyaca Santuario de Flora y Fauna, 
Iguaque Cabafia Chaina, 5°25'N 73°27'W, 
2,600 m, Malaise trap, 17 May—5 Jun. 
2001, P. Reina, No. M1758 (BOGOTA). 
Paratypes: 14 6; 3 d: same data as holo- 
type: 5 6: 30 Apr—17 May 2001, No. 
Mil7533"6) 63 21° Jun—6 Jul) 2001, No. 
M1971 (BOGOTA, DAVIS). 

Etymology.—The species is named after 
the collection site in montane Colombia. 

Discussion.—Among the species that 
lack reddish coloration and lack a medial 
clypeal projection in the male two species, 
montanus and pecki, have the apicolateral 
angles of the clypeal projection forming a 
right to obtuse angle, as opposed to acute. 
Merithynnus montanus can be distinguished 
in the male from pecki by the saddle-shaped 


589 


clypeus (seen in profile) and aedeagus with- 
out a ventral lobe before the apical strap. 


Merithynnus nasalis Kimsey, 
new species 
(Figs. 8, 19; 43, 53) 


Male.—Body length 14-17 mm. Head 
(Figs. 8, 19): Clypeus polished, impunc- 
tate, except along dorsal margin (punctures 
small, nearly contiguous), apicomedially 
truncate, with broad flattened triangular 
area below medial tooth or projection; an- 
tennal lobes rounded; mandible broadly an- 
gulate medially, with long apical tooth and 
smaller subapical one; area between eye 
margin and antennal socket with dense 
small punctures, | PD apart; antennal lobes 
angulate; frons finely shagreened, punctures 
larger than on lower face, contiguous to 0.5 
PD apart; vertex finely shagreened, with 
punctures 0.5—1.0 PD; hindocellus 2.5 ocel- 
lar diameters from eye margin; flagellomere 
I 2.5—2.0X as long as broad; flagellomere 
If 3.5 as long as broad; flagellomere XI 
6X as long as broad. Mesosoma: Pronotum 
without transverse anterior carina, finely 
shagreened, punctures tiny, 1—3 PD apart: 
scutal punctures 1—4 PD apart medially, 
nearly contiguous laterally; scutellum near- 
ly impunctate medially, 1-2 PD apart lat- 
erally; metanotum impunctate; propodeum 
finely shagreened, punctures 2—4 PD apart, 
except impunctate above petiolar socket, 
petiolar grooves smooth, without septa; me- 
sopleuron finely shagreened, dorsal margin 
of scrobal sulcus not bulging, punctures |— 
2 PD apart; legs unmodified. Metasoma: 
Terga and sterna finely shagreened, punc- 
tures shallow, 2—4 PD apart; hypopygium 
narrowly ligulate apically, dorsal surface ir- 
regularly sculptured. Genital capsule (Figs. 
43, 53): Paramere less than twice as long 
as broad, bending ventrally, ventral half 
concave, apically bidentate in dorsal and 
lateral views, with submedially bulging in- 
ner margin, marked by conspicuous row of 
pits; aedeagus short, with heavily sclero- 
tized dorsal lobe, apical lobe with small, 


membranous lateral lobe. Color: Black, 


590 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


with yellow markings, and red on legs and 
metasoma, clypeus with yellow stripe along 
apical margin extending dorsally over me- 
dial projection, mandible yellow medially; 
antennal lobe with yellow spot; inner eye 
margin with two spots, posterior eye margin 
with basal yellow stripe extending dorsally 
from eye margin near midpoint to vertex; 
pronotum with broad transverse yellow 
stripe and large lateral spot; scutum with 
medial stripe and two small lateral spots; 
tegula yellow; scutellum with large medial 
and small anterolateral yellow spots; metan- 
otum yellow medially; propodeum with 
large U-shaped sublateral stripe; mesopleu- 
ron with dorsal U-shaped yellow spot and 
posterior yellow spot; coxae with anterior 
yellow spot; femora with ventral yellow 
stripe; forefemur blackish red to black, mid 
and hindfemora red; tibiae red; tarsi red to 
yellow; metasomal terga and sterna with 
large, irregular yellow spot; tergum I black- 
ish basally; tergum V black, with large yel- 
lowish lateral spot, sternum V black, with 
small lateral yellow spot; segments VI, VII 
blackish; wing membrane yellow-tinted, 
veins brown. Vestiture: Sparse, golden. 

Female.—Unknown. 

Type material.—Holotype 6: COLOM- 
BIA: Boyaca, Santuario de Flora y Fauna 
Iguaque, 5°25'N 73°27'W, 2,820 m, Malaise 
trap, 17 Mar—lApr. 2000, P. Reina, No. 
M130 (BOGOTA). Paratypes: 18 36; 5 4, 
same data as holotype; 5 d, 2-19 Apr. 
2000, No. M31; 8 6, 28 Feb.—-17 Mar. 
2000, No. M9 (BOGOTA, DAVIS). 

Etymology.—The species is named for 
the sharp, noselike projection on the clyp- 
eus. 

Discussion.—This is one of the three 
species (acuminatus, nasalis and paradox- 
us) with red coloration and a medial clypeal 
projection in the male. Merithynnus nasalis 
can be distinguished from acuminatus by 
the flat to concave apicomedial clypeal 
margin, versus medially lobate one in acu- 
minatus. Both of these species have a slen- 
der, ligulate hypopygium; other Merithyn- 
nus have the hypopygium broader and more 


horse hoof-like or parallel—sided. The male 
genitalia of nasalis are unusual. The para- 
mere is short, less than twice as long as 
broad and bends ventrally. The ventral sur- 
face is concave. In addition, the paramere 
is apically bidentate, with the inner margin 
submedially bulging, marked by conspicu- 
ous row of pits. The aedeagus is short, with 
a heavily sclerotized dorsal lobe, and the 
apical strap has a small, membranous lateral 
lobe. 


Merithynnus osorioi Kimsey 
(Figs. 9, 20, 30-31, 34, 44, 54) 


Merithynnus osorioi Kimsey 1991:77. Ho- 
lotype male: Venezuela: Yaracuy, La 
Puerta (GAINESVILLE). 


Male.—Body length 16 mm. Head (Figs. 
9, 20): Clypeus nearly impunctate apically, 
dorsally punctures contiguous to 0.5PD 
apart, with elongate medial truncation, trun- 
cation apex medially slightly emarginated, 
with acute lateral angles, sides strongly in- 
curved; area between antennal socket and 
eye margin with punctures 0.5 PD apart; 
mandible slender, with elongate apical 
tooth, large subapical tooth and large sub- 
basal angle on inner margin; vertex and 
frons punctures contiguous to 0.5 PD apart, 
smaller and sparser along lower eye mar- 
gin; hindocellus 3 ocellar diameters from 
eye margin; flagellomere I 2.5 as long as 
broad; flagellomere II 3X as long as broad. 
Mesosoma: pronotum polished with tiny 
shallow punctures 2—4 PD apart; scutal 
punctures 0.5—2.0 PD apart; scutellum pol- 
ished, punctures 2—5 PD apart; metanotum 
3—4 PD apart, becoming denser laterally; 
propodeum finely shagreened, with sparse, 
shallow punctures (punctures 0.5—1.0 PD 
apart laterally), nearly impunctate medially; 
mesopleural punctures 0.5—1.0 PD apart. 
Metasoma: Finely shagreened, punctures 
1—3 PD apart; hypopygium broadly round- 
ed apically, laterally notched (Fig. 31). 
Genital capsule (Figs. 44, 54): Paramere 
abruptly narrowed apically, ventral margin 
deeply indented subapically; aedeagal shaft 


VOLUME 107, NUMBER 3 


short, with large ventral lobe before apical 
strap. Color: Black, with yellow markings; 
clypeus ventromedially yellow, mandible 
yellow medially; antennal lobe yellow; eye 
with two yellow spots on inner margin, one 
on posterior margin; vertex with short lat- 
eral yellow stripe; pronotum with trans- 
verse, yellow medial stripe, interrupted me- 
dially, with small yellow lateral spot; scu- 
tum and scutellum with small yellow me- 
dial and lateral spots; propodeum with 
comma-shaped yellow spot and lateral yel- 
low spot; mesopleuron with three yellow 
spots; coxae with dorsal yellow spot; fem- 
ora yellow ventrally; metasomal terga I-IV 
and sternum III each with small lateral yel- 
low spot; wing membrane yellow-tinted, 
marginal cell darker than rest of wing; veins 
brown. Vestiture: Sparse and erect, silvery 
ventrally, becoming brown dorsally. 

Female.—Body length 10 mm. Head: 
Mandible slender, edentate; frons polished, 
punctures 4—6 PD apart. Mesosoma (Fig. 
30): Pronotal disk subquadrate, convex me- 
dially, without declivity before collar, an- 
terolateral corners of disc elevated, with 
dorsal tuft of erect setae; propleuron convex 
in profile; propodeum with long dorsal sur- 
face, sharply declivous to posterior face, 
juncture marked by sharp thorn-like medial 
projection. Metasoma: Tergum VI pygidi- 
um coarsely, irregularly punctate, covered 
by long, erect, dense setae, apically 5- 
lobed, with shallow apicomedial lobe; ster- 
num VI apical plate forming half circle, 
without dorsal emargination (Fig. 34). Col- 
or: Dark, reddish brown. 

Material examined.—VENEZUELA: 
Yaracuy, La Puerta (only the type series 
was seen). 

Discussion.—Males of Merithynnus oso- 
rioi most closely resemble those of bogo- 
tanus and turbulentus. In the males, osorioi 
can be distinguished from these species by 
the laterally acute apical projection (obtuse 
or forming a right angle in bogotanus and 
turbulentus) of the clypeus, finely shagree- 
ned propodeum and no yellow spot between 
the ocelli. 


591 


Merithynnus paradoxus Kimsey, 
new species 
(Pigss LOA 35 2123358455555) 


Male.—Body length 10-17 mm. Head 
(Figs. 10, 21): Clypeus dorsally punctate 
and shagreened, ventrally polished and 
nearly impunctate, with acute medial pro- 
jection, otherwise concave medially, pro- 
duced apicomedially into narrow upturned, 
apically truncate lobe; frons and vertex sha- 
greened, with coarse contiguous punctures; 
antennal lobes apically angulate; hindocel- 
lus 2.5 MOD from eye margin and 3 MOD 
from posterior head margin; mandible with- 
out subbasal tooth, with submedial angle on 
inner margin, broadest submedially; flagel- 
lomere I 3.5 as long as broad; flagello- 
mere II 4X as long as broad; flagellomere 
IX 5x as long as broad. Mesosoma: Sur- 
faces of pronotum, mesopleuron, scutum 
and propodeum shagreened, with dense 
contiguous punctures; scutellum and metan- 
otum polished, with punctures 1—2 PD 
apart; forecoxa with long digitate apical 
lobe (Fig. 13); hindtrochanter subtriangular 
in profile, with angular apicoventral corner. 
Metasoma: Segments finely shagreened, 
with punctures 1—3 PD apart. Genital cap- 
sule (Figs. 45, 55): Paramere curved ven- 
trally, parallel-sided, broadly rounded api- 
cally, with acute apicoventral tooth in lat- 
eral view, slender, with apical thumb-like 
lobe pointing toward midline in dorsal 
view; aedeagus with large, sclerotized dor- 
sal lobe. Color: Black, with yellowish 
markings, face with two broadly separated 
yellow spots along inner eye margin and 
two along outer eye margin; one (some- 
times two) yellow spots on antennal lobe, 
clypeus ventrally yellow; mandible with 
small medial yellow spot; pronotum with 
yellow stripe along posterior margin and 
small lateral spot; tegula with yellow spot; 
mesopleuron with oblique yellow stripe; 
coxae with dorsal yellow spot; scutum and 
scutellum each with small medial yellow 
spot; metanotum whitish medially; propo- 
deum with slender, sublateral longitudinal 


192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


stripe; fore and midfemur black dorsally, 
becoming redder ventrally; trochanters 
blackish; rest of leg red; metasomal seg- 
ments I-IV red, with yellow medially, seg- 
ment I black dorsobasally; segment V 
black, with yellow lateral tergal spot; seg- 
ments VI and VII black; wing membrane 
evenly yellow-tinted, veins yellowish 
brown. Vestiture: Pale golden ventrally, be- 
coming dark brown on vertex and thoracic 
dorsum. 

Female.—Body length 7-8 mm. Head: 
Mandible short, slender, edentate; frons 
coarsely dentate, punctures contiguous, be- 
coming sparser toward vertex, with 2—4 
long setae; scape ventrally with row of 
stout, erect setae. Mesosoma (Fig. 32): 
Pronotum with anterior collar nearly as 
long as elevated disk, disk anteriorly trilo- 
bate, medial lobe acute, punctures 1—5 PD 
apart; scutellum elevated, ovoid, with con- 
tiguous punctures; scutum and metanotum 
barely visible; propodeum without dorsal 
surface, dorsal half concave, ventral part 
slightly convex, with dense, long, erect se- 
tae. Metasoma: Tergum I abruptly decli- 
vous anteriorly, anterior surface with dense, 
erect, long setae, with transverse subapical 
row of pits; tergum II with transverse, U- 
shaped, submedial carina, followed by large 
transverse subapical ridge, coarsely sculp- 
tured between, posterior margin flared; ter- 
ga III-V with transverse submedial punctate 
stripe, punctures contiguous to | PD apart, 
with subapical, broadly W-shaped trans- 
verse groove before flat, smooth apical rim; 
apical surface of tergum VI carina-edged, 
carina sharply angulate, with lateral brush 
of setae, apex bilobate, with submedial lon- 
gitudinal row of setae (Fig. 35). Color: 
Yellowish brown to dark brown. 

Type material—Holotype 6: COLOM- 
BIA: Santuario de Flora y Fauna Iguaque, 
La Planada, 5°25N 73°27W, 2,850 m, P. 
Reina, Malaise trap, 28 Feb.—Mar. 2000, 
No. M6; Malaise trap (BOGOTA). Para- 
types: 231 5, 4 2; same locality as holo- 
type: 5 6, 19 Apr—6 May 2000, No. M57; 
5 3, 28 Feb.—Mar. 2000, No. M6; 5 6, 11 


Oct._1 Nov. 2000, No. M751; 5 6, 1-13 
Nov. 2000, No. M827; 6 3, 28 Feb.—16 
Mar. 2000, No. M40; 6 6, 9-23 Mar. 2001, 
No. M1513; 7 6, 2-19 Apr. 2000, No. 
M32: 7 6, 7-21 Jan-- 2001, No: M125i25 
3, 8-25 Jun. 2000, No. M189; 7 6 21 Jan— 
7 Feb. 2000, No. M1249; 6 6, 2—22 Aug. 
2001, No. M2023; 5 6, 17 Aug.—1 Sep. 
2000, No. M518; 5 6, 25 Jun—13 Jul 
2000, No. M248; 7 6, 23 Sep.—11 Oct. 
2000, No. M754; 6 6, 4—21 Dec. 2000, No. 
M1079; 6 3, 2-19 Apr. 2000, No. M28; 4 
6, 28 Feb—17 Mar. 2000, No. M7; 5 6, 
13-30 Jul. 2000, No. M381; 5 46, 21 Dec.— 
7 Jan. 2001, No. M1070; 4 3, 1—23 Sep. 
2000, No. 616; 6 5, 6—23 May 2000, No. 
M73; Boyaca, Cabana, Mamarramos, 
5°25N 73°27W, 2,855 m, Malaise trap: 1 d, 
16 Mar—1 Apr. 2000, M123; 1 d, 1-23 
Sep. 2000, No. M614; 4 3, 16 Mar.—1 Apr. 
2000, No. M124; 2 6 1-19 Apr. 2000, No. 
M23; 2 dg, 1-19 Apr. 2000, No. M24; 1 6, 
21 Dec 2000-7 Jan. 2001, No. M1072; 
Boyaca, Cerro Pan de Azucar, 5°25’N 
73°27'W, 3,300 m: 6 6, 16 Apr.—2 May 
2001, No. M1739; 4 3, 1 @, 2-22 Aug. 
2001, No. M2023; 5 6, 10—28 Jun. 2001, 
No. M1834; 5 6,1 2, 18 Jul.—2 Aug. 2001, 
No. M1967; 7 63, 12 Sep.-13 Oct. 2001, 
No. M2197; 6 6, 28 Jun.—18 Jul. 2001, No. 
M1965; 7 6, 22 Aug.—12 Sep. 2001, No. 
M2067; 4 6, 18 May—10 Jun. 2001, No. 
M1767; 5 3, 27 Mar—16 Apr. 2001, No. 
M1517; 9 3, 9-23 Mar. 2001, No. M1513; 
6 3d, 2-18 May 2001, No. M1754; Boyaca, 
5°25N 73°27W, 2,820 m, Malaise trap: 6 6 
28 Feb.—17 Mar. 2000, No. 9; 1 6, 2 @ 17 
Mar.—1 Apr. 2000, No. M126; 2 6, 17 
Mar.—-1 Apr. 2000, No. M130; 3 6, 16 
Mar.—1 Apr. 2000, No. M128; 4 6, 2-19 
Apr. 2000, No. M27; 6 6, 28 Feb._16 Mar. 
2000, No. M5; 4 6, 28 Feb.—17 Mar. 2000, 
No. M7; 5 3, 28 Feb.-17 Mar. 2000, No. 
M8; Norte de Santander Parque Nacional 
Natural, Tama, El Arenal, 7°26N 72°26W, 
1,250 m, Malaise trap, A. Cortes, M291: 2 
3, 7-15 Aug. 1998: (BOGOTA, DAVIS). 
Etymology.—The name, paradoxus, is 
Greek (masculine) for strange, paradoxical, 


VOLUME 107, NUMBER 3 


referring to the odd modifications of the 
clypeus and forecoxa. 

Discussion.—The structure of the male 
clypeus and foreleg will immediately distin- 
guish this species from all others in the ge- 
nus. Otherwise it is one of three species, 
including acuminatus and nasalis, with red- 
dish markings and a tooth-like medial pro- 
jection on the clypeus. The female can be 
distinguished by the anteriorly trilobate 
pronotum, apically bilobate pygidium and 
dorsally concave propodeum. 


Merithynnus pecki Kimsey 
(Figs. 11, 22, 36-37, 46, 56) 


Merithynnus pecki Kimsey 1991:75. Holo- 
type male; Venezuela: Mérida, El Valle, 
15 km east Mérida (OTTAWA). 


Male.—Body length 11-17 mm. Head 
(Figs. 11, 22): Clypeus flat apically, in- 
dented medially, with broad, medially 
emarginate apex, lateral angle of apex 
forming 90° to slightly obtuse angle, punc- 
tures 2—4 PD apart apically, becoming con- 
tiguous dorsally; antennal lobe flattened an- 
teriorly, not strongly projecting above clyp- 
eal margin; frons and vertex with contigu- 
ous punctures; hindocellus 2.6—2.7 ocellar 
diameters from eye margin; flagellomere I 
2.5 as long as broad; flagellomere II 3.5 
as long as broad; flagellomere XI 5X as 
long as broad; mandible slender, with small 
subapical tooth, without subbasal angle, 
broadest subbasally. Mesosoma: Pronotum 
finely shagreened, densely punctate, punc- 
tures contiguous to 0.5 PD apart; scutal 
punctures 0.5—1.0 PD apart medially, be- 
coming contiguous laterally; scutellar punc- 
tures 2-4 PD apart medially, becoming 
nearly contiguous laterally; metanotal punc- 
tures 1-2 PD apart; propodeum finely sha- 
greened, punctures obscure, 1—2 PD apart; 
mesopleuron finely shagreened, punctures 
0.5—1.0 PD apart. Metasoma: Integument 
finely shagreened, punctures 0.5—2.0 PD 
apart; hypopygium apically ovoid, dorsal 
surface smooth. Genital capsule (Figs. 46, 
56): Paramere broadly rounded apically, 


593 


with small apicoventral tooth, outer surface 
markedly convex, ventral margin sinuous to 
lobed medially; aedeagus about half as long 
as paramere, without dorsal lobe, with ven- 
tral membranous lobes before apical strap. 
Color: Black, with yellow markings; clyp- 
eus ventrally yellow; mandible medially 
yellow, antennal lobe yellow, inner eye 
margin with yellow stripe extending to top 
of eye; lower posterior eye margin with 
large yellow spot; vertex with transverse 
yellow stripe, interrupted medially; prono- 
tum with transverse medial yellow stripe 
and anterolateral yellow spot; scutum with 
two small yellow posteromedial spots and 
lateral yellow stripe; scutellum with two 
yellow medial spots and anterolateral yel- 
low spot; metanotum medially yellow, with 
lateral yellow spot; propodeum with com- 
ma-shaped, sublateral yellow spot and an- 
terolateral spot; mesopleuron with anterior 
U-shaped yellow spot and small ovoid pos- 
terior yellow spot; coxae with dorsal and 
ventral yellow spots; femora yellow ven- 
trally; tarsi dark brown; metasomal tergum 
I with yellow basolateral spot and trans- 
verse subapical yellow stripe; sternum I 
with small lateral yellow spot; tergum II 
with transverse yellow submedial stripe; 
terga III—V and sterna I-IV with large lat- 
eral yellow spot; wing membrane yellow- 
tinted, veins yellowish brown. Vestiture: 
sparse, silvery. 

Female.—Body length 8—9 mm. Head: 
Mandible long, slender, edentate; frons 
coarsely punctate, punctures large, 1-2 PD 
apart, with tiny punctures in interspaces, 
with 2—5 scattered long setae. Mesosoma 
(Fig. 37): Finely, densely shagreened; 
pronotum with anterior collar three-fourths 
as long as elevated disk, disk anteriorly bi- 
lobate, evenly convex, punctures I1—2 PD 
apart; scutellum elevated, about as long as 
broad, punctures 1—2 PD apart; propodeum 
with dorsal surface longer than scutellum, 
deeply concave posteromedially. Metaso- 
ma: Finely, densely shagreened; tergum | 
narrowed anteriorly, subconical, declivous 
anteriorly, anterior surface with dense, erect 


594 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


setae, with transverse indented, subapical 
row of pits; tergum I with transverse, U- 
shaped submedial carina, followed by large 
transverse subapical ridge, coarsely sculp- 
tured between, posterior margin flared; ter- 
ga III-V each with transverse submedial 
punctate stripe, punctures contiguous to | 
PD apart, with subapical broadly W-shaped 
transverse groove before flat, smooth apical 
rim; apical surface of tergum VI carina- 
edged, posteriorly coarsely and irregularly 
punctate, with submedial longitudinal row 
of setae, marginal carina sharply angulate, 
with lateral brush of setae, apex with large 
triangular medial projection, with adjacent 
tuft of setae (Fig. 36). Color: Yellowish 
brown to dark brown. 

Material examined.—50 6, 5 @: VE- 
NEZUELA: Mérida, El Valle, 15 km e Mé- 
rida; Tabay Mucuy; S. Truchicola; 20 km 
se Azulita; La Carbonerra, Campo Elias; 19 
km nw Las Cruces (BUENOS AIRES, 
CAMBRIDGE, OTTAWA, DAVIS). 

Discussion.—The most distinctive fea- 
ture of pecki is the broad, apically flattened 
and bilobate male clypeus. The female can 
be distinguished by the nearly flat propleura 
and pygidial plate with a large, single apical 
projection. 


Merithynnus turbulentus (Turner) 
(Figs. 12, 23, 47, 57) 


Scotaena turbulentus Turner 1908:71. Ho- 
lotype male; Venezuela: Caracas (OX- 
FORD). 


Male.—Body length 12-13 mm. Head 
(Figs. 12, 23): Clypeus nearly impunctate, 
conspicuously produced apicomedially, 
with apex sharply bidentate, concave me- 
dially; area between antennal socket and 
eye margin with small punctures 0.5—1.0 
PD apart; frons and vertex with punctures 
0.5 PD apart; hindocellus 4 ocellar diame- 
ters from eye margin; antennal lobe round- 
ed; flagellomeres I-II 3.5 as long as broad; 
flagellomere XI 6.5X as long as_ broad; 
mandible with long acute apical tooth, sub- 
apical notch medially and subbasally an- 


gulate, broadest submedially. Mesosoma: 
Pronotum polished, nearly impunctate; scu- 
tum polished, punctures 1—2 PD apart me- 
dially, 0.5—1.0 PD apart laterally; scutellum 
impunctate medially, laterally and anteri- 
orly punctures 1—2 PD apart; metanotum 
impunctate; propodeum polished, with 
small punctures 3—4 PD apart laterally, 
nearly impunctate medially above petiolar 
socket; mesopleuron polished, punctures 1— 
2 PD apart. Metasoma: Integument dense- 
ly, finely shagreened, punctures shallow, 2— 
4 PD apart; hypopygium apically broadly 
rounded, with dorsal surface irregularly, 
transversely wrinkled. Genital capsule 
(Figs. 47, 57): Paramere slender, elongate 
and tapering apically in lateral view, elon- 
gate and apically spoon-shaped in dorsal 
view; aedeagus without dorsal lobe, less 
than half as long as paramere. Color: black, 
with yellow markings; clypeus yellow ven- 
trally, inner eye margin with yellow stripe 
extending to hindocellus, interrupted me- 
dially; antennal lobe yellow, mandible yel- 
low medially, posterior eye margin with 
yellow stripe along lower three-fourths, at- 
tached medially to transverse yellow stripe 
traversing vertex; pronotum with transverse 
medial yellow stripe, interrupted medially 
and large lateral yellow spot, scutum with 
posteromedial yellow spot and yellow lat- 
eral stripe, scutellum with two large medial 
and lateral yellow spots, metanotum yellow 
medially and laterally, propodeum with 
large comma-shaped yellow sublateral and 
dorsolateral spots; forecoxa anteriorly yel- 
low, mid and hindcoxae each with small 
dorsal and ventral yellow spots; femora 
with yellow ventral stripe, rest of legs 
black; metasomal terga I-IV with small, lat- 
eral yellow spot, sterna H—IV with small 
lateral yellow spots; wing membrane yel- 
low-tinted, veins brown. Vestiture: Silvery. 

Female.—unknown. 

Material examined.—2 6: VENEZUE- 
LA: Caracas (holotype); COLOMBIA: Val- 
le, Peas Blancas, Feb. 1975 (DAVIS, OX- 
FORD). 

Discussion.—Male 


turbulentus most 


VOLUME 107, NUMBER 3 


closely resemble male bogotanus and can 
be distinguished from that species by the 
darkly stained forewing marginal cell, the 
clypeus with the apical projection straight- 
sided (not rounded or convex as in bogo- 
tanus), and the yellow stripe along the inner 
eye margin not reaching beyond the ocelli. 


ACKNOWLEDGMENTS 


This study was made possible by the col- 
lecting efforts of Mike Sharkey and the Al- 
exander von Humboldt Biological Resourc- 
es Research Institute, Bogota, Colombia, 
the loan of type specimens by Frank Koch, 
Lionel Stange and John Huber, and trans- 


595 


portation of one of the types by Lubomir 
Masner. The Colombia project was funded 
by the U.S. National Science Foundation, 
grant No. DEB 9972024. 


LITERATURE CITED 


Kimsey, L. S. 1991. Additional new genera and spe- 
cies of South American Thynnine wasps. Psyche 
98:71—-80. 

Turner, R. E. 1908. Notes on the Thynnidae with re- 
marks on some aberrant genera of Scoliidae. 
Transactions of the Entomological Society of Lon- 
don 56: 63-87. 

. 1910. On the Thynnidae and Scoliidae col- 

lected in Paraguay. Zoologische Jahrbiicher, Ab- 

teilung fiir Systematik 29: 179-227. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 596-603 


A NEW SPECIES OF DOUGLASIIDAE (LEPIDOPTERA) FROM THE 
EASTERN NEARCTIC 


T. L. HARRISON 


Department of Entomology, University of Illinois, 320 Morrill Hall, 505 South Good- 
win Avenue, Urbana, IL 61801, U.S.A. (e-mail: tharriso@uiuc.edu) 


Abstract.—Tinagma gaedikei, n. sp. (Lepidoptera: Douglasiidae), is described from 
adult specimens collected in central Illinois, USA. The new species belongs to a lineage 
that includes three Nearctic species of Tinagma Zeller from California and northern Mex- 
ico, plus several Palearctic species, including Tinagma balteolellum (Fischer von Rosler- 


stamm). 


Key Words: 


Microlepidoptera, biogeography, taxonomy, Tinagma, powelli, californi- 


cum, mexicanum, Boraginaceae, Echium, Hydrophyllaceae, Phacelia 


Douglasiidae is a small (ca. 25 described 
species worldwide) family of microlepidop- 
tera occurring primarily in the Palearctic 
and Nearctic regions, with one representa- 
tive species each in the Neotropical, Ori- 
ental, and Australian regions (Heppner 
1984, 1991; Nielsen 1996; Davis and Rob- 
inson 1999; Powell et al. 1999). Gaedike 
(1974) published a revision of Palearctic 
Douglasiidae, and three additional Palearc- 
tic species have since been described (Gae- 
dike 1987, 1991). Nearctic Douglasidae 
were revised by Gaedike (1990). Biological 
information on the immature stages of 
Douglasiidae is scant; larvae of the few 
species that have been reared are leaf min- 
ers, flower petiole miners, and stem borers, 
utilizing Rosaceae, Lamiaceae, Boragina- 
ceae, and possibly Hydrophyllaceae (Hepp- 
ner 1987, Gaedike 1990, Powell et al. 
999): 

Especially because of the paucity of de- 
finitive knowledge of the immature stages 
of Douglasiidae, placement of the family 
within Lepidoptera is problematic. Heppner 
and Duckworth (1983) and Heppner (1987, 
1991) assigned it to Yponomeutoidea; Kyr- 


ki (1984) to Tineoidea; Heppner (1998) to 
Tineoidea, Series Gracillariiformes; and 
Davis and Robinson (1999) to Gracillarioi- 
dea. A point of particular uncertainty is the 
status of the pupa. Heppner (1987) stated 
that douglasiid pupae “‘are reported to re- 
main in the pupal chamber at adult eclo- 
sion” (which implies yponomeutoid asso- 
ciation), whereas Davis and Robinson 
(1999) said of the pupa, “reportedly with 
abdominal tergal spines (Kyrki 1984) and 
partially extruded from cocoon prior to 
adult eclosion” (which supports placement 
in Tineoidea/Gracillarioidea). Confirmed 
rearings of Douglasiidae, with preserved 
immatures and detailed biological notes, are 
needed to shed light on the taxonomic af- 
finities of this family. 

Gaedike (1990) observed that, on basis 
of genital morphology, known Nearctic 
Douglasiidae (all of which are placed in the 
genus TJinagma Zeller) divide into two 
groups, each of which Gaedike informally 
named on basis of a Palearctic species of 
Tinagma with which each respective Nearc- 
tic group shows affinity. The 7inagma per- 
dicellum Zeller group (three western and 


VOLUME 107, NUMBER 3 


Si) 


Figs. 1-2. Tinagma gaedikei. 1, Adult moth, dorsal aspect. Scale bar = 5.0 mm. 2, Male genitalia, caudal 
aspect, right valve reflected laterally, left valve removed; te, tegumen; ae, aedeagus; an, anellus; ju, juxta; vi, 


vinculum; va, valve. Scale bar = 0.5 mm. 


one eastern species in the Nearctic) has no 
acuminate process at the apex of the male 
valve and no large, serrately-margined 
sclerotized plate associated with the ostium 
bursae of the female. Known larval food 
plants of the 7. perdicellum group in the 
Nearctic include species of Rosaceae 
(Heppner 1987). 

The Tinagma balteolellum (Fischer von 
Roéslerstamm) group (three Nearctic species 
from California and northern Mexico) has 
an acuminate process at the apex of the 
male valve and a large, serrately-margined 
sclerotized plate associated with the ostium 
bursae of the female. Confirmed larval food 


plant associations have not been published 
for the Nearctic species of this group, but 
two of those species have been collected as 
adults on plants of the genera Cryptantha 
Lehmann (Boraginaceae) and/or Phacelia 
Jussieu (Hydrophyllaceae), and Gaedike 
(1990) speculated that these are probably 
the larval food plants of the associated 
moths. The taxonomically-affiliated Pale- 
arctic douglasiid, 7. balteolellum, feeds as 
a larva on viper’s bugloss, Echium vulgare 
Linnaeus (Boraginaceae). 

In late April and early May 2004, I col- 
lected adults of an undescribed species of 
Douglasiidae in east-central Illinois, USA. 


598 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Genital morphology and association of the 
adult with flowers of Phacelia clearly align 
this insect with the 7. balteolellum group, 
known previously in the Nearctic only from 
California and northern Mexico. Because 
this moth represents a substantial change in 
our knowledge of the biogeography of the 
species group to which it belongs, and be- 
cause I wish to inform lepidopterists of the 
presence of this insect (especially in the 
hope that subsequently it will be reared), 
the species is described below. 


Tinagma gaedikei Harrison, new species 
(Figs. 1-12) 


Adult (Fig. 1).—Male and female similar 
in appearance, not showing sexual dichro- 
matism as some other members of the 7. 
balteolellum species group. Mean forewing 
length 3.1 mm (n = 10, range = 2.6—3.3 
mm); mean alar expanse 6.3 mm (n = 10, 
range = 5.0—6.8 mm). 

Head: Smoothly scaled, dorsal surface 
shining dark gray, anterior 30 percent dark 
gray with slightly pale-tipped scales (= 
base of each scale dark gray, apex slightly 
paler gray); antenna uniformly dark gray, 
without pronounced pale annulations; face 
shining whitish gray; labial palpus shining 
whitish gray, second segment smoothly 
scaled, narrowly ringed apically with dark 
gray, and with a minute ventrolateral tuft at 
apex; maxillary palpus rudimentary, not ev- 
ident; haustellum well developed, membra- 
nous, not scaled; compound eye well de- 
veloped; lateral ocellus prominent. 

Thorax: Dorsal surface shining dark 
gray; collar dark gray with slightly pale- 
tipped scales; wing venation typical for 
Douglasiidae, as illustrated by Covell 
(1984); vestiture on basal half of dorsal sur- 
face of forewing dark gray with slightly 
pale-tipped scales, and with strong coppery 
reflections in some angles of illumination; 
at apical edge of this dark-gray area, across 
entire width of wing, scales lack pale tips, 
giving impression of a narrow dark fascia; 
basal dark-gray area delimited apically by 
a narrow, perpendicularly-transverse white 


fascia occupying entire width of wing, wid- 
ening slightly in area posterior to fold; basal 
margin of fascia slightly sinuate, apical 
margin nearly straight; apical half of wing 
beyond white fascia dark gray with promi- 
nently pale-tipped scales (= base of each 
scale dark gray, apex much paler gray), giv- 
ing the area a uniformly speckled appear- 
ance, without metallic reflection; a very 
small patch of white scales (absent in some 
specimens) may be present in this area mid- 
way between anterior and posterior margins 
at about 0.75 length of wing; fringe dark 
brownish gray with dark-tipped scales 
forming two distinct lines; ventral surface 
of wing dark gray, with pattern of dorsal 
surface faintly discernible; hindwing and its 
fringe uniformly dark gray on dorsal and 
ventral surfaces; ventral surface of thorax 
shining whitish gray; on all legs, coxa, tro- 
chanter, and femur shining whitish gray, 
tibia and basal tarsomere dark gray with 
slightly pale-tipped scales, all other tarso- 
meres shining whitish gray ringed with dark 
gray at apices; meso- and metatibiae each 
with a small dorsoapical tuft projecting 
over base of tarsus; metatibia with a dorsal 
fringe of long, whitish hairlike scales along 
entire length of segment. 

Abdomen: Dorsal surface shining dark 
gray; ventral surface shining whitish gray. 

Male genitalia and associated structures 
(Figs. 2—7): Structures are interpreted 
based on characterizations given by Klots 
(1956). Valvae symmetrical; valve (Figs. 2— 
3) broadly subtriangular; apex of valve with 
an acuminate articulated process, 3.3X as 
long as its width at base, lying folded over 
medial surface of valve and projecting an- 
terad to 0.8 length of valve; ventral mar- 
gin of valve nearly straight, appearing as a 
uniformly narrow fold bearing long hairlike 
setae; apical 65 percent of dorsal margin of 
valve nearly straight, lying at 60° angle to 
ventral margin, and lined with a row of ap- 
proximately 20 short thickened setae plus a 
few long flattened setae, the thickened setae 
increasing gradually in length from basal to 
apical ends of row; basal 35 percent of dor- 


VOLUME 107, NUMBER 3 599 


Figs. 3-7. Tinagma gaedikei, male genitalia and associated structures. 3, Left valve, medial aspect. 4, Teg- 
umen, ventral aspect, flattened. 5, Vinculum, ventral aspect, flattened. 6, Aedeagus, anellus and juxta, left lateral 
aspect. 7, Seventh and 8" abdominal segments, cut longitudinally and spread flat to show external surfaces of 
terga, pleura, and sterna. Scale bar = 0.25 mm for Figs. 3-5, 0.5 mm for Figs. 6—7. 


600 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


sal margin of valve devoid of setae, shal- 
lowly concave in an even curve, attenuated 
into a prominent anterodorsal process; teg- 
umen (Figs. 2, 4) a dorsal hoodlike struc- 
ture, narrowing laterally as it curves ven- 
trad, its apices extending into close associ- 
ation with those of vinculum, the two struc- 
tures together forming a sclerotized “ring” 
with which the valvae articulate; tegumen 
lightly sclerotized, composed of a large, an- 
terior uniformly-punctate element and a 
small, medial rounded lobe, not punctate, 
projecting posterad from posterior margin 
of large element; vinculum (Figs. 2, 5) a 
ventral sclerotized band, curving dorsad, its 
apices coming into close association with 
those of tegumen; with anterior margin pro- 
duced into a short saccus (Fig. 5); each lat- 
eral arm narrowest immediately laterad of 
saccus, then gradually broadening laterally, 
culminating in a trifid lateral margin, with 
anterior lateral process more pronounced 
and heavily sclerotized than posterior two 
processes (Fig. 5); juxta (Figs. 2, 6) a me- 
dial, transverse rectangular plate lying in 
close association with posterior margin of 
vinculum; anellus (Figs. 2, 6) two flattened 
plates lying in close contact with aedeagus 
on either side; juxta and anellus connected 
by two narrow, curved processes (appar- 
ently extensions of the anellus); aedeagus 
(Fig. 6) without cornuti, approximately 10 
as long as width at base, narrowing gradu- 
ally to apex, width at apex approximately 
0.25 width at base; abdominal segment 8 
(Fig. 7) showing two male-specific modifi- 
cations: first, each pleural area with a small 
domelike lobe bearing 15—20 narrowly- 
clavate processes; and second, tergum with 
two lateral, triangular sclerotized patches, 
the anterior point of each triangle produced 
into a narrow tapering rod extending anter- 
ad to posterior margin of tergum 7; sternum 
of segment 8 uniformly sclerotized, unmod- 
ified. 

Female genitalia (Figs. 8—12): Oviposi- 
tor membranous; posterior apophyses fila- 
mentous, each slightly thickened near pos- 
terior end; anterior apophyses slender, each 


with a flat triangular enlargement at poste- 
rior end; a flat, broadly-fusiform sclerotized 
plate (Fig. 9) associated with ostium bursae; 
lateral margins of plate entire, posterior 
margin (Fig. 10) truncate and very finely 
serrate; anterior 45 percent of plate envel- 
oped in a membranous, longitudinally- 
striate invagination, widest at its posterior 
margin (which is narrowly sclerotized ven- 
trally), narrowing anteriorly at 65° angle 
and ultimately converging on a narrow, dif- 
ferentiated rim at anterior margin of scler- 
otized plate representing point of emer- 
gence of ductus bursae; ductus bursae light- 
ly sclerotized in posterior half of length, 
membranous in anterior half, anterior limit 
of sclerotized region containing five or six 
small sclerotized spinules (Fig. 11); corpus 
bursae membranous, elongate-ovoid; sig- 
num (Fig. 12) a stellate patch of approxi- 
mately 30 aciculate sclerotized rods, some 
of which may be sloughed off into interior 
of corpus bursae in mated specimens; ab- 
dominal segment 8 without modification. 

Type material.—Holotype male: Collect- 
ed as diurnal adult on flower of Phacelia 
purshii, USA: Illinois, Coles County, 
TI1IN, R9E, Sec. 11, 29-IV-2004, T. Har- 
rison (USNM, = National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, DC, USA). Allotype female: 
Same data as for holotype, except collected 
30-IV-2004 (USNM). Paratypes: 4 6,4 9; 
same data as for allotype except | d and 1 
2 collected 6-V-2004, 1 ¢ collected 8-V- 
2004, and | 6 collected 11-V-2004 
(Deutsches Entomologisches Institut, Miin- 
cheberg, Germany; Illinois Natural History 
Survey, Champaign, Illinois, USA; and pri- 
vate collection of James R. Wiker, Green- 
view, Illinois, USA). 

Etymology.—The species is named for 
Dr. Reinhard Gaedike, in recognition of his 
many years of dedicated study of the global 
fauna of Douglasiidae. 

Biology.—Adults of 7. gaedikei were 
collected in late April and early May in me- 
sic deciduous forest on flowers of Phacelia 
purshii Buckley (Hydrophyllaceae). Moths 


VOLUME 107, NUMBER 3 601 


Figs. 8-12. Tinagma gaedikei, female genitalia. 8, Genitalia, ventral aspect; pl, papillae anales; pa, posterior 
apophysis; aa, anterior apophysis; si, signum; sp, sclerotized plate associated with ostium bursae; mi, membra- 
nous invagination enveloping anterior region of sclerotized plate; ar, anterior rim of sclerotized plate; db, ductus 
bursae; cb, corpus bursae. 9, Sclerotized plate associated with ostium bursae. 10, Detail of posterior margin of 
sclerotized plate associated with ostium bursae. 11, Posterior region of ductus bursae. 12, Signum. Scale bar = 
1.0 mm for Fig. 8, 0.5 mm for Fig. 9, 0.125 mm for Fig. 10, and 0.25 mm for Figs. 1 1—12. 


602 


were collected between 1000 and 1200 
Central Standard Time (observations were 
not made at other times of day), in condi- 
tions ranging from bright sunlight to heavy 
overcast. The pristine condition of collected 
adults and the presence of a spermatophore 
in one dissected female indicate that adults 
emerge and mate in late April and early 
May. Nothing else is known of the life cy- 
cle of this insect. 

Geographic range.—Tinagma gaedikei 
has been recorded only from the type lo- 
cality, Coles County, Illinois, USA. 

Diagnosis.—The only known eastern Ne- 
arctic species of Douglasiidae other than T. 
gaedikei is Tinagma obscurofasciella 
(Chambers), which is placed in the 7. per- 
dicellum group (Gaedike 1990). Tinagma 
gaedikei differs from T. obscurofasciella in 
size, color pattern of the forewing, and gen- 
ital morphology of both genders; the latter 
two characters in T. obscurofasciella were 
illustrated by Covell (1984) and Gaedike 
(1990), respectively. 

The TJ. balteolellum group contains, in 
addition to 7. gaedikei, three Nearctic spe- 
cies, Tinagma powelli, Tinagma californi- 
cum, and Tinagma mexicanum, all of which 
were described by Gaedike (1990). Fore- 
wing coloration of 7. gaedikei differs from 
that of the three southwestern Nearctic spe- 
cies, as described by Gaedike (1990). In 
genital morphology, 7. gaedikei lacks the 
setose globular development seen at the 
apex of the valve of 7. mexicanum (the fe- 
male of which is unknown). 7inagma gae- 
dikei differs markedly from 7. powelli and 
T. californicum in female genital morphol- 
ogy. The lateral margins of the ostial plate 
in T. gaedikei are entire (or very finely and 
sparsely serrate near posterior margin of 
plate), the posterior margin of the plate is 
uninterruptedly transverse, and the plate is 
widest at midlength. In 7. powelli, the lat- 
eral margins of the posterior half of the 
plate are coarsely and densely serrate, and 
the posterior margin of the plate is divided 
by a deep medial cleft. In 7. californicum, 
the plate is widest at its posterior margin, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


which is jaggedly quadrifid. Also, the geo- 
graphic range of T. gaedikei probably does 
not overlap those of the other three Nearctic 
species of this complex. 


ACKNOWLEDGMENTS 


I thank two reviewers for helpful com- 
ments on the manuscript. 


LITERATURE CITED 


Covell, Jr, C. V. 1984. A Field Guide to the Moths of 
Eastern North America. Houghton Mifflin, Bos- 
ton, 496 pp. 

Davis, D. R., and G. S. Robinson. 1999. The Tineoidea 
and Gracillarioidea, pp. 91-118. In Kristensen, N. 
P., ed. Lepidoptera, butterflies and moths, Vol. 1: 
Evolution, systematics, and biogeography. Hand- 
book of Zoology 4(35): 1-491. 

Gaedike, R. 1974. Revision der palaéarktischen Doug- 
lasiidae (Lepidoptera). Acta Faunistica Entomo- 
logica Musei Nationalis Pragae 15: 79-102. 

. 1987. Beitrag zur Kenntnis der palaarktischen 

Douglasiidae (Lepidoptera): Tinagma klimeschi 

sp. n., aus Rhodos. Nota Lepidopterologica 10: 

158-162. 

. 1990. Revision der nearktischen Douglasiidae 

(Lepidoptera). Beitrage zur Entomologie 40: 287— 

300. 

. 1991. Neue und seltene Douglasiidae. Deut- 
sche Entomologische Zeitschrift (Neue Folge) 38: 
19-25. 

Heppner, J. B. 1984. Douglasiidae, p. 57. Jn Heppner, 
J. B., ed. Atlas of Neotropical Lepidoptera, 
Checklist: Part I, Micropterigoidea-Immoidea. Dr 
W. Junk Publishers, The Hague, 112 pp. 

1987. Douglasiidae (Yponomeutoidea), pp. 

408—409. In Stehr, E W., ed. Immature Insects. 

Kendall/Hunt, Dubuque, 754 pp. 

. 1991. Faunal regions and the diversity of Lep- 

idoptera. Tropical Lepidoptera 2(Supplement 1): 

1-85. 

. 1998. Classification of Lepidoptera. Part I, In- 
troduction. Holarctic Lepidoptera 5(Supplement 
1): 1-148 + index 1-6. 

Heppner, J. B., and W. D. Duckworth. 1983. Dougla- 
siidae, p. 27. In Hodges, R. W., T. Dominick, D. 
R. Davis, D. C. Ferguson, J. G. Franclemont, E. 
G. Munroe, and J. A. Powell, eds. 1983. Check 
List of the Lepidoptera of America North of Mex- 
ico. London: E. W. Classey and Wedge Entomo- 
logical Research Foundation, 284 pp. 

Klots, A. B. 1956. Lepidoptera, pp. 115—130. In Tux- 
en, S. L., ed. Taxonomist’s Glossary of Genitalia 
in Insects. Ejnar Munksgaard, Copenhagen, 284 
Pp- 

Kyrki, J. 1984. The Yponomeutoidea: A reassessment 


VOLUME 107, NUMBER 3 603 


of the superfamily and its suprageneric groups. Powell, J. A., C. Mitter, and B. Farrell. 1999. Evolu- 


Entomologica Scandinavica 15: 71—84. tion of larval food preferences in Lepidoptera, pp. 


Nielsen, E. S. 1996. Douglasiidae, p. 46. Jn Nielsen, 403-422. In Kristensen, N. P., ed. Lepidoptera, 
E. S., E. D. Edwards, and T. V. Rangsi, eds. butterflies and moths, Vol. 1: Evolution, system- 
Checklist of the Lepidoptera of Australia. CSIRO atics, and biogeography. Handbook of Zoology 


Division of Entomology, Canberra, 426 pp. 4(35): 1-491. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 604-622 


ANOPHELES (ANOPHELES) LESTERI BAISAS AND HU 
(DIPTERA: CULICIDAE): NEOTYPE DESIGNATION AND DESCRIPTION 


LEOPOLDO M. RUEDA, RICHARD C. WILKERSON, AND CONG LI 


Department of Entomology, Walter Reed Army Institute of Research, 503 Robert Grant 
Avenue, Silver Spring, MD 20910-7500, U.S.A. (LMR email: ruedapol @si.edu; RCW: 
wilkersonr@si.edu; CL: licong @si.edu) 


Abstract.—The Asian malaria vector, Anopheles (Anopheles) lesteri Baisas and Hu, 
1936, is described with illustrations of the larval and pupal stages, adult female, and the 
male genitalia. Taxonomic and related literature records, diagnostic features, distribution, 
rDNA ITS2 sequence, bionomics, and medical importance of An. lesteri are included. A 
neotype female for the species from near the original type locality in Luzon, Philippines, 


is designated. 


Key Words: 
malaria 


Because of the recent increase of human 
malaria cases in South Korea (Feighner et 
al. 1998, Lee et al. 1998), there is a serious 
need to clarify the identity of the possible 
mosquito vectors. Misidentifications of vec- 
tor species often lead to inadequate under- 
standing of the epidemiology of disease 
transmission and inadvertently affect con- 
trol measures. Anopheles (Anopheles) les- 
teri Baisas and Hu, 1936, may be the most 
significant vector of malarial parasites in 
Korea, Taiwan, Japan (particularly Okina- 
wa), and central and northern China (Har- 
rison 1973). It is one of the 27 species of 
the Hyrcanus Group of Anopheles (Anoph- 
eles) having an Oriental or eastern Palearc- 
tic distribution (Ramsdale 2001, http:// 
Www.mosquitocatalog.org). It may have a 
potential role in malarial and filarial para- 
site transmission and disease outbreaks in 
countries where it occurs. Recently, Wilk- 
erson et al. (2003) demonstrated that An. 
anthropophagus Xu and Feng, the most 1m- 
portant vector malaria vector in eastern 
China, is actually a junior synonym of An. 
lesteri. 


Anopheles lesteri, Culicidae, taxonomy, description, neotype, mosquitoes, 


Anopheles lesteri was described by Bais- 
as and Hu (1936: 214) as An. hyrcanus vat. 
lesteri from 1 male ot M1—8) and 1 female 
(lot M1—12), with corresponding larval and 
pupal exuviae, from Santa Mesa, Manila, 
Luzon, Philippines (Jose P. Ingal, coll. 2 
March 1936). Many specimens (or “‘coty- 
pes’) were collected from Calauan, Laguna 
in 1935, but type specimens were selected 
from the Santa Mesa specimens collected in 
1936. The syntypes or type specimens 
(“‘types” and “‘cotypes’”’, collected from 
Santa Mesa and Calauan, Laguna, Luzon), 
were supposed to be deposited in the Phil- 
ippine National Museum, Manila. Other 
specimens from the same batch were to be 
deposited in the National Museum of Nat- 
ural History, Washington, DC, and Henry 
Lester Institute of Medical Research, 
Shanghai, China (Baisas and Hu 1936). 
Knight and Stone (1977: 22) noted that the 
type specimens could not be found. Our in- 
quiries were also unsuccessful in search of 
these specimens in possible depositories in 
Manila and Shanghai. Specimens of An. 


VOLUME 107, NUMBER 3 


lesteri found in the National Museum of 
Natural History include 2 slides with the 
following labels: slide # 1- right label light 
yellow, with reddish border: ““M 1-13 An. 
hyr. var. lesteri Sta. Mesa, Manila Mar. 5, 
1936”, left label: ““Anopheles (An.) lesteri 
lesteri Baisas & Hu det. B.A. Harrison’’; 
slide # 2—right label light yellow, with red- 
dish border: *““F171—50 An. lesteri cotype- 
male Calauan, Laguna July 15, 1935.” 
However, larval and pupal exuviae, partic- 
ularly those mounted on slide # 2, are in 
bad conditions (i.e., dark unknown mount- 
ing media, cracked specimens, etc.), and 
most morphological characters are difficult 
to see under the compound microscope. 

Also, the original adult description (Bais- 
as and Hu 1936) is not sufficiently complete 
for accurate identification of the species, 
and no detailed descriptions of the larval 
and pupal stages or of the male terminalia 
of specimens from the type locality (Luzon) 
have been made. In view of this, it becomes 
imperative to provide detailed descriptions 
of various life stages and to designate a 
neotype for this important species. 

In the present paper, a neotype female 
from near the original type locality is des- 
ignated for An. lesteri and associated ribo- 
somal DNA internal transcribed spacer 2 
(rDNA ITS2) sequence provided. Descrip- 
tions and illustrations are provided for the 
adult female and male, pupa and larva of 
this species from the same type locality. 


MATERIALS AND METHODS 


Except for wing spot nomenclature, 
which is taken from Wilkerson and Peyton 
(1990), for wing venation terms from Belk- 
in (1962), and for pupal abdominal dark 
marks, the terminology and abbreviations 
of Harbach and Knight (1980, 1982) are 
used for the morphological characters and 
illustrations. Abbreviations used are as fol- 
low: E, egg; G, genitalia; L, larva; Le, lar- 
val exuviae; NE, non-existent; P, pupa; Pe, 
pupal exuviae; var., variety. An asterisk fol- 
lowing the abbreviation of a given life stag 


= 
indicates that at least part of the life stage 


605 


was illustrated in the publication cited. Col- 
lection codes of the most recent collections 
consist of a country code in capital letters 
followed by a collection number (e.g., PH 
9—1 is an individual from collection 9 from 
the Philippines; a specimen number lower 
than 100 is used if there are associated lar- 
val and pupal exuviae, and equal or greater 
than 100 if there are no associated larval 
exuviae). 

DNA isolation and sequencing. DNA 
was isolated by phenol-chloroform extrac- 
tion, as described by Wilkerson et al. 
(1993), from a leg of the adult neotype fe- 
male, a leg of a second female, and 2 entire 
males, minus genitalia, from new type lo- 
cality in Calauan, Laguna. Direct sequenc- 
ing was carried out as described in Wilk- 
erson et al. (2004) using their primers. The 
beginning and end of the rDNA ITS2 was 
estimated as in Cornel et al. (1996). 


TAXONOMIC TREATMENT 


Anopheles (Anopheles) lesteri 
Baisas and Hu 


(Figs. 1—4) 


Anopheles yesoensis Tsuzuki 1901: 717 
(nomen dubium). 

Anopheles jesoensis Tsuzuki 1902: 286 (no- 
men dubium). 

Anopheles hyrcanus var. lesteri Baisas and 
Fin el9 3.62022 98( Gee Aa ee ES) aailiypes 
Santa Mesa, Manila (Luzon), Philippines 
(NE), other specimens/*‘cotypes”’: Ca- 
lauan, Laguna (Luzon); Ohmori 1957: 
209 (6*, E*); Ohmori 1959: 222 (P*). 

Anopheles (Anopheles) lesteri: Otsuru and 
@Ohmoriel960:<475(6 ASP ise 
taxonomy; bionomics; distribution, Ja- 
pan: Honshu [Hyogo, Mie, Niigata, Ya- 
maguchi], Kyushu [Kumamoto, Kagosi- 
ma, Oita, Fukuoka, Saga, Nagasaki]); 
Whang 1962: 39 (distribution, Korea: 
Tansan, Wondang, Guidandong, Yong- 
jueup); Reid 1968: 105 (type form); Ca- 
gampang-Ramos and Darsie 1970: 14 
(identification key); Basio 1971: 36 (dis- 
tribution, Philippines: widely found in 


606 


Luzon including Manila, Pampanga); 
Basio and Reisen 1971:60 (L, distribu- 
tion, Guam); Tanaka 1971: 4 (distribu- 
tion, Japan: Ryukyu Islands); Harrison 
1973: 4 (taxonomy); Baisas 1974: 50 
(?*, PR L*, E; taxonomy); Tanaka et al. 
1979: 83 (6* 9*, P*, L*, E; taxonomy, 
bionomics, distribution, Japan: Hokkai- 
do, Honshu, Kyushu, Ryukyu Archipel- 
ago [Amami, Okinawa Gunto, Yaeyama 
Gunto]); Rueda et al. 2004 (distribution, 
China: Hong Kong). 

Anopheles (Anopheles) lesteri lesteri: Chau 
1982 (distribution, China: Hong Kong). 
Anopheles (Anopheles) lesteri anthropoph- 
agus Xu and Feng 1975: 81, 97 (¢*, 3*, 

pP*, L*, E*; taxonomy). 

Anopheles (Anopheles) anthropophagus: 
Ma 1981: 11 (key; distribution, China: 
Fukien, Kiangsi, Kiangsu, Kwangsi, 
Kweichow, Shanghai, provinces south of 
Yantze River); Wilkerson et al. 2003: 1 
(new synonym of /esteri). 


Other literature records.—Otsuru 1949: 
139 (as possible malaria vector, Japan); Ot- 
suru and Ohmori 1960: 33 (bionomics, Ja- 
pan); Kamimura 1968: 15 (as possible ma- 
laria vector, Japan); Reisen et al. 1972: 319 
(distribution, Guam); Zhang et al. 1980: 
140 (as experimental vector of Vietnam 
monkey malarial parasite, near Plasmodium 
cynomolgi); Xu et al. 1981: 265 (scanning 
electron micrographs [SEMs] of adults, pu- 
pae, eggs, China); Takai et al. 1984: 251 
(hybridization, Japan); Zhang et al. 1987: 
191 (vectorial capacity for malayan filaria- 
sis, Sichuan, China); Xu et al. 1988: 247 
(control using insecticide-treated bed net, 
Guangxi, China); Ma and Wang 1988: 65 
(salivary gland chromosome, China); Wang 
and Zheng 1989: 175 (blood meals, Guizh- 
ou, China); Ye and Zhu 1989: 256 (enzyme 
electrophoresis, China); Dapeng et al. 1996: 
100 (as vector of P. falciparum, and chem- 
ical vector control, Xinyang, China); Li et 
al. 1991: 8 (DNA-restriction fragment 
length differences, China); Liu et al. 1991: 
147 (as vector of malayan filariasis, Fujian, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


China); Niu et al. 1992: 267 (DNA probe); 
Chen et al. 1994: 27—30 (trace and macro 
elements in hemolymph); Cheng et al. 
1995: 321, (control using insecticide-treated 
bed net, Henan, China); Gu et al. 1966: 34 
(distribution, China); Shahjehan 1996: 205 
(DNA probes, China); Song and Peng 
1996: 161 (control using mermithid nema- 
todes, Sichuan, China); Xu et al. 1997: 807 
(as vector of filariasis, Henan, China); 
Sleigh et al. 1998: 265 (as P. vivax vector, 
Henan, China); Xu et al. 1998: 135—136 (as 
vector of P. vivax; control using insecticide- 
treated bed net and residual spraying, Hu- 
bei, China); Kim et al. 1999: 181 (seasonal 
prevalence, South Korea); Zizhao et al. 
1999: 240-242 (as vector of P. falciparum 
malaria, China); Lee et al. 2000: 77 (PCR, 
presence of P. vivax circumsporozoite pro- 
tein, South Korea); Ma et al. 2000a: 325 
(PCR assay and rDNA-ITS2 sequencing, 
China); Ma et al. 2000b: 36 (rDNA-ITS2, 
Korea); Burkett et al. 2001: 196, 2002: 45 
(trap attractants, South Korea); Huang et al. 
2001: 340 (habitat and distribution, Hubei, 
China); Coleman et al. 2002: 244 (presence 
of P. vivax circumsporozoite protein, South 
Korea); Min et al. 2002: 198; Shin et al. 
2002: 41 (vector competence to P. vivax, 
Korea); Toma 2002: 7 (distribution review, 
Ryukyu Archipelago, Japan); Wilkerson et 
al. 2003: 1 (species molecular confirmation, 
tDNA-ITS2, China, Philippines, South Ko- 
rea; note on geographical range, China). 


ORIGINAL DESCRIPTION 


In support of previous and present inter- 
pretations of the name /esteri, the original 
description given by Baisas and Hu (1936) 
is as follows. “‘Dark and pale scales of 
wings well contrasted. Costa—Jet black ex- 
cepting for the preapical and subcostal pale 
spots. The subcostal spot is composed of 
from 10 to 22 pale scales. No scattered pale 
scales elsewhere on the dark portions of the 
costa. Subcosta—Invariably dark excepting 
for 1 to 4 pale scales at the apex, which 
form a part of the subcostal spot. Vein 1— 
Preapical pale spot distinct and complete. 


VOLUME 107, NUMBER 3 


Subcostal spot usually incomplete, seldom 
complete, and more rarely absent. Mid pale 
spot usually small and incomplete, some- 
times absent. Sector pale spot usually small 
and incomplete. A few scattered pale scales 
are sometimes present on the dark area be- 
tween the preapical and subcostal spots but 
these are not as many as those found in ni- 
gerrimus. A few pale scales towards the 
base below the presector dark spot, which 
do not, however, form definite spots. Vein 
2—Stem mainly pale with some greyish or 
dark scales on lateral borders. Anterior fork 
dark with complete preapical pale spot. 
Posterior fork dark with a pale spot at 
about, or a little below the middle. Vein 3— 
A definite dark area, at base, and another at 
apex; apical half of intervening area with 
median squames mainly pale; lateral 
squames mainly dark; basal half mainly 
pale with a few scattered dark scales. Vein 
4—Stem dark or mainly dark towards base 
below cross viens, mainly pale towards 
apex. Forks dark at bases and apices, the 
intervening portion mainly pale but lateral 
squames with fair distribution of dark 
scales. Vein 5—Apical half or more of stem 
pale, sometimes with a few scattered dark 
scales. A definite dark area a little below 
middle, followed by a mixture of dark and 
pale scales, the pale ones sometimes pre- 
dominating or occupying the whole area ex- 
cepting the extreme base where a few dark 
scales are located. Anterior fork mainly 
dark with the usual dark spots ill-demar- 
cated. Sometimes the basal and sub-basal 
dark areas are formed on the pale portion 
towards the apex. Posterior fork pale ex- 
cepting for the apical dark spot. Vein 6— 
Pale with a dark area at middle, and another 
one at the apex. In some specimens, a few 
dark scales are sometimes scattered on the 
pale portion towards the base. Fringe—Pale 
spot at apex of wing involves variable por- 
tion of area opposite vein | and vein 4.1. 
Pale spot opposite vein 5.2 absent in all 
males and females examined. Humeral 
vein—Usually with 2 or 3 dark scales. Re- 


607 


migium—Mainly dark-scaled with a few 
pale scales on the anterior border.” 


SUPPLEMENTAL DESCRIPTION FROM 
THIS STUDY 


Female (Fig. 1).—Integument dark 
brown with silvery or grayish pollinosity. 
The following measurements and counts, 7 
= 6, except when indicated. Head: Inter- 
ocular space with 10—11 (nm = 3) long, pale 
setae intermixed with long and small, nar- 
row, appressed white scales; vertex, occiput 
and upper portion of postgena with numer- 
ous erect, truncate scales; patch of white 
scales on the middorsal portion of vertex; 
patch of dark brown to black scales on lat- 
eral portion of vertex, occiput and upper 
portion of postgena; long dark brown to 
black setae on ventral portion of postgena. 
Clypeus bare dorsally, with dark scales lat- 
erally. Pedicel of antenna with 6—11 (n = 
4) small, dorsolateral, narrow to broad, 
grayish white spatulate scales, and 3 short, 
dark ventrolateral setae; flagellomere | with 
numerous narrow to broad white scales; re- 
maining flagellomeres with a few scattered 
narrow to broad white scales. Scales of 
maxillary palpus slender, spatulate, mostly 
dark brown to black with intermixed dark 
brown setae; narrow band of white scales 
at base of palpomeres 3 and 4, and at base 
and apex of palpomere 5; apical white band 
of palpomere 5 slightly shorter than other 
basal palpomere white bands; base of max- 
illary palpus dorsally with single long, erect 
dark seta; length of maxillary palpus 1.75— 
1.98 mm (mean = 1.86 mm); ratio of length 
of each of palpomeres 2—5 to total length 
of palpus, 2 = 0.33—0.34 (mean = 0.34), 3 
= 0.32—0.37 (mean = 0.35), 4 = 0.17—0.23 
(mean = 0.20), 5 = 0.1-0.12 (mean = 
0.11); ratio of combined palpomeres 2—5 to 
total length of palpus, 0.99—1.0 (mean = 
0.99); ratio of combined palpomeres 4, 5 to 
total length of palpus, 0.27—0.35 (mean = 
0.31); palpus 1.15—1.26 (mean = 1.22) for- 
efemur length. Proboscis dark-scaled, ex- 
cept labellum light brown; base of probos- 
cis with long, erect dark setae and scales; 


608 


ie 


we 


Fig. 1. Anopheles lesteri, adult female, habitus. 


proboscis length 1.75—1.95 mm (mean = 
1.86 mm, n = 3); proboscis 0.99—1.01 
(mean = 1.0, n = 3) palpus length. Thorax: 
Scutum dark brown, with gray pollinosity; 
2 submedian longitudinal lines on anterior 
area; a pair of indistinct black spots near 
antealar area; median anterior promontory 
with patch of intermixed narrow, short and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


= 


(GUANA) Ws 


long pale yellow scales. Darker lateral areas 
of scutum with longer dark setae. Scutal 
fossa, antealar area and supraalar area 
slightly pale pollinose. Scutellum dark, 
slightly pale dusted, with 16—22 shorter and 
14-19 long setae, short setae intermixed 
pale yellow and dark brown, long setae 
dark brown. Antepronotum with 12—14 


VOLUME 107, NUMBER 3 


609 


Table 1. Anopheles lesteri: descriptive statistics for ratios of veins C and R-R, wing spot lengths/length of 


wing measured from humeral crossvein*. 


——————— a 


Wing Spot 


Range Mean + SD 


SS rss 


Costa (C) 


Basal dark to sector dark (BD+PHD+HD+PD+SD) 
Subcostal pale (SCP) 
Preapical dark (PD) 
Preapical pale (PP) 
Apical dark (AD) 
Vein R-R, 
Basal dark to presector dark (BD+PHD+HD+PSD) 
Sector pale (SP) 
Sector dark (SD) 
Subcostal pale (SCP) 
Preapical dark (PD) 
Preapical pale (PP) 
Apical dark (AD) 


0.69--0.74 0.72 + 0.02 [0.73] 
0.03—0.07 0.05 + 0.01 [0.05] 
0.27-0.31 0.29 + 0.01 [0.29] 
0.03—0.05 0.04 + 0.01 [0.03] 
0.04—0.06 0.05 + 0.01 [0.04] 
0.38-0.41 0.40 + 0.01 [0.38] 
0.04—0.13 0.08 + 0.04 [0.12] 
0.21-0.26 0.24 + 0.02 [0.21] 
0.05—0.06 0.05 + 0.00 [0.06] 
0.24-0.29 0.26 + 0.02 [0.25] 
0.04—0.06 0.05 + 0.01 [0.06] 
0.03—0.05 0.04 + 0.01 [0.03] 


* Six wings, from the neotype and alloneotype, and 3 individuals; [ 


dark brown setae. Pleuron brown to dark 
brown; upper proepisternum with 3 or 4 se- 
tae, without scales; prespiracular area with 
2 or 3 setae, without scales; prealar area 
with 4 or 5 setae, without scales; upper me- 
sokatepisternum with 3 or 4 setae, without 
scales; lower mesokatepisternum with 4 or 
5 setae, without scales; upper mesepimeron 
with 4 or 5 setae, without scales. Legs: 
Fore- and midlegs dark-scaled except white 
scales dorsally on apex of tibia; complete 
narrow apical pale bands on tarsomeres |— 
3, and very narrow apical dorsal pale patch 
on tarsomere 4; apical bands on tarsomeres 
2 and 3 longest, complete ventrally, about 
0.1 length of tarsomere; pale scales on tar- 
somere 4 dorsally and laterally only, not 
connected ventrally; basal pale band on tar- 
somere 5 absent. Hindlegs dark-scaled as 
fore- and midlegs, except white scales on 
tarsomeres 1—3 dorsally and laterally only, 
not ventrally. Forefemur length 1.40—1.72 
mm (mean = 1.53 mm, n = 6), ratio of 
forefemur length to proboscis length 0.79— 
0.88 (mean = 0.82). Midfemur length 
1.74—2.12 mm (mean = 1.94 mm), ratio of 
midfemur length to proboscis length 0.93— 
1.12 (mean = 1.04). Hindfemur length 
2.01—2.2 mm (mean = 2.11 mm), ratio of 


], neotype female. 


hindfemur length to proboscis length 1.07— 
1.21 (mean = 1.14). 

Wing (Table 1): Length (measured from 
humeral cross vein to wing tip, excluding 
fringe) 2.8—3.18 mm (mean = 3.05 mm). 
Dark scales brown to black, pale wing 
scales white and pale yellow. Costa (C) 
dark-scaled with small subcostal pale spot 
(SCP, mean = 0.15) and preapical pale spot 
(PP, mean = 0.12); remigium dark scaled; 
humeral crossvein and arculus without scale 
patch. Subcosta (Sc) dark-scaled with few 
scattered spatulate white scales from base 
to sector dark (SD). Radius (R) to R, dark- 
scaled except 3 pale spots (SP, SCP and 
PP), scattered white spatulate scales from 
base to presector dark (PSD), and a stripe 
of white scales on SD before SCP; length 
of white stripe 0.3-0.38 mm (mean = 
0.33); base of R, dark-scaled; bifurcation of 
R, and R, dark-scaled; tips of R,, Rs, R; 
and R,,; with pale fringe. Media (M) dark- 
scaled with pale area before fork; bifurca- 
tion of M ,,, and M,,, dark scaled; tips of 
M,,, and M,,, with dark fringe. Cubitus 
(Cu) with basal dark spot, length 0.18—0.28 
(mean = 0.24); Cu, with 4 dark spots, 
length of first basal spot 0.1—0.18 mm 
(mean = 0.14), second spot 0.23—0.25 mm 


610 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(mean = 0.24), third spot 0.25—0.45 mm 
(mean = 0.33), distal fourth spot 0.13—0.3 
mm (mean = 0.21); Cu, with distal dark 
spot only, length 0.18—0.25 mm (mean = 
0.21); tips of Cu, and Cu, with dark fringe. 
Anal vein (1A) with 2 dark spots, basal spot 
length 0.23—0.25 mm (mean = 0.24), distal 
spot length 0.25—0.35 mm (mean = 0.31), 
tip of 1A with dark fringe. Halter: Scabel- 
lum, pedicel and capitellum dark brown 
with grayish pollinosity. Abdomen: Terga 
and sterna dark brown to black with grayish 
pollinosity, covered with pale brown to 
golden brown setae. For neotype female, 
descriptive statistics for ratios of costal and 
R-R, wing spot lengths/length of wing mea- 
sured from humeral crossvein are shown in 
brackets in Table 1. 

Male (Figs. 2C, D).—As in female ex- 
cept for following sexual differences. Max- 
illary palpus 0.94—0.98 length of proboscis 
(mean = 0.97; n = 4 for this and following 
measurements except where indicated), 
apex of palpomere 3 and all of palpomeres 
4 and 5 enlarged. Maxillary palpus with 
dark brown and white scales, dorsal surface 
of all segments with white scales; palpom- 
ere 2 with slightly erect dark brown scales 
at base and light gray scales from middle 
to apex; palpomere 3 dark-scaled with long 
yellowish to light brown setae at apex; pal- 
pomere 4 pale yellow to dark brown-scaled 
with narrow basal white band, inner surface 
with long yellowish-brown to light brown 
setae; palpomere 5 pale brown-scaled with 
narrow basal white band, lateral surface 
with white scales and numerous dark brown 
short setae. Proboscis length 2.65—2.9 mm 
(mean = 2.69 mm), dark brown-scaled. 
Anal vein with single dark spot. Tergum IX 
(width, 2.92 mm) bare, with pair of elon- 
gate caudally directed capitate lobes; length 
of lobe from base to tip 0.82 distance be- 
tween 2 lobes; middle of lobe narrower, 
0.45 width of lobe tip and 0.42 width of 
lobe base. Gonocoxite 1.91—2.13X as long 
as wide at widest point, widest about 0.08 
from base; dorsal (postrotational) surface 
with many long setae distally, slender fu- 


siform and spatulate scales and numerous 
small spicules proximally; ventral surface 
as dorsal but with lateral scales and nume- 
rous longer spicules; mesal parabasal spine 
(parabasal |) stout, borne on slightly raised 
base; parabasal 2 stout with slender tip; pa- 
rabasal | base 0.07—0.15 from base of gon- 
ocoxite; parabasal 2 base 0.14—0.15 from 
base of gonocoxite; internal seta slender, 
about as long as parabasal 2, base 0.41— 
0.46 distance from base of gonocoxite. 
Gonostylus widened at base and narrowed 
toward middle and tip, base 2.27 wider 
than middle or tip; gonostylus 0.47 length 
of gonocoxite; gonostylus 8.89 longer 
than gonostylar claw. Claspette. Dorsal 
lobe of claspette with 3 closely appressed 
setae of about equal length; tips of 2 lateral 
setae curved and bluntly rounded; tip of 
middle seta slightly curved and round; tip 
of composite structure club-shaped. Ventral 
lobe of claspette with 2 long subapical se- 
tae, most apical much longer than other. 
Both ventral and dorsal lobes, and areas in 
between them, with numerous spicules. Ae- 
deagal leaflets 4 per side; 2 most mesal leaf- 
lets broadest, with broad, thin, nearly trans- 
parent inner margins; other leaflets with 
narrow, thin, nearly transparent inner mar- 
gins. 

Pupa (Figs. 2A, B).—Position and de- 
velopment of setae as figured; range and 
modal number of branches, and number of 
branches of neotype female as in Table 2. 
Integument darkly pigmented. Exuviae co- 
lorless to dark brown. Cephalothorax: Me- 
sothoracic wing with checkered dark 
stripes; metathoracic wing pigmented on 
middle, ventral spiracular sensilla distinct. 
Trumpet with dark thickend areas bearing 
saw-toothed or serrate edge, meatus with 
simple cleft, and its subbasal part with nu- 
merous spinules; trumpet length 0.30—0.45 
mm (mean = 0.35 mm, n = 10 for this and 
following measurements and counts except 
where indicated), width 0.16—0.23 mm 
(mean = 0.13 mm, measured at base of pin- 
na), index 1.36—2.5 (mean = 1.73); trach- 
eoid area 0.45 length of trumpet. Abdomen: 


VOLUME 107, NUMBER 3 611 


ae ay 
C ae 8 


fi 
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ERDAS 
\ 


Fig. 2. Anopheles lesteri. (A) Pupa, cephalothorax. (B) Pupa, metathorax and abdomen, left side dorsal, right 
side ventral. (C) Male genitalia. (D) Tergum IX. Abbreviations used include: AADM = anterior accessory dark 


mark, ADM = anterior dark mark, CT = cephalothorax, GL = genital lobe, Pa = paddle, PADM = posterior 
accessory dark mark, PDM = posterior dark mark. 


S OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


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VOLUME 107, NUMBER 3 


Abdominal tergum I with 2 anterior dark 
marks (ADM), and 2 elongate posterior 
dark marks (PDM); each PDM narrows 
mesally at base, with maximum width 
(0.016—0.02 mm, mean = 0.019) towards 
distal tip, length 0.14—0.31 mm (mean = 
0.26, n = 10), about 0.12—0.28 (mean = 
0.24, n = 10) width of abdominal segment 
I, and longer than the longest branch of seta 
1-I. Abdominal terga II—-VII with 1 ADM, 
2 anterior accessory dark marks (AADM), 
and 2 posterior accessory dark marks 
(PADM); VIII with 1 ADM and no AADM 
and PADM;; cuticular reticulations distinct 
on II-IV; spinules scattered mostly on an- 
terior 0.37 of dorsal and lateral sides of VII 
and VIII. Seta 1-I fan-like with 15—23 acic- 
ulate dendritic branches; 6-I with 1-3 
branches; 7-I with 2 or 3 branches; 9-I with 
1 or 2 branches. Setae 1, 5-II-VII well de- 
veloped; 1-V 1.04—1.31 (mean = 1.16, n = 
4) length of 5-V; 1-VI 0.83—1.09 (mean = 
0.97, n = 4); 1-VII 1.84—2.22 (mean = 
1.09, n = 6); 3-VI aligned with and mesal 
of 1-VI unlike on other segments; 8-I-II ab- 
sent; 9-I simple, single or forked; 9-II very 
short, simple, spine-like; 9-III short, with 
slightly rounded tip, 1.50—4.00 (mean = 
2.51) length of 9-II; 9-IV strong, lightly 
pigmented and slightly pointed, 01.00—2.80 
(mean = 1.83) length of 9-III; 9-V-VIII 
long, lightly pigmented and slightly point- 
ed; 9-V 1.00—1.50 (mean = 1.68) length of 
9-IV; 9-VI 0.81—1.33 (mean = 1.02) length 
of 9-V; 9-VII 1.00—1.34 (mean = 1.11) 
length of 9-VI; 9-VIII 0.80—1.06 (mean = 
0.96) length of 9-VII; 9-VI 1.11—1.39 
(mean = 1.27, n = 5) length of segment 
VI; 9-VII 1.05—1.18 (mean = 1.13, n = 5) 
length of segment VI; 9-VIII 0.85—0.92 
(mean = 0.90, n = 5) length of segment 
VIII. Seta 9-I positioned near anterolateral 
edge of tergum; 9-II on the posterolateral 
edge of tergum; 9-III-VIII near posterolat- 
eral edge of tergum. Segment VII 1.05—1.19 
(mean = 1.12, n = 5) length of segment 
VI; segment VIII 1.11—1.39 (mean = 1.26, 
n = 5) length of segment VI; segment VIII 
1.05—1.18 (mean = 1.13, n = 5) length of 


613 


segment VII. Segment VII 0.85—0.92 (mean 
= 0.90, n = 5) width of segment VI (width 
at posterior margins); segment VIII 0.69— 
0.92 (mean = 0.84, n = 5) width of seg- 
ment VI; segment VIII 0.75—1.01 (mean = 
0.93, n = 5) width of segment VII. Width/ 
length of segment VI 2.29—2.88 (mean = 
2.48, n = 5), VII 1.84—2.21 (mean = 1.99, 
n = 5), VIII 1.23-1.85 (mean = 1.65, n = 
5). Paddle length 0.70—0.75 mm (mean = 


0.72 mm, n = 5), width 0.15—0.34 mm 
(mean = 0.22 mm, n = 5), length/width 


ratio 2.24—4.86 (mean = 3.73, n = 5); re- 
fractile index 0.34—0.67 (mean = 0.54, n = 
5); paddle seta 1-Pa simple or forked (2—4 
apical branches), length 0.14—0.18 mm 
(mean = 0.16, n = 9); 2-Pa simple or 
forked (2 apical branches), length 0.40— 
0.14 mm (mean = 0.11); length of 1-Pa 
1.11—3.80 (mean = 1.66, n = 9) length of 
2-Pa. Width/length of genital lobe of female 
1.33-1.43 (mean = 1.37, n = 3), male 
0.87—0.89 (mean = 0.88, n = 2): numerous 
spicules present on subapical and apical 
margins of genital lobe of female, absent in 
male. 

Larva (Fig. 3).—Position and develop- 
ment of setae as figured; range and modal 
number of branches and number of branch- 
es of neotype female as shown in Table 3. 
Head: Length 0.68—0.71 mm (mean = 
0.70, n = 4), width 0.66—0.76 mm (mean 
0.71, n = 4). Antennal length 0.25—0.29 
mm (mean = 0.26, n = 8), slightly tapered 
toward apex, 4.57—6.00 (mean = 5.27, n = 
8) longer than wide; with spicules longer 
and more numerous ventrally and in vicin- 
ity of seta 1A; spicule length 0.01—0.02 mm 
(mean = 0.02, n = 12). Seta 1-A with 9- 
13 (mode = 10, n = 8) branches, length 
0.16—0.24 mm (mean = 0.19, n = 8), in- 
serted 0.11—0.22 mm (mean = 0.14, n = 8) 
from base of antenna; 2-A single, pointed, 
length 0.10—0.20 mm (mean = 0.13, n = 
8); 3-A single, pointed, length 0.05—0.17 
mm (mean = 0.08, n = 7); 4-A with 6-8 
branches (mode = 8, n = 7); 5-A short, 
spine-like, 0.06—0.17 (mean = 0.13, n = 8) 
length of seta 1-A; 6-A_ spine-like about 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


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Anopheles lesteri, larva. (A) Head, left side dorsal, right side ventral. (B) Dorsomentum (Dm). (C) 


Fig. 3. 
Thorax and abdominal segments I-VI, left side dorsal, right side ventral. (D) Pecten plate (PP) and pecten spines. 


(E) Abdominal segments VIII—X, side view. 


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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


1.6X longer than seta 5-A. Seta 2-C single 
1.69—2.37 (mean = 1.92, n = 3) length of 
3-C; seta 2-C close to mate of opposite side 
0.002—0.006 mm (mean = 0.004, n = 4); 
3-C densely dendritic with 32—68 main 
branches (mode = 42), 0.10—0.14 (mean = 
0.12, n = 8) length of 2-C, clypeal index 
(distance between bases 2-C and 3-C on | 
side/distance between bases of 2-C) 11.33— 
40.00 (mean = 27.83, n = 4). Thorax: Seta 
1-P with 1—3 branches (mode = 1, n = 8); 
9—-10,12-P single; 9—12-P setal support 
plate spine length 0.04 mm. Setae 9—12-M 
single; 9-M 3.12—12.67 (mean = 9.48, n = 
3) length 10-M; 9—12-M setal support plate 
spine length 0.01—0.02 mm (mean = 0.02, 
n = 4). Setae 9-10-T single; 9-T 1.18—1.31 
(mean = 1.24, n = 4) length of 10-T; seta 
12-T with 1—3 branches; 9—12-T setal sup- 
port plate spine length 0.02 mm (n = 2); 
13-T with 3 branches. Abdomen. Seta 1-I 
with 9-14 branches (mode = 12, n = 8); 
1-II 10—15 branches (mode = 13, n = 8). 
Seta 1-II—VII palmate with well-developed 
leaflets, each leaflet with short filament; O- 
II-VI and 14-II—VIII weakly developed; 
0,8,14-I, 14-II absent or rare; 3-II—III,V-VI 
single; 3-I, IV,VII branched. Seta 1-X sin- 
gle, 1.30—2.40 (mean = 1.81, n = 5) length 
of saddle; 1-X inserted on saddle. Saddle 
with minute, sparse spicules on lateral sur- 
face. Integument of posterior margin of seg- 
ment X with strongly developed dark 
brown to black spicules. Spiracular appa- 
ratus. Pecten plate with 12—18 spines; ar- 
rangement of spines alternating long and 
short, with 7 or 8 (mode = 7, n = 6) long 
spines and 5—11 (mode = 9, n = 6) short 
spines; long spines 1.27—11.67 (mean = 
3.86, n = 37) length of short spines. Two 
posterolateral spiracular lobe plates present, 
each plate with elongate, slender, sclero- 
tized projection from inner caudal margin. 

Type material.—Neotype female with as- 
sociated slide-mounted larval and pupal ex- 
uviae and DNA of a midleg of female; 
reared from a larva collected from a ditch 
on hill with slow flowing clear water, 
28.8°C, pH 6.79, salinity 0.07 ppt, conduc- 


tivity 0.15 mS, data as follows: ““Tanque’’, 
Calauan, Laguna, Luzon, Philippines, L. M. 
Rueda Coll. 29 July 2002, 14°08'44’N, 
121°18'54’E, collection and specimen no. 
PH 9-7. Deposited in the National Museum 
of Natural History, Smithsonian Institution, 
Washington, DC. (WRBU ACC No. 1729). 
Alloneotype male with slide-mounted gen- 
italia, extracted DNA of combined head, 
thorax and abdomen, and associated slide- 
mounted pupal exuviae with collection no. 
PH 9-6 and same collection data as neo- 
type female. The morphological descrip- 
tions of the head, thorax and abdomen of 
the alloneotype male were recorded before 
being processed for DNA. We were unable 
to collect any specimens from the original 
type locality of Santa Mesa, Manila, Luzon, 
because it is now a highly urban area totally 
lacking typical larval habitats. Baisas and 
Hu (1936) noted that many cotypes of An. 
lesteri were collected from Calauan, Lagu- 
na, Luzon, about 50 km south of Santa 
Mesa. This locality remains rural and we 
were able to collect specimens from Ca- 
lauan for the present study. Morphological 
data in Tables 1—3 are based on measure- 
ments of the neotype, alloneotype and as- 
sociated specimens collected from Calauan 
in 2002. 

Other material examined.—-247 speci- 
mens in the National Museum of Natural 
History, Smithsonian Institution, Washing- 
ton DC, consisting of 54 2, 29 3, 76 pupal 
exuviae, 85 larval exuviae, and 2 ¢6 geni- 
talia. PHILIPPINES, LUZON: Province of 
LAGUNA, Calauan, same collection data 
as female neotype, PH 9—3, 1 F PeLe; PH 
9-8, 1 2 Pele; PH 9-11, 1 6 G Pele; 17 
April 1930, Lot 77-19, 1 35; 20 Apr. 1930, 
ot 77-192 ese lO Wane 1938 othe 
Os Satuly. 19S1R om 2475 3) oe ale Sep 
IOS, 2 OL 3 Os sl Seay 1932, WEE ine 
coll., Lot K317i, 1 ¢. Province of MIN- 
DORO ORIENTAL, Ordovilla, 0.5 km W. 
Victoria, seepage spring, B. Harrison and 
Kol coll., 17 July 1969, P58-127, 1 Pe; B. 
Harrison coll., 19 July 1969, P61-37, 1 
LePe; 1969, B. Harrison and Kol coll., 


VOLUME 107, NUMBER 3 


iL 
61 
Wi 
181 
241 
301 
SAL 


aattagaagt 
tcgtgcataa 
aaggccgcga 
gttctaggca 
EIEIEIESNEIEIEIE AL 
Caagagactg 
tccgtgcaga 


ttggcaaaca 
aggtgtaaga 
agacagacaa 
ggcgcgcecct 
aaaattgagg 
gaatcggaag 
acgactggaa 


gaaaactacg 
gagatctcgt 
gtagtaaaca 
gacgctgtgt 
PalalgacEeee 
ttgaacaacg 
gatgcaagtt 


cagtgattgg 
cgatcgcttg 
gcagcagatg 
gtagatggag 
G\EKNSIBIEIC(CIE ie 
gaacactcta 
ctacctgaga 


tgetggtcac 
catctcggaa 
EqQEECCegqcg 
CAaggegEeceE 
cgagatagtg 
ttagcaaaca 
atcattatca 


617 


cacgtcacgg 
CEEGEgEEga 
attggcggaa 
(eee eieche 
gaatgggctg 
ctacccagaa 
cttacgagtg 


421 
Fig. 4. 


aggccactcg gtggtcaa 


Internal transcribed spacer 2 (ITS2) sequence of the ribosomal DNA for An. /esteri from the type 


locality, Calauan, Laguna, Philippines. GenBank accession number AY375469. 


SEAMP Acc. 233, P44-1, 1 2; P58-36, 1 
6s Peres, Il 23 esr I veo Jeera Le) be 
1 6; P58-127, 1 d; Caminawit Pt., 30 Dec. 
1944, G. H. Pena coll., No. C-41, 1 M. 
MOUNTAIN PROVINCE, Baguio, 12 June 
1945, 32MSU#140, 1 2; S. E. Sheilds coll., 
10 Aug. 1945, 24MSU#432, 12 2, 4 6; 
Trinidad, May 1945, J. G. Franclemont, 4 
2, 1 3. Province of NUEVA ECIJA, Mu- 
noz, Rozeboom, Knight and Laffoon coll., 
8 Aug. 1945, #1153, 3 LePe. Province of 
PAMPANGA, Olongapo (Santa Rita), Roz- 
eboom, Knight and Laffoon coll., 10 Aug. 
1945, #1115, 4 PeLe; #1116, 1 PeLe. Prov- 
ince of RIZAL, Camp Nichols, PVT Car- 
raway coll., 14 Dec. 1924, 1 92; 18 Dec. 
1945, P469, salty fishpond with algae, | d; 
Mandaluyong, 17 Dec. 1945, P468, rice 
field, i 2. VISAYAS: Province of LEYTE, 
Tacloban, Rozeboom, Knight and Laffoon 
coll., 16 Aug. 1945, No. 1713, 1 LePe, 1 
2, 1 6; Southern Leyte, 2 Dec. 1944, O. 
H. Graham, 4 2, 1 6d; Palo Alto, 1945, 1 
2; 8 Jan. 1945, 32MSU#384a, 1 ¢; 1 Feb. 
1945, 32MSU#P430, 1 6; 11 Mar. 1945, 
32MSU#474, 1 2. Province of NEGROS 
OCCIDENTAL, La Carlota, 31 Jan. 1930, 
W. V. King coll., Lot 124-16, 1 d.; Silay, 
3 Feb. 1930, Lot 137, 1 6. Province of SA- 
MAR, Osmena, Rozeboom, Knight and 
Laffoon coll., 8 Sept. 1945, #1334.30, prog- 
eny brood, 9 Le, 29 PeLe; #1334.34, 2 Le, 
12 PeLe; #1348.3, 13 PeLe; #1348.6, 4 
PeLe; 1945, Rozeboon, Knight, Laffoon 
coll., No. 1348.5, 3 M:; San Antonio, 29 
Nov. 1944, J. H. Paullos coll., No. 506, | 
2; 1 Dec. 1944, J. H. Paullos coll., No. 507, 
I) 2: Dec. 1944, 2:2, 3.6. 


Molecular characterization—DNA was 
extracted from a midleg of the neotype fe- 
male (PH9-—7), the whole body (excluding 
genitalia) of the alloneotype male (PH9—6), 
a midleg of a female (PH9-—3), and the 
whole body (excluding genitalia) of a male 
(PH9—11). Ribosomal DNA ITS2 sequenc- 
es are the same for all 4 (GenBank acces- 
sion number AY375469 (PH9-—6; Fig. 4). 
Other sequences in GenBank that match 
these sequences are as follow: under the 
name An. anthropophagus, Acc. Nos. 
AF384172, AJ004941, AF543860; An. les- 
teri Korea, Acc. No. AY375468); An. les- 
teri China (locally identified as An. anthro- 
pophagus), Acc. No. AY 375467. 

Distribution.—China (Hong Kong, south 
and central areas of the mainland, extending 
west to 105°E longitude and north to 43°N), 
Guam, Japan (including Ryukyu Islands), 
Korea, Philippines (Luzon: Laguna, Manila, 
Mindoro Oriental, Mountain Province, 
Nueva Ecija, Pampanga, Rizal; Visayas: 
Leyte, Negros Occidental, Samar). 

Medical importance.—Anopheles lesteri 
is a human biter and is considered a prin- 
cipal vector of malaria in southern China 
(Beales 1984, Chow 1991, Ho et al. 1962, 
Ma 1981) and other areas of the country (as 
anthrophophagus, Tang et al. 1991). It is 
suspected of being a primary vector in Ja- 
pan and Korea (Kamimura 1968, Otsuru 
1949, Tanaka et al. 1979). Natural infection 
rates of An. lesteri in the 1960s were 1.9 to 
14.4 times greater than An. sinensis in Chi- 
na (Gu et al. 1966). In the Philippines and 
Guam, its biting habits are unknown, and it 
is not known to transmit malarial parasites. 


618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


It (as anthropophagus) has a strong prefer- 
ence for human blood, and plays an impor- 
tant role in the transmission of filariasis and 
malaria in central and south China (Xu and 
Feng 1975). Harrison (1973) suggested a 
need to determine the distribution, behavior, 
and malarial and filarial parasite suscepti- 
bilities of An. lesteri throughout its range. 
This species, instead of An. sinensis, may 
be the more significant vector in Taiwan, 
Okinawa, Japan, Korea, and central and 
northern China. 

Bionomics.—The larvae of An. lesteri 
are found in a variety of habitats including 
freshwater pools, margins of ponds, lakes, 
areas preferably not affected by salt tides 
(Baisas 1974), and ditches with slowly 
flowing clear water in hilly areas in the 
Philippines. In Japan, the larvae occur in 
marshes, sluggish streams, ground pools, 
ponds, rice fields, and other impounded wa- 
ters (Tanaka et al. 1979). Unlike An. sinen- 
sis, An. lesteri prefers places that are cool 
and shaded. Adult populations of An. lesteri 
reach their peaks during the summer season 
in Hokkaido (Kamimura 1976), and during 
June and October in Honshu and Kyushu, 
Japan (Otsuru and Ohmori 1960). The spe- 
cies is more frequently found in coastal 
areas than inland. In Hong Kong, it com- 
monly occurs in hilly areas and grassy 
fields (Chau 1982). In Guam, An. lesteri 
larvae were found in a carabao wallow 
(Basio and Reisen 1971). Adults of An. les- 
teri, An. sinensis and other anophelines 
were collected in cow sheds and living 
rooms of houses in villages during malaria 
surveys in Korea (Whang 1962). Anopheles 
lesteri has been confused with An. sinensis 
and other members of the Hyrcanus Group, 
and some published records of its distribu- 
tion and bionomics are not accurate, partic- 
ularly in Japan, Korea and China. 


DISCUSSION 


Although Baisas and Hu (1936) provided 
the original description of An. lesteri, it was 
not sufficient for accurate identification of 
the species. This resulted in misidentifica- 


tions of the species in many parts of its geo- 
graphical range in Asia, particularly Korea, 
Japan and China. The morphological infor- 
mation in this paper, coupled with rDNA 
ITS2 sequence (Wilkerson et al. 2003), will 
help in solving those problems. 
Anophleles lesteri has the following di- 
agnostic features. Adult female. Maxillary 
palpus with palpomere 3 having narrow 
basal pale band about as wide as pale bands 
of other palpomeres; vein Cu2 without api- 
cal pale fringe spot (unlike sinensis, sine- 
roides, pullus); subcostal pale (SCP) spot 
narrow; humeral crossveins without scales 
(unlike pseudosinensis); midcoxa without 
upper patch of pale scales (unlike sinensis); 
hindtarsomeres 2—4 with narrow apical pale 
bands, hindtarsomere 4 without basal pale 
band (unlike peditaeniatus). Adult male. 
Male genitalia with dorsal lobe of claspette 
having 3 closely appressed setae of about 
equal length. Aedeagus with 4 leaflets on 
each side; 2 most mesal leaflets with broad- 
er transparent inner margins than other leaf- 
lets. Tergum IX bare, with pair of caudally 
directed elongate capitate lobes. Pupa. 
Trumpet with thick and serrate rim. Wing 
with checkered dark stripes. Setae 9-IIJ—-VIU 
single, with narrowly rounded apex; seta 5- 
V with 13—24 branches. Larva. Setae 2-C, 
3-P, 3, 5-M single; 3-C with 32—68 branch- 
es; 4-M with 3—5 erect branches; 9-M more 
than 3 times the length of 10-M; 9-M about 
1.5 times longer than 10-T; 5-III with 5—8 
branches; 9-III with 6—11 branches; 13-IV 
with 3—5 branches; 1-X strong, single about 
2 times or more length of saddle; pecten 
with 7 or 8 long spines and 5—11 short 
spines. Reid (1953) and Harrison (1973) 
provided useful diagnostic pupal and adult 
morphological characters to separate An. 
lesteri from An. sinensis. Harrison and 
Scanlon (1975) also listed several charac- 
ters of all life stages of 10 species of the 
Hyrcanus Group found in Thailand. They 
also discussed extensively the morphologi- 
cal taxonomy of the Lesteri Subgroup. 
Comparisons of pupal and larval characters 
of An. lesteri and related species of the Hyr- 


VOLUME 107, NUMBER 3 


canus Group from China and other areas of 
Asia are needed to further clarify species 
differences. Other morphological features 
described in this paper for larvae, pupae 
and adults may be helpful for separating 
An. lesteri from related species. 
Morphological similarities in all stages, 
along with intraspecific variation of many 
species in the Hyrcanus Group, have lead 
to much confusion in Korea, Japan and Chi- 
na (Tanaka et al. 1979), and possibly in oth- 
er areas of Asia where they occur. Wilker- 
son et al. (2003) suggested that the best way 
to infer conspecificity of populations across 
large geographic areas is to compare spec- 
imens from type localities. Based on the 
combination of published and their newly 
generated rDNA ITS2 sequences, Wilker- 
son et al. (2003) found that An. lesteri from 
South Korea and An. anthropophagus from 
Jiangsu Province, China, are the same as 
An. lesteri from near its type locality in the 
Philippines (Calauan, Laguna, Luzon). 
Anopheles anthropohagus, considered a 
major malaria vector in central and north 
China, is actually An. lesteri, not a separate 
species. With that finding, they placed An. 
anthropophagus in synonymy with its se- 
nior synonym, An. lesteri. Any morpholog- 
ical features previously thought to differ- 
entiate An. anthropophagus and An. lesteri 
are evidently variable characters of a single 
species. What is called An. lesteri in China 
(as reported by Gao et al. 2004: 7, 9) is 
actually an unknown species when com- 
pared with the work of Wilkerson et al. 
(2003). Several molecular studies (e.g., Li 
et al. 1991: Ma et al. 2000a, b; Gao et al. 
2004) were conducted but were unable to 
clarify the taxonomic identity of An. lesteri 
found in China and Korea. With the collec- 
tion of An. lesteri specimens from near the 
type locality in Luzon, Philippines, Wilk- 
erson et al. (2003) were able to compare the 
specimens with those from Korea and Chi- 
na, and they concluded that the so called 
An. anthropophagus from China, and the so 
called An. lesteri from Korea are conspe- 
cific with An. lesteri from the Philippines. 


619 


With the designation of the neotype and 
detailed descriptions of various life stages 
of An. lesteri based on specimens collected 
near the type locality, future systematic 
studies may be conducted using various 
methods, including morphological, molec- 
ular or biochemical. With the identity of An. 
lesteri resolved, the effectiveness of malaria 
vector control practices could be further im- 
proved. As suggested by Harrison (1973), 
additional information is needed on the dis- 
tribution, behavior, and malarial and filarial 
parasite susceptibilities of vector species 
throughout their ranges. A vector species 
may be a more significant parasite vector in 
one geographical area than in others. For 
example, An. lesteri is considered an im- 
portant malaria vector in China and Korea, 
but not in the Philippines. Furthermore, be- 
cause several species of the Hyrcanus 
Group are involved in the transmission of 
malaria and filariasis in Asia, there is a need 
to revise the taxonomy of the whole group 
and to further clarify the identities of the 
cryptic species, particularly the vectors, in 
the group. 


ACKNOWLEDGMENTS 


We thank Bruce A. Harrison, Yiau-Min 
Huang and Maria Anice M. Sallum for re- 
viewing the manuscript and providing in- 
valuable advice. Special thanks to the fol- 
lowing: Taina Litwak for her illustrations; 
Bel Rueda, Buddy Buenavista, Lerma Bue- 
navista, Benjie Puma, and Modesta Coro- 
nado-Puma for their help in collecting mos- 
quitoes. Jim Pecor provided technical sup- 
port in curation of specimens and prepara- 
tion of slide mounts. This research was 
performed under a Memorandum of Under- 
standing between the Walter Reed Army In- 
stitute of Research and the Smithsonian In- 
stitution, with institutional support provided 
by both organizations. The opinions and as- 
sertions contained herein are those of the 
authors and are not to be construed as of- 
ficial or reflecting the views of the Depart- 
ment of the Army or the Department of De- 
fense. 


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PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 623-626 


TWO BACTERIAL PATHOGENS OF HELICOVERPA ARMIGERA (HUBNER) 
(LEPIDOPTERA: NOCTUIDAE) 


MusTAFA YAMAN, IRFAN ASLAN, ONDER CALMASUR, AND FIKRETTIN SAHIN 


(MY) Department of Biology, Faculty of Arts and Sciences, Karadeniz Technical Uni- 
versity, 61080 Trabzon, Turkey; (IA, OG, FS) Atatiirk University, Faculty of Agriculture, 
Plant Protection Department, 25240 Erzurum, Turkey (e-mail: calmasur@atauni.edu.tr, 
ondercal 1 @hotmail.com) 


Abstract.—Bacterial pathogens of Helicoverpa armigera (Hiibner) (Lepidoptera: Noc- 
tuidae) were investigated. Two different pathogenic bacteria were isolated from unhealthy 
and dead larvae. They were identified as Pantoea agglomerans (Ewing and Fife) and 
Alcaligenes piechaudii Kiredjian et al. on the basis of fatty acid methyl ester (FAMEs) 
analysis and carbon utilization profiles by using Microbial Identification and Biolog Mi- 
croplate Systems. Laboratory experiments carried out to determine insecticidal activities 
of these isolates showed that Pantoea agglomerans and Alcaligenes piechaudii have 
95.0% and 98.75% mortality on third-instar larvae of H. armigera, respectively, after 14 
days. This is the first study to demonstrate that Pantoea agglomerans and Alcaligenes 
piechaudii are pathogenic bacterial flora of H. armigera. These bacteria may have great 


potential for use in biological control of H. armigera. 


Key Words: 


The genus Helicoverpa consists of sev- 
eral pest species, and among them Helcov- 
erpa armigera (Hiibner) is one of the most 
economically important pests in Turkey. It 
is an especially notable pest of sorghum, 
maize, cotton, chickpeas, and tomatoes 
(Turkish Ministry of Agriculture 1995). 
Chemical substances utilized to control this 
pest have hazardous effects in the environ- 
ment. As the use of chemical pesticides is 
a social issue, the objectives of nutrition, 
health, and environmental quality can be 
addressed more efficiently by the imple- 
mentation of integrated pest management 
techniques (IPM) rather than through cur- 
rent crop protection practices (Norgard 
1976). 

Increasing problems with synthetic insec- 
ticides have spurred the search for alterna- 
tive pest management strategies that would 


Helicoverpa armigera, bacterial pathogen, microbial control 


reduce reliance on synthetic insecticides. 
Biological control of plant pests is an alter- 
native control method in lieu of chemical 
pesticides. Recently, several micro-organ- 
isms have been isolated and identified from 
insects as potential biological control agents 
(Weiser et al. 2002, Yaman 2003, Yaman 
and Radek 2003). 

Studies on natural enemies of H. armi- 
gera have focused on pathogenic viruses 
(Tuan and Hou 1988, Teakle and Byrne 
1989, Parnell et al. 1999, Narayanan 2002), 
Bacillus thuringiensis (Bt) (Navon et al. 
1990), and microsporidia (Tsai et al. 2003). 
However, very little is known about other 
bacterial pathogens limiting its populations. 
We present results of a study on the isola- 
tion, identification, and insecticidal effect of 
two new pathogenic bacteria for H. armi- 
gera in Turkey. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


MATERIALS AND METHODS 


Collection of H. armigera larvae.—Dur- 
ing spring and summer 2001 and 2002, H. 
armigera larvae were collected from chick- 
pea plants grown in the Agricultural Ex- 
periment Station in the vicinity of Erzurum, 
Turkey. The temperature range in the field 
was approximately 20—25°C. Dead and liv- 
ing larvae exhibiting characteristic disease 
symptoms (no feeding, slow moving, and 
color changes on the body) were selected 
and transported to the laboratory in sterile 
tubes within | h after collection. Dead lar- 
vae found in the field were put into sterile 
tubes to prevent possible contamination. All 
dead and unhealthy larvae collected were 
kept in a refrigerator for 2 h to be cooled 
then used for microbial isolation. 

Isolation of bacterial strains and culture 
conditions.—After macroscopic examina- 
tion, microscopic examination was done to 
be sure of bacterial infection. Dead larvae 
were surface sterilized in 70% alcohol 
(Poinar 1978). For this, a small vessel was 
filled with 10 ml of the disinfectant, the lar- 
va was put in it, and the vessel was corked 
and shaken for 2 minutes. The larva then 
was rinsed three times in sterile water in the 
same way in three clean vessels. After cut- 
ting the cuticle with sterile scissors, a drop 
of the fluid content was taken with an in- 
oculating loop, diluted 100 times with ster- 
ile water, and spread on nutrient agar plates. 
Plates were incubated at 28°C to 37°C for 
2-3 d. After the incubation period, plates 
were examined and bacterial colonies were 
selected (Lipa 1975, Thiery and Frachon 
1997). Selected colonies were purified by 
subculture on plates. The two most preva- 
lent colony types of bacteria were selected 
and purified on nutrient agar plate by sub- 
culturing. Bacterial strains were maintained 
for long-term storage in nutrient broth with 
15% glycerol at —86°C for further tests. 

Identification of bacterial strains.—All 
isolated bacterial strains were identified 
based on fatty acid profiles determined us- 
ing the Microbial Identification System 


(Hewlett-Packard 6890A, Palo Alto, CA) 
with TSBA (Tripticose Say Broth Agar) da- 
tabase in the Sherlock Microbial Identifi- 
cation System software package (MIDI, 
Microbial ID, Inca, Newark, DE) and car- 
bon substrate utilization fingerprints ana- 
lyzed by the Biology GN and GP database 
with Microlog software in Biolog Micro- 
plate Systems (Biolog Inc., Hayward, CA). 
The isolates were stored at the Department 
of Plant Protection, Faculty of Agriculture, 
Ataturk University. 

Bioassays of bacterial isolates —Bioas- 
says were performed on third-stage instars 
of 20 larvae of H. armigera for each bac- 
terial isolate to determine their insecticidal 
activity. Helicoverpa armigera larvae dam- 
age chickpea by feeding on leaves and 
grains. Therefore, the larvae were fed with 
chickpea leaves and grains sprayed with the 
bacterial suspension (Dalmage 1981, 
McGuire et al. 1997). The bacterial isolates 
were grown in nutrient broth at 28°C over- 
night. The bacterial suspension for each 
pathogen was prepared in sterile water at 
the concentration of 10° CFU/ml and then 
immediately used for bioassay study. The 
control group was fed with chickpea leaves 
and grains sprayed with sterilized water. 
After 48 h, the larvae received fresh diet 
every 24 h. The larvae were tested for each 
bioassay during 14 d. The bacteria were 
readily re-isolated from dead larvae and 
identified as original isolates used for in- 
oculation by MIS. All larvae tested were 
kept at 26°C and 60% RH on a 12:12 h 
photo regime. Observations were carried 
out daily and dead larvae were removed im- 
mediately. All bioassays were repeated 4 
times on different days. Data were evalu- 
ated using Abbott’s formula (Abbott 1925). 

Data analysis.—Univariate using SPSS 
11.0 software was used to determine if 
there was a statistically significant differ- 
ence in insecticidal effect between the in- 
secticidal activities and time. The results 
showed significant difference at P < 0.05 
levels. 


VOLUME 107, NUMBER 3 


625 


Table 1. Results of multiple comparison with mean and standard error of time. Values followed by different 
letters in the same column differ significantly at P < 0.05. 


Pantoea agglomerans 


Alcaligenes piechaudii 


Time (Days) N Mean SE Mean SE 
5 4 36;259ce% 4,732 AS 56% 5,543 
10 4 77,50b** 6,614 83,75b** 3,145 
14 4 95 ,00a** 2,041 OS Sane 1,250 
Control 4 1,25d** 1,250 2,50d** 1,443 
= 12) << (OOS), 


RESULTS AND DISCUSSION 


Recent studies on natural enemies of H. 
armigera have been focused on pathogenic 
viruses (Tuan and Hou 1988, Teakle and 
Byrne 1989, Parnell et al. 1999, Narayanan 
2002) and Bacillus thuringiensis Berliner 
and microsporidia (Tsai et al. 2003). In this 
study, we isolated two different nonspore- 
forming bacteria from living and dead lar- 
vae of H. armigera. Fatty acid analysis 
identified the bacterial pathogens as Pan- 
toea agglomerans (Ewing and Fife) and Al- 
caligenes piechaudii Kiredjian et al. with 
similarity incidence of 53% and 89%, re- 
spectively. The identity of Panoea agglom- 
erans at species level and Alcaligenes pie- 
chaudii at genus level was confirmed by 
Biolog GN Microplate with similarity in- 
cidence of 78% and 44%, respectively. 

Different species of the genus Alcalige- 
nes such as A. recti, A. faecalis, and A. 
odorans are found in insects (Lipa and Wi- 
land 1972, Bucher 1981). Majumder et al. 
(1955) isolated Bacillus thuringiensis vat. 
thuringiensis from H. armigera. Lipa and 
Wiland (1972) isolated four bacteria, Aero- 
bacter cloaceae (Jordan), Alcaligenes recti 
(Packer and Vishniac), Escherichia freundii 
(Braak), and Escherichia coli (Migula) 
from H. armigera, but they did not test their 
pathogenicity on H. armigera. They gave 
an extensive list of bacteria and bacterial 
diseases recorded in Noctuidae. Our records 
are the first for Pantoea agglomerans and 
Alcaligenes piechaudii from Noctuidae. 
This study adds two pathogenic bacteria to 
the bacterial microflora of H. armigera and 


confirms that the bacterial microflora of H. 
armigera consists principally of nonspore- 
forming bacteria with one exception, Bacil- 
lus thuringiensis var. thuringiensis. 

Results of the bioassays carried out for 
determining the insecticidal effect of the 
isolated bacteria showed that Pantoea ag- 
glomerans and Alcaligenes piechaudii have 
95.00% and 98.75% mortality on the larvae 
of H. armigera, respectively, after 14 days 
(Table 1). Koch postulate studies showed 
that both of the bacterial strains caused sim- 
ilar disease symptoms on H. armigera lar- 
vae, which were lack of appetite, slowing 
and then failure in movement, and discol- 
oration (blackish color) on the body. Lipa 
and Wiland (1972) tested pathogenicity of 
their bacteria isolated from H. armigera 
against third-instar larvae of Agrotis c-ni- 
grum and Agrotis segetum. The highest 
pathogenic bacterium showed 67% and 
92% mortality after two days for Agrotis c- 
nigrum and Agrotis segetum, respectively, 
and among them was Alcaligenes recti iso- 
lated from diseased larvae of H. armigera 
(Lipa and Wiland 1972). Our isolates 
showed high pathogenicity (95.00% and 
98.75%) against H. armigera (Table 1). 
These are the first recorded 
forming bacteria showing such high path- 
ogenicity to H. armigera. We isolated them 
from the hemocoel of H. armigera. Poten- 


non-spore- 


tial pathogens multiply and cause death of 
insects after they penetrate the hemocoel in 
small numbers (Lipa 1975). 

It is important that Pantoea agglomerans 
and Alcaligenes piechaudii have sufficient 


626 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


insecticidal effects on the larvae of H. ar- 
migera, because biological control studies 
involving bacteria of H. armigera usually 
focus on B. thuringiensis (Navon et al. 
1990). These bacteria can consequently be 
natural suppressors against H. armigera 
populations. 


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PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 627-641 


DESCRIPTIONS OF ADDITIONAL NEW WORLD TRICHOGRAMMATIDAE 
(HYMENOPTERA): THE GENUS NICOLAVESPA AND A NEW 
SPECIES OF HAECKELIANIA 


JOHN D. PINTO 


Department of Entomology, University of California, Riverside, CA 92521, U.S.A. 
(e-mail: jpinto@ucr.edu); current address: P.O. Box 2266, Waldport, OR 97394, U.S.A. 


Abstract.—New taxa of Trichogrammatidae from the New World are described. In- 
cluded is the new genus Nicolavespa with two new species, N. theresae and N. luiseno, 
and a new species of Haeckeliania, H. sperata. Nicolavespa theresae is widespread in 
North America; the closely related N. luiseno is known only from southern California. 
Haeckeliania is one of the most speciose genera of Trichogrammatidae although few 
species have been described. Haeckeliania sperata, from Dominica, attacks eggs of Cur- 
culionidae and is a potential biological control agent against the Diaprepes root weevil. 


Key Words: 
root weevil 


Described below are three new species of 
Trichogrammatidae. Two belong to the new 
genus WNicolavespa, a group ranging 
throughout much of North America as far 
south as Costa Rica. Although relatively 
widespread and sometimes locally abun- 
dant, the genus has not been collected pre- 
viously. All specimens are derived from rel- 
atively recent collections. The hosts of Ni- 
colavespa are unknown. The third species 
described belongs to the unrelated genus 
Haeckeliania Girault. The purpose of this 
isolated species description is to provide a 
name for a species from Dominica associ- 
ated with an on-going biological control 
project by Jorge Pefia (University of Flori- 
da) to suppress the Diaprepes root weevil 
(Curculionidae) on citrus and other crops in 
Florida (Hall et al. 2001). It also provides 
the opportunity to call attention to the char- 
acteristics and considerable species diver- 
sity of Haeckeliania, a genus heretofore 
known only from a single described species 
in the New World. 


Hymenoptera, Trichogrammatidae, Nicolavespa, Haeckeliania, Diaprepes 


Descriptive terminology follows earlier 
papers (Pinto 2004, Pinto and George 
2004). Acronyms for antennal sensilla and 
setae, based largely on terms used for 77ri- 
chogramma (Olson and Andow 1993, Pinto 
1999), include APB (aporous sensillar tri- 
chodea B, socketed), PLS (sensilla placo- 
dea), BPS (basiconic peg sensilla), FS (fla- 
gelliform setae or multiporous pitted sen- 
silla trichodea A, unsocketed), UPP (uni- 
porous pit pore sensilla trichodea D), and 
APA (aporous setae A, unsocketed). The 
UPP refers to a single sensillum at the apex 
of the female club. The flagellum of the an- 
tenna includes anellar (A), funicular (F) and 
club (C) segments. The medial and lateral 
surfaces of the antennae refer to ‘inner’ and 
‘outer’ facing surfaces, respectively, when 
they are directed forward. Acronyms for 
wing venation are SMV (submarginal vein), 
PM (premarginal vein), MV (marginal 
vein) and SV (stigmal vein) (Fig. 9). As in 
earlier papers, OL refers to ovipositor 
length and HTL to hind tibial length. Body 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


length measurements are taken from card- 
mounted material; all other measurements 
are from specimens on slides. Card-mount- 
ed specimens were dried with Hexame- 
thyldisilazane ((HMDS’, Heraty and Hawks 
1998). Quantitative data for several mea- 
surements are reported as means followed 
by the range. 


Nicolavespa Pinto, new genus 


Type species: Nicolavespa theresae Pinto, 
n. sp. 


Diagnosis.—Antenna (Figs. 4—7) with 
two funicular segments and three club seg- 
ments; Fl short, transverse, closely ap- 
pressed to F2; F2 more elongate, subcy- 
lindrical; club of female with a distinct api- 
cal process. Foretibia without spines on 
dorsal surface. Forewing (Fig. 9) with setal 
tracts distinct including RS1; SV well de- 
fined. Male genitalia (Figs. 12—15) with ae- 
deagus indistinct from genital capsule api- 
cally but with apodemes present basally; 
volsellae well developed, subspatulate api- 
cally; parameres apparently absent; with a 
pair of strong ventral setae at base of vol- 
sellae. 

Description.—Female (Figs. 1—2). Head 
(Fig. 3): Relatively broad, as wide as great- 
est mesosomal width. Eyes purple. Antenna 
with anellus and funicle each with two seg- 
ments (Fig. 5), club three segmented; sur- 
face of pedicel smooth, that of funicle and 
base of club longitudinally wrinkled; A2 
partially fused to Fl; funicular segments 
closely appressed, F1 distinctly wider than 
long and much shorter than F2; F2 elon- 
gate, subcylindrical; last club segment with 
an elongate apical process (Fig. 6). Maxil- 
lary palp one-segmented. Mesosoma (Fig. 
8): Midlobe of mesoscutum and scutellum 
each with two pair of moderately long se- 
tae. Propodeal disk not produced. Meso- 
phragma notched apically. Foreleg without 
prominent spines on dorsal surface. Fore- 
wing (Fig. 9) moderately broad, slightly 
less than 2 as long as wide; MV straight, 
contacting wing margin its entire length, 


gradually increasing in width from base to 
apex, ca. 1.5 as long as PM, ending 
abruptly at juncture with SV; PM narrowly 
separated from MY, angling posteriorly and 
forming a narrow costal cell; SV distinct, 
ca. half length of MV, constricted between 
stigma and MV. Hind wing with three setal 
tracks. Metasoma: Ovipositor moderately 
elongate, distinctly longer than hind tibia, 
not produced appreciably beyond apex of 
metasoma. Hypopygium short. 

Male. Similar to female except antennal 
club without an apical process (Fig. 7); last 
metasomal sternum broadly divided medi- 
ally. Genitalia (Figs. 12—15) short, subcon- 
ical, tapering to apex, with aedeagus indis- 
tinct from genital capsule in apical half but 
apodemes distinct basally; anterodorsal ap- 
erture, wider than long, confined to basal 
third of capsule; ventral surface with sub- 
spatulate volsellae, the volsellae with a pre- 
sumed single digital spine laterally; para- 
meres apparently absent (see Discussion); a 
pair of strong ventral setae at base of vol- 
sellae; a short apically bifid protrusion dor- 
sally near apex (Fig. 12). 

Etymology.—Derived from the name of 
my father, the late Nicola Pinto, and vespa, 
Latin for wasp. Gender, feminine. 

Discussion.— Nicolavespa is superficial- 
ly similar to Chaetostricha. Both are rather 
slender, with moderately broad wings and 
similar venation. As in Nicolavespa the an- 
tenna of Chaetostricha consists of a two- 
segmented funicle and a three-segmented 
club. Also in both the F1 is short, transverse 
and appressed to a longer, subcylindrical 
F2. Unlike Nicolavespa, Chaetostricha 
lacks an apical process on the female club, 
usually has a dorsally spinose foretibia, a 
narrower head, and its male genitalia are 
considerably more reduced: the aedeagus is 
completely fused to the genital capsule, ae- 
deagal apodemes are not present, and vol- 
sellae also are absent (see Viggiani 1971). 

Similarity notwithstanding, Nicolavespa 
is not close to Chaetostricha. In the family 
classification of Viggiani (1971), based 
largely on male genitalia, Nicolavespa is as- 


VOLUME 107, NUMBER 3 


629 


Figs. 1-2. Nicolavespa 2 (venter in focus). 1, N. theresae. 2, N. luiseno. 


signed to the Trichogrammatinae. Chaetos- 
tricha is a member of the Oligositinae as 
currently defined. Instead, Nicolavespa ap- 
pears allied with Lathromeris. In both, the 
antenna has seven flagellar segments and 
the female club bears an apical process. 
Also, wing dimensions and venation are 
similar. Lathromeris is separated by its five 
segmented club, lack of an RSI setal track 
on the forewing, and presence of a well de- 
veloped hypopygium (Doutt and Viggiani 
1968). Although similar in general appear- 
ance, the male genitalia of Lathromeris ap- 
pear to lack volsellae. 

Certain structures of the male genitalia in 
Nicolavespa are not readily homologized. 
Thus, it is not clear if parameres are pres- 
ent. They either are absent or represented 
by the pair of spines at the base of the vol- 
sellae (Fig. 13). The bifid subapical protru- 
sion on the dorsum of the capsule (Fig. 12) 
has not been noted in other trichogramma- 
tids. Because it is not observed under the 


light microscope its presence in other taxa 
would easily escape detection. 

Nicolavespa runs to couplet 37 with 
Chaetostricha and Zagella in the key to 
North American Trichogrammatidae (Pinto 
1997). It is separated from both by the elon- 
gate apical process on the female club, and 
the presence of aedeagal apodemes. 


Nicolavespa theresae Pinto, new species 
(Figs. 1, 4—7, 9-10, 12-14) 


Description.—Female. Body length 0.70 
(0.60—0.85) mm (n = 10); HTL = 0.20 
(0.15—0.24) mm (n = 11); metasoma length 
(excluding slight ovipositor extension) Ca. 
1.75 that of mesosoma; metasoma grad- 
ually narrowing posteriorly, attenuate in 
apical half. Co/or: Variable (see Variation) 
but type series dark brown except as fol- 
lows: upper face and vertex orange brown; 
antenna light brown, apical club segment 
distinctly darker; ventral half of epimeron 
yellow; notauli, metanotum, propodeal disk, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


metasoma with basal two terga, and poste- 
rior portion of terga VI and VII yellow 
brown to light brown; legs lighter at base 
of femora and at base and apex of tibiae, 
tarsi lighter; forewing moderately infuscate 
behind venation. Head: Almost as long as 
wide; scrobes extending half the distance 
from toruli to median ocellus. Antenna nar- 
row, arising at level of ventral margin of 
eye; relative length/width of scape, pedicel, 
funicle and club as follows: 4.2 (3.8—4.7), 
2.0 (1.7-2.1), 1.5 (1.3-1.8), 4.6 (4.35.0); 
scape narrow, elongate, narrower in apical 
half; pedicel ca. 0.5X length of scape; F1 
twice as wide as long and slightly less than 
half length of F2, funicular segments close- 
ly appressed, subequal in width, funicular 
width ca. 0.75 that of club; club narrow, 
subfusiform, C1 and C2 subequal in length, 
C3 narrowly subconical, ca. twice length of 
Cl; apical process of club elongate, com- 
prising ca. a third total length of C3. Dis- 
tribution of antennal sensilla on funicle and 
club as follows: Fl and F2—APB and 
APA; C1—FS, APB, APA and 1-2 PLS, 
C2—FS, APA (on few individuals) and 2 
PLS; C3— FS, 4 PLS, apical process with 
1 short APB near base and a short, stout 
UPP at apex; apex of each funicular and 
club segment with subglobose BPS, a rel- 
atively broad circle of shallowly impressed 
surface surrounding base of each BPS. 
Maxillary palp with apical peglike sensil- 
lum elongate, cylindrical, ca. 0.4% length 
of terminal seta. 

Mesosoma: Midlobe of mesoscutum al- 
most as wide as long, ca. 1.6 as long as 
scutellum; midlobe and scutellum with 1m- 
bricate sculpturing; sculpted cells varying 
in dimension, subrectangular laterally, nar- 
row and elongate medially, most cells with 
low transverse ridges within, imbrication 
most pronounced anteriorly on midlobe. 
Forewing moderately broad, widest near 
apex, 1.9X as long as wide, fringe setal 
length 0.15—0.20 wing width (see Varia- 
tion); venation occupying 0.5 wing length; 
relative length of SMV, PM, MV and SV 
= 9s tl Sts Vemoderatelyiconstacted 


between stigma and MV; stigma angled to- 
ward wing apex; PM with two setae; MV 
with six dorsal setae (three marginal, long; 
three posterior, shorter) and five-six ventral 
setae. Forewing membrane moderately 
densely setose, with 14—15 linear setal 
tracks, with a field of ca. 20 alar acanthae 
on ventral surface at base. Hind wing with 
three setal tracks, posterior-most track with 
shorter setae, and not quite attaining apex 
of wing. 

Metasoma: Last tergum relatively elon- 
gate, ca. half as long as wide, with cercal 
setae usually not reaching ovipositor apex 
(Fig. 10). Ovipositor elongate (Fig. 1), oc- 
cupying almost entire length of metasoma 
(in slide-mounted specimens ovipositor 
length subequal to length of metasoma to 
apex of tersum: Vill); OL/HTE-= 2:42 
(2.0—2.8) (m = 15). 

Male. As in female except as follows: 
Body less elongate, 0.4—0.6 mm; HTL = 
0.17 (0.14—0.20) mm (n = 9); metasoma of 
similar width throughout, apex truncate, not 
attenuate; metasoma shorter, length equal to 
or only slightly greater than that of meso- 
soma. Antenna with pedicel slightly longer, 
0.6—0.7 scape length; club without an apical 
process and with fewer PLS (one present or 
absent from Cl, one on C2, two on C3); 
club shorter due to absence of apical pro- 
cess, C3 ca. 1.25 length of Cl. Forewing 
with longer fringe, longest setae ca. 0.25 
width of wing. Genitalia 0.49 (0.46—0.53) 
(n = 5) length of hind tibia; genital capsule 
0.34 (0.3—-0.40) (n = 8) as wide as long 
(Fig. 14). 

Types.—Holotype 2, allotype ¢. UNIT- 
ED STATES. California: Riverside Co.; 
Santa Rosa Plateau Ecological Reserve; 
33°32.538'N, 117°14.758'W; vii-30/viii-14- 
2001; Malaise trap #1; PEET Survey; de- 
posited in the National Museum of Natural 
History, Washington, DC. Paratypes desig- 
nated as follows: 4 2 (data as holotype); 
36, 6 2 (data as holotype except— 
33°32.489'N, 117°14.652'W; Malaise trap 
#3): 1 2 (data as holotype except— 
33°32.524'N, 117°14.644'’W; Malaise trap 


VOLUME 107, NUMBER 3 


631 


Figs. 3-8. 


Nicolavespa. 3, N. luiseno, head. 4—7, N. theresae, antenna. 4, Medial view (2). 5, Anellar and 


funicular segments (medial, 2). 6, Apex of club showing apical process and UPP sensillum inserted at apex 


(2). 7, Medial view (¢). 8, N. luiseno, mesosoma (° ). 


#2). Paratypes deposited in The Natural 
History Museum, London; the Canadian 
National Collection, Ottawa; the University 
of California, Riverside (Department of En- 
tomology); and the University of Naples, 
Portici. All except three paratypes mounted 
on slides in Canada balsam. Chaparral veg- 
etation characterizes the type locality. 


Etymology.—Named for my mother, 
Theresa Aloisio Pinto. 

Geographic distribution.—A widespread 
species. Known from California, Texas and 
United States, the Turks 
and Caicos Islands, and Costa Rica. 


eastern south to 


Variation.—Color varies geographically 


in N. theresae. Untortunately, this variation 


32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


could be adequately examined only in the 
minority of specimens mounted on cards. 
The most obvious variation is in the color 
of the midlobe of the mesoscutum. Speci- 
mens from southern California are darkest, 
with a homogeneous dark brown mesoscu- 
tum. Northern California material (Solano 
Co.) also are dark but most specimens have 
a pair of narrowly separated, yellow-brown 
vittae in the anterior half of the disk; the 
non-vittate area of the disk also is a lighter 
brown than the general surface color in 
southern California material. The scutal vit- 
tae are absent in specimens from western 
Texas but present in all material examined 
east of there (eastern Texas, Illinois, South 
Carolina, Maryland). Specimens from Gua- 
temala are similar to eastern U.S. specimens 
except that the head below the eyes also is 
lighter. Material from Costa Rica and Chia- 
pas is lightest. In these specimens the dor- 
sum of the mesosoma is almost entirely yel- 
lowish; a single Costa Rican specimen, 
however, is darker, similar to those from 
Guatemala. The head below the eyes also 
is lighter in Costa Rican specimens. Carded 
material from intermediate areas in Mexico 
was not available for study. 

The range of variation in oOvipositor 
length is considerable in this species (see 
above). Specimens with the shortest ovi- 
positor are found in the southern part of the 
range. The average OL/HTL ratio is 2.4. In 
certain specimens from the south (e.g., 
Guerrero, Turks and Caicos Islands) the ra- 
tio is 2.0 or only slightly greater. Variation 
is not obviously geographically clinal how- 
ever, since in Costa Rica, the southern-most 
region of distribution, the ovipositor is rel- 
atively long (OL/HTL = 2.3-2.7, n = 3). 

The length of the forewing fringe setae 
may also vary geographically. In specimens 
from the United States the longest fringe 
setae are 0.15 the wing width or less. In 
Mexico and Central America the fringe se- 
tae are longer, ca. 0.2 the wing width. 

Material studied.—41 ¢ (17 on slides), 
77 & (36 on slides) as follows: COSTA 
RICA. Alajuela: PN Arenal (La Peninsula, 


600 m), 10°27'N, 84°45’'W; 11-25-03; 
sweep; 3 2; J. Noyes. PN Arenal (Sendero 
Pil6n, 600 m) 10°27'N, 84°43'W; 11-26-03; 
sweep; 2 ¢; J. Noyes. Reserva Rincon 
Forestal (Est. Caribe, 400 m); 10°53’N, 
85°18'W; ii-19/20-03; 1 2; J. Noyes. He- 
redia: Puerto Viejo, 3 km S (OTS—La Sel- 
va, 100 m); ix-92; 1 9; PB. Hanson. Puntar- 
enas: Golfo Dulce, 3 km SW Rincon de 
Osa (10 m); ix/xi-89; 1 2; P. Hanson. R. E 
Golfo Dulce; 24 km W Piedras Blancas 
(200 m); iv/v-92, xii-92, viii/ix-93; 3 2; P. 
Hanson. GUATEMALA. Petén (2.0 km E 
Rikal)-xne27- 8804S ae eeleasallle: 
MEXICO. Chiapas: Ocozocoautla (El 
Aguacero, 1,800’—2,000’); viii-8-90; 1 2; J. 
Woolley. Guerrero: Acapulco, 30 km N: 
viii-6-84; sweep; | 2; G. Gordh. Xochi- 
pala, 6.2 mi. SW; vii-13-85; 2 2; J. Wool- 
ley/G. Zolnerowich. Jalisco: Guadalajara, 
17 mi. N; vii-6-84; 1 3d, 1 2; J. Woolley. 
Morelos: Tlayacapan; x-29-82; sweep; | 
6; A. Gonzalez. Sinaloa: Villa Union, 5.5 
mi. E (on Durango Hwy); ix-3-88; sweep; 
1 2; A. Sanders. TURKS AND CAICOS 
ISLANDS. Country record only; u1-17/24- 
91; pan trap; 1 ¢, 2 2; S. Rigby. UNITED 
STATES. Arizona: Patagonia (Santa Cruz 
Co.); vi-16-1994; Malaise trap; 1 6; E. 
Wilk/B. Brown. California: Altadena; viii/ 
ix-91: 1 2: R. Crandall. Santa Rosa Plateau 
Ecological Reserve (Riverside Co.); 4 6, 
12 2; (see Types). San Pedro (Defense Fuel 
Support Point) (Los Angeles Co.); 13°46’N, 
118°18'W; v-26/vii-6-02, vi-6/vili-18-02; 
Malaise trap in coastal sage scrub; 5 6,3 2; 
J. George. Winters, 11 km W (Cold Canyon 
Reservoir) (Solano Co.); viii-1/15-94; Mal- 
aise trap in live oak woods; 24 6, 22 2; L. 
Kimsey. Georgia: Sapelo Island; ix/xi-19- 
87; flight intercept trap; 1 2; BRC Hyme- 
noptera Team. //linois: Centralia (Carl Hall 
Park); ix-7-93; sweep at edge of woods; 1 
2; J. Pinto. Chicago (Indian Boundary For- 
est Preserve); vili-24-83; sweep in deep 
woods; 2 @; J. Pinto. Maryland: Patuxent 
Research Station (Prince Georges Co.); vii- 
27/vili-3-86, vili-3/9-86; Malaise trap; 3 &; 
D. Wahl. Port Republic; vii/ix-86; flight in- 


VOLUME 107, NUMBER 3 


Figs. 9-13. 


at tergum VIII, arrow ‘b’ at cercal setae). 11, N. /uiseno (arrows ‘a’ and ‘b’ as in Fig. 10). 12 


6 genitalia. 12, Dorsal view (arrow 


Nicolavespa. 9, Forewing, N. theresae (veins as follows: SMV = submarginal, PM 
ginal, MV = marginal, SV = stigmal). 10-11, Metasomal tergum VIII (dorsal, 2). 10, N. theresae (arrow ‘a 


‘a’ at apodeme of aedeagus, arrow 


= premar- 


13, N. theresae, 
‘b’ at bifid protrusion at apex). 13, 


Ventral view (arrow ‘a’ at presumed digital spine, arrow ‘b’ at apex [digitus?] of volsella). 


tercept trap; 1 2; M. Sharkey/Munroe. 
Oklahoma: Red Oak (Latimer Co.); 1x-93; 
flight intercept trap; | 2; K. Stephan. South 
Carolina: Pendleton (Tanglewood Spring, 
225 m); 34°39’N, 82°47’W; vii-30/viii-20- 
87, ix-115/30-87; Malaise trap; 3 2; J. 


Morse. Texas: Bastrop State Park; x-7/21- 
90, x-21/29-90; 2 9; R. Wharton. Big 
Bend Ranch (Presidio Co.); 29°29'40"N, 
104°06’00"W; x-27/xi-1-89; 1 2%; D. 
Judd. Big Bend Ranch; 29°27'30'N, 
104°01'04"W;  x-28/x-1-89; Malaise trap 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


14 19 


Figs. 14-15. Nicolavespa ¢ genitalia (dorsal). 14, 
N. theresae. 15, N. luiseno. 


‘nr. fl. composite’ 1 2; D. Judd. Big Bend 
Ranch SNA (La Saucedo, 3.5 mi. SE) (Pre- 
sidio Co.); vi-18/23-90; yellow pan trap; | 
2; J. Woolley/G. Zolnerowich. Rio Grande 
State Park (Singing Chaparral Trail); vi-19- 
86; 1 2; J. Woolley. 

Comments.—The separation of N. ther- 
esae from N. luiseno is based primarily on 
Ovipositor length, length of the last meta- 
somal tergum in females, and relative pro- 
portions of the male genital capsule. Details 
are given below. 

The single male from Patagonia, Arizo- 
na, is questionably placed here. Its larger 
size (HTL = 0.19 mm) suggests N. there- 
sae. The genital capsule is not positioned 
adequately for measurement. 


Nicolavespa luiseno Pinto, new species 
Geis, 28), ti, il, Ws) 


Description.—As in N. theresae except 
as follows: Female. Smaller, body length 
0.55 (0.5—0.7) mm (@ = 11); HTL = 0.19 
(0.18—0.21) mm (n = 9); metasoma length 
(excluding slight ovipositor extension) less 


than 1.50 that of metasoma; metasoma 
more abruptly narrowed posteriorly, not 
gradually narrowing and attenuate in apical 
half. Color within range of variation of WN. 
theresae (see Variation). Forewing fringe 
setae somewhat longer, longest 0.20 (0.16— 
0.23) (n = 7) wing width. Last tergum of 
metasoma (Fig. 11) considerably shorter, 
less than a third as long as wide, with cercal 
setae reaching apex of ovipositor. Oviposi- 
tor much shorter (Fig. 2), only occupying 
ca. half length of metasoma (in slide- 
mounted specimens ovipositor length is ca. 
half length of metasoma to apex of tergum 
ViITh): OLE — 6016S 1F7)s@e—" 8): 

Male. Body length 0.35 mm (n = 2). 
HTL = 0.14 (0.13—0.17) mm (n = 5). Gen- 
italia length 0.48 (0.4—0.5) (n = 5) that of 
hind tibia; genital capsule wider, 0.43 
(0.41—0.45) (n = 5) as wide as long (Fig. 
1S). 

Types.—Holotype &, allotype 5. UNIT- 
ED STATES. California: Riverside Coun- 
ty; Menifee Valley (hills on W side); 
33°39'N, 117°13'’W; 1,800’ el.; vi-14/28- 
1995; Malaise trap; John D. Pinto, collec- 
tor; deposited in the National Museum of 
Natural History, Washington DC. Six @ 
and 5 6 with same data as holo- and ailo- 
type designated as paratypes. Paratypes de- 
posited in The Natural History Museum, 
London; the Canadian National Collection, 
Ottawa; the University of California, Riv- 
erside (Department of Entomology); and 
the University of Naples, Portici. All types 
are on slides, mounted in Canada balsam. 
Mixed chaparral and coastal sage scrub 
vegetation characterizes the type locality. 

Etymology.—Luiseno, the aboriginal 
people originally inhabiting the known 
range of this species in Riverside Co. 

Geographic distributionKnown from 
three localities in southern California. 

Material examined.—20 6 (18 on 
slides), 97 @ (45 on slides). UNITED 
STATES. California: Granite Mountains 
Reserve, Granite Cove (4,200’) (San Ber- 
nardino Co.); 34°48’N, 115°39'W; v-14/17- 
1994; flight intercept trap; 1 @; J. Pinto/G. 


VOLUME 107, NUMBER 3 


Platner. Lake Skinner, NE of (ca. 1,570’) 
(Riverside Co.); 33°36'N, 117°02’W; iv-24/ 
v-8-97, v-7/21-96, v-8/22-97, v-21/vi-5/96, 
v-22/vi-5-97, vi-4/18-96, vi-16/30-98, vi- 
18/vii-2-96, vi-18/vii-2-98, vii-16/30-98, 
xi-6/20-97, xi-20/x1i-4-97; Malaise traps in 
coastal sage scrub; 5 6, 45 @; J. Pinto. 
Menifee Valley (hills on W side, 1,800’) 
(Riverside Co.); 33°39'N, 117°13’W; iv-30- 
96, vi-7/13-95, vi-14/28-95, vi-20/vii-12- 
95, vi-28/vii-12-95, vii-12-95, vii-19/viii-1- 
95, vili-1-95, x-18/x1i-2-95; Malaise traps 
in mixed chaparral/coastal sage scrub; 15 
3, 50 2; J. Pinto (see Types). 

Comments.—WNicolavespa luiseno is very 
similar to N. theresae. It is separated by its 
shorter ovipositor and less obviously atten- 
uate metasoma (cf. Figs. 1—2). In N. luiseno 
the ovipositor is considerably less than 
twice the length of the hind tibia. In N. ther- 
esae, its length is twice or more the hind 
tibial length. Also the length of metasomal 
tergum VIII is considerably greater in N. 
theresae than in N. luiseno and tergal shape 
differs as well (cf. Figs. 10—11). Males are 
difficult to separate. The only difference I 
have found is the greater width/length ratio 
of the genital capsule in N. luiseno (see 
above and cf. Figs. 14—15). 

Treating Nicolavespa luiseno as a species 
rather than a race of N. theresae was a dif- 
ficult decision. Arguing for species treat- 
ment is the occurrence of both forms in 
similar habitats (chaparral and coastal sage 
scrub) and in close proximity in southern 
California (within 15 km of each other), as 
well as the absence of character overlap 
throughout the range of both. Interestingly, 
material of N. theresae from southern Mex- 
ico and Central America most closely ap- 
proaches N. luiseno with regard to ovipos- 
itor length, body size and forewing fringe 
length. Unfortunately, relatively few speci- 
mens are available from this area, and little 
material has been collected from interme- 
diate regions of northern and central Mex- 
1cO. 

Nicolavespa can be relatively abundant 
in certain localities of southern California. 


635 


More extensive collecting is required to de- 
termine if the two species are syntopic. 


Haeckeliania Girault 


Haeckeliania Girault 1912: 97. Type spe- 
cies: Haeckeliania haeckeli Girault, by 
original designation. 


Diagnosis.—Female. Antenna (Fig. 18) 
with two anelli and a five-segmented club; 
club widest at Cl, Cl subequal in length to 
C2 or somewhat shorter but never anelli- 
form, with at least some PLS separated 
from surface for much of their length; club 
often terminating in an apical process. Max- 
illary palp two-segmented (Fig. 17). Fore- 
wing relatively broad (Fig. 26), length of 
venation usually less than half that of wing; 
MV gradually broadened to apex and end- 
ing abruptly at juncture with SV; SV well 
developed, shorter than MV; PM slightly 
shorter to, more commonly, slightly longer 
than MV, angled posteriorly from MV to 
form a well-developed costal cell. Forewing 
disk in almost all species with distinct lon- 
gitudinal setal tracks including an RSI. 
Midlobe of mesoscutum and scutellum each 
with two pair of setae (Fig. 24); notauli usu- 
ally broadly emarginate. Dorsum of first 
metasomal tergum (petiolar segment) tra- 
versed by a row of erect, sharply pointed 
denticles (Fig. 25). 

Male. As female except antenna with all 
PLS attached to surface at base only, seti- 
form (Fig. 22). Genitalia as in Trichogram- 
matini (sensu Viggiani 1971) (Figs. 29-31), 
with a distinct aedeagus, parameres and 
volsellae; aedeagus often longer than geni- 
tal capsule; genital capsule open its entire 
length on dorsal surface. 

Remarks.—Haeckeliania males were un- 
available to Viggiani (1971, 1984) for his 
classification of Trichogrammatidae based 
on male genitalia. The discovery of males 
(Pinto 1997) clearly places the genus in his 
Trichogrammatini (Trichogrammatinae). 
This is an assemblage of basal genera, 
probably paraphyletic, in which the aedea- 
gus and genital capsule are separate (Vig- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


giani 1971). Within this group Haeckeliania 
is phenetically most similar to Ophioneuris, 
Hispidophila and Brachyufens (see Doutt 
and Viggiani 1968 for separation; also see 
Pinto 1997 for separation from other North 
American genera). The presence of a trans- 
verse row of denticles on the petiolar seg- 
ment (Fig. 25), a feature not previously not- 
ed in the Trichogrammatidae, is not unique 
to Haeckeliania but characterizes several 
other Trichogrammatini genera (Tricho- 
gramma, Trichogrammatoidea, Paratricho- 
gramma, Australufens, Xenufens, Tricho- 
grammatomyia, Brachyufens, Brachyia, So- 
ikiella). The structure is not known to occur 
in the other tribes of the family. 

The form of antennal sexual dimorphism 
in Haeckeliania is uncommon in the Tri- 
chogrammatidae. Although basic structure 
is similar in both sexes (five club segments, 
funicle absent), the surface of the flagellar 
segments in males is dominated by elon- 
gate, setiform PLS (Fig. 22). In females 
there are fewer PLS and most are attached 
to the surface for much of their length (Fig. 
18). Similar surface ultrastructure of these 
sensilla in the two sexes indicates homol- 
ogy. In both, the surface is longitudinally 
grooved with numerous pores. In this way 
they are easily distinguished from the rel- 
atively smooth FS, the only other setiform 
sensilla of equivalent length on the anten- 
nae of Trichogrammatidae (Fig. 21). Nu- 
merous setiform PLS also occur on the an- 
tenna of Soikiella and Trichogrammatoidea 
males (Velten and Pinto 1990, Pinto 1999). 

Eight species of Haeckeliania have been 
described to date (Noyes 2001). This in- 
cludes four from Australia, three from Asia 
and only a single species, H. minuta Vig- 
giani, from the New World. This modest 
number fails to adequately portray the di- 
versity of the genus. The examination of 
collections of Trichogrammatidae from 
throughout the world suggests that Haecke- 
liania is one of the largest genera in the 
family. It is likely that no more than 5—10% 
of the fauna is described. It is particularly 
diverse and abundant in Australasia and the 


New World tropics. It also occurs in Africa 
and Asia. The genus has yet to be recorded 
from Europe, and I am aware of only two 
or three undescribed species in the United 
States. 

Most species of Haeckeliania are uni- 
formly dark brown in color, compact and 
gibbous in shape. In this section of the ge- 
nus the metasoma is broadly rounded api- 
cally and subequal in length to the meso- 
soma. Haeckeliania minuta, described from 
Brazil by Viggiani (1992), belongs to this 
section. Haeckeliania sperata, described 
below, represents a less common phenotype 
characterized by a lighter body color (gen- 
erally light yellow or yellow orange, and 
marked with brown), and a more slender, 
apically attenuate, and elongate body (me- 
tasoma clearly longer than mesosoma). H. 
sperata is the only species of this type for- 
mally recognized to date but additional un- 
described New World species have a similar 
facies. 

Haeckeliania sperata and a few other un- 
described New World species are known to 
parasitize the eggs of Curculionidae. 


Haeckeliania sperata Pinto, new species 
(Figs. 16—27, 29-31) 


Description.—Female. Body relatively 
narrow, elongate, length 0.5 (0.5—0.6) mm 
(n = 9); metasoma 1.4—1.5X length of me- 
sosoma, attenuate to apex. Color: Largely 
yellow orange with pronotum brown; me- 
tasomal terga II-IV predominantly brown, 
suffused with yellow, more apical terga 
brown along lateral margins only; legs and 
head beneath eyes light yellow; antenna 
with scape and pedicel light yellow, flagel- 
lum light yellow brown. Head: Toruli 
placed above ventral margin of eyes (Fig. 
16). Antenna with length/width ratio of 
scape, pedicel and club 2.3—1.2—3.6; scape 
moderately wide its entire length, ca. 1.6X 
length of pedicel; pedicel bead shaped, its 
surface weakly ridged transversely; club 
subconical, its length 1.3—1.4x that of 
scape and pedicel combined, C1 articula- 
tion point with anelli acentric; club widest 


VOLUME 107, NUMBER 3 637 


Figs. 16-23. Haeckeliania sperata. 16, Head. 17, Maxilla (venter). 18-23, Antenna. 18, 2 (lateral). 19, First 
anellar segment and base of club (@, lateral) (second anellar segment not visible in lateral view). 20, Club apex 
2 (arrow at apical process with setiform UPP sensillum inserted at apex, lateral). 21, Antennal sensilla (° ) 
(arrow ‘a’ at placoid sensillum [PLS]: surface grooved and pitted; arrow *b’ at flagelliform seta [FS]: surface 


appearing smooth). 22, d (medial). 23, Club apex ¢ (as in Fig. 20, except medial). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


at Cl and C2, gradually tapering to apex 
with an elongate apical process; Cl and C2 
subequal in length, C3—C5 increasingly 
elongate, C5 ca. 3X the length of Cl with 
apical process comprising ca. % its length. 
PLS characteristics and distribution on club 
as follows: Cl with one PLS curving from 
basal to dorsal surface, its apical third ex- 
tending beyond surface; C3 and C4 each 
with two PLS, one extending entire length 
of segment with apical third or more sepa- 
rated from surface, the other setiform, at- 
tached only at base; C5 with three-four 
PLS, one extending entire length of seg- 
ment with only apical fourth free of surface, 
the other two attached at base only and ex- 
tending well beyond tip of apical process. 
Other structures on antennal club include at 
least one large subglobose BPS at apex of 
each segment; one APB near base of Cl, 
near apex of C3 and just below apex of api- 
cal process; a few FS on C3-—C5; several 
APA on Cl1-—C3; and a setiform UPP in- 
serted at tip of apical process (this sensil- 
lum subequal in length to that of apical pro- 
cess itself). Mandible with two strong basal 
teeth and one or two weaker teeth apically. 
Maxillary palp with segment II ca. half 
length of I, apical sensillum slightly longer 
than II. 

Mesosoma: Midlobe of mesoscutum 
1.2X as long as scutellum; dorsum of me- 
sosoma weakly sculptured, surface divided 
into elongate reticulae, reticular spaces nar- 
rower on scutellum. Propodeal disk elevat- 
ed, slightly produced, arcuate posteriorly, 
slightly longer than metanotum. Forewing 
0.55X as wide as long, fringe setal length 
0.15—0.17 wing width, venation extend- 
ing almost to half length of wing; PM 
slightly shorter or subequal in length to MV, 
both veins of similar width, two PM cam- 
paniform sensilla well separated from one 
another; SV ca. 0.8 length of MV; disk with 
ca. 17 longitudinal setal tracks, most setae 
associated with a track; costal cell with 5— 
8 setae on ventral surface. 

Metasoma: Ovipositor extending only 
slightly beyond apex of metasoma; OL/ 


HTL = 2.1-—2.4 (n = 6) (HTL of measured 
specimens = 0.15—0.17 mm); gonoplac 
comprising ca. Y; length of ovipositor; gon- 
opophyses extending slightly anterior to 
gonangulae. 

Male. As in female except metasoma 
with minimal dark coloration (almost en- 
tirely orange) and broadly rounded apically, 
not attenuate; also, characteristics of anten- 
na differ as follows: club narrower and lon- 
ger, 4.8-5.2 (n = 2) as long as wide, ca. 
1.6X length of scape and pedicel combined, 
segments more loosely articulated, with a 
longer and more broadly based apical pro- 
cess comprising almost half length of C5 
(see below); PLS all modified into setiform 
structures, attached to surface at base only 
(4, 4, 3, 2, 3 in number on C1—C5, resp.), 
longest ca. 3X greatest club width; APB ab- 
sent from Cl and C2; surface without APA; 
FS on C4 only, 2 in number; apical process 
with a very long spinelike sensillum at apex 
(= UPP?) subequal in length to that of C5 
itself. 

Genital capsule elongate, basal section 
very poorly sclerotized and difficult to dis- 
cern in slide-mounted material; aedeagus 
slightly longer than capsule and 1.2 
length of hind tibia; apodemes comprising 
ca. 0.4 total aedeagal length; parameres and 
volsellae extending to same level at apex of 
capsule; parameres bifid apically; intervol- 
sellar process absent. 

Types.—Holotype 2, and allotype 6. 
DOMINICA. Grand Bay, Agricultural Sta- 
tion; 1-6-2004; ex. Diaprepes eggs on Cit- 
rus; N. Commodore; deposited in the Na- 
tional Museum of Natural History, Wash- 
ington, DC. Paratypes include 6 6 and 27 
2, data same as holo- and allotype; depos- 
ited in the Canadian National Collection, 
Ottawa; The Museum of Natural History, 
London; the Florida State Collection of Ar- 
thropods, Gainesville; and the University of 
California, Riverside (Department of Ento- 
mology). All types except 1 6 and 6 2 
paratypes are mounted on slides in Canada 
balsam. All type material emerged from the 
same host egg mass. 


VOLUME 107, NUMBER 3 639 


Figs. 24-31. Haeckeliania. 24—27, H. sperata. 24, Mesosoma. 25, Base of metasoma (arrow at row of 
denticles on petiolar segment). 26, Forewing. 27, Venation of forewing (dorsal) with arrows at disjunct cam- 
paniform sensilla at apex of premarginal vein. 28, Haeckeliania sp., as in Fig. 27, except the two campaniform 
sensilla at apex of premarginal vein adjacent (see arrow). 29-30, H. sperata, d genitalia. 29, Dorsal (arrow ‘a’ 
to aedeagal apodeme at base, arrow ‘b’ to apex of aedeagus. 30, Venter. 31, Detail of apex (ventral), with arrow 
to apically bifid paramere (note volsellar digital spines beneath [dorsal to] parameres). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Etymology.—Sperata (Latin: hoped for). 

Geographic distribution.—Known only 
from Dominica and Costa Rica. Targeted 
for possible release in Florida (Jorge Pena, 
personal communication). 

Material examined.—11 6 (10 on 
slides), 43 2 (32 on slides). COSTA RICA. 
Allajuela: PN Arenal, La Peninsula (600 
m); 10°27'N, 84°45'W; 11-25-2003; sweep; 
1 2; J. Noyes. DOMINICA. Grand Bay, 
Agricultural Station (see types); 7 5, 28 @. 
Grand Bay; iv-20 & iv-28-2003; ex Dia- 
prepes sp. eggs on Citrus sp.; 1 d, 10 9; 
J. Pefia/C. McCoy/R. Hill. “Syndicate & 
Grand Bay’’; iv-29-2003; ex. Diaprepes sp. 
or Pachnaeus eggs on Citrus sp. 3 6, 4 2; 
JaPenaetrale 

Comments.—As already summarized, 
Haeckeliania sperata is distinguished from 
the vast majority of congeners by the lighter 
body color, and its relatively slender body. 
These features also distinguish it from H. 
minuta, the only other member of the genus 
described from the New World (Viggiani 
1992). Whereas most Haeckeliania are en- 
tirely brown and robust, in H. sperata the 
mesosoma is predominantly yellow orange, 
as is much of the metasoma, and its body 
shape is relatively slender. In addition, in 
the majority of Haeckeliania the marginal 
vein is somewhat shorter and wider than the 
premarginal vein. In H. sperata the two 
veins are similar in length and width. 

A small number of undescribed New 
World species have a facies similar to H. 
sperata. The following combination of 
characters will separate H. sperata from 
these: Forewing with the two campaniform 
sensilla on PM distinctly separated from 
one another (Fig. 27); metanotum slightly 
shorter than propodeal disk (Fig. 24); fe- 
male antennal club with a setiform apical 
sensillum (UPP) (Fig. 20); ovipositor length 
between 2 and 2.5 as long as hind tibia; 
male antennal club with an apical process 
(Fig. 23). In one presumably close relative 
of H. sperata from Central America the 
campaniform sensilla are similarly disjunct 
but the sensillum inserted at the apex of the 


apical process in females is much more ro- 
bust, cylindrical in shape, and truncate at 
its apex. Another Central American and 
Carribean species has an extremely long 
ovipositor (ca. 3.5 as long as its hind tib- 
1a). 

Two females from Florida (Gainesville 
and Covington) and several from Chiapas 
appear to represent a species closest to H. 
sperata. The primary differences are color 
(see below), the contiguous campaniform 
sensilla on the PM (as in Fig. 28), a longer 
metanotum (1.5—-2x length of propodeal 
disk), and an apically truncate rather than 
attenuate apical sensillum on the club. 
Clearly associated males are unknown. 

All similar species are separable from H. 
sperata by color. In H. sperata the light- 
colored areas are deep yellow orange; in re- 
lated species they are light yellow. Also, the 
dorsum of the metasoma in females of H. 
sperata is brown basally but suffused with 
yellow orange, and the apical terga are pre- 
dominantly yellow orange with brown lat- 
eral markings. In similar forms all except 
the apical two metasomal terga are uni- 
formly brown and the apical ones are en- 
tirely light yellow. 


ACKNOWLEDGMENTS 


Jorge Pena and Rita Duncan provided all 
Dominican specimens of Haeckeliania sper- 
ata. Gary Platner provided technical assis- 
tance. This study was supported primarily by 
a grant from the USDA (NRIICGP) (2001- 
35316-11012; J. Pinto, PI) and an NSF 
PEET grant (BSR-9978150; J. Heraty, PI). 


LITERATURE CITED 


Doutt, R. L. and G. Viggiani. 1968. The classification 
of the Trichogrammatidae (Hymenoptera: Chalci- 
doidea). Proceedings of the California Academy 
of Sciences (4" series) 35: 477-586. 

Girault, A. A. 1912. Australian Hymenoptera Chalci- 
doidea—The family Trichogrammatidae with de- 
scriptions of new genera and species. Memoirs of 
the Queensland Museum 1: 66-116. 

Hall, D., J. Pena, R. Franqui, R. Nguyen, P. Stansly, 
C. McCoy, S. Lapointe, R. Adair, and B. Bullock. 
2001. Status of biological control by egg parasit- 
oids of Diaprepes abbreviatus (Coleoptera: Cur- 


VOLUME 107, NUMBER 3 


culionidae) in citrus in Florida and Puerto Rico. 
BioControl 46: 61—70. 

Heraty, J. and D. Hawks. 1998. Hexamethyldisila- 
zane—A chemical alternative for drying insects. 
Entomological News 109: 369-374. 

Noyes, J. S. 2001. Interactive catalogue of world Chal- 
cidoidea. CD-ROM. The Natural History Muse- 
um. 

Olson, D. M. and D. A. Andow. 1993. Antennal sen- 
silla of female Trichogramma nubilale (Ertle and 
Davis) [sic] (Hymenoptera: Trichogrammatidae) 
and comparisons with other parasitic Hymenop- 
tera. International Journal of Insect Morphology 
and Embryology 22: 507-520. 

Pinto, J. D. 1997. Trichogrammatidae. Chap. 22, pp. 
726-752. In Gibson, G. A. P., J. T. Huber, and J. 
B. Woolley, eds. Annotated Keys to the Genera of 
Nearctic Chalcidoidea (Hymenoptera), NRC Re- 
search Press, Ottawa, Canada, 794 pp. 

. 1999 (1998). The systematics of the North 

American species of Trichogramma (Hymenop- 

tera: Trichogrammatidae). Memoirs of the Ento- 

mological Society of Washington, No. 22, 287 pp. 

. 2004. A review of the genus Doirania (Hy- 

menoptera: Trichogrammatidae) with a description 


641 


of a new species from North America. Proceed- 
ings of the Entomological Society of Washington 
106: 352-360. 

Pinto, J. D. and J. George. 2004. Kyuwia, a new genus 
of Trichogrammatidae (Hymenoptera) from Afri- 
ca. Proceedings of the Entomological Society of 
Washington 106: 531-539. 

Velten, R. K. and J. D. Pinto. 1990. Soikiella Nowicki 
(Hymenoptera: Trichogrammatidae): Occurrence 
in North America, description of a new species, 
and association of the male. The Pan-Pacific En- 
tomologist 66: 246-250. 

Viggiani, G. 1971. Ricerche sugli Hymenoptera Chal- 
cidoidea XXVIII. Studio morfologico comparati- 
vo dell’armatura genitale esterna maschile dei Tri- 
chogrammatidae. Bollettino del Laboratorio di 
Entomologia Agraria ‘Filippo Silvestri’ di Portici 
29: 181-222. 

. 1984. Further contribution to the knowledge 

of the male genitalia in the Trichogrammatidae 

(Hym. Chalcidoidea). Bollettino del Laboratorio 

di Entomologia Agraria ‘Filippo Silvestri’ di Por- 

tici 41: 173-182. 

1992. New species of Trichogrammatidae 

(Hymenoptera: Chalcidoidea) from South Ameri- 

ca. Redia 75: 253-265. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 642-651 


PREDATORY BEHAVIOR OF REPIPTA FLAVICANS STAL 
(HEMIPTERA: REDUVIIDAE), A NATURAL ENEMY OF DIABROTICINA 
(COLEOPTERA: CHRYSOMELIDAE) 


SAGRARIO GAMEZ-VIRUES AND ASTRID EBEN 


Departamento de Ecologia Funcional, Instituto de Ecologia, A. C., Km 2.5 Antigua 
Carretera a Coatepec No. 351, 91070. Xalapa, Veracruz, México (e-mail: sagrariogamez@ 
hotmail.com) 


Abstract.—Predation bioassays were conducted to evaluate the predatory behavior of 
Repipta flavicans Stal (Hemiptera) under laboratory conditions. In the field, this predator 
was identified as a natural enemy of the Diabroticina beetle, Acalymma blomorum Munroe 
and Smith (Coleoptera: Chrysomelidae). Predator specificity, functional response, and the 
potential tritrophic effect of the secondary compounds (cucurbitacins) sequestered by prey 
were determined. An effect of the elytra coloration of the prey was not observed, but in 
some cases prey body size seemed to have had an effect on predation. The results of the 
tritrophic level assay suggested that cucurbitacins sequestered by A. blomorum did not 
have a repellent or toxic effect on this predator. By contrast, plant secondary compounds 
present in prey increased the longevity of R. flavicans females. Cucurbitacins were present 
in the predators after ingesting A. blomorum. 


Resumen.—Se realizaron bioensayos de depredacién para evaluar el comportamiento 
predatorio de Repipta flavicans Stal (Hemiptera: Reduviidae) bajo condiciones de labor- 
atorio. En campo, este depredador fue identificado como enemigo natural de la Diabro- 
ticina Acalymma blomorum Munroe and Smith (Coleoptera: Chrysomelidae). Ademas, se 
determin6 su especificidad, respuesta funcional y el efecto tritr6fico de los compuestos 
secundarios (cucurbitacinas) obtenidos por las presas. No se observ6 que la coloraci6n de 
los élitros, de los escarabajos utilizados, tuviera un efecto sobre el depredador, sin em- 
bargo, en algunos casos el tamafio corporal parece haber afectado la elecci6n del mismo. 
Los resultados sugieren que las cucurbitacinas retenidas por A. blomorum no tuvieron un 
efecto repelente 0 t6xico en el depredador. Por el contrario, la presencia de los compuestos 
secundarios en las presas incrementaron la longevidad de las hembras R. flavicans y 
estuvieron presentes en los depredadores después de que ingirieron presas A. blomorum. 


Key Words: Acalymma blomorum, tritrophic effect, cucurbitacins, Diabroticina, preda- 


tion, Repipta flavicans 


Harpocaptorine species (Hemiptera: Red- 
uviidae) are common predators on other in- 
sects throughout the Neotropical Region 
(Schuh and Slater 1995). Repipta flavicans 
Stal is distributed in the central states of 
Mexico. In a previous paper, it was reported 


for the first time as a predator of Acalymma 
blomorum Munroe and Smith in its natural 
habitat (Gamez-Virués et al. 2003). This 
prey species belongs to the Diabroticina 
beetles (Chrysomelidae) which are native to 
Mexico and Central America (Webster 


VOLUME 107, NUMBER 3 


1895). Their main economically important 
hosts are maize, beans, and cucurbits. For 
that reason, several species of Acalymma 
are considered important pests (Burkness 
and Hutchison 1998), and numerous studies 
have been conducted on these pests (Brust 
and Foster 1995, Kuhlman and Van der 
Burgt 1998, Lance 1988). 

Nevertheless, little information exists 
about the natural enemies of Diabroticina in 
their native habitat. The reduviid Castolus 
tricolor Champion was recognized as a 
predator of Diabroticina in Costa Rica 
(Risch 1981). Eben and Barbercheck (1996) 
reported the spider Oxyopes salticus Hentz 
(Oxyopidae) and three species of Reduvi- 
idae as predators of A. blomorum and Dia- 
brotica balteata LeConte in Veracruz, Mex- 
ico. Studies on the use of biocontrol agents 
against Diabroticina species were per- 
formed with entomopathogenic nematodes 
(Jackson 1996), entomopathogens fungi 
(Tallamy et al. 1998), and recently with 
flies of the family Tachinidae (Zhang et al. 
2004). Furthermore, Howe et al. (1976) re- 
ported for the first time that ingestion of 
cucurbitacins by Diabroticina can be related 
to protection against birds and other insec- 
tivorous vertebrates. The hypothesis of 
chemical defense has since been studied 
with a diverse array of natural enemies, 
however, no clear pattern has been detected 
(Ferguson and Metcalf 1985, Nishida ei al. 
1992, Brust and Barbercheck 1992, Barber- 
check et al. 1995). 

The objective of this study was to char- 
acterize the predatory behavior and speci- 
ficity of R. flavicans on Diabroticina bee- 
tles. The effect of cucurbitacins present in 
the diet of the beetle prey was also evalu- 
ated for this predatory bug. 


MATERIALS AND METHODS 


Predators.—Repipta flavicans adults 
were collected in the field where they were 
observed preying on A. blomorum. They 
were found in particularly high numbers on 
two cucurbit species: Cucurbita okeecho- 
beensis ssp. martinezii L. Bailey, a wild bit- 


643 


ter cucurbit, and C. moschata (Lam.) Poiret, 
an edible cucurbit. Predatory bugs were 
kept under laboratory conditions (25 + 3°C; 
13:11, L:D). They were provided daily with 
adults of D. balteata, D. porracea Harold 
and larvae of Anastrepha ludens (Loew) 
(Diptera: Tephritidae) as food. A laboratory 
colony was initiated with the eggs depos- 
ited by the field-captured bugs, using meth- 
ods described by Gdmez-Virués et al. 
(2003). Nymphs used later in bioassays 
were kept away from Diabroticina species 
and were fed only with A. ludens larvae. 

Diabroticina prey.—For all bioassays, 
adults Diabroticina beetles were used as 
prey. Beetles were collected on the same 
cucurbits than predators. Some beetles were 
fed with fruits of Cucurbita pepo L., a cul- 
tivar free of cucurbitacins, and artificial diet 
(Branson et al. 1975) during 28 d. This al- 
lowed excretion of previously sequestered 
bitter compounds (treatment | = without 
cucurbitacins, SC). Other beetles were fed 
with bitter fruits of C. 0. martinezii, that are 
rich in cucurbitacins, for a period of 7 to 
10 d (Andersen et al. 1988) (treatment 2 = 
with cucurbitacins, CC). In addition, five 
other Diabroticina species collected in the 
field on the same hosts were used in bio- 
assays: A. innubum (Fabricius), A. trivitta- 
tum Mannerheim, Cerotoma atrofasciata 
Jacoby, D. balteata, and D. tibialis Baly. 
Beetles were kept on the SC diet for at least 
28 d before they were used in bioassays. 
The six Diabroticina species were separated 
by species and kept in cages of 30 cm xX 
30 cm X 30 cm, under laboratory condi- 
tions. 

Functional response of R. flavicans.— 
Predation of R. flavicans was evaluated ac- 
cording to prey density. Naive females were 
used as predators. Each female was offered 
1, 2, 5, 10, 20, or 30 individuals of A. blo- 
morum as prey. Predation was evaluated af- 
ter | h, 8 h, and 24 h. Transparent plastic 
containers (5 1) with a twigs to simulate a 
natural environment were used in the assay. 
Twelve repetitions were made for each prey 
density. Results were analyzed by one-way 


644 


ANOVA (P < 0.05) with SigmaStat™ sta- 
tistical software version 2.0 (Jandel Scien- 
tific 1992-1997). 

Predation bioassays.—Twenty-four hours 
before each bioassay, bugs were placed in- 
dividually in 250 ml cages and were de- 
prived of food. Bioassays concluded when 
bugs attacked a beetle or after 48 h. As a 
control, five A. blomorum fed on SC diet 
and five A. blomorum fed on CC diet were 
placed in plastic containers (250 ml) and 
their mortality without the presence of a 
predator was recorded after 48 h. 

1) Specificity. This was evaluated by ob- 
serving whether the predatory behavior of 
R. flavicans was affected by coloration of 
the elytra or the body size of its prey. Six 
Diabroticina species, that coexist as adults 
on the same Cucurbita spp., were used. 

Coloration: Two Diabroticina species 
were offered to each predator in three com- 
binations: one A. blomorum and another 
species with a) similar coloration: A. trivit- 
tatum, or b) different coloration: C. atro- 
fasciata or D. balteata. 

Body size: Two beetles were offered to 
each predator in choice tests: one A. innub- 
um, with similar coloration that A. blomo- 
rum, and one D. tibialis, both species of 
greater size than A. blomorum. For these 
assays, ten female and ten male R. flavicans 
from the laboratory colonies were used. 
The results were analyzed with a chi-square 
test (P<0.05, SigmaStat). 

2) Effect of the diet of A. blomorum on 
predation by R. flavicans. Two A. blomo- 
rum, one of each diet treatment, were of- 
fered to each predator in a choice situation. 
The bioassay was finished when the pred- 
ator attacked one of the beetles or after a 
period of 48 h. Twenty females collected in 
the field, and 10 females and 10 males ob- 
tained in laboratory were used. The results 
were analyzed with a t-test for independent 
samples (P < 0.05, SigmaStat). 

3) Effect of the diet of A. blomorum on 
the longevity of R. flavicans. Each predator 
was offered two A. blomorum fed on SC 
diet or two A. blomorum fed on CC diet 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


every 48 h, during their entire adult life- 
time. Twenty-four hours after providing the 
prey, the number of predated beetles and 
the survival of each bug were recorded. 
Only R. flavicans females raised under lab- 
oratory conditions were used. Nine repeti- 
tions per treatment were made. The results 
were analyzed with a t-test for independent 
samples (P<0.05, SigmaStat). 

Qualitative detection of cucurbitacins in 
A. blomorum and R. flavicans.—A variation 
of the method established by Halaweish and 
Tallamy was used (1993). Cucurbitacins 
were detected with thin layer chromatog- 
raphy (TLC). The samples were obtained 
by grinding 15 A. blomorum beetles of each 
treatment in 3 ml ethanol (70%). Each ex- 
tract was filtered and the solid phase was 
discharged. To the extract, 2 ml hexane 
were applied, it was shaken slowly in a se- 
paratory funnel, and the hexane phase was 
discharged (2X). Afterwards, 2 ml chloro- 
form were applied, and the ethanolic phase 
was discharged. Finally, the extract was 
evaporated until the solvent was eliminated 
and a yellowish residue was obtained. In 
order to determine if the bugs acquired the 
plant compounds sequestered by their prey, 
both females that were fed during their 
adult life exclusively with A. blomorum fed 
on SC diet and females fed with A. blo- 
morum fed on CC diet were used (bioassay 
3). All dead Repipta females per treatment 
were stored individually in ethanol (70%). 
Extracts were obtained with the same meth- 
od as above described for A. blomorum. 
The samples of A. blomorum and R. flavi- 
cans were applied on silica gel 60 TLC 
plates (without fluorescent indicator, 
MERCK). Plates were developed with ethyl 
acetate: toluene (6:4). Plates were observed 
under UV light to 254 nm. The retention 
factor (Rf) of each visible compound was 
determined and compared with standard 
values for cucurbitacin B, D, and I. 


RESULTS 


Functional response of R. flavicans.— 
Predation was proportional to prey density 


VOLUME 107, NUMBER 3 


645 


20 30 


Y 3.0 
{5°} 

= 

ac) 
S25 
Ss 

Q 
10 
Re 

: 

Dd) 

Qo. 

wey 1S 
(cD) 

wd 

s 
SO 
N 

a) 

— 

(D) 

oS 

= 015 
S) 

fees 

(Dd) 

a 2) 

E 0.0 
F 1) 2S 10 

Number of beetles offered 

Fig. 1. 


Acalymma blomorum, under laboratory conditions. 


(Fig. 1), but eventually the rate of con- 
sumption remains constant regardless of in- 
creases in prey density. Predators consumed 
a maximum of 2.4 + 0.86 (SE) prey in 24 
h. A functional response curve of type II 
(Holling 1959) was observed. 

Predation bioassays.—An immediate ef- 
fect of cucurbitacins on the predators was 
not observed. Predators did not reject A. 
blomorum fed on CC diet. No mortality was 
observed in the control beetles. Females 
and males responded in a similar pattern to 
the prey species offered. However, signifi- 
cant preferences were obtained only for fe- 
males, for the pair, A. innubum vs. D. tibi- 
alis, males did not prey on D. tibialis (Fig. 
Dy. 

1) Specificity. Coloration: When com- 
paring predation on A. blomorum and A. tri- 
vittatum no significant difference was de- 
tected. Nevertheless, greater predation was 


Functional response of Repipta flavicans females in 1 h, 8 h and 24 h, to different densities of 


observed on A. blomorum than on C. atro- 


fasciata (P = 0.005). Finally, when A. blo- 


morum and D. balteata were offered, pred- 
ators preferred D. balteata (P = 0.025, Fig. 
2). No significant difference in predatory 
behavior between female and male bugs 
was detected. 

Body size: No preference for one of the 
larger species, A. innubum and D. tibialis 
was found (Fig. 2). Also, no difference in 
numbers of beetles preying upon the small- 
er or the larger species was found. Never- 
theless, female R. flavicans preyed upon 
significantly greater number of beetles than 
did males (P = 0.025). 

2) Effect of the diet of A. blomorum on 
predation by R. flavicans. No difference be- 
tween treatments was found. Neither be- 
tween predators collected in the field or 
raised in the laboratory, nor between fe- 
males and males. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


3) Effect of the diet of ‘A. blomorum on 
the longevity of R. flavicans. The longevity 
recorded for female predators that ate A. 
blomorum fed on the SC diet was 66 + 12. 
8 d, whereas individuals which preyed on 
A. blomorum fed on the CC diet was 103 
+ 12.3 d. The difference between treat- 
ments was significant (P < 0.05). Also, true 
bugs consumed more individuals of A. blo- 
morum fed on the CC diet (83%) than of 
A. blomorum fed on the SC diet (75%, P = 
0.0044). 

Quantitative detection of cucurbitacins in 
A. blomorum and R. flavicans.—In chro- 
matograms obtained from A. blomorum fed 
on the SC diet two spots were detected, one 
with Rf = 0.45 and a second with Rf = 
0.14. In extracts from A. blomorum fed on 
the CC diet one spot with Rf value of 0.53 
was obtained. From the extracts of R. flav- 
icans fed with A. blomorum fed on the CC 
diet we observed one spots with Rf = 0.53. 
As standards we obtained for cucurbitacin 
B: Rf = 0.72, for cucurbitacin D: Rf = 
0.41, and for cucurbitacin I: Rf = 0.55. 


DISCUSSION 


In our study Repipta flavicans responded 
positively to prey density, but quickly be- 
came satiated, and was not specialized on 
Acalymma spp. Even though R. flavicans 
responded positively to prey density, its 
predatory behavior did not seem to be den- 
sity dependent. An explication for this ob- 
servation might be that the probability of 
success in the first attack is related to prey 
density. However, after the first prey is con- 
sumed, this predator seemed to be some- 
what saturated and its subsequent prey cap- 
tures were slower (Holling 1959). In addi- 
tion, due to the long handling time neces- 
sary for capture and extraoral digestion, the 
average number of prey consumed was only 
2.4 beetles in our 24 h assays, as well as in 
the 48 h longevity assays. Laboratory ex- 
periments by O’Neil (1997) showed, that 
the functional response of the Podisus ma- 
culiventris (Heteroptera: Pentatomidae) in- 
creased according to the density of Leptin- 


otarsa decemlineata (Say) (Coleoptera: 
Chrysomelidae). This study, however, was 
done with the nearly immobile larvae of 
that species, and like in our study, handling 
time, rather than prey availability, limited 
the number of prey items that a predator 
can consume. 

In the field, individuals of R. flavicans 
were observed to prey only on A. blomo- 
rum, although several Diabroticina species 
coexisted on the same hosts. 

Under laboratory conditions, when com- 
paring predation on A. blomorum and A. tri- 
vittatum, no significant difference was 
found between prey species, perhaps due to 
the similarity in coloration of their elytra 
and in body size. Nevertheless, R. flavicans 
females preferred A. blomorum over C. 
atrofasciata, and both, females and males 
preferred D. balteata. Therefore, the pref- 
erence of D. balteata over A. blomorum ob- 
served in our bioassays was surprising. A 
factor that could have affected predation on 
these two species, was their activity level. 
According to observations in the laborato- 
ry, individuals of D. balteata were more ac- 
tive than A. blomorum. Once the bioassay 
was initiated, D. balteata beetles were con- 
stantly moving, which probably attracted 
the attention of the predator, since it walked 
behind the prey or waited until the beetle 
came within reach. By contrast, A. blomo- 
rum adults stayed motionless when they 
discovered the presence of the predatory 
bugs (Gamez-Virués et al. 2003). 

When A. innubum and D. tibialis (both 
species are larger than A. blomorum) were 
offered, females preyed upon more beetles 
than did males. Also, males did not suc- 
cessfully attack D. tibialis. Female R. flav- 
icans are larger than males (Gamez-Virués 
et al. 2003) and may have increased nutri- 
tional requirements due to egg production. 
The resulting costs might reflect the greater 
voracity of female bugs. Nevertheless, 
males tried to capture individuals of D. tib- 
ialis, but due to their size and perhaps also 
their weight, beetles managed to escape be- 
fore the bug could introduce its stylets into 


VOLUME 107, NUMBER 3 647 


HO sas: 'ner Onder Dane Orncnor stAstov Oe trSqe aliO 


U-0 


Fig. 2. Number beetles of Acalymma blomorum (Ab), A. innubum (Ai), A. trivittatum (At), Cerotoma atro- 
fasciata (Ca), Diabrotica balteata (Db), and D. tibialis (Dt), predated by female and male Repipta flavicans in 
a choice bioassays. Significant difference (P < 0.05) between Diabroticina species are indicated with an aster- 
isk (*). 


the intersegmental membranes of the prey’s = morum have distinct longitudinal ridges and 
abdomen. In addition, D. tibialis and C. a reticular pattern with punctures. Those 
atrofasciata beetles have shiny, wax-cov- morphological characteristics might allow 
ered elytra, whereas the elytra of A. blo- Repipta bugs to capture them with greater 


648 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


facility. Further experiments including bee- 
tles from other families are necessary to de- 
termine if this predator is truly a generalist. 

Cucurbitacins are evidently aimed to re- 
pel vertebrate herbivores (DaCosta and 
Jones 1971). Some invertebrates like naive 
preying mantids rejected Diabroticina bee- 
tles that had fed during seven days on 
squash fruit with high concentrations of cu- 
curbitacins (Ferguson and Metcalf 1985). 
Contrary to the chemical defense hypothe- 
sis, the presence of cucurbitacins in the bee- 
tles of our study did not affect the predatory 
behavior of R. flavicans. There are several 
studies that did not find negative effects of 
secondary plant compounds on species of 
the third trophic level (Barbercheck 1993, 
Down et al. 2003, Karimzadeh et al. 2004). 
On the other hand, Mitchell et al. (2004) 
observed that oviposition behavior of a par- 
asitoid wasp was significantly slower due to 
secondary compounds in host eggs, but 
time for drilling, oviposition, and marking 
was unaffected as well as progeny emer- 
gence, longevity, or sex ratio. 

Apparently R. flavicans benefited from 
the cucurbitacins in its prey, since its lon- 
gevity was enhanced. It might be possible, 
that this predatory bug uses cucurbitacins 
sequestered from prey beetles for its own 
chemical defense. It is well known that har- 
pactorine species emit volatile secretions 
with pungent scents as chemical defense 
(Ambrose 1999). In addition, numerous 
predacious Hemiptera have been observed 
to complement their diet with plant mate- 
rial, which can accelerate nymphal devel- 
opment, and increase longevity and fecun- 
dity (Ambrose 1999). Our observations 
agree with the data reported by Ambrose, 
since the longevity of R. flavicans was 
greater in individuals that ate A. blomorum 
fed on the CC diet. Furthermore, in the lab- 
oratory, we observed R. flavicans eating 
fruits of both cucurbit hosts. In a previous 
paper we described its laboratory biology 
and established a small scale rearing meth- 
od (Gamez-Virués et al. 2003). If R. flavi- 
cans obtain a direct effect from cucurbita- 


cins, we would expect to collect greater 
number of R. flavicans on C. o. martinezit 
(bitter cucurbit), but this does not agree 
with our field observations. We collected 
0.54 individuals of R. flavicans per hour on 
C. moschata against 0.01 individuals on C. 
o. martinezii (unpubl. data). In addition, 
numbers of beetles collected on C. moscha- 
ta were significantly greater than on C. o. 
martinezii (Gamez-Virués and Eben, 2005). 
Perhaps R. flavicans reacted more the abun- 
dance of its prey than to the non volatile 
secondary compounds that characterize 
many wild cucurbits. Moreover, perhaps 
beetles fed on CC diet adquire more nutri- 
ents than those fed on SC diet due to their 
compulsive behavior to cucurbitacins (Met- 
calf 1986). As consequence, R. flavicans 
reared on A. blomorum fed on CC diet ob- 
tained dietary benefits that improved its 
longevity. This hypothesis remains to be 
further investigated. 

To our knowledge, no studies have been 
published that demonstrate a direct benefi- 
cial effect of non-volatile secondary com- 
pounds sequestered by herbivores on pred- 
ators or parasitoids. Spiteller et al. (2000) 
found that beet armyworm oral secretions 
contain volicitin, a compound partly pro- 
duced by the insect, partly metabolized 
from plant sequestered linolenic acid. Upon 
wounding, volicitin elicits volatile secretion 
in maize. Gentry and Dyer (2002) reported 
that caterpillars containing unpalatable 
chemicals were preferentially parasitezed. 
Those compounds then attract natural ene- 
mies of the herbivore. 

According to our field observations, we 
know that R. flavicans are able to eat A. 
blomorum beetles found on wild, cucurbi- 
tacin-producing Cucurbita species as well 
as on non-bitter cultivars. Possibly as a re- 
sult of the longterm coexistence of these 
species in their native habitat, this predator 
has adapted to the plant secondary com- 
pounds as well. This hypothesis might be 
further supported by our findings that these 
true bugs sequester cucurbitacins from their 
prey. More experiments are required to un- 


VOLUME 107, NUMBER 3 


derstand potential tritrophic level effects in 
the described interaction. 

This study is the first that used a preda- 
tory hemipteran to test the chemical defense 
hypothesis in Diabroticina. Reduviidae 
have sucking mouth parts and use extraoral 
digestion. Perhaps when using invertebrate 
predators with a buccal apparatus different 
from true bugs, the rejection of A. blomo- 
rum with sequestered cucurbitacins would 
be evident, as has been reported in earlier 
studies for other Diabroticina species (Fer- 
guson and Metcalf 1985, Tallamy et al. 
1997, Tallamy et al. 1998). Ferguson and 
Metcalf (1985) reported that 100 % of Aca- 
lymma vittatum Fabricius collected in the 
field had extractable concentrations of cu- 
curbitacins in their body. In Mexico, where 
numerous wild bitter cucucrbit species can 
be found, the percentage of Acalymma spp. 
that retain cucurbitacins under natural con- 
ditions is not known. We are currently gath- 
ering these data. 

The compounds detected in the whole- 
body extracts of both beetle treatments 
and of R. flavicans chromatographically 
matched those of the cucurbitacins detected 
in the fruit extracts (Rf = 0.53, cucurbitacin 
I). It has been reported (Metcalf et al. 1982) 
that C. 0. martinezii produces cucurbitacin 
E and I. The cucurbitacins with Rf value of 
0.45 might have been another cucurbitacin: 
J or K glycoside reported for a closely re- 
lated subspecies, C. okeechobeensis (Met- 
calf et al. 1982) or metabolites of the com- 
pound (Halaweish and Tallamy 1993), but 
standards were not available for confirma- 
tion. We are currently isolating the cucur- 
bitacin profile of C. 0. martinezii (unpubl. 
data). The chromatograms obtained from 
the whole body extracts of A. blomorum 
reared on the SC diet show that beetles 
maintain sequestered cucurbitacins longer 
than 28 d. Therefore, we were not able to 
compare between beetles with and without 
cucurbitacins, but rather made a compari- 
son between beetles with different concen- 
trations of sequestered compounds. In an 
earlier study, it was reported that A. vitta- 


649 


tum excreted 67% of sequestered cucurbi- 
tacins after 48 h (Ferguson et al. 1985, An- 
dersen et al.1988). No further experimental 
data have yet been published on the length 
of time cucurbitacins can be stored in the 
body of Diabroticina beetles. 

Our results suggest that due to the slow 
predation and quick satiation, R. flavicans 
can probably not be used as_ biocontrol 
agent against Diabroticina in the field, but 
maybe in a greenhouse situation. Neverthe- 
less, it might be important to conserve this 
predator for a multi-species approach to in- 
tegrated pest management. 


ACKNOWLEDGMENTS 


We thank L. Cervantes P. and E. Barrera 
for the identification of the Repipta flavi- 
cans and their suggestions for maintaining 
the laboratory colony. We acknowledge the 
help of E. Juarez and A. Arcos with the 
chromatography. A. Martinez and C. Her- 
nandez-Osorio help with the statistical anal- 
ysis. L. A. Dyer, C. Paris, and the two an- 
onimous reviewers for their helpful sugges- 
tions. This research was funded by CON- 
ACYT project No. 35501 V to A- Eben, 
and partially with the scholarship CONA- 
CYT 162421 to S. Gamez-Virués. 


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5-16. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 652—656 


A NEW SPECIES OF PORRICONDYLA (DIPTERA: CECIDOMYITIDAE) FROM 
SOUTHWESTERN PENNSYLVANIA 


JOHN D. PLAKIDAS 


111 Emerson Avenue, Pittsburgh, PA 15215, U.S.A. (e-mail: johnplakidas@comcast. 
net) 


Abstract.—A species new to science, Porricondyla recondita Plakidas (Diptera: Ceci- 
domyiidae: Porricondylinae), is described and illustrated. The larva of this new species 
is noteworthy in that it lacks a spatula but has an analogous structure on the dorsum of 
the prothorax, a prothoracic collar, possibly used for tunneling through substrate. The 


prothoracic collar is unique among Porricondylinae. 


Key Words: 


Porricondylinae are a subfamily of gall 
midges whose larvae feed exclusively on 
fungal mycelium (Mamaev and Krivoshei- 
na 1965, Plakidas 1999). The decay habitats 
in which they live include soil, leaf litter, 
fallen trees, aborted flower buds, shelf fungi 
and any other long-lived decaying sub- 
strates where fungi can proliferate. Recent 
publications (Spungis 1981, 1985, 1987, 
1989, 1992, 2002; Dallai et al. 1996; Gagné 
2004) are useful in understanding the biol- 
ogy and systematics of the Porricondylinae. 

Larvae of the new species, Porricondyla 
recondita Plakidas, were found feeding in 
brown-rot decay beneath loose bark of a 
fallen red maple tree. The causative agents 
of brown-rot are fungi capable of utilizing 
cellulose and leaving behind lignin which 
is brown in color (Pugh 1980, Richards 
1974). These larvae are believed to feed on 
the mycelial growth produced by the brown 
rotting fungus growing on the decaying 
wood. Fifteen larvae, presumed to be sib- 
lings, were collected in all. Of these, three 
were slide mounted and the remainder were 
cultured in plastic petri dishes moistened 
with paper towels. Three males and three 
females were reared and slide mounted in 
euparal. 


Cecidomyiidae, Porricondyla, gall midges, prothoracic collar 


Porricondyla recondita Plakidas, 
new species 
(Figs. 1-13) 


Adult.—Color: Eyes black; antenna 
slate gray; thorax and abdomen brown; legs 
brown, covered with black setiform scales; 
wings hyaline, fringed with black hairs. 

Male: Head: Palpiger + 4-segmented 
palpus (Fig. 1), labellum with setae distally, 
eye bridge lacking, frons with 3—4 setae per 
side (Fig. 1). Antenna: Scape, pedicel and 
14-flagellomeres, circumfila on all flagel- 
lomeres consisting of a single loop (Fig. 2). 
Thorax (Fig. 3): Dorsocentral and dorso- 
lateral setae present. Katepisternum bare, 
anepisternum with one seta, anepimeron 
with 2 or 3 setae. Legs: Tarsal claws on all 
legs with a basal tooth, empodium rudi- 
mentary. Wing (Fig. 4): Length 2 mm, 
CuA, vein to wing margin, CuA, vein ab- 
sent. Abdomen: Tergites 1—8 and sternites 
2-8 lightly sclerotized. Genitalia (Fig 5): 
Cercus long, narrow, tapering to pointed 
apex; gonocoxites and gonostyli covered 
with setae and setulae, apical inner margin 
of gonostyli with 5—7 sclerotized teeth. 
Gonocoxal apodemes extending to anterior 
margin of ninth tergite. Aedeagus longer 


VOLUME 107, NUMBER 3 


653 


Figs. 1-8. 


than gonostyli; parameres lightly sclero- 
tized basally and fused apically forming a 
shield (tegmen) ventrad to aedeagus. 
Female: Head: Palpiger + 4-segmented 
palpus, labellum with setae distally, eye 
bridge lacking, frons with 2—3 setae per 
side. Antenna: Scape, pedicel and 12-fla- 
gellomeres, circumfila on all flagellomeres 
consisting of two interconnected loops (Fig. 
6). Thorax (Fig. 7): Dorsolateral and dor- 
socentral setae present; anepisternum and 
katepisternum bare; anepimeron with a sin- 
gle seta. Legs: Tarsal claws on all legs with 
a basal tooth, empodium rudimentary. 
Wing: As in male. Abdomen: Tergites 1-6 


Porricondyla recondita. 1, Palpus. 2, Male 3™ flagellomere. 3, Male thorax, lateral. 4, Wing. 5, 
Male genitalia. 6, Female 3" flagellomere. 7, Female thorax, lateral. 8, Female abdomen, segments 9-10. 


and sternites 2—6 lightly sclerotized; seg- 
ments 7 and 8 membranous, protrusible. 
Ninth and 10" abdominal segments (Fig. 8): 
9" tergite lightly sclerotized with dorsal and 
lateral setae; 9" sternite membranous with 
setae posteriorly and densely covered with 
setae situated at anterior margin; cercus 2- 
segmented, covered with setae and setulae, 
first segment rectangular, second segment 
ovoid and shorter than first; hypoproct 
ovoid, situated below cerci, covered with 
setulae and with a few setae distally. Sper- 
mathecae membranous, not visible on slide- 
mounted cleared or uncleared specimens. 
Larva (Fig. 9).—Length 3 mm, tawny or- 


654 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


i, es : 
SS g 
A : ¥ 
/ = ee ee 
spiracle SNe 
12 sclerotized setae 
Figs. 9-13. 


thoracic 
Spiracle 


rows of 
sclerotized 
teeth 


Porricondyla recondita. 9, Larva, dorsal. 10, Head capsule, dorsal. 11, Prothoracic collar. 12, 


8" & 9% abdominal segments, dorsal. 13, Pupa, ventrolateral. 


ange, cylindrical. Head (Fig. 10): Dark 
brown with elongate posterior apodemes, 
antenna longer than wide, with a minute pa- 
pilla situated distally and with sclerotized 
basal collar; spatula absent. Thorax: Seg- 
ments ventrally with 4 groups of 3 lateral 
papillae, 1 of each group with seta and 2 
without setae. Sclerotized collar (Fig. 11) 
situated on dorsal surface of prothorax be- 
tween and extending laterally just beyond 
thoracic spiracles comprising 5 centrally lo- 
cated tooth-like processes. Six dorsal papil- 
lae present, each with a coniform seta an- 
terior to collar and at least 2 pleural coni- 
form setae situated at each posterior edge 
of prothorax. Abdomen: Segments 1-7 
with 6 dorsal and 2 pleural papillae each 


with a short seta. Spinule fields situated on 
dorsal and ventral anterior margin of meso- 
and metathorax and all abdominal seg- 
ments. Anterior ventral papillae on the first 
7 abdominal segments situated inside spi- 
nule fields near posterior margin. Spiracles 
on 8" abdominal segment situated near dor- 
sal posterior margin and elevated above 
body wall (Fig. 12). Four setiform dorsal 
papillae situated between spiracles. Termi- 
nal segment shorter than 8"; 3 simple anal 
papillae situated on either side of anal pore; 
dorsal surface with 8 terminal papillae, 3 
pair setiform, 1 pair large, pigmented, con- 
iform, hooklike (Fig. 12). 

Pupa (Fig. 13).—Illustrated from a fe- 
male pupal skin. Head with sclerotized an- 


VOLUME 107, NUMBER 3 


tennal horns situated on apical inner margin 
of antennal sheaths; face with 2 pairs of mi- 
nute setiform papillae anterior to clypeal 
sheath; thoracic spiracle elongate, blunt; ab- 
domen 9-segmented; cuticle covered with 
spicules; spiracles situated on lateral mar- 
gins of segments 2—7; a single row of scler- 
otized teeth situated on dorsal anterior mar- 
gin of segments 2—8; last segment bilobate. 

Types.—Holotype, male, Pennsylvania, 
Allegheny Co., 15 km NE Pittsburgh, 
emerged 15 May 2000; allotype female and 
pupal skin, same data as holotype; paratype 
larva same pertinent data as holotype. All 
deposited in the National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, DC. Other specimens: 2 lar- 
vae, 2 males and | female all slide mount- 
ed, in the author’s collection. 

Etymology.—The specific name, recon- 
dita, means hidden, and refers to the cryptic 
life of the larva. 

Discussion.—Using the key from Parnell 
(1971) males key to couplet 4 and superfi- 
cially resemble P. unidentata, but with 
three specific differences. Porricondyla re- 
condita lacks heavily pigmented parameres 
in the male genitalia, the CUA, vein is ab- 
sent, and the apical margin of the gonostyli 
has 5—7 teeth, whereas P. unidentata has 
heavily pigmented parameres, has the 
CUA, vein present, and the apical margin 


of the gonostyli has 2—3 teeth. Females of 


P. recondita differ from all other females 
with the CUA, vein absent, 12 flagello- 
meres all with circumfila and the sperma- 
theceae unpigmented. The larva, which 
lacks a spatula, has a heavily sclerotized 
prothoracic dorsal collar, a condition unique 
in the Porricondylinae. Only one other ge- 
nus, Ledomyia (Cecidomyiinae), has larvae 
with a dorsal collar (Gagné 1985, Larew et 
al. 1987). In Ledomyia the dorsal collar is 
similar in appearance to that of P. recon- 
dita. However, differences do occur be- 
tween larvae of the two genera. All Ledo- 
myia larvae possess a spatula and have 2 
setiform papillae between the spiracles on 
the 8" abdominal segment. Porricondyla 


655 


recondita larvae lack a spatula and have 4 
setiform papillae between the spiracles, the 
latter being a subfamily characteristic. 


ACKNOWLEDGMENTS 


I am grateful to Dr. R. J. Gagné Research 
Entomologist, Emeritus, Systematic Ento- 
mology Laboratory, USDA, for his helpful 
comments which improved the quality of 
this paper. I also thank Dr. Voldemars Spun- 
gis, Faculty of the Department of Zoology 
and Animal Ecology, University of Latvia, 
who reviewed and made helpful comments 
on the first draft of this paper. 


LITERATURE CITED 


Dallai, R., P. Lupetti, EF Frati, B. M. Mamaey, and B. 
A. Afzelius. 1996. Characteristics of sperm ultra- 
structure in the gall midges Porricondylinae (In- 
secta, Diptera, Cecidomyiidae), with phylogenetic 
considerations on the subfamily. Zoomorphology 
116: 85—94. 

Gagné, R. J. 1985. Descriptions of new Nearctic Ce- 
cidomylidae (Diptera) that live in xylem vessels 
of fresh-cut wood, and a review of Ledomyia 
(s.str). Proceedings of the Entomological Society 
of Washington 87: 116—134. 

. 2004. A Catalog of the Cecidomyiidae (Dip- 
tera) of the World. Memoirs of the Entomological 
Society of Washington No. 25, 408 pp. 

Larew, H. G., R. J. Gagné, and A. Y. Rossman. 1987. 
Fungal gall caused by a new species of Ledomyia 
(Diptera: Cecidomyiidae) on Xylaria enterogena 
(Ascomycetes: Xylariaceae). Annals of the Ento- 
mological Society of America 80: 502—507. 

Mamaey, B. M. and N. P. Krivosheina. 1965. The Lar- 
vae of the Gall Midges. ix + 293 pp. (English 
translation, 1993. A.A. Balkema Publishers, 
Brookfield, Vermont.) 

Parnell, J. R. 1971. A revision of the Nearctic Porri- 
condylinae (Diptera: Cecidomyiidae) based large- 
ly on an examination of the Felt types. Miscella- 


neous Publications, Entomological Society of 
America 7: 275-348. 

Plakidas, J. D. 1999. Identification of the North Amer- 
ican Porricondyline Larvae (Diptera: Cecidomyi- 

Loyalfield Publishing, Pittsburgh, Pennsyl- 
vania, 129 pp. 

jeipyednys (Gy, apo Les MFO} 
Transactions of the British Mycological Society 
75: 1-14. 

Richards, B. N. 
tem. Longman, Harlow, 

V. 1981. New species of gall midges from the 


idae). 


Strategies in fungal ecology. 


1974. Introduction to the Soil Ecosys- 
Essex. 
Spungis, 


tribes Oligotrophini and Porricondylini. Latvijas 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Entomologs 24: 43—55. (In Russian with English . 1989. A revision of the European gall midge 


summary.) species of the genus Camptomyia Kieffer (Dip- 
. 1985. Gall midges of the subtribe Diallactina tera: Cecidomyiidae). Latvijas Entomologs 32: 

(Diptera: Cecidomyiidae) in Latvia. Latvijas En- 54-74. (In Russian with English summary.) 

tomologs 28: 38-53. (In Russian with English . 1992. A revision of the European gall midges 

summary.) of the tribe Winnertziini. Latvijas Entomologs 
. 1987. Gall midges of the subtribe Dicerurina Supplementum V, 39 pp. 

(Diptera: Cecidomyiidae) in Latvia. Latvijas En- . 2002. A check-list of Latvian Porricondylinae 


tomologs 30: 15-42. (In Russian with English (Diptera: Cecidomyiidae) with notes on new rec- 
summary.) ords. Latvijas Entomologs 39: 56—60. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 657-662 


NOTES ON THE LIFE OF DR. CLARA SOUTHMAYD LUDLOW, PH.D., 
MEDICAL ENTOMOLOGIST (1852-1924) 


TERRY L. CARPENTER 


Armed Forces Pest Management Board, Office of the Deputy Under Secretary of 
Defense for Installations and Environment, The Pentagon, Washington, DC 20301-3400, 
U.S.A. (e-mail: terry.carpenter@osd.mil) 


Abstract.—Dr. Clara Southmayd Ludlow, the first woman known to have published 
extensively on the taxonomy of mosquitoes and their occurrence in relation to the inci- 
dence of mosquito-borne diseases, forged a notable career in medical entomology in as- 
sociation with the United States Army. Though much is known about her accomplish- 
ments, knowledge of her professional and personal life has been less well documented. 
New information about her life and three previously unpublished photographs of her are 


presented in this article. 


Key Words: 
ical entomology 


Dr. Clara Southmayd Ludlow is a signif- 
icant figure in the history of entomology, in 
professional and personal terms, as the first 
woman known to have published extensive- 
ly on the taxonomy of mosquitoes and their 
occurrence in relation to the incidence of 
mosquito-borne diseases (Knight and Pugh 
1974, Neilson 1987). Not only did she 
forge a career in medical entomology dur- 
ing a time when women were rare among 
the ranks of entomologists, she did so in 
association with the military, where the 
presence of women was even rarer. 

Dr. Ludlow’s life and career have been 
addressed by Kitzmiller (1982), with excel- 
lent supplementation by R. A. Ward (Kitz- 
miller and Ward 1987) and Patterson 
(2003), works that are recommended for de- 
tails necessary to fully understand the sig- 
nificance of her career as an entomologist. 

Though the record of her professional 
life can be illuminated through her papers, 
Kitzmiller and Ward (1987) note that ““we 
know almost nothing of the first 45 years 


Clara Southmayd Ludlow, entomologist, mosquito taxonomy, military, med- 


of Clara Ludlow’s existence.” This work 
adds some detail to the knowledge of that 
period, and her later life. A brief summary 
of her life and career is necessary here to 
place in context the additional information 
discovered. 

Dr. Ludlow was born on 26 December 
1852 at Easton, Pennsylvania, the eldest 
child of Jacob Rapalje and Anna Mary 
(Hunt) Ludlow (Cameron 1924-1925). Dr. 
Ludlow’s ancestry was evidently of some 
pride to her, as she devoted a significant 
portion of her entry in the Biographical Cy- 
clopedia of American Women—a_ source 
not cited by previous biographers—to ge- 
nealogical detail (Cameron 1924—1925). 
Parental and sibling families were certainly 
important in her life, bringing professional 
associations that led to opportunities that 
might have been impossible to acquire oth- 
erwise. 

On 8 June 1860, Dr. Ludlow was enu- 
merated in the 1860 United States federal 
census in her parents’ household in Easton, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
University, Kayser Photographic Collection, PA-131). 


Pennsylvania, at the age of 7, the eldest of 
four children and only daughter (National 
Archives 1860 Federal Census). I have not 
been successful in locating her in the 1870 
federal census, when she would have been 
age 17. It is probable that she accompanied 
her parents to Knoxville, Tennessee, im- 
mediately after the Civil War, and they are 
reported to have returned to Easton, Penn- 


0559 ob 
“6 CyjcAGO 


Clara S. Ludlow, ca. 1879-1889 (University Archives, The Gelman Library, The George Washington 


sylvania, by 1870 (Kitzmiller and Ward 
1987); therefore, they may have been 
missed by the census officials because they 
were in transit. 

In 1877, Dr. Ludlow enrolled in the New 
England Conservatory of Music, from 
which she graduated in 1879 (Kitzmiller 
and Ward 1987). Her whereabouts from this 
time until about 1897 have been only spec- 


VOLUME 107, NUMBER 3 


ulative. The census of 1880 enumerates her 
on 8 June 1880, residing at the Monticello 
Female Seminary near Alton, Illinois, a sin- 
gle woman, her occupation listed as ‘““Music 
[Teacher]? (National Archives 1880 Fed- 
eral Census). Most of the 1890 federal cen- 
sus was destroyed, and no record of her is 
found in the surviving fragments of that 
enumeration. Her entry in the Biographical 
Cyclopedia of American Women (Cameron 
1924—1925) states that “for many years she 
made music her profession, teaching and 
doing a certain amount of concert work.” 
Her professional records, deposited at the 
National Museum of Health and Medicine 
in Washington, D.C., begin with the year 
1889, suggesting that at some point in the 
late 1880s, she began to turn to science as 
an avocation or perhaps vocation (National 
Museum of Health and Medicine 2004). 

In 1897, she was a student at Mississippi 
Agricultural and Mechanical College (now 
Mississippi State University) in Starkville, 
Mississippi. On 20 June 1900, she was enu- 
merated in the federal census residing on 
the campus of Mississippi A&M as a single 
woman, with no occupation listed (National 
Archives 1900 Federal Census). Enumer- 
ated nearby was Dr. George W. Herrick (or 
Herrich), professor of biology at the col- 
lege, reportedly an influential figure in the 
development of her interest in mosquitoes 
(Kitzmiller and Ward 1987). 

She graduated from Mississippi A&M in 
1900 with the degree of Bachelor of Sci- 
ence in Agriculture (Kitzmiller and Ward 
1987). In 1901, she was awarded the Master 
of Arts degree in Botany by Mississippi 
A&M, reportedly 31 years before a gradu- 
ate program was formally offered by that 
institution (Kitzmiller and Ward 1987). In- 
terestingly, one of her preserved works is a 
folder of drawings of Viola species (the vi- 
olets) in the Edward L. Greene Papers of 
the University of Notre Dame Archives, 
perhaps obtained by Professor Greene when 
he worked in the Washington, D.C., area as 
a faculty member of the Botany Department 
of the Catholic University of America 


659 


(1885-1904) or in his capacity as an asso- 
ciate in botany at the Smithsonian Institu- 
tion (1904-1909) (University of Notre 
Dame Archives 2004), either of which po- 
sitions might have brought him into profes- 
sional correspondence with Dr. Ludlow. 

After graduation in 1901 with her Mas- 
ters of Arts degree, Dr. Ludlow traveled to 
Manila, Philippines, to visit a brother who 
was Stationed there as an artillery officer in 
the United States Army (Kitzmiller and 
Ward 1987). Approximately one year later, 
she returned to the States with her brother, 
who had contracted an illness, but during 
her stay in Manila, she began an association 
with military medicine that would endure 
for the rest of her life (Kitzmiller and Ward 
1987, National Museum of Health and 
Medicine 2004). 

In 1904, she was Lecturer on mosquitoes 
and disease at the Army Medical Museum 
in Washington, D.C. By 1907, she was 
Demonstrator of Histology and Embryolo- 
gy at George Washington University in 
Washington, D.C. (Calvert 1924), where 
she received her Doctor of Philosophy de- 
gree in 1908. Her doctoral dissertation was 
entitled ““The Mosquitoes of the Philippine 
Islands: The Distribution of Certain Species 
and Their Occurrence in Relation to the In- 
cidence of Certain Diseases’? (Calvert 
1924). She remained on the faculty of 
George Washington University, where in 
1909 she was Instructor of Histology and 
Embryology (Calvert 1924). She surpassed 
yet another milestone in 1908 when she was 
elected to active membership in the Amer- 
ican Society of Tropical Medicine, the first 
woman and the first non-physician scientist 
member of the society (Burke 2003). 

In April of 1910, she was enumerated by 
the 1910 federal census in the District of 
Columbia, Washington, residing at 133 15th 
Street, age 58, unmarried, her occupation 
listed as ‘‘Scientist, Prev[entive] Medicine” 
(National. Archives 1910 Federal Census). 
From 1916 through 1920 she 
Anatomist at the Army Medical Museum 
(Calvert 1924), now the National Museum 


served as 


6! PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. 
The George Washington University, Kayser Photographic Collection, PA-132). 


of Health and Medicine, on the campus of 
Walter Reed Army Medical Center in 
Washington, D.C. Her records at the mu- 
seum, consisting of correspondence, notes, 
reports, logbooks, and other research ma- 
terials, state that her research centered on 
identifying mosquitoes, including a project 
working with specimens sent in from mili- 
tary posts that resulted in the production of 
a Museum film, ““Mosquito Eradication,” 
in 1918 (National Museum of Health and 
Medicine 2004). 

In January 1920, Dr. Ludlow was enu- 
merated in the 1920 federal census residing 
in the District of Columbia, at 1415/1435 
15th Street, age 68, unmarried, and her oc- 
cupation is listed as ““Anatomist, Govt. Ser- 
vice’? (National Archives 1920 Federal 
Census). During 1920, she became the mu- 
seum’s Chief Entomologist, a position she 


Clara S. Ludlow at work in her room, ca. 1905—1911 (University Archives, The Gelman Library, 


held until her death (Calvert 1924, National 
Museum of Health and Medicine 2004). 

Dr. Ludlow passed away on 28 Septem- 
ber 1924 in Washington, D.C. (Calvert 
1924) and is interred in the Arlington Na- 
tional Cemetery, in Section 2, Grave No. 
3843, beside her father, who was Surgeon 
of the Ist Regiment, Pennsylvania Volun- 
teers, United States Army, in the American 
Civil War. Her grave is located under a 
large oak below the Old Amphitheater ad- 
jacent to Arlington House. Her stone states 
only her name and date of death, and 
“daughter of Jacob’’—hardly an adequate 
memorial for this great lady. 

Dr. Ludlow is included in a bibliography 
of biographies of entomologists published 
in 1945 (Carpenter 1945), but the cited 
obituary, published in the Journal of the 
Washington Academy of Sciences, states 


VOLUME 107, NUMBER 3 


661 


Fig. 3. 


Clara S. Ludlow, Alumna and Faculty, ca. 1905-1911 (University Archives, The Gelman Library, 


The George Washington University, Kayser Photographic Collection, PA-130). 


only that “[h]Jer work was mainly in con- 
nection with the disease prevention activi- 
ties of the army” (Anonymous 1924) and 
does not mention her pioneering role as an 
entomologist. The failure of her peers to 
fully memorialize her remarkable achieve- 
ments may be in some part due to her re- 


portedly irascible personality (Kitzmiller 


and Ward 1987); as well, it may reflect the 
sexism of early twentieth century America. 

Until now, only one likeness of Dr. Lud- 
low was known to exist, the one published 
in the issue of Mosquito Systematics dedi- 
cated to her in 1987 (Neilson 1987). In my 
research into the life of Dr. Ludlow, I dis- 
covered three previously unknown photo- 
graphs of her in the George Washington 
University Archives, Kayser Photographic 
Collection, in Washington, D.C. These 
three photographs, which provide signifi- 
cant visual insight into her life, are repro- 


duced herein, as Figs. 1—3, with the per- 
mission of the Gelman Library, Special 
Collections Department/University Ar- 
chives, Washington, D.C. 


ACKNOWLEDGMENT 


The author acknowledges with thanks the 
encouragement and technical support of 
Richard G. Robbins, Defense Pest Manage- 
ment Information Analysis Center, Armed 
Forces Pest Management Board, Washing- 
Worn IDC. 


LITERATURE CITED 


Anonymous. 1924. Scientific notes and news. Journal 
of the Washington Academy of Sciences 14: 418. 
Burke, D. S. 2003. American Society of Tropical Med- 
icine and Hygiene Centennial Celebration Ad- 
dress. Accessed on the Worldwide Web at URL 
<http://www.astmh.org/address.pdf>, 8 August 
2004. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Calvert, P. P. 1924. Obituary. Entomological News 35: 
379-380. 

Cameron, M. W., ed. 1924-1925. Biographical Cyclo- 
pedia of American Women. Halvord Publishing 
Co., New York, pp. 189-190. 

Carpenter, M. M. 1945. Bibliography of biographies of 
entomologists. American Midland Naturalist 33: 
62. 

Kitzmiller, J. B. 1982. Anopheline Names, Their Der- 
ivations and Histories. Thomas Say Foundation 8: 
316-321. 

Kitzmiller, J. B. and R. R. Ward. 1987. Biography of 
Clara Southmayd Ludlow 1852-1924. Mosquito 
Systematics 19: 251—258. 

Knight, K. L. and R. B. Pugh. 1974. A bibliography 
of mosquito writings of C. S. Ludlow and D. W. 
Coquillett. Mosquito Systematics 6: 214—219. 

National Archives, 1860 Federal Census of the United 
States, Micropublication T-653, Roll 1147, Book 
1, Pennsylvania, Northampton County, Easton, 
Bushkill Ward, stamped page 252, 8 June 1860. 

National Archives, 1880 Federal Census of the United 
States, Micropublication T-9, Roll 233, Book 1, 
Illinois, Madison County, Godfrey Township, 
Enumeration District 1, page 11A, 8 June 1880. 


National Archives, 1900 Federal Census of the United 
States, Micropublication T-623, Roll 823, Book 1, 
Enumeration District 86, Sheet 21A, Oktibbeha 
County, Beat 1, Agricultural & Mechanical Col- 
lege, 20 June 1900. 

National Archives, 1910 Federal Census of the United 
States, Micropublication T-624, Roll 149, Part 2, 
District of Columbia, Washington City, Enumer- 
ation District 8, Sheet SA, 21—22 April 1910. 

National Archives, 1920 Federal Census of the United 
States, Micropublication T-625, Roll 205, District 
of Columbia, Washington City, Enumeration Dis- 
trict 9, Sheet 3A, 3—5 January 1920. 

National Museum of Health and Medicine. 2004. Lud- 
low Entomology Records, 1889-1924. National 
Museum of Health and Medicine, Walter Reed 
Army Medical Center, Washington, D.C. 

Neilson, L. T. 1987. Editor’s corner. Mosquito System- 
atics 19: 259. 

Patterson, G. 2003. War & entomology, Clara Ludlow, 
early female mosquito taxonomist. Wing Beats 14: 
14-16. 

University of Notre Dame Archives. 2004. Edward 
Lee Greene Papers (GRE), 607 Hesburgh Library, 
Notre Dame, Indiana. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 663-670 


NEW SPECIES OF THE GENUS POLYCENTROPUS CURTIS 
(TRICHOPTERA: POLYCENTROPODIDAE) FROM MEXICO 


R. BARBA-ALVAREZ AND J. BUENO-SORIA 


Instituto de Biologia, UNAM. Apdo Postal 70-153, Mexico 04510 D. E (e-mail: 


bueno @servidor.unam.mx) 


Abstract.—Five new species of the genus Polycentropus are described and illustrated 
from collections made in Mexico: Polycentropus aliciae n. sp., Polycentropus mixteco 
n. sp., Polycentropus ibarrai n. sp., Polycentropus dianae n. sp., and Polycentropus 


giovanae Nn. sp. 


Key Words: 


The genus Polycentrus is well represent- 
ed in Mexico and Central America as re- 
corded by Holzenthal and Hamilton (1988) 
and Flint et al. (1999). Barba-Alvarez 
(1991) registered 27 species for Mexico and 
recently Chamorro-Lacayo (2003) de- 
scribed three more new species from Cen- 
tral America. However, because of the great 
dispersion of aquatic resources over a wide 
variety of mountains stream habitats in 
Mexico, it is quite probable that many more 
undescribed species still are present, as was 
pointed out by Hamilton (1986) in his re- 
vision of the New World species of Poly- 
centropus. All new species described in this 
paper belong to the Gertschi group, and are 
placed into the various species complexes 
proposed by Hamilton (1986). 

During the identification process we 
compared affinities between the new spe- 
cies herein described. The most important 
characters we used to segregate the species 
into species complexes are the number of 
spines in the endotheca of the phallus and 
the shape of the preanal and inferior ap- 
pendages, as defined by Hamilton (1986). 
Terminology employed in the description of 
the new species follows Hamilton (1986). 
The material herein described will be de- 


Trichoptera, Polycentropus, new species, Mexico 


posited at the Coleccién Nacional de Insec- 
tos (CNIN) at the Universidad Nacional 
Autonoma de México and at the National 
Museum of Natural History, Smithsonian 
Institution, Washington, DC. (NMNH). 


Polycentropus aliciae Barba-Alvarez and 
Bueno-Soria, new species 
(Fig. 1) 


Diagnosis.—This species belongs in the 
picana complex proposed by Hamilton 
(1986) and related to Polycentropus zanclus 
Flint (1980), described from Guatemala. 
They are similar in the elongate shape of 
the inferior appendages. However, Polycen- 
tropus aliciae differs from P. zanclus by the 
longer, strongly dorsally curved postero- 
ventral process of the preanal appendages, 
giving it a sickle-shape. In P. zanclus the 
mesoventral process is shorter, rodlike and 
slightly curved ventrally. 

Description.—Male: Length of forewing 
8 mm. Wings dark brown covered with mi- 
nute gold spots; antenna, legs, and body 
ventrally stramineous. Male genitalia: Seg- 
ment IX with anterior margin slightly 
oblique, posterior margin slightly produced 
medially. Segment X membranous. Inter- 
mediate appendages reduced to a pair of 


164. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


vill Int. app. 


Fig. 1. 


Polycentropus aliciae, male genitalia. 1a, Lateral. 1b, Dorsal. lc, Ventral. 1d, Phallus, lateral. le, 


dorsal. 1f, Subphallic sclerite. int. app. :eq intermediate appendages; inf. app. = inferior appendages; pre. app. 
= preanal appendages; mv. p. = mesoventral process; spi. = spines; ph. sc. = phallic sclerite. 


short, rodlike structures covered with mi- 
crosetae at base of preanal appendages. Pre- 
anal appendages in lateral view almost 
square, posterodorsally rounded; postero- 
ventrally produced into elongate, upturned, 
hooklike processes; in posteroventral view 
apices appear slightly heliciform. Inferior 
appendages elongate, slightly upcurved; in 
lateral view with a basal thumblike lobe, 
narrowing apically, each with basomesal 
spinelike process highly sclerotized. Phallus 
with a group of eight small spines into a 
phallic membrane; in lateral view with 
short phallobase and apicoventral lobe 
strongly curved anteroventrad; in ventral 


view apicoventral lobe trilobed, mesal lobe 
elongate; phallic sclerite long and weakly 
sclerotized; subphallic sclerite large and 
forming a complete ring around phallobase. 

Female: Unknown. 

Type material—Holotype: 6. MEXICO: 
Veracruz, Rio Jamapa, 5 km NE Coscoma- 
tepec, 29-i-1984, J. Bueno (CNIN). Para- 
types: Chiapas, 25 km from Lagunas de 
Montebello, 2-iv-1981, C. R. Beutelspacher, 
6 3 (CNIN), 2 6 (NMNH); Union Juarez, 
ca. Tapachula, 22-iv-1983, elev. 1,700 m, 
Bueno, Barrera & Brailovsky, 3 6 (CNIN). 
Oaxaca: Portillo del Rayo, 1,540 m, 1-xii- 
82, A. Ibarra & M. Garcia, 3 ¢ (CNIN). 


VOLUME 107, NUMBER 3 


Vill 


Fig. 2. 
Dorsal. 


Etymology.—We dedicate this species to 
the memory of Alicia Rodriguez-Palafox, a 
young Mexican entomologist devoted to the 
study of Hymenoptera, who unfortunately 
recently died. 


Polycentropus mixteco Barba-Alvarez 
and Bueno-Soria, new species 
(Fig. 2) 


Diagnosis.—This species is a member of 
the halidus complex by the long interme- 
diate appendages and the presence of one 
pair of small spines in the phallus. Polycen- 
tropus mixteco resembles to Polycentropus 


665 


Polycentropus mixteco, male genitalia. 2a, Lateral. 2b, Dorsal. 2c, Ventral. 2d, Phallus, lateral. 2e, 


ibarrai, n. sp., by the dorsally directed lobe 
of the inferior appendages. Polycentropus 
mixteco can be distinguished from P. tbar- 
rai by the digitate dorsal lobe of preanal 
appendages, the curved midbasal portion of 
the intermediate appendages and the square 
shape of the inferior appendages in ventral 
aspect. The former structure in ventral view 
appears long, narrow and with bifid apex in 
P. ibarrai. 

Description.—Male: Length of fore- 
wing, 8 mm. Color dark brown; antenna, 
legs and body ventrally stramineous. Male 
genitalia: Segment IX with anterior margin 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


slightly curved, posterior margin slightly 
produced at midlength. Segment X mem- 
branous. Intermediate appendages, in lateral 
view, elongate, slightly sinuous, with acute 
apices; in dorsal view, well separated from 
each other, basal half curved mesad, apical 
half curved lateral. Preanal appendage in 
lateral and dorsal view, thumblike, setose 
and shorter than intermediate appendage; 
mesoventral process caudally directed, 
short and bluntly rounded. Inferior append- 
age in lateral view with a high, narrow, dor- 
solateral flange, rounded dorsally; dorsally- 
directed caudomesal point exposed in lat- 
eral aspect; in ventral aspect square, mesal 
margin straight, caudomesal point promi- 
nent, rounded. Phallus, in lateral view, with 
endothecal membrane bearing a pair of 
small spines; in dorsal view phallic sclerite 
ovate with two dark lateral spines, apico- 
ventral process short. 

Female: Unknown. 

Type material —Holotype: 6. MEXICO: 
Oaxaca, La Esperanza, route 175, ca. Valle 
Nacional, 18.iv.1983, A. Ibarra and M. 
Garcia (CNIN). 

Etymology.—tThe species epithet alludes 
to the Mixteco Indians, inhabitants of the 
region in the State of Oaxaca, where the 
type was collected. 


Polycentropus ibarrai Barba-Alvarez 
and Bueno-Soria, new species 
(Fig. 3) 


Diagnosis.—This species is a member of 
the halidus complex designated by Hamil- 
ton (1986), based on the presence of only 
one pair of spines in the phallus. Polycen- 
tropus ibarrai slightly resembles P. mixte- 
co, N. sp., in the digitate lobe of the inferior 
appendages, in lateral view. Polycentropus 
ibarrai can be distinguished from P. mix- 
teco by the bilobulate mesoventral process 
of the preanal appendage, the straight shape 
of the intermediate appendage in dorsal as- 
pect and the long, narrow, and bifid apex of 
the inferior appendage in ventral aspect. 

Description.—Male: Length of fore- 
wing, 8 mm. Color in alcohol light brown; 


antenna, legs and body ventrally strami- 
neous. Male genitalia: Segment [X with an- 
terior margin slightly convex, posterior 
margin slightly produced medially. Seg- 
ment X membranous. Intermediate append- 
ages, in lateral view elongate cylindrical, 
slightly curved ventrally; in dorsal view 
over each other. Preanal appendage, in lat- 
eral view slightly produced into a stout, lat- 
eral process almost triangular, with rounded 
apex; mesoventral processes bilobate, inter- 
nal lobule of mesoventral processes situated 
laterally to phallobase; in dorsal view mes- 
ally bearing sclerotized cylindrical process- 
es with apical setae. Inferior appendage, in 
lateral view, with a well-developed dorso- 
lateral, narrow, flange, ventral surface ap- 
pearing rectangular with apex truncate; in 
ventral view with erected mesal lobe with 
apex bifid. Phallus with apicoventral lip 
short, endotheca with a basal pair of short, 
parallelside, curved spines; in dorsal view 
phallic sclerite ovate with two dark lateral 
stripes; apicoventral process short. 

Female: Unknown. 

Type material.—Holotype: 6: MEXICO: 
Hidalgo, Hixtlahuaco, Hotel Campestre 
Conchita, 20°53.025’N, 98°42.140’W, el. 
1,400 m, 25.xi.1998, E. Barrera and A. 
Ibarra (CNIN). 

Etymology.—We name this species in 
honor of Adolfo Ibarra, collector at the In- 
stituto de Biologia, UNAM. 


Polycentropus dianae Barba-Alvarez and 
Bueno-Soria, new species 
(Fig. 4) 


Diagnosis.—This species is another 
member of the picana complex, similar in 
overall appearance to Polycentropus azte- 
cus Flint 1967, particularly by the rounded 
shape and mesal teeth of the inferior ap- 
pendages. However, P. dianae, can be dis- 
tinguished from P. aztecus by the preanal 
appendages, with the ventrally curved, pos- 
terodorsal process, the slightly elongated 
posterior margin of [IX segment, and the 
presence of a pair of large basal spines in 
the endothecal membrane. 


VOLUME 107, NUMBER 3 


3c 


Fig. 3. 
Dorsal. ap. bif. = apex bifurcated. 


Description.—Male: Length of forewing 
7 mm. Color dark brown, forewing with 
small white spots and groups of golden se- 
tae. Male genitalia: Segment IX in lateral 
view with anterior margin slightly rounded, 
posterior margin distinctly produced at mid- 
length. Tergum X membranous. Intermedi- 
ate appendages cylindrical; in dorsal aspect 
thin and slightly divergent at apex; in lateral 
aspect slightly curved ventrad, apex with 
small, pale setae. Preanal appendages in lat- 
eral view svelte, with long, posterodorsal 
process, strongly curved ventrad with acute 
apex; mesoventral processes produced into 


spl. 


667 


int. app. 


pre. app. > rx 
i 


ph. sc. 


Polycentropus ibarrai, male genitalia. 3a, Lateral. 3b, Dorsal. 3c, Ventral. 3d, Phallus, lateral. 3e, 


small, rounded lobe; in dorsal aspect stout, 
thumblike. Inferior appendage in_ lateral 
view nearly oval, dorsal margin narrow and 
rounded, posterior margin rounded; in ven- 
tral view ovoid, mesal margin divergent, 
curved, bearing toothlike process at mid- 
length. Phallus in lateral view, with long 
apicoventral process, broad basally, apically 
narrowed; endothecal membrane with two 
elongate spines basally; apically with two 
sets of three lateral spines; phallic sclerite, 
cylindrical with posterior margin indented; 
subphallic sclerite, a simple, slightly curved 
bar. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Spee ee 


Af 


Fig. 4. Polycentropus dianae, male genitalia. 4a, Lateral. 4b, Dorsal. 4c, Ventral. 4d, Phallus, lateral. 4e, 


Dorsal. 4f, Subphallic sclerite. 


Female: Unknown. 

Type material—Holotype: ¢. MEXICO: 
Nuevo Leon, Mpio. Zaragoza, El Salto, 
9.xi.1995, R. Barba, 1 (CNIN). Paratypes: 
Nuevo Leo6n, Mpio. Zaragoza, El Salto, 
9.x1.1995, R. Barba, 1 d (CNIN); Mpio. 
Zaragoza, Plan del Cerrito, 20.iv.1995, A. 
Contreras, 1 ¢ (CNIN), | d6 (NMNH); 
Mpio. Santiago, Potrero Redondo, 
10.v.1985, A. Contreras, 2 6 (CNIN). 


Etymology.—We have the pleasure of 
dedicating this species to Diana Fernandez 
de Barba, wife of the senior author. 


Polycentropus giovannae Barba-Alvarez 
and Bueno-Soria, new species 
(Fig. 5) 


Diagnosis.—This new species is close to 
Polycentropus encera Denning and Sykora, 
1971, based on the ovoid shape of the in- 


VOLUME 107, NUMBER 3 


———— a aN 


fe 


Fig. 5. 
Se, Phallus, lateral. 5f, Dorsal. 


ferior appendages and is a member of the 
bartolus complex of the Gertschi group as 
defined by Hamilton (1986). It is slightly 
related to Polycentropus encera Denning 
and Sykora, 1971, by the similar ovoid 
shape of the inferior appendages. Polycen- 
ropus giovannae can be distinguished by 
the upturned intermediate appendages, 
which in Polycentropus encera are straight 


and curved ventrad, by the rectangular 


669 


| pre. app. 


pre. app. 


ph. sc. 
spl. 


Polycentropus giovannae, male genitalia. 5a, Lateral. 5b, Dorsal. 5c, Ventral. Sd, Posteroventral view. 


shape and larger size of the preanal ap- 
pendages and by the the ventromesal pro- 
cess which is apically acute and curved 
ventrad. 

Description.—Male: Length of fore- 
wing, 8 mm. Color dark brown with a line 
of golden spots on dorsal, ventral and pos- 
terior margins of forewing. Male genitalia: 
Segment [IX with anterior margin slightly 
convex, posterior margin slightly sinuate. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Tergum X membranous. Intermediate ap- 
pendages, in lateral view, cylindrical with 
acute, upturned apex; in dorsal view, with 
rounded basal lobes, meeting medially, api- 
ces directed laterally. Preanal appendages, 
in lateral view rectangular, posterodorsally 
rounded, mesoventral process apically acute 
and curved ventrad. Inferior appendages, in 
lateral view, with a narrow, rounded, dorsal 
lobe, with mesal ridge well developed; in 
ventral view, with lateral digitate lobe; ap- 
icomesal processes angulated with a short, 
truncate apex. Phallus, in lateral view, with 
apicoventral lip short, pointed, and ventral- 
ly directed with a long, slightly curved 
spine; phallic sclerite elongate; subphallic 
sclerite U-shaped. 

Female: Unknown. 

Type material—Holotype: d: MEXICO: 
Oaxaca, route 175, La Esperanza, ca Valle 
Nacional, 18.iv.1983, A. Ibarra and M. 
Garcia (CNIN). 

Etymology.—We have the pleasure of 
dedicating this species to Giovanna Barba- 
Fernandez, daughter of the senior author. 


ACKNOWLEDGMENTS 


We thank Oliver S. Flint Jr. for review 
and suggestions on the manuscript; Nancy 
Adams, Collection Manager Specialist for 


her help during the stay of J. Bueno-Soria 
at the National Museum of Natural History, 
Smithsonian Institution and the anonymous 
reviewers for their time and effort to im- 
prove the manuscript. 


LITERATURE CITED 


Barba-Alvarez, R. E. 1991. Revisi6n Taxonomica del 
Género Polycentropus Curtis para México (Tri- 
choptera:Polycentropodidae). Tesis para obtener el 
Titulo de Bidlogo. Facultad de Ciencias, UNAM, 
98 pp. 

Chamorro-Lacayo, M. L. 2003. Seven New species of 
Polycentropodidae (Trichoptera) From Nicaragua 
and Costa Rica. Proceedings of the Entomological 
Society of Washington 105(2): 484—498. 

Denning D. G. 1971. A new genus and a new species 
of Trichoptera. The Pan-Pacific Entomologist 47: 
202-210. 

Flint, O. S. Jr. 1981. Studies of Neotropical caddisflies, 
X XIX: The Trichoptera genus Polycentropus (Tri- 
choptera: Psychomyiidae). Journal of the Wash- 
ington Academy of Science 70: 148-160. 

Flint, O. S. Jr, R. W. Holzentha]J, and S. C. Harris. 
1999. Catalog of the Neotropical Caddisflies (In- 
secta:Trichoptera). Ohio Biological Survey, Co- 
lumbus, Ohio, 239 pp. 

Hamilton, S. W. 1986. Systematics and Biogeography 
of the New World Polycentropus sensu stricto 
(Trichoptera: Polycentropodidae). Ph.D. Disserta- 
tion, Clemson University, Clemson, South Caro- 
lina, 257 pp. 

Holzenthal, R. W. and S. W. Hamilton. 1988. New spe- 
cies and records of Costa Rican Polycentropus 
(Trichoptera: Polycentropodidae). Journal of the 
New York Entomological Society 96(3): 332-334. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 671-685 


WEEVILS OF THE GENERA ARCHOCOPTURUS HELLER AND ZYGOPSELLA 
CHAMPION: SIBLING SPECIES AND MIMETIC HOMOPLASY 
(COLEOPTERA: CUCURLIONIDAE: CONODERINAE) 


HENRY A. HESPENHEIDE 


Department of Ecology and Evolutionary Biology, University of California, Los An- 
geles, CA 90095-1606, U.S.A. (e-mail: henryh@biology.ucla.edu) 


Abstract.—Study of recent collections and types shows that the genus Archocopturus 
Heller is both more and less diverse than previously thought. Specimens included under 
the name Archocopturus regalis (Boheman) are a complex of species that are superficially 
very similar, probably because they participate in a single mimicry complex whose models 
are flies of the genus Medetera (Dolichopodidae). The true Archocopturus regalis is ap- 
parently limited to northern South America. Four species are described from Central 
America—A. laselvaensis, n. sp., A. medeterae, n. sp., A. championi, n. sp., and A. 
minutus, n. sp.—and others are known from South America. Archocopturus pulchellus 
Hustache is transferred to the previously monotypic genus Zygopsella (n. comb.) and new 
records are given for Zygopsella ruficauda Champion. Archocopturus basalis Hustache is 
transferred to Macrocopturus (n. comb.). Mimicry of Medetera in Central America in- 
volves at least 28 species in seven genera and most species are undescribed. Mimicry 
selects either for convergent evolution and homoplasy among less closely related species 
or selects against divergence of closely related, sibling species. Such homoplasy may be 
a significant part of tropical biodiversity. 


Key Words: Diptera, Dolichopodidae, Macrocopturus, Medetera, mimicry 


Heller (1895) based the genus Archocop- 
turus on the species Copturus regalis Bohe- 
man, described originally from Cayenne 
(French Guiana), and he cited material from 
*“Amazonas”’ and Peru as that species. Hus- 
tache (1932) described the other two spe- 
cies currently placed in Archocopturus from 
Guadeloupe. Champion (1906) included 
material from México to Panama under 
Heller’s name and described the genus Zyg- 
opsella which he distinguished only from 
Archocopturus, implying a close relation- 
ship of the two genera. 

Examination of collections of what were 
apparently Archocopturus regalis (Bohe- 
man) (sensu Champion) from La Selva Bi- 
ological Station, Costa Rica, showed small 


but consistent differences in size, morphol- 
ogy and coloration among the specimens. 
Dissection revealed that these differences 
were correlated with striking differences in 
male genitalia and that two very similar but 
distinct species were represented. Exami- 
nation of specimens of yet another distinct 
species from México and acquisition of ma- 
terial of more than one species from South 
America led to examination of the type of 
Archocopturus regalis in order to associate 
the name with one of the species. At this 
time it does not appear that the true A. re- 
galis occurs in Central America or Panama. 
Unfortunately, I have seen too little mate- 
rial—especially of males—from South 
America to be able to treat species from that 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


area at this time, although it is clear that 
several additional undescribed species oc- 
cur there also. It is suggested below that the 
small external differences between species 
is a further example of “‘mimetic homopla- 
sy,” due to their participation in a common 
mimicry complex (see below, also Hespen- 
heide 1996b). 

Additionally, study of one Hustache 
(1932) type and material from Guadeloupe 
of the other of his Archocopturus showed 
that neither species from that island belongs 
in the genus. One of them represents the 
second species known from the genus Zyg- 
opsella and the other is transferred here to 
Macrocopturus Heller. 

The following collection codens are used 
throughout the text: AMNH, American Mu- 
seum of Natural History, New York, NY; 
BMNH: The Natural History Museum, 
London, England; CASC, California Acad- 
emy of Sciences, San Francisco, CA, 
U.S.A.; CHAH: Henry A. Hespenheide, 
University of California, Los Angeles, CA, 
U.S.A.; CMNC: Canadian Museum of Na- 
ture, Ottawa, Canada; CWOB: Charles W. 
O’Brien, Tallahassee, FL, U.S.A.; EAPZ, 
Escuela Agricola Panamericana Zamorano, 
Tegucigalpa, Honduras; EMEC, University 
of California, Berkeley, CA, U.S.A.; 
GBFM, Universidad de Panama; INBC: In- 
stituto Nacional de Biodiversidad, Santo 
Domingo de Heredia, Costa Rica; LACM, 
Museum of Natural History of Los Angeles 
County, Los Angeles, CA, U.S.A.; MCZ, 
Museum of Comparative Zoology, Harvard 
University, Cambridge, MA, U.S.A.; 
MNHN: Musée d’ Histoire Naturelle, Paris, 
France; MUCR, Universidad de Costa Rica, 
San Pedro, Costa Rica; NHRS, Naturhis- 
toriska Riksmuseet, Stockholm, Sweden; 
SEAN, Museo Entomoldgico, Léon, Nica- 
ragua; STRI: Smithsonian Tropical Re- 
search Institute, Ancon, Panama; TAMU, 
Texas A & M University, College Station, 
TX, U.S.A.; UNAM, Universidad Nacional 
Autonoma de México; USNM: National 
Museum of Natural History, Smithsonian 
Institution, Washington, DC, U.S.A. 


Archocopturus Heller 


Archocopturus Heller 1895: 56. Type spe- 
cies: Copturus regalis Boheman, by orig- 
inal designation. 


Heller characterized Archocopturus by 
the opalescent blue or blue-green scale-like 
setae on the densely punctate pronotum and 
by the large punctures in the elytral striae. 
In addition, the first and second funicular 
segments are subequal, the mesosternum is 
unmodified, and only the posterior femora 
are dentate beneath. Champion (1906) also 
includes “‘the very large approximate eyes, 
separated by a narrow lanciform space 
above, the basally widened rostrum, 
...fand] the narrow antennal club, with 
rather long second joint.” 

Characters.—The close similarity of and 
small differences between species of Archo- 
copturus may be due to mimetic homoplasy 
(Hespenheide 1996; see below). The pos- 
session of opalescent blue-green scalelike 
setae are a distinctive feature of a putative 
mimicry complex whose models are doli- 
chopodid flies of the genus Medetera Fi- 
scher von Waldheim (Hespenheide 1973, 
1995), and are shared by species in several 
other conoderine genera that are considered 
part of this complex, namely Macrocoptu- 
rus Heller, Hoplocopturus Heller, Coptu- 
romimus Heller, Zygopsella Champion, and 
two genera that appear to be undescribed. 
The occurrence of opalescent setae in spe- 
cies of several genera was noted by Heller 
in his original description and cannot there- 
fore be a generic character as interpreted by 
Hustache (1932). 

Within the genus Archocopturus, species 
can be quite similar, a situation which has 
been termed ‘‘sibling species’? (Mayr 
1963). Although male genitalia can be quite 
distinctive, separating species by external 
characters can be difficult. Although subtle, 
the color and patterns of setae are usually 
distinctive. The coloration and patterns of 
setae on the elytra, the front and base of the 
rostrum, and on the epimeron and meso- 
and metepisterna are usually characteristic, 


VOLUME 107, NUMBER 3 


if subtly so. The tooth on the posterior fem- 
ora differs from very small to strong, as do 
the carinae on the femora. 


KEY TO ARCHOCOPTURUS SPECIES 


1. Elytra with relatively uniform distribution of 
setae on disc and no medial sutural spot, gla- 
brous at lateral margins, apices separately 
broadly rounded (Fig. 4); length < 2.5 mm; 
Costa Rica to Panama (South America?) ... . 

A. minutus, Nn. sp. 

— Elytra with more complex pattern of transverse 
fascia and medial sutural spot, apices separate- 
ly more or less angulate (Figs.1—3); length > 
3.0 mm. 

2. Elytral intervals narrower than striae, outer in- 
tervals subcarinate; face from above middle of 
eyes to antennal insertions on rostrum densely 
covered with yellowish (male) or opalescent 
blue (female) setae; México and Guatemala 

SES SEA GS RS ee et A. championi, n. sp. 

— Elytral intervals as wide as or wider than striae, 
never subcarinate; lower face and base of ros- 
trum with sparse white setae or only small ar- 

CAS Of GOS SALAS saccococcconnggocens 3 

3. Areas of opalescent blue scales on sides of 
pronotum separate for entire length, not joined 
in middle at apex (Fig. 1); design of white and 
yellow-brown setae on elytra very distinct; 
scales on epimeron pale yellow; tooth on pos- 
terior femur very small; length < 3.5 mm; Cos- 
ta Rica, Panama ........ A. laselvaensis, n. sp. 

— Areas of opalescent blue scales on sides of 
pronotum joined in middle at apex (Figs.2—3); 
design of setae on elytra more or less distinct; 
scales on epimeron dark orange brown; tooth 
on posterior femur large; length > 4.0 mm, 
México to South America 

4. Lower face and base of rostrum with sparse 
white setae; elytral intervals distinctly wider 
than striae; design of mostly white setae on 
elytra indistinct; México to Panama 

RED AAU A RaW ee Gh lated lone! oan A. medeterae, n. sp. 

— Lower face and base of rostrum with “‘mous- 
tache” of oblique lines of dense white setae 
from midline immediately below eyes to base 
of rostrum; elytral intervals subequal in width 
to striae; design of white and brown setae on 
elytra distinct; South America (French Guiana) 

3 6 5882 6 lot: Bidld o ol ere A. regalis (Boheman) 


Archocopturus laselvaensis Hespenheide, 
new species 
(Figs. 1, 5) 


Holotype male.—Black, except dark red- 
dish brown on antennae, spot on basal 2 of 


673 


each elytron between suture and humerus, 
and middle and posterior femora and tibiae; 
uniformly and densely covered with setae 
beneath and on legs, much more sparsely 
so above, rostrum glabrous except for sides 
at base. Color pattern complex: setae white 
on head, beneath, and on legs, setae on 
pronotum scale-like, opalescent blue on up- 
per sides and in triangle along midline at 
base; elytra with few small opalescent blue 
setae at bases of intervals 1—3 and behind 
scutellum; a few white setae anterior to hu- 
meri, on intervals 1—2 for basal %4, in spot 
on intervals 1—2 just beyond middle and ex- 
tending along interval | to apex; setae pale 
brown and sparse in striae on apical %, 
denser in transverse band at basal % on in- 
tervals 3-8 and in patch beyond apical % 
on intervals 2—4, extending to apex on in- 
terval 2; setae pale yellowish on meso- and 
metepisterna and anterior 4% of epimeron; 
3.4 mm long, 1.6 mm wide (Fig. 1). 

Pronotum with lateral margins very 
weakly arcuate, inconspicuously carinate 
along midline on basal %, somewhat wider 
than long. Elytra about % broader than 
pronotum, elongate, about % longer than 
wide, apices separately angulate at apex of 
interval 3. Abdomen with first ventral ster- 
nite with very shallow, medial, oval de- 
pression beyond middle. Posterior femur 
carinate, carina less distinct on basal '%, 
with small acute tooth on inner margin at 
apical ¥;. Aedeagus narrow, apex strongly 
deflexed in lateral view, apex narrowly 
rounded in dorsal view (Fig. 5). 

Allotype female.—As male, but abdo- 
men with first ventral sternite very weakly 
convex; 3.55 mm long. 

Holotype male.—Costa Rica: Heredia 
Pr., EF La Selva, 3 km S Pto. Viejo, 10°26'N 
84°01'W, 09.04.1983, H.A. Hespenheide 
(INBC). 

Allotype female.—Costa Rica: Heredia 
Pr., La Selva Biol. Sta., 3 km S Pto. Viejo, 
10°26’N_ 84°, 03.08.1993, M/01/164 (IN- 
BIOCRI002266014, INBC), 

Paratypes—COSTA RICA: Heredia Pr. 


same data as holotype but 29.07.1976, 


74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-4. Central American Archocopturus species. a, Dorsal habitus—left side with pattern of setae in- 
dicated (pronotum with areas of opalescent setae cross-hatched; elytra with pale brown setae stippled, white 
setae open), right side with reddish brown ground color stippled, otherwise black. b, Front of head and base of 
rostrum; all figures to same scale. 1, A. laselvaensis. 2, A. medeterae. 3, A. championi. 4, A. minutus. 


VOLUME 107, NUMBER 3 


27.03.1984 (2, CHAH); La Selva Biol. Sta., 
3 km S Pto. Viejo, 10°26’N 84°01’W, 16, 
23.07.1992, 18, 20, 23.07.1994, 23.08.1999, 
10.07.1994, Heliocarpus, 02.08.1996, balsa, 
14.07.1998, balsa log (15, CHAH), 19— 
24.07.1992, G. Wright, Malaise trap, second 
growth, SOC 1000 (2, CHAH), 15.03.1993, 
parcelas sucesionales, M/01/032 (1, INBC, 
INBIOCRIO002265677), 03.08.1993, parcelas 
sucesionales, M/01/164, (3, INBC, INBI- 
OCRI002266015—6, —9), La Selva Biol. Sta., 
nr Pto. Viejo, 08-17.08.1987, J. Brambila, 
Malaise (2, CWOB), E La Selva, 3 km S Pto. 
Viejo, 10°26'N 84°01'W, Est. Biol. La Selva, 
50-150 m, 10°26’N 84°0O1’W, 09.1992, 
P. Hanson & C. Godoy (1, MUCR), 
15.07.1993, M/01/152 (1, INBC, INBI- 
OCRIO002260837, INBC), 16.10.1995, M/O1/ 
471 (1, INBC, INBIOCRIO0O02300589), 
08.1992, Huertos (1, INBC, INBI- 
OCRIO01217035), 03.07.1993, FOT/07/28, 
Goethalsia meiantha (1, INBC, INBI- 
OCRI002068870), 24.08.1995, L.M. La- 
Pierre, Heurtos (1, INBC, INBI- 
OCRIO002055923), 25.08.1995, L.M. La- 
Pierre, STR 1,100 m (1, INBC, INBI- 
OCRI002055721), 3 km S Pto. Viejo, 
OTS-La Selva, 100 m, 10.1992, P. Hanson, 
Malaise trap (1, CWOB), 1 km NW Biol. 
Sta. La Selva, 50 m, 31.08.1998, C.W. & 
L.B. O’Brien (1, CWOB); Alajuela Prov., 
20 km S Upala,11—21.10.1991, ED. Parker 
(1, CWOB), Bijagua, 29.07.1990, W.F 
Chamberlain (1, TAMU), Est. Biol. San Ra- 
mon, 900 m, 07-08.1995, P. Hanson (1, 
MUCR); Guanacaste Pr., Buena Vista, Ho- 
tel Borinquen, 15.05.2003, J. & A. Rifkind, 
P Gum (2, CHAH); Prov. Limon, Sector 
Corocori, 30 km N de Cariari, Finca E. Ro- 
jas, 150 m, L-N-286000-567500, 03.1994, 
E. Rojas (2, INBC, INBIOCRIO001740728, 
—698), 04.1994, E. Rojas (5, INBC, IN- 
BIOCRIO0O01786147, —164-6, -—168), 
09.1994, E. Rojas (1, INBC, INBI- 
OCRI001996544), R.B. Hitoy Cerere, 
Send. hacia Rompe Pecho cerca Casa de 
madera, 100—200 m, L-N-184700-644200 
22—29.01.2002, W. Arana Gy; 
INB0003433272, INBC); Est. Hitoy Cer- 


675 


ere, R. Cerere, Res.Biol. Hitoy Cerere, 100 
m, L-N-184200-643300, 10.1992, G. Car- 
ballo (1, INBC, INBIOCRI000906668); 
Puntarenas Prov., R.E Golfo Dulce, 3 km S 
Rincon, 10 m, 10—12.1990, P. Hanson, Mal- 
aise trap (3, CWOB, MUCR), 24 mi NW 
Villa Neilly, 09.07.1974, O’Brien & Mar- 
shall (2, CWOB); San José Prov., 2—5 km 
W Ciudad Colon, 200—2,700', 05.12.1995, 
J. Rifkind, H. Lezama, on felled tree (1, 
CHAH). HONDURAS: Cortes: 18 km N 
Confradia, 05.08.1977, C.W. & L. O’Brien 
& Marshall (1, CWOB), | km W El Jaral, 
Finca Fe, 700 m, 31.07.1977, O’Briens & 
Marshall (1, CWOB). NICARAGUA: 
Chontales, T. Belt (3, BMNH). PANAMA: 
Tole, Champion (1, BMNH), Panama Prov., 
Canal Zone, Madden Forest, mi 5.0, 
09°07'N 79°38'W, 29.07.1971, W. Bivin (1, 
CHAH), Madden Dam, 09°13’'N 79°38’W, 
02.08.1971, H.A. Hespenheide, balsa 
t{ree]f[all] (4, CHAH), 09.08.1971, H.A. 
Hespenheide, Ochroma (3, CHAH), Ft. 
Sherman, 19.01.1980, D. Engleman (1, 
CWOB), Ft. Sherman, 09°21’N 79°59’W, 
21.07.1977, H.A. Hespenheide (1, CHAH), 
7 km W Margarita, 09°20'N 79°58’W (1, 
CHAH), 7 km S Gatun Lock, 22.05.1978, 
C.W. & L.B. O’Brien & Marshall (1, 
CWOB), Pan American Hwy, 30 km E 
Canita, 15—29.06.1992, J. & K. Ribardo (2, 
CSAGC)F Gero; biCampanas49n227 008 
27.05.1981, E. Giesbert (1, CHAH), Cerro 
Campana, 800 m (2, CHAH), Cerro Cam- 
pana, 850 m, 08°40’ N 79°56’ W (1, 
CHAH), 56 km E Chepo, 24.05.1981, D. 
Engleman, on woodfall (2, CWOB). In ad- 
dition to borrowed material, Paratypes from 
CHAH deposited in AMNH, CMNC, 
LACM, GBFM, NHRS, USNM. 
Etymology.—Named for the La Selva 
Biological Station where the holotype and 
allotype were collected, and location of the 
Arthropods of La Selva (ALAS) project. 
Discussion.—This species is somewhat 
smaller and less common than the very sim- 
ilar following species and is usually easily 
recognized by the pale yellowish setae on 
the epimeron and meso- and metepisterna. 


676 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Specimens vary in length from 2.7 to 4.0 
mm (mean = 3.55 mm, N = 83). 


Archocopturus medeterae Hespenheide, 
new species 
(Figs. 2, 6) 


Holotype male.—Black, except dark red- 
dish brown spot on basal % of each elytron 
between suture and humerus, and middle 
and posterior femora and tibiae; uniformly 
and moderately densely covered with setae 
beneath and on legs, much more sparsely 
so above, rostrum glabrous except for base 
below eyes. Color pattern complex: setae 
white and moderately dense on head, be- 
neath, and on anterior legs, sparser on mid- 
dle and posterior legs, setae on pronotum 
scalelike, opalescent blue on upper sides 
and across apex behind collar and in narrow 
ellipse along midline on basal %; elytra with 
very few small opalescent blue setae at ba- 
ses of intervals 2—3 and behind scutellum; 
few oval white setae anterior to humeri and 
on intervals 6—8 behind humeri, more 
densely on intervals and striae 1—2 for basal 
%, on interval 3 for basal ¥, in spot on in- 
tervals 1—2 just beyond middle, and on in- 
tervals 1—2 for apical %; long white hair- 
like setae in striae beyond the middle to 
apical ¥.; scattered pale brown setae else- 
where; setae dark orange brown on epime- 
ron, Mesepisternum and metepisternum; 4.3 
mm long, 2.0 mm wide (Fig. 2). 

Pronotum with lateral margins subparal- 
lel except at base and apex, inconspicuously 
carinate along midline, somewhat wider 
than long. Elytra about % broader than 
pronotum, elongate, about % longer than 
wide, shallowly transversely depressed be- 
hind scutellum; apices separately rounded- 
angulate at apex of interval 3. Abdomen 
with first ventral sternite somewhat flat- 
tened. Posterior femur carinate for entire 
legth, with large acute tooth on inner mar- 
gin at apical 7%, middle femur carinate for 
apical %4, anterior femur for apical 4%. Ae- 
deagus moderately broad, apex broadly 
rounded in dorsal view and slightly de- 
flexed in lateral view (Fig. 6). 


Allotype female.—As male, but abdo- 
men with first ventral sternite very weakly 
convex; 3.9 mm long. 

Holotype male.—Costa Rica: Heredia 
Pr., La Selva Biol. Sta., 3 km S Pto. Viejo, 
10°26'N 84°O1’W, 24.06.1991, H.A. Hes- 
penheide (INBC). 

Allotype female.—Costa Rica: same data 
as holotype but 25.07.1996 (INBC). 

Paratypes.—COSTA RICA: Heredia Pr., 
La Selva Biol. Sta., 3 km S Pto. Viejo, 02— 
04.1993, P. Hanson, Malaise trap (1, 
MUCR), same data as holotype but 25, 
26.07, 03,11.08.1996, 10.07.1994, on trunk 
of Ochroma lagopus (17, CHAH); 28.07— 
02.08.1992, G. Wright, Malaise trap, sec- 
ond growth, SOC 1000 (1, CHAH), Here- 
dia Pr, E La Selva, 3 km S Pto. Viejo, 
NOP ASIN SAO WE PSODMNSEO, 0) 
14.07.1982, 12.04.1984, 03.07.1985, 
29.03.1987, H.A. Hespenheide (8, CHAH), 
01.04.1987, M.M. Chavarria D. (1, 
CHAH), Heredia Pr., La Selva Biol. Sta., 3 
km S Pto. Viejo, 10°26’N 84°01'W, 
15.04.1991, H.A. Hespenheide (1, INBC, 
INBIOCRIO001216260), 15.04.1993, parce- 
las sucesionales, M/01/064 (2, INBC, IN- 
BIOCRIO02259218, —9), 15.07.1993, par- 
celas sucesionales, M/01/152, (2, INBC, 
INBIOCRI002260835-6), KE La Selva, 3 km 
S Pto. Viejo, 10°26’N 84°01'W, 25.03.1987 
(1, INBC, INBIOCRI0O01216775), Est. 
Biol. La Selva, 50-150 m, 10°26'N 
84°O1'W, 08.1992, Huertos (4, INBC, IN- 
OOO NANTSS, =o, —=S/, =O2D)- 
09.1992 (1, INBC, INBIOCRIO01222019), 
18.05.1993, parcelas sucesionales, M/0O1/ 
096, (1, INBC, INBIOCRI0022661231), 
03.08.1993, M/01/164, (1, INBC, INBI- 
OCRI002292826), 30.06.1995, M/01/387, 
(2, INBC, INBIOCRI002286946, —51), 
01.08.1995, M/01/411, (1, INBC, INBI- 
OCRI002289268), 05.03.1998, M/18/ 
703, Borde Suampo (1, INBC, INBI- 
OCRI002739513), 16.04.1998, M/18/706 
(1, INBC, INBIOCRI002283483), 
30.04.1998, M/18/707 (1, INBC, INBI- 
OCRIO002739680), 04.08.1999, M/19/728, 
Bosque secundario (2, INBC, INBI- 


VOLUME 107, NUMBER 3 


677 
0.5 mm 
Figs. 5-8. Male aedeagi of Archocopturus, dorsal (left) and lateral (right) views; all figures to same scale. 
5, A. laselvaensis. 6, A. medeterae. 7, A. champion. 8, A. minutus. 
OCRI002622972-3), 23.08.1999, M/19/ 04.10.1999, M/19/732 (2, INBC, INBI- 


729 (5, INBC, INBIOCRI002620363, 
—376, —386, —427, —464), 06.09.1999, M/ 
18/730 (7, INBC, INBIOCRI002620957, 
—=967; —10118,,—024; —216, =285, +342), 


OCRI002625011, —020), 01.11.1999, M/ 
19/734 (4, INBC, INBIOCRI002727105, 
0850-238; (257) SOS 199s cave 


LaPierre, dead Artocarpus altilis (5, INBC, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


INBIOCRIO02055822, —842—4, —855); 3 
km S Pto. Viejo, OTS-La Selva, 100m, 
11.1992, P Hanson, Malaise trap (1, 
CWOB), Pto. Viejo, OET La Selva, 100 m, 
01.1993, Hanson & Godoy (1, MUCR), 11 
km SE La Virgen, 450-550 m, 10°20'N 
84°14’W, Transect, 17—19.03.2003, M. 
Sharkey, Malaise, (1, INBC, 
INB0003237456), Horg. Sarapiqui, A.C. 
Cordillera Volcanica Central, 50 m, L-N- 
262100-542350, 07—08.1992, R. Vargas (1, 
INBC, INBIOCRIO01973853), Est. Mag- 
sasay, PN. Braulio Carillo, 200m, L-N- 
264600, 531000, 05.1991, M. Zumbado (2, 
INBC, INBIOCRIO01310609—10), L-N- 
264600, 531100, 12.1990, R. Aguilar (1, 
INBC, INBIOCRI0O00701597), Alajuela 
Prov., 20 km S Upala, 25—31.10.1990, ED. 
Parker (1, CWOB); Limon Prov., Zent, 
30.05.1956, E Lara, on cacao (1, BMNH), 
Prov. Limon, Manzanillo, RNFS Gandoca 
y Manzanillo, 0-100 m, L-N-398100- 
610600, 06—27.01.1993, EA. Quesada (1, 
INBC, INBIOCRIO01295981), Est. Hitoy 
Cerere, R. Cerere, Res.Biol. Hitoy Cerere, 
100 m, L-N-184200-643300, 07.1992, G. 
Carballo (1, INBC, INBIOCRIO00709964), 
R.B. Hitoy Cerere, Valle de La Estrella, 
100—200 m, L-N-184600-643400, 12.11— 
07.12.1993, G. Carballo, Malaise (1, INBC, 
INB0003496384); Sector Corocori, 30 km 
N de Cariari, Finca E. Rojas, 150 m, L-N- 
286000-567500, 03.1994, E. Rojas (1, 
INBC, INBIOCRIO01740994), 04.1994, E. 
Rojas (6, INBC, INBIOCRIO01786167, 
— 169-72, —198), Sector Corocori, 30 km 
N de Cariari, Finca de E. Rojas, A.C. Tor- 
tuguero, 150m, L-N-286000-567500, 
01,1994, E. Rojas (1, INBC, INBI- 
OCRIO01855910), Sector Corocori, 30 km 
N de Cariari, Finca de E. Rojas, 150 m, L- 
N-286000-567500, 07,1993, E. Rojas (1, 
INBC, INBIOCRI0O01699673), Sector Cer- 
ro Corocori, Fca de E. Rojas, 150m, L-N- 
286000-567500 26.03—24.04.1994, E. Ro- 
jas (2, INBC, INBIOCRI000735275, 
—767567); Prov. Puntlarenas]., Rancho 
Quemado, Peninsula de Osa, 200 m, L-N- 
292500, 511000, 10.1991, E Quesada (1, 


INBC, INBIOCRI000540684); Punt. Pr., 
vic. Rincon, Osa Pen., 28.07.1991, ET. Ho- 
vore (1, CHAH); San Jose Pr., 4 km N San 
Isidro, 29.07.1991, ET. Hovore (2, CHAH). 
BELIZE: Cayo, 22 mi SE Belmopan, 
18.08.1977, C.W. & L. O’Brien & Marshall 
(2, CWOB), 24 mi SE Belmopan, 16, 
18.08.1977, C.W. & L. O’Brien & Marshall 
(17, CWOB). HONDURAS: Atlantida, PN 
Pico Bonito, El Portillo, 29.06.2001, R. 
Turnbow (1, CWOB); Yoro, Ocotillo, 
30.07.95, R.D. Cave (1, EAPZ). MEXICO: 
San Luis Potosi, Huichihuayan, 18.06.1941, 
H.S. Dybas (1, CWOB), 8 mi N Huichi- 
huayan, 20.06.1941, H.S. Dybas (2, 
CWOB); Tabasco, 8 mi W Cardenas, 
07.10.1976, Cate & Clark (1, TAMU); Ve- 
racruz, Tezonapa, 08.08.1941, H.S. Dybas 
(1, CWOB), Penuela, 15.07.1941, H.S. Dy- 
bas (1, CWOB), Los Tuxtlas Biol. Sta. 
UINIZNIMIS® 250s iim, USLOS NOSIS, Co es Ik. 
O’Brien & G. Marshall (2, CWOB), Est. 
Biol Los Tuxtlas, 18°35’N 95°05'W, 02, 
04.05.1991, H.A. Hespenheide (2, CHAH), 
01—09.07.1988, J.A. Chemsak (1, EMEC), 
Veracruz, 9.5 mi SW Catemaco, 1,800’, 
16.07.1959, B. & B. Valentine (1, CHAH), 
Toxpam, Salle Coll. (3, BMNH). NICA- 
RAGUA: Rio San Juan Pr, Refugio Bartola, 
16 km ESE EI Castillo, 10°58/59'N 84°20/ 
21’W, 03.05.1999, H.A. Hespenheide (1, 
SEAN). PANAMA: Bocas del Toro Pr., 2— 
5 km W Almirante, road to Ojo de Agua, 
09°17'N 82°26'W, 06.07.1974, T.-L. Erwin, 
D.R. Whitehead (1, CWOB), Colon, Achi- 
ote Road, 26.09.1982, D. Engleman (1, 
CWOB), Canal Zone, Ft. Sherman, 
09°21'N 79°59'W, 21.07.1977, H.A. Hes- 
penheide (1, CHAH), Fort Sher- 
man,18.02.1992, ET, Hovore (2, CHAH), N 
shore Gatun Lake, 02.06.1984, E. Giesbert 
(1, CHAH), Margarita, Rancho Ramos, 
02.04.1984, D. Engleman (5, CWOB), Pan- 
ama Pr, 56 km E Chepo, 24.05.1981, D. 
Engleman, on woodfall (13, CWOB), 80 
km E Chepo, 09—20.05.1981, J.E. Wappes 
(1, CWOB), Canal Zone, Barro Colorado 
Iss) 1 O9S1OAN 7950. War 08: O79 70: 
05.04.1973, 01.06.1977 H.A. Hespenheide 


VOLUME 107, NUMBER 3 


(5, CHAH), Barro Colorado Is., 
22.01.1959, H. Dybas (1, CWOB), H. Wol- 
da, 18—23.01, 01—06.02.1987, 1B window 
trap, 18—23.12.1988, 08-13 [2], 30—31.01, 
14—-19.02, 27—30.03.1989, 3A window trap, 
04—09.01.1987, 13-18, 20—25, 27-—30.03, 
03-08, 10-15, 17—22.04, 01—06.05.1988, 
3B window trap (2, CWOB), H. Wolda, 
03.07.1977, 20.03, 19.08.1978, uv trap 1 (3 
m. high), Canal Zone, 7 km S Gatun Lock, 
22.05.1978, C.W. & L.B. O’Brien & Mar- 
shall (1, CWOB), Panama Prov., Pan Amer- 
ican Hwy, 30 km E Canita, 15—29.06.1992, 
J. & K. Ribardo (16, CASC), Panama Pr., 
12 km N El Llano, El Llano-Carti Rd., 
24.01.1993, ET. Hovore (1, CHAH), Pma. 
Prov., 9 km SE Bayano Bridge, 9°10'N 
78°46’ W, 08.09.1974, H.P.Stockwell (2, 
BMNH), [Chiriqui Prov.], Bugaba, Cham- 
pion (1, BMNH), Bugaba, 800-—1,500 ft. 
Champion (1, BMNH). In addition to bor- 
rowed material, Paratypes from CHAH will 
be deposited in AMNH, CMNC, CNCI, 
GBFM, LACM, MCZ, MNHN, NHRS, 
UNAM, STRI, USNM. 

Etymology.—Named for the putative 
models (Medetera spp., Dolichopodidae) of 
which this species and most Archocopturus 
are mimics (see discussion below). 

Discussion.—This is the largest and most 
widespread species in Central America, 
usually easily recognized by the dark or- 
ange brown setae on the epimeron and 
meso- and metepisterna. Specimens vary in 
length from 3.1 to 5.0 mm (mean = 4.32 
mm, N = 210). 


Archocopturus championi Hespenheide, 
new species 
(Bigs) 3247) 


Holotype male.—Black, except dark red- 
dish brown in irregluar transverse spot on 
each elytron behind humerus and before 
middle, antenna, and middle and posterior 
femora and tibiae; uniformly and densely 
covered with setae beneath and on anterior 
legs and below and lateral to eyes, sparser 
on middle and posterior legs, in distinct pat- 
tern above, rostrum glabrous except for 


679 


base above antennal insertions. Color pat- 
tern complex: setae small and yellowish on 
head, small and white on the legs, very 
large and white beneath, setae on pronotum 
scale-like, opalescent blue on upper sides 
and across apex behind collar and along 
midline, more broadly so before scutellum; 
elytra with very few small opalescent blue 
setae at bases of intervals 2—3; white setae 
anterior to humeri and in transverse fascia 
on intervals 6—10 behind humeri, on inter- 
vals and striae 1—3 for basal %, in chevron- 
like spot on intervals 1—2 and stria | at mid- 
dle, continuing on interval 1 to apex, and 
in trasverse fascia at apical % across inter- 
vals 1—4, continuing on interval 2 to apex; 
setae pale brown in trasverse fascia on in- 
tervals 3—6 at basal %; setae orange brown 
on epimeron, mesepisternum and anterior 2 
of metepisternum; 3.0 mm long, 1.4 mm 
wide (Fig. 3). 

Pronotum with lateral margins subparal- 
lel except at base and apex, inconspicuously 
carinate along midline, somewhat wider 
than long. Elytra about % broader than 
pronotum, elongate, about %4 longer than 
wide, striae broader than intervals, intervals 
5—10 subcarinate; apices separately, sharply 
angulate at apex of interval 3. Abdomen 
with first ventral sternite broadly, shallowly 
depressed. Posterior femur carinate for en- 
tire length, with very small obtuse tooth on 
inner margin beyond middle, middle femur 
carinate for apical 4, anterior femur for api- 
cal %. Aedeagus narrow, attenuate, apex 
abruptly deflexed in lateral view and nar- 
rrowly truncate in dorsal view (Fig. 7). 

Allotype female.—As male except setae 
on base of rostrum sparser and with opal- 
escent blue setae at sides and between bases 
of eyes; abdomen with first ventral sternite 
slightly convex; 3.3 mm long. 

Holotype male.—Guatemala: Pantaleon, 
1700 ft., Champion (BMNH). 

Allotype female.—Guatemala: same data 
as holotype (BMNH). 

Paratypes—GUATEMALA: same data 
as holotype (10, BMNH). MEXICO: Vera- 


80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


cruz, El Fortin, 02.07.1947, H.S. Dybas (1, 
CWOB). 

Etymology.—This species is named in 
honor of the collector of the holotype and 
author of the section on the “‘Zygopina”’ 
(now Conoderinae) in the Biologia Centra- 
li-Americana, G. C. Champion (1906). 

Discussion.—The extensive and dense 
area of setae on the face—and the opales- 
cent setae of the females—is very distinc- 
tive. The slender genitalia with the apically 
abruptly deflexed and truncate apex are 
quite different from those of other Archo- 
copturus and very similar to those of spe- 
cies of Cylindrocopturus Heller related to 
C. elongatus Champion. A very similar 
species has been seen from Rio de Janiero, 
Brasil. Specimens vary in length from 2.9 
to 3.5 mm with females averaging larger 
(males, mean = 3.11 mm, N = 5; females, 
mean = 3.35 mm, N = 8). The Mexican 
specimen is figured. 


Archocopturus minutus Hespenheide, 
new species 
(Figs. 4, 8) 


Holotype male.—Black, except dark red- 
dish brown on antenna, much of disc of 
each elytron behind scutellum and from su- 
ture halfway to lateral margins, more ex- 
tended laterally behind humerus, and mid- 
dle and posterior femora and tibiae; uni- 
formly and densely covered with setae be- 
neath, on legs and on head below eyes, 
much more sparsely so above, rostrum gla- 
brous from just above antennal insertions. 
Color pattern complex: setae white on head, 
beneath, and on legs, setae on pronotum 
scalelike, opalescent blue on upper sides; 
elytra with few white setae anterior to hu- 
meri, more on intervals 1—4 and included 
striae for basal ¥% and in transverse fascia 
on intervals 6—10 behind humeri; setae nar- 
row, elongate and white in striae 1—4 (and 
5 apically) on apical %4, setae hairlike and 
pale brown on intervals 1—5 (and 6 apical- 
ly); setae pale yellowish on mesepisternum 
and anterior % of metepisternum; 2.3 mm 
long, 1.1 mm wide (Fig. 4). 


Pronotum with lateral margins arcuately 
rounded, not carinate, coarsely punctate, 1— 
Y% wider than long. Elytra together about % 
broader than pronotum, about 7, longer 
than wide, apices separately broadly round- 
ed. Abdomen with first ventral sternite with 
small, glabrous, shallow, medial, oval de- 
pression beyond middle. Posterior femur 
carinate, with very small acute tooth on in- 
ner margin beyond middle, middle femur 
carinate, anterior femur for apical 4%. Ae- 
deagus gradually attenuate, apex deflexed 
in lateral view, apex narrowly rounded in 
dorsal view (Fig. 8). 

Allotype female.—As male, but abdo- 
men with first ventral sternite convex; uni- 
formly setose, 2.4 mm long. 

Holotype male.—Costa Rica: Heredia 
Pr., La Selva Biol. Sta., 3 km S Pto. Viejo, 
10°26'N 84°01'W, 05.05.1993, Conceveiba 
pleiostoma, FOT/04/ (INBC, INBI- 
OCRI002267368). 

Allotype female.—Costa Rica: same data 
as holotype (INBC, INBIOCRI002267367). 

Paratypes.—COSTA RICA: Heredia Pr., 
same data as holotype but 23.07.1998, H.A. 
Hespenheide, t[ree] flall] Protium pittieri 
(1, CHAH), 01.11.1993, Parcelas sucesion- 


ales, M/0%48 (1, INBC,  INBI- 
OCRIO002259263) 05.03.1993 Carapa gut- 
anensis, FOT/02/ (1, INBC, INBI- 


OCRI002267366); same data as holotype 
but ‘‘Flinca] La Selva,’ 11, 31.07.1982, 
H.A. Hespenheide, t[ree] f[all] Protium- 
Ochroma (5, BMNH, CHAH), 14.07.1982, 
H.A. Hespenheide, Protium t[ree] f[all] (2, 
CHAH); same data as holotype and “50— 
150 m, INBio-OET” 08.1992, (1, INBC, 
INBIOCRIO01217035), 14.01.1993, ex Vi- 
rola koshnyi, FVK/01/02 (1, INBC, INBI- 
OCRI001222049), 04.09.1993, Virola 
koshnyi, FVK/11/01, 07 (2, INBC, INBI- 
OCRI002068228, —260486), 09.11.1993, 
Virola koshnyi, FVK/15/05 (1, INBC, IN- 
BIOCRIO001240838), 15.10.1994, Virola 
koshnyi, FVK/24/36, 38 (2, INBC, INBI- 
OCRI002286363, —293533), 03.09.1993, 
Tapirira guianensis, FOT/10/04, 06, 07, 08, 
09 10, 15, 17 (10, INBC, INBIOCRI- 


VOLUME 107, NUMBER 3 


002068484, —485, —490, —494, —500, 
ON — OD lio 524) il 278961 ), 
08.01.1994, Pentaclethra macrloba, FPM/18/ 
37 (1, INBC, INBIOCRIO02260305), 
28.12.1999, Tachigalia costaricensis, FOT/ 
41/13 (1, INBC, INBIOCRI002725101), 
13.11.1997, M/18/690, Borde suampo (5, 
INBC, INBIOCRIO02282913, —922, 
=O50), Saas SVS WZ EGS I Vlesy/ 
693, Borde suampo (2, INBC, INBI- 
OCRI002283068, —070); Prov. Puntar- 
enas, Golfito, Res Ptal Golfo Dulce, Est 
Agujas, 250—350 m, 09.04.2000, A. Azo- 
feifa Interseccion, L.S.-276750_526550 
(1, INBC, INB0003086077); Puntarenas 
Prov., R.E Golfo Dulce, 3 km S Rincon, 10 
m, 09.1991, P. Hanson, Malaise trap (1, 
CWOB), Puntarenas Prov., R.E Golfo Dul- 
ce, 24 km W Piedras Blancas, 200 m, 10, 
11.1990, P. Hanson, Malaise trap (2, 
CWOB, MUCR), [Prov?] Hamburg Farm, 
Reventazon, ebene Limon, 30.05.1935, E 
Nevermann (1, USNM). PANAMA: 
C[anal] Z[one], Barro Colo[rado] Is, 
08.1946, Zetek, 5238 (1, USNM). In addi- 
tion to borrowed material, Paratypes from 
CHAH will be deposited in GBFM. 

Etymology.—Named for the small size 
of this species. 

Discussion.—As interpreted here, this is 
the most distinct species of Archocopturus 
in its size, the pattern of setae on the elytra, 
and in lacking the angulate elytral apices. 
The sexes do not differ significantly in 
length, and specimens vary in length from 
2.15 to 2.85 mm (mean = 2.46 mm, N = 
42). Almost half of the specimens have 
been collected at La Selva by canopy fog- 
ging of five different tree species, suggest- 
ing that the larval host is a canopy liana or 
epiphyte. Single specimens of two other 
species have been seen that are very similar 
to A. minutus, one of which is probably dif- 
ferent enough to require a separate, new ge- 
nus. 


Archocopturus regalis (Boheman) 
(Fig. 9) 

Copturus regalis Boheman 1845: 105. 

Archocopturus regalis: Heller 1895:56. 


681 


Diagnosis.—Black, except dark reddish 
brown on antenna, and middle and posterior 
femora; uniformly and densely covered 
with setae beneath, somewhat more sparse- 
ly so on legs, in distinct patterns above, ros- 
trum glabrous except for sides at extreme 
base. Color pattern complex: setae white on 
head, denser in distinct ‘“‘moustache”’ from 
the inner angles of the eyes to basal angles 
of rostrum, beneath, and largely on legs, se- 
tae on pronotum scalelike, opalescent blue 
on upper sides, joining in middle at apex, 
and in narrow triangle along midline at 
base; elytra with few small opalescent blue 
setae at bases of intervals and striae 1—2 
and behind scutellum; a few white setae an- 
terior to humeri, on intervals 1—3 for basal 
%, on interval 4 just at base, in spot on in- 
tervals 1—2 just beyond middle and extend- 
ing along interval | to apex; setae pale 
brown and sparse in striae on apical % ex- 
cept at apex, denser in transverse band at 
basal 4% on intervals (and striae) 3—10 and 
in patch beyond apical % on intervals 2—4, 
extending to apex on interval 2; setae pale 
yellowish on femora and on epimeron, me- 
sepisternum and anterior *% of metepister- 
num; 4.3 mm long, 1.9 mm wide. 

Pronotum with lateral margins very 
weakly undulate to subparallel, not carinate, 
about as wide as long. Elytra about % 
broader than pronotum, elongate, about 
longer than wide, apices separately angulate 
at apex of interval 3. Posterior femur cari- 
nate, with acute tooth on inner margin at 
apical ¥;; middle and anterior femora cari- 
nate for apical '. 

Holotype.—‘“*893/Cayen/Coll. Chevrol./ 
14/Archocopturus regalis (Boh.) determ: 
K.M. Heller,” by monotypy (NHRS). 

Discussion.—The sex of the type was not 
determined, but it is probably a female. Al- 
though Heller cites material from **Ama- 
zonas”> and Peru, these almost certainly 
represent species distinct from A. regalis as 
no Amazonian specimens seen by me 
match the unique type of that species. It is 
the only species I have seen with mous- 
divergent patches of 


tachelike narrow, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 9. Archocopturus regalis. a, Dorsal habitus of holotype (as in Figs. 1—4). b, Front of head and base of 


rostrum. 


dense white setae above the base of the ros- 
trum. 


Zygopsella Champion 


Zygopsella Champion 1906: 42. Type spe- 
cies: Zygopsella ruficauda Champion, by 
monotypy. 


Champion differentiated Zygopsella from 
Archocopturus because the former pos- 
sessed “‘much smaller, inferiorly acuminate 
eyes, [a] narrow, acuminate antennal club, 
and ... equally unidentate femora.’ Each 


elytron also possesses a more or less pro- 
nounced dorsoventrally flattened and api- 
cally rounded lobe that projects from the 
apex from the second through the fifth ely- 
tral intervals. 


Zygopsella ruficauda Champion 
Zygopsella ruficauda Champion 1906: 42. 


Specimens examined.—COSTA RICA: 
Prov. Guanacaste, Est. Palo Verde, PN. 
Palo Verde, 10 m, L-N 259000, 888400, 
25.03—21.04.1992, M. Ortiz (INBC, INBI- 


VOLUME 107, NUMBER 3 


OCRIO00786987); Prov. Heredia, 16 km 
SSE La Virgen, 1,050—-1,150 m, 10°16'’N 
84°05’W, 21.03.2001, Transect, 11/TN/16/ 
016 CUNBC, INB0003209199). GUATE- 
MALA: Vera Paz, Senahu (2 Syntypes, 
BMNH); Esc., 10 km S_ Esquintla, 
20.06.1985, W.E. Clark (CWOB). MEXI- 
CO: Veracruz, Est. Biol. de Los Tuxtlas, 
160 m SNM, 09.07.1995, A. Ibarra 
(UNAM). PANAMA, Panama Pr., Cerro 
Jefe, 700 m, 27.03.1976, H.P. Stockwell 
(STRD 

Discussion.—The distribution of this 
species is puzzling, with a very small num- 
ber of specimens known from localities that 
are widely separated geographically and 
very different ecologically. Most are from 
wet middle elevation sites on the Caribbean 
slopes, but the Palo Verde specimen is from 
a Pacific lowland site with a strong dry sea- 
son. 


Zygopsella pulchella (Hustache), 
new combination 


Archocopturus pulchellus Hustache 1932: 
28. Syntypes: “Guadeloupe: Trois-Rivi- 
éres, Gourbeyre, Dufau; Clairieres, vers 
600m, Dufau,”’ not seen. 


Specimen examined.—GUADELOUPE: 
BasseTerre, Bras-David, Rte. de la Tra- 
Vemscue OSs 1965s" C.W. ce IE B: ‘© Brien 
(CWOB, det. C. O’Brien). 

Discussion.—Hustache (1932) described 
two species from Guadeloupe in the genus 
Archocopturus, basing their assignment to 
the genus on the subequal first and second 
funicular segments and the possession of 
metallic blue-green scalelike setae on the 
pronotum, neither of which is characteristic, 
as discussed above. ““Archocopturus” pul- 
chellus is very close to Zygopsella ruficau- 
da, differing only in details in the pattern 
of setae. 


Macrocopturus basalis (Hustache), 
new combination 


Archocopturus basalis Hustache 1932: 29 


683 


Specimens examined.—GUADE- 
LOUPE: BasseTerre, Sofaia, 6 km SW Ste. 
Rose, 26.05.1985, C.W. & L.B. O’Brien 
(CWOB, det. C. O’Brien, ““compared with 
type’), Trois-Rivieres, Gourbeyre, Dufau 
(syntype, MNHN). PUERTO RICO: Mari- 
cao Forest, 2—3,000 ft., 30.05—02.06.1938, 
Darlington (MCZ). 

Discussion.—The other species Hustache 
misassigned to Archocopturus is transferred 
here to Macrocopturus Heller. Its diminu- 
tive size superficially suggests the genus 
Eulechriops Faust, but the mesosternum of 
basalis is unmodified. The genus Macro- 
copturus as currently constituted is clearly 
not a monophyletic group, but is very 
large—over 100 species are known from 
the La Selva Biological station (Hespenhei- 
de, unpublished)—and will require exten- 
sive further study. The specimen from 
Puerto Rico is the first record from the An- 
tilles other than Guadeloupe. 


DISCUSSION 


Flies as models for mimicry: Medetera 
ecology and behavior.—Earlier I suggested 
that a recurring color pattern of red head, 
variegated black pronotum, and pale elytra 
among a number of unrelated beetle groups 
might be explained as mimicry of flies 
(Hespenheide 1973). Flies with red eyes 
were hypothesized to be avoided by bird 
predators because of the difficulty of cap- 
turing them. Many of the beetles were con- 
oderine weevils which commonly were ob- 
served perching on the sides of tree trunks. 
Although I discussed Archocopturus as part 
of that complex, I later (Hespenheide 1995) 
distinguished a second complex of beetle 
mimics that had the pronotum metallic blue 
(rather than variegated black) and usually 
lacked the red and suggested that these 
were mimics of dolichopodid flies of the 
genus Medetera. 

Flies of the genus Medetera are predators 
as larvae on the larval forms of wood-bor- 
ing insects and some species often are very 
common on recently-fallen trees or perched 
on the sides of standing trees in the vicinity 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


of tree falls (Bickel 1985, 1987). Many con- 
oderine beetles are wood borers as larvae 
and are found in the same situations. I have 
previously pointed out the importance of 
the association of model and mimic in the 
same microhabitat (Hespenheide 1996a). 
Adult Archocopturus in particular are often 
seen at tree falls, and some individuals 
show a characteristic behavior of rapidly 
palpitating their tarsi; on one occasion I 
have observed similar behavior by a species 
of Medetera. 

Other participants.—At least two other 
species complexes of conoderine weevils 
participate in the mimicry of Medetera in 
Central America, Macrocopturus lampro- 
thorax (Heller) and Copturomimus caeru- 
leotictus Champion. As with Archocoptu- 
rus, both have the metallic blue setae on the 
pronotum and both are complexes of sibling 
species, rather than a single species. Ma- 
crocopturus lamprothorax is a complex of 
at least 18 species differing in subtle details 
of pattern and male genitalia; Copturomi- 
mus caeruleotinctus is a complex of at least 
three species. As with ‘Archocopturus re- 
galis,”’ there are only two names for at least 
21 species. Specimens of two additional un- 
described species that may represent unde- 
scribed genera also are known to me. 

Another mimicry complex in conoderine 
weevils involves metallic blue setae on the 
base of the pronotum and red setae on the 
anterior portion (Hoplocopturus scintillans 
Champion complex). Despite the presence 
of the metallic blue setae, this complex 
seems to be associated with the specific mi- 
crohabitat of aroid roots rather than tree 
trunks and probably does not have Mede- 
tera species as models (Hespenheide 
1996a). 

Mimicry and biodiversity.—The effect of 
selection in the evolution of mimicry is to 
increase the similarity of mimic to models 
and, consequently, of mimics to one anoth- 
er, creating mimetic homoplasy when more 
distantly related species independently 
evolve a mimetic pattern. Alternatively, se- 
lection for mimetic resemblance will pre- 


vent significant divergence in cases when 
an already-mimetic species undergoes spe- 
ciation, creating sibling species. Without 
detailed morphological and/or genetic anal- 
yses it is unknown, of course, which resem- 
blances between species actually are due to 
convergent evolution for mimicry (mimetic 
homoplasy) and which are due to ancestry 
(sibling species). In either case, close mor- 
phological similarity means that true differ- 
ences between species will be overlooked 
in poorly studied taxa, which has certainly 
been the case for the highly diverse Neo- 
tropical insect fauna, so that many taxa are 
undescribed. For the taxa involved in the 
mimicry of Medetera, all four Central 
American Archocopturus are undescribed, 
and 21 of 24 other species and two of five 
other genera; that is, 89% of the species and 
3 of the genera. 

Much of the material available for this 
study has come from the Arthropods of La 
Selva (ALAS) Project at La Selva Biolog- 
ical Station in Costa Rica. Of the 28 species 
and six genera known to me and hypothe- 
sized to mimic Medetera, 21 species and 
four of the genera have come from ALAS 
or La Selva collections. Intensive collection 
efforts at other sites will doubtless yield 
many additional taxa, especially in South 
America. It is daunting to contemplate how 
little we know of tropical insects (Hespen- 
heide 2001). 


ACKNOWLEDGMENTS 


In addition to the curators at the collec- 
tions which loaned specimens, cited earlier, 
Chris Lyal, Sharon Shute, and Max Barclay 
assisted during visits to The Natural History 
Museum, London, and Bert Viklund ar- 
ranged the loan of the type of Archocop- 
turus regalis from Stockholm. Charles W. 
O’Brien shared expertise and generously al- 
lowed study in his personal collection. Dan- 
iel J. Bickel, Australian Museum, Sydney, 
Australia, provided information on Mede- 
tera. Margaret Kowalczyk prepared the fi- 
nal illustrations. I have been supported 
through the ALAS Project at La Selva (Na- 


VOLUME 107, NUMBER 3 


tional Science Foundation grants BSR 
9025024, DEB 9401069, DEB 9706976, 
and DEB-0072702), by small grants from 
the UCLA Academic Senate, and exten- 
sively by personal funds. 


LITERATURE CITED 


Bickel, D. J. 1985. A revision of the Nearctic Mede- 
tera (Diptera: Dolichopodidae). United States De- 
partment of Agriculture, Agricultural Research 
Service Technical Bulletin No. 1692 i—v, 109 pp. 

. 1987. A revision of the Oriental and Austra- 
lasian Medetera (Diptera: Dolichopodidae). Re- 
cords of the Australian Museum 39: 195-259. 

Boheman, C. H. 1845. Jn Schoenherr, C. J., Genera et 
species curculionidum, Vol. 8, pt. 2: 1- 504. 

Champion, G. C. 1906. Curculionidae, Curculioninae, 
Zygopina. In Godman and Salvin, Biologia Cen- 
trali-Americana, Insecta, Coleoptera, IV, 5: 1-130. 

Heller, A. 1895. Zygopiden-Studien. II, mit besonderer 
Berticksichtigen der Gattung Copturus. Abhan- 
dlungen und Berichte des k6niglichen zoologisch- 


685 


en und anthropologish-Ethnographischen Muse- 
ums zu Dresden 11: 1—70. 

Hespenheide, H. A. 1973. A novel mimicry complex: 
Beetles and flies. Journal of Entomology (Lon- 
don), A, 48: 49-S6. 

. 1995. Mimicry in the Zygopinae (Coleoptera: 

Curculionidae). Memoirs of the Entomological 

Society of Washington No.14, pp. 145-154. 

. 1996a. The role of plants in structuring com- 

munities of mimetic insects, pp. 109-126. In, Gib- 

son, A., ed. Neotropical Biodiversity and Conser- 
vation, Mildred Mathias Botanical Garden, Los 

Angeles, California. 

. 1996b. Clytrine chrysomelids as models of 

mimicry complexes, pp. 227—239. Jn Jolivet, P. H. 

L. and M. L. Cox, eds., Chrysomelidae Biology, 

2 Ecological Studies,. SPB Academic Publishing, 

The Hague. 

. 2001. Beetles, pp. 351—358. Jn S. Levin, ed. 
Encyclopedia of Biodiversity, Vol. 1. Academic 
Press, San Diego, California. 

Hustache, A. 1932 [‘*1931’’]. Curculionides de la Gua- 
deloupe, troisieéme partie. Faune des Colonies 
Frangaises. 142 pp. 

Mayr, E. 1963. Animal Species and Evolution. Har- 
vard University Press, Cambridge, Massachusetts. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 686-692 


THE TAXONOMIC PLACEMENT OF SEVERAL NEW WORLD AND 
ORIENTAL GASTERUPTIID WASPS (HYMENOPTERA: GASTERUPTIIDAE) 


JOHN T. JENNINGS AND DAviD R. SMITH 


(JTJ) Centre for Evolutionary Biology and Biodiversity, and School of Agriculture and 
Wine, Waite Campus, The University of Adelaide, SA 5064, Australia (e-mail: 
john.jennings @ adelaide.edu.au); (DRS) Systematic Entomology Laboratory, PSI, Agri- 
cultural Research Service, U.S. Department of Agriculture, % National Museum of Nat- 
ural History, Smithsonian Institution, PO. Box 37012, MRC-168, Washington, DC 20013- 
7012, U.S.A. (e-mail: dsmith @sel.barc.usda.gov) 


Abstract.—The generic placement of several New World and Oriental gasteruptiid 
wasps is reviewed. Based on a number of character states, including non-overlapping 
mandibles (when at rest), the presence of a trochantellus (prefemur) and an exserted 
Ovipositor, and the pattern of the veins in the forewing, five species currently in Pseu- 
dofoenus are transferred to Gasteruption L.; namely G. longiceps (Kieffer 1910), n. 
comb., G. ceylonensis (Enderlein 1912), n. comb., G. maculicorne Cameron 1887, re- 
vised comb., G. sericeum Cameron 1887, revised comb., and G. tenuicolle Schletterer 
1885, revised comb. In addition, G. longiceps (Kieffer 1910) is a junior secondary hom- 
onym of G. longiceps Kieffer 1904 and is renamed G. austini, n. name, and Pseudofoenus 


neotropicus Schrottky 1906 is placed as incertae sedis. 


Key Words: 


Members of the family Gasteruptiidae 
are predator-inquilines of solitary bees and 
wasps, and are currently divided into two 
subfamilies, Gasteruptiinae, comprising the 
single genus Gasteruption L., and Hyptio- 
gastrinae comprising two genera, Hyptio- 
gaster Kieffer and Pseudofoenus Kieffer 
(Jennings and Austin 2002). 

Pseudofoenus was described by Kieffer 
(1902) to accommodate three Gasteruption 
species (P. unguiculatum (Westwood), P. 
unguicularae (Smith), and P. pedunculatum 
(Schletterer)), all from New Zealand. In his 
revision of 1912, Kieffer included 16 spe- 
cies, while the catalog by Hedicke (1939) 
included 17 species. However, both Kieffer 
and Hedicke had a broad concept of Pseu- 
dofoenus that failed to distinguish conver- 
gent wing venation among members of the 


Hymenoptera, Gasteruptiidae, Gasteruptiinae, Gasteruption, Pseudofoenus 


Gasteruptiidae (see Jennings and Austin 
2002). 

In the 1950s, a number of taxa described 
in Pseudofoenus were transferred to Gas- 
teruption. Pasteels (1956) transferred P. 
azurescens Szépligeti, P. braunsi Kieffer, 
P. keifferi Szépligeti, P. latigenalis (Schlet- 
terer), P. nasutus Szépligeti, and P. vires- 
cens (Enderlein); Pasteels (1957) trans- 
ferred P. fluvialis Turner, P. cylindricus 
Turner, and P. isthmalis Turner; and Pas- 
teels (1958) transferred P. manilensis Kief- 
fer. 

Crosskey (1962), in a comprehensive re- 
assessment of the Gasteruptiidae, provided 
much needed stability to the generic clas- 
sification that lasted for 40 years. Based pri- 
marily on wing venation, he included only 
five species from New Zealand in Pseudo- 


VOLUME 107, NUMBER 3 


foenus; P. crassipes (Smith), P. nocticolor 
Kieffer, P. pedunculatus (Schletterer), P. 
unguicularis (Smith), and P. unguiculatus 
(Westwood). He also transferred P. angus- 
tatus Kieffer from Mexico to Gasteruption. 
Later, P. kaweahensis (Bradley) from North 
America was transferred to Gasteruption by 
Carlson (1979). Jennings and Austin (1994) 
revised Pseudofoenus but largely adapted 
Crosskey’s concept of the genus. Interest- 
ingly, the four included species, P. crassi- 
pes, P. nocticolor, P. pedunculatus, and P. 
unguiculatus, represent the male and female 
of only two sexually dimorphic species, but 
to date it has not been possible to properly 
associate them. 

Following a substantial phylogenetic re- 
vision of the mainly Australian subfamily 
Hyptiogastrinae, Jennings and Austin 
(2002) redefined the limits of Pseudofoenus 
to include all hyptiogastrine wasps with a 
short, robust, upwardly curved ovipositor, 
normally hidden by ovipositor sheaths. 
They included 78 species from Australia, 
New Zealand, Papua New Guinea, south- 
western Pacific islands (Fiji, New Caledon- 
ia, and Vanuatu), and South America. 
Those hyptiogastrines with an exserted ovi- 
positor were included in the Australian ge- 
nus Hyptiogaster (Jennings and Austin 
1997a). 

However, several New World and Ori- 
ental taxa previously included in Pseudo- 
foenus by Hedicke (1939) have not been 
examined since, and their generic position 
has remained in doubt. They are: P. ma- 
culicornis (Cameron) and P. sericeus 
(Cameron) from Guatemala, P. longiceps 
Kieffer and P. neotropicus Schrottky from 
Paraguay, and P. tenuicollis (Schletterer) 
from Mexico. Further, and overlooked by 
Hedicke (1939), the generic placement of 
P. ceylonensis Enderlein, 1912, from Sri 
Lanka is also in doubt. Here, we present the 
results of a re-interpretation of these taxa 
and discuss their placement in the Gaster- 
uptiidae. 


687 


METHODS, TERMINOLOGY AND 
ABBREVIATIONS 


Specimens were observed either under a 
Zeiss light microscope or a NZ16 Leica 
light microscope equipped with a Nikon 
DXM 1200 digital camera. Images were 
processed with Auto-Montage 4.02. 

Terminology for wing venation follows 
the modified Comstock-Needham system 
after Sharkey (1988), with some modifica- 
tions, including Achterberg’s (1979) no- 
menclature for cells (Fig. 1; also see Jen- 
nings and Austin 1994). 

Abbreviations for institutional reposito- 
ries cited in this paper are: BMNH (The 
Natural History Museum, London, United 
Kingdom) and DEI (Deutsches Entomolo- 
gisches Institut, Miincheberg, Germany). In 
an attempt to locate types, we contacted 
many other collections that may have been 
the repository of type material. 


KEY TO SUBFAMILIES OF GASTERUPTIIDAE 


The Gasteruptiidae are currently divided 
into two subfamilies, Gasteruptiinae, com- 
prising the single species-rich genus Gas- 
teruption L. and Hyptiogastrinae compris- 
ing two genera, Hyptiogaster and Pseudo- 
foenus (see Jennings and Austin 2002 for 
diagnoses and discussion of the two sub- 
families). The two subfamilies may be iden- 
tified using the following key. 


1. Mandibles long and broadly overlapping when 
in closed position; trochantellus (prefemur) ab- 
sent; first discal cell formed by forewing vein 
1-Rs+M intersecting basal cell about one-quar- 
ter to one-third distance from M+Cu (see Fig. 
Ic) or, rarely, first discal cell absent and veins 
I1-Rs+M and 1-Cu(b) fused to form 
Rs+M-+Cu(b), veins 1-M and m-cu absent (see 
Fig. 1b) [New Zealand hyptiogastrines only]; 
female subgenital sternite simple; hind tro- 
chanter with or without a groove; ovipositor 
usually short and hidden at rest (Pseudofoen- 
us), or exposed and at least 0.25 length of me- 
tasoma (Hyptiogaster) Hyptiogastrinae 

— Mandibles short and not broadly overlapping 
when in closed position; trochantellus (prefe- 


mur) usually present (Figs. 2a, b), sometimes 
indicated by a slightly differentiated basal 


swelling; first discal cell formed by forewing 


88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ere 


1-R1 


1-R1 


= 


b 


Fig. 1. Forewings of Gasteruptiidae. a, Gasteruption sp. from Australia. b, Pseudofoenus crassipes from 
New Zealand. c, P. thoracicus (Guérin-Menéville) from Australia. | = marginal cell, 2 = submarginal, 3 = 
discal, 4 = subdiscal, 5 = costal, 6 = basal, 7 = subbasal, 8 = plical. Scale bar = 1 mm. 


VOLUME 107, NUMBER 3 


vein 1-Rs+M forming node at 1-Rs, M+Cu 
and 1-Cu, vein 1-M absent (see Fig. la) [sev- 
eral species with 1m-cu absent, or nearly so, 
such that the first discal cell is triangular]; fe- 
male subgenital sternite notched or slit; hind 
trochanter with a groove; ovipositor long, at 
least 0.5 length of metasoma (Gasteruption) 
Gasteruptiinae 


TREATMENT OF SPECIES 


Gasteruption austini Jennings and 
Smith, new name 


Pseudofoenus longiceps Kieffer 1910: 242; 
Kieffer 1912: 207; Hedicke 1939: 46. 


Holotype.—Missing. Kieffer (1910) de- 
scribed a female from Villa Morra, Para- 
guay, 19 Dec. 1904, collected by J. D. An- 
isits. 

Based on Kieffer’s (1910) original de- 
scription, the forewing venation of P. lon- 
giceps is atypical of the majority of Gas- 
teruption in lacking the first discal cell, and 
thus resembles the New Zealand taxa P. 
crassipes (Smith) (Fig. 1b) and P. pedun- 
culatus (Schletterer). Crosskey (1962) and 
Jennings and Austin (1994) discuss the pos- 
sible evolutionary pathways leading to the 
loss of the first discal cell. Other than those 
taxa from New Zealand, all hyptiogastrines 
have the first discal cell formed by forewing 
vein 1-Rs+M intersecting the basal cell 
about one-quarter to one-third the distance 
from M+Cu (Jennings and Austin 2002). 
Kieffer (1910) also indicated that the ovi- 
positor is exserted and about the same 
length as the metasoma. Given the combi- 
nation of having an exserted ovipositor and 
the pattern of the wing venation, we trans- 
fer P. longiceps to Gasteruption, n. comb. 
As the species epithet is a junior secondary 
homonym of G. longiceps Kieffer 1904 
from Western Australia, we propose the 
new replacement name P. austini, after 
Prof. Andrew Austin, The University of 
Adelaide. 


Gasteruption ceylonensis (Enderlein), 
n. comb. 


Pseudofoenus ceylonensis Enderlein 1912: 
388; Hedicke 1939: 45. 


689 


levee, 2. 
ceylonensis, 2, holotype. b, G. sericeum, 
trochantellus (prefemur). t = trochantellus. Scale bars; 
a = 0.5 mm, b = 1.0 mm. 


Hind trochanter and femur. a, Gasteruption 


O 


?, showing 


Holotype.—@, “‘Penilankulam (illeg.), 
Ceylon, W. Horn, [18]99” (DEI). Poor con- 
dition; most of the antennal segments and 
metasoma missing. Enderlein (1912) indi- 
cated that the holotype was deposited in Vi- 
enna, but this was an error. 

Although the forewing venation of this 
species is atypical of most Gasteruption in 
that the first discal cell is absent (see Fig. 
1b), there is, however, a distinct trochantel- 
lus on the hind leg (Fig. 2a) and the man- 
dibles do not overlap at rest. Unfortunately, 
the ovipositor is missing on the holotype, 
but Enderlein (1912) indicated in the orig- 
inal description that it is clearly exserted 
beyond the tip of the metasoma. Based on 
these character states, we transfer it to Gas- 
teruption. 


Gasteruption maculicorne Cameron, 
revised comb. 


Gasteruption maculicorne Cameron 1887: 
424, pl. 18, fig. 5; Schletterer 1889: 475. 
Pseudofoenus maculicornis: Kieffer 1912: 
207; Hedicke 1939: 46. (Note: Bradley 
1909: 109 stated maculicorne, but in er- 
rata on p. 194, he corrected it to seri- 
ceum, therefore, maculicorne on p. 109 
is a mistake.) 


10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Holotype.— ‘‘Calderas, Guatemala, 
Champion” (BMNH 3.a.198). Poor condi- 
tion; most of the antennal segments, right 
forewing, and metasoma missing. Left fore- 
wing glued to card. 

Cameron (1887) described this species in 
Gasteruption, but Kieffer (1912) transferred 
it to Pseudofoenus. It clearly belongs in 
Gasteruption based on the forewing vena- 
tion, which is typical of the majority of 
Gasteruption (see Fig. la), the presence of 
a trochantellus on the hind leg, and man- 
dibles that do not overlap at rest. Although 
the ovipositor is missing on the holotype, 
Cameron’s figure (1887: pl. 18, fig. 5) 
shows that it is exserted well beyond the tip 
of the metasoma. On the basis of these 
character states, we return this species to 
Gasteruption. 


Gasteruption sericeum Cameron, 
revised comb. 


Gasteruption sericeum Cameron 1887: 424, 
pl. 18, figs. 3, 4; Schletterer 1889: 473; 
Kieffer 1902: 10. 

Rhydinofoenus sericeus: Bradley 1909: 39. 

Pseudofoenus sericeus: Bradley 1908: 109, 
194 (as maculicorne on p. 109, corrected 
to sericeus on p. 194); Kieffer 1912: 208; 
Hedicke 1939: 46. 


Syntypes.—Missing. Described from 
both sexes; thus, Cameron had two or more 
specimens, but no holotype was designated. 
The type locality is San Ger6nimo, Guate- 
mala. 

This species, originally described in Gas- 
teruption, was transferred to Pseudofoenus 
by Bradley (1908) and then to Rhydino- 
foenus Bradley by Bradley (1909), based 
largely on the lack of forewing vein 1m-cu, 
resulting in the absence of the first discal 
cell. Rhydinofoenus was later synonymised 
with Pseudofoenus (Hedicke 1939). Al- 
though the syntypes are apparently missing, 
there are three specimens in the BMNH, 1 
2 and 2 6 from San Ger6énimo, Guatemala, 
collected by Champion. Because Cameron 
(1887) described both sexes, these with 


Cameron’s determination label in his hand- 
writing undoubtedly are part or all of the 
type series and may not have been separat- 
ed as types in the BMNH. 

Although the forewing venation of this 
species is atypical of most Gasteruption 
(see Fig. 1b), it has a distinct trochantellus 
on the hind leg (Fig. 2b) and the mandibles 
do not overlap at rest, both characteristic of 
Gasteruption. The original description and 
Cameron’s figure (1887: pl. 18, fig. 3) clear- 
ly indicate that the ovipositor is 1.5X as 
long as the metasoma (Cameron 1887). On 
the basis of these character states, we return 
this species to Gasteruption. 


Gasteruption tenuicolle Schletterer, 
revised comb. 


Gasteruption tenuicolle Schletterer 1885: 
291: Schletterer 1889: 471; Kieffer 1902: 
10. 

Pseudofoenus tenuicollis: Szépligeti 1903: 
8655) Kiefter 1912-9 205-sHedickegl9s9- 
46. 


Holotype.—Missing. Schletterer (1885) 
described a female from “‘Mexiko (Orizaba, 
Bilimek).”’ 

This species was originally placed in 
Gasteruption (Schletterer 1885) but later 
transferred to Pseudofoenus by Szépligeti 
(1903). The original description (Schletterer 
1885) indicates that the ovipositor is ex- 
serted, and based on this we transfer this 
species back to Gasteruption. 

In the DEI collection, there is a female 
from San José, Costa Rica, collected by 
Heinr. Schmidt, 2.x.[19]27 and labelled as 
tenuicolle. The forewing venation of this 
specimen is typical of most Gasteruption in 
that the first discal cell is formed by vein 
1-Rs+M resulting in a node at 1-Rs, M+Cu 
and 1-Cu, and vein I-M is absent (see Fig. 
la). In addition, it has a trochantellus on the 
hind leg and mandibles that do not overlap 
at rest, and an exserted ovipositor. Although 
we are not convinced that the specimen has 
been correctly identified, it clearly belongs 
to Gasteruption. 


VOLUME 107, NUMBER 3 


Pseudofoenus neotropicus Schrottky 


Pseudofoenus neotropicus Schrottky 1906: 
62; Kieffer 1912: 206; Hedicke 1939: 46. 


Holotype.—Missing. Described from a 
male collected 28 Jan 1905, from ‘Villa 
Encanacion,” Paraguay. 

This species was originally placed in 
Pseudofoenus (Schrottky 1906), presum- 
ably based on forewing venation, 1.e., the 
absence of vein I|m-cu resulting in the ab- 
sence of the first discal cell. This form of 
wing venation is, however, not found in ei- 
ther of the two South American species of 
Pseudofoenus, P. deletangi (Schletterer) or 
P. infumatus (Schletterer), both of which 
have a typical hyptiogastrine wing pattern 
(see Fig. la) (Jennings and Austin 1997b, 
2002). In all probability, this species be- 
longs in Gasteruption. However, as the ho- 
lotype is missing, no extant specimens can 
be found that match the original descrip- 
tion, and the lack of evidence relating to, in 
particular, wing venation and whether the 
ovipositor is exserted, we place this species 
in incertae sedis. 


ACKNOWLEDGMENTS 


We thank the curators of the BMNH and 
DEI collections for access to types and oth- 
er material, and the curators of many other 
collections for trying to locate type mate- 
rial. We also thank Andy Austin and Nick 
Stevens, The University of Adelaide, and 
Thomas J. Henry, Systematic Entomology 
Laboratory, USDA, Washington, DC, for 
their comments on the manuscript. 


LITERATURE CITED 


Achterberg, C. van 1979. A revision of the subfamily 
Zelinae auct. (Hymenoptera, Braconidae). 
Tijdschrift voor Entomologie 122: 241—479. 

Bradley, J. C. 1908. The Evaniidae, ensign- flies, an 
archaic family of Hymenoptera. Transactions of 
the American Entomological Society 34: 101-— 
194. 

. 1909. A new genus and two new species of 
Foeninae from Sonoran California. (Hym.). Deut- 
sche Entomologische Zeitschrift 1909: 37-41. 

Cameron, P. 1887. Insecta. Hymenoptera (families 


691 


Tenthredinidae-Chrysididae), Vol. 1. Jn Godman 
and Salvin, Biologia Centrali-Americana, 1883-— 
1900, 473 pp., 20 pls. 

Carlson, R. W. 1979. Superfamily Evanioidea, pp. 
1109-1118. In Krombein, K. V., P. D. Hurd, Jr, 
D. R. Smith, and B. D. Burks, eds. Catalog of 
Hymenoptera in America North of Mexico, Vel. 
1. Symphyta and Apocrita (Parasitica). Smithson- 
ian Institution Press, Washington, D.C. pp. 1-— 
1198. 

Crosskey, R. W. 1962. The classification of the Gas- 
teruptiidae (Hymenoptera). Transactions of the 
Royal Entomological Society of London 114: 
377-402. 

Enderlein, G. 1912. Uber einige Evaniiden und Ste- 
phaniden des Deutschen Entomologischen Muse- 
ums (Hym.). Entomologische Mitteilungen 1: 
388-391. 

Hedicke, H. 1939. Gasteruptiidae. /n Hedicke, H., ed. 
Hymenopterorum Catalogus, Pars 11, 54 pp. Dr. 
W. Junk, ’s’Gravenhage. 

Jennings, J. T. and A. D. Austin. 1994. Revision of the 
genus Pseudofoenus Kieffer (Hymenoptera: Gas- 
teruptiidae), a hyptiogastrine wasp genus endemic 
to New Zealand. Invertebrate Taxonomy 8: 1289— 
1303. 


1997a. Revision of the Australian genus 
Hyptiogaster Kieffer (Hymenoptera, Evanioidea, 
Hyptiogastrinae), with descriptions of seven new 
species. Journal of Natural History 31: 1533- 
1562. 

. 1997b. Revision of Aulacofoenus Kieffer (Hy- 
menoptera: Gasteruptiidae), hyptiogastrine wasps 
with a restricted Gondwanic distribution. Inverte- 
brate Taxonomy 11: 943-976. 

. 2002. Systematics and distribution of world 


hyptiogastrine wasps (Hymenoptera: Gasterupti- 
idae). Invertebrate Systematics 16: 735-811. 
Kieffer, J. J. 1902. Hymenoptera, Fam. Evaniidae. /n 
Wytsman, P., ed. Genera Insectorum, Fasc. 2, 13 
pp., | plate. Bruxelles. 
1904. Description de Stéphanides et 
d’Evaniides nouveaux. Bulletin de la Société 
d’Histoire Naturalle de Metz, 2nd Série 11: 1- 
30. 


1910. Beschreibung einer neuen Evaniidae. 
Zoologische Jahrbiicher Abteilung fiir Systematik 
Okologie und Geographie der Tiere 29: 242. 

. 1912. Evaniidae. Das Tierreich 30: 1—431. 


Pasteels, J. J. 1956. Révision du genre Gasteruption 


(Hymenoptera, Evanoidea, Gasteruptionidae). II. 
Espéces mélanésiennes. Nova Guinea (N.S.) 7: 
207-248. 

. 1957. Révision du genre Gasteruption (Hy- 


menoptera, Evanoidea, Gasteruptionidae). Espe- 


ces australiennes. Memoires Institut Royal des 


Sciences Naturelles de Belgique 56: 1—125. 


2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 1958. Révision du genre Gasteruption (Hy- 
menoptera, Evanoidea, Gasteruptionidae). V. Es- 
péces Indo-Malaise. Bulletin et Annales de la So- 
ciété Royale Entomologique de Belgique 94: 169— 
DNS. 

Schletterer, A. 1885. Die Hymenopteren-Gattung Gas- 
teruption Latr. (Foenus aut.). Verhandlungen der 
K. K. Zoologisch-Botanischen Gesellschaft in 
Wien 35: 267-326. 

. 1889. Die Hymenopteren-gruppe der Evani- 


den. Annalen des K. K. Naturhistorichen Hof- 
museum, Wien 4: 107-180, 289-338, 373-546. 

Schrottky, C. 1906. Neue Evaniiden aus Paraguay 
(Hym.). Zeitschrift fiir Systematische Hymenop- 
terologie und Dipterologie 6: 56—62. 

Sharkey, M. 1988. Ichneumonoid wing venation. Ich- 
news 11: 2-12. 

Szépligeti, V. 1903. Neue Evaniiden aus der sammlung 
der ungarischen national-museums. Annales Mu- 
sei Nationalis Hungarici 1: 364—395. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 693-699 


ALDERFLY (NEUROPTERA: SIALIDAE) FLIGHT PERIODS, SEX RATIOS, 
AND HABITAT USE IN A VIRGINIA FRESHWATER TIDAL MARSH, 
LOW FOREST, AND THEIR ECOTONES 


EDWARD M. BARROwS, ANNE M. MCINTYRE, AND OLIVER S. FLINT. JR. 


(EMB, AMM) Laboratory of Entomology and Biodiversity, Department of Biology, 
Reiss Building Suite 406, Georgetown University, Box 571229, Washington, D.C. 20057- 
1229, U.S.A. (e-mail: barrowse @ georgetown.edu); (OSF) Department of Entomology, 
Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A. (e-mail: flinto@si.edu) 


Abstract.—Six Malaise traps, run from April 1998 through December 1999, obtained 
3306 adult Sialis iola Ross (34% females) and one male S. mohri Ross in Dyke Marsh 
Preserve (DMP), Virginia. They flew from early April through early June. The abundance 
of S. iola significantly varied among a low forest, freshwater tidal marsh, and the forest- 
marsh ecotone, sometimes being significantly more common in the ecotone than other 
habitats. Malaise traps can be used efficiently to survey and monitor sialids in DMP and 
similar places. A thorough monitoring effort in the Preserve should include at least two 
traps in the three habitats and monitoring throughout sialid flight periods and the Preserve. 


Key Words: 
low forest 


This study concerns sialids (Neuroptera: 
Sialidae, Alderflies) in a freshwater tidal 
marsh and adjacent low forest in the Mid- 
Atlantic region of the U.S.A. We provide 
information about adult sialid abundances 
and flight periods in three main habitats of 
Dyke Marsh Preserve (DMP), Virginia— 
low forest, freshwater tidal marsh, and the 
ecotone between them, based on Malaise- 
trap samples. The National Park Service 
(NPS) requires information on DMP spe- 
cies, their abundances, and their habitat 
uses, in order to manage the Preserve prop- 
erly. A sialid study is particularly crucial at 
this time because Virginia may use a pes- 
ticide to control mosquitoes that carry the 
West Nile Virus. Such mosquito control 
could have marked deleterious effects on 
many DMP organisms. 

Sialids can be abundant in aquatic and 
adjacent habitats where they are predators 


Neuroptera, Sialidae, alderflies, flight periods, sex ratios, freshwater marsh, 


of other arthropods and food for vertebrates 
and other organisms including fish (Azam 
and Anderson 1969). Bowlker (1747) wrote 
in The Art of Angling, “‘the Orle Fly ... is 
the best Fly to Fish with after the May 
Flyes are gone” (reference in the Oxford 
English Dictionary 1971). ““Orle” is an old 
name for “alder.” 

There are 24 North American sialid spe- 
cies. Females lay egg masses of from about 
200 through 900 eggs on objects over water 
such as bridges, culverts, and vegetation 
(Azam and Anderson 1969, Arnold and 
Drew 1987, Brigham 1982, Canterbury and 
Neff 1980). After hatching, the larvae fall 
into the water where they live under stones, 
vegetation, and other objects. Sialid larvae 
are aquatic predators of small aquatic in- 
sects (Pritchard and Leischner 1973). Fully 
developed larvae crawl out of the water 
onto land, and each forms an earthen cell 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


forest 


Potomac River 


— ecotone 


Fi 


F2 El 


| tidal gut — 


path —. 


Fig. 1. 


M2 M1 


I 


marsh 


100 m 


Locations of the Malaise traps in Dyke Marsh Preserve, Virginia. Ei and E2 are positions of the 


ecotone traps; Fl and F2, forest traps; and M1 and M2, marsh traps. 


in soil where it pupates (Ross 1937). Adults 
usually occur near water and live only a 
few days (Murnane 2004). Males are mod- 
erately fast fliers and more active during the 
day than at night. They have soft mouth- 
parts and may not feed (Azam and Ander- 
son 1969), or feed much, as adults (Ross 
O37). 

We report that our sampled sialids were 
over 99% Sialis iola Ross, a little-studied, 
abundant aquatic insect. They flew from 
early April through early June, and were 
markedly different in abundances and sex 
ratios among Malaise traps and habitats. 
Further we discuss the use of such traps for 
surveying and monitoring sialids. 


MATERIALS AND METHODS 


We collected sialids in DMP from April 
1998 through December 1999 using six 
Townes-style Malaise traps (Townes 1972) 
in Dyke Marsh Preserve (DMP), part of the 
George Washington National Parkway, Vir- 
ginia, administered by the NPS (Johnston 
2000). The 380-acre Preserve, on the west- 
ern shore of the Potomac River in Fairfax 


County, Virginia, contains the largest re- 
maining freshwater tidal marsh in the 
Washington, D.C., area (Johnston 2000). 
The Preserve has experienced marked deg- 
radation due to alien invasive organisms, 
shoreline erosion due to boat wakes and 
storms, and water pollution and associated 
harmful algal blooms. 

Two traps were placed in each of three 
habitats—low forest, freshwater tidal 
marsh, and the ecotone between them (Fig. 
1). The six traps were in a broad transect 
that ran east and west. The ecotone (defined 
as 10 m on each side of the forest-marsh 
edge) ran about 200 m approximately 
north-northeast and south-southwest in our 
sampling area. We oriented each trap so that 
its longitudinal axis ran east and west and 
its collecting head faced due east. The for- 
est traps were about 50 m west of the eco- 
tone, and the marsh traps averaged about 60 
m east of the ecotone. Forest trap 1 (F1) 
was about 30 m north-northeast of forest 
trap 2 (F2), ecotone trap 1 (El) was about 
30 m north-northeast of ecotone trap 2 (E2), 
and marsh trap 1 (M1) was about 30 m east 


VOLUME 107, NUMBER 3 


east 


collecting head 


Wt) «he 


ne Oe 
sa 


tidal gut 


Fig. 2. 
temporarily on the trap’s base. 


of marsh trap 2 (M2). The ecotone traps 
were in the edge of the open marsh (where 


they usually experienced about 0.3 m of 


water during high-tide periods) and were 
about 3 m from the edge of the forest. The 
mid-point location of the forest traps is 
38.77194°N 77.05083°W; ecotone traps, 
38.77139°N 77.05056°W; and marsh traps, 
38.77172°N 77.04990°W. 

Each trap was 1.2 m wide, 1.7 m long, 
1.0 m high at its back and 2.0 m high at its 
front, the location of its collecting head 
(Fig. 2; Barrows and Kjar, 2005; keyword: 
Malaise trap, images of our study traps in 
each habitat). Each trap had |-mm?* mesh, 
nylon gauze; 61-mm wide, black crab-cage 
wire; a supporting metal frame; and a col- 
lecting head. We spray-painted trap gauze 
and supporting frames black in an attempt 
to decrease their visibility to sialids and hu- 
man park visitors. The crab-cage wire en- 
circled the base of each trap and prevented 
objects such as snapping turtles and drift- 


== guide pole 


F Ce BELT CT. Patil eite vial wi 
Beer TeTT ih 


bd 
: ine htt mae lier | 


A buoyant Townes-style Malaise trap in the marsh at low tide, 


695 


west 


SERA” 
| 


ae vie 
ah ' oth ri "s 


1 fh ten hari 


12 April 1998. Two backpacks are 


wood from tearing trap gauze. Each trap 
was mounted on a floating platform, 1.2 by 
1.8 m, that rose up to | m when the tide 
entered the Preserve’s marsh. Vertical metal 
poles kept traps in place as they moved up 
and down. Forest traps were not buoyant 
because their forest sites did not flood dur- 
ing our study period, but can flood as high 
as 2.6 m during hurricanes. Sialids flew or 
crawled into the top part of a trap’s col- 
lecting head, and then became entrapped in 
its bottom part filled with 95% ethanol. All 
traps ran during our entire 21-month sam- 
pling period, except the marsh traps. We re- 
moved them from December 1998 
through late March 1999, because possible 
that time could 


late 
flooding during have de- 
stroyed them. 

Sialid samples are from five intervals in 
1998 (5 April—6 June) and four intervals in 
1999 (11 April-6 June). We did not find 
sialids in our samples during other sam- 
pling periods. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Number of sialids 


Habitats and years 


Fig. 3. 


Both female and male Sialis iola were more common in the ecotones than in the other habitats in 


both years. Black parts of the bars represent females; gray parts, males. The forest traps had one female and 
one male in 1998 (which are not shown on the graph) and no sialids in 1999. 


To look for possible differences in the 
number of sialids per day among habitats, 
we used repeated-measures analysis of var- 
iance (ANOVA) and the Student-Newman- 
Keuls test (SPSS, Inc. 2003). To look for 
possible differences between observed sex 
ratios and expected 1:1 sex ratios within a 
habitat or interval, we used Preacher’s 
(2003) online Chi-square test program. 


RESULTS AND DISCUSSION 


Traps obtained 3307 sialids comprising 
3306 Sialis iola and one male S. mohri 
Ross during our 2-yr study. The S. mohri 
was from a sample from 12—28 April 1998 
from an ecotone trap. Our traps were in a 
limited area of the Preserve, and we do not 
know whether S. mohri is rare throughout 
the Preserve. The sialids that were previ- 
ously know from the Washington, D.C., 
area (defined as Montgomery and Prince 
Georges counties, Maryland; Arlington and 
Fairfax counties, Virginia; and Washington, 
D.C.) are Sialis aequalis Banks (which is 
known from MD and VA), S. americana 
(Rambur) (DC, MD), S. infumata Newman 
(MD, VA); S. iola Ross (DC, MD), S. itas- 


ca Ross (DC, MD, VA), S. joppa Ross 
(MD), S. vagans Ross (VA), and S. velata 
Ross (DC, MD, VA) based on Ross (1937), 
Flint (1964), Tarter et al. (1978), Whiting 
(1991), and material in the collection of the 
National Museum of Natural History, 
Smithsonian Institution. Sialis mohri was 
previously recorded from Minnesota 
through New Brunswick south though New 
Jersey and Pennsylvania in eastern U.S. and 
south through Tennessee, Mississippi, Ar- 
kansas, and Oklahoma in central U.S., but 
not from Virginia. Sialis iola was not pre- 
viously known from the Washington, D.C., 
area. 

There were 1,168 S. ‘ola in the samples 
in 1998 and 2,138 in 1999. The observed 
yearly differences in this species’ abun- 
dance might be the result of natural fluc- 
tuations in its population size. Both females 
and males were most common in the eco- 
tone compared to the marsh and forest 
based on raw data from 1998 samples, 1999 
samples, and 1998 and 1999 samples com- 
bined (Fig. 3). The 1998 males and the 
1999 females were significantly more com- 
mon in the ecotone than the forest or marsh 


VOLUME 107, NUMBER 3 


697 


Table 1. The number of females, males, and both sexes combined varied among habitats based on Malaise- 


trap samples from Dyke Marsh Preserve, Virginia.* 


1998 1999 Combined P 

Group F 12 F P F P 
Females + males 8.312 0.060 6.345 0.084 10.581 =0.010 
Females 4.220 0.134 162.765 0.001 17.835 =0.001 
Males 11.970 0.037 2.708 0.213 9.687 =0.010 


* The degrees of freedom for all repeated-measures ANOVAs is 2. We combined probabilities using the 


process described by Sokal and Rohlf (1969). 


(Table 1, P = 0.05, repeated-measures AN- 
OVA and Student-Newman-Keuls test). 
The statistical analyses may have not found 
other actual differences in abundances 
among habitats, because of large abundance 
variances of trap samples within a habitat. 
Females alone, males alone, and both sexes 
combined showed significantly different 
abundances among the three habitats when 
the P values of both years were combined 
(Table 1). 

Sialis iola flew from April through early 
June in both years (Figs. 4a, b), a shorter 
span than that recorded for the entire range 
of the species which is 1 April through 29 
July (Tarter et al. 1978, Whiting 1991). In 
1998, the number of females peaked in the 
samples of 10-17 May and 17—26 May, and 
the number of males peaked in the sample 
of 12—28 April. In 1999, both females and 
males were most common in the sample of 
9-23 May. Flight periods suggest that S. 
iola was protandrous in both years. 

Sampled sex ratios of S. iola were sig- 
nificantly male biased in 1998, 1999, and 
both years combined (Table 2). This bias 
may be due to an actual preponderance of 
males in the Preserve, or, if this species has 
a true 1:1 adult sex ratio, a greater tendency 
for the traps to catch males rather than fe- 
males. As in other species, S. iola males 
may fly more than females and, therefore, 
be more likely to be trapped than females. 
For example, in two western U.S. species, 
Sialis californica Banks and S. rotunda 
Banks, adult males are more active than fe- 
males and swarm near vegetation along 
shores and walk up and down plants (Azam 


and Anderson 1969). Females are usually 
heavily egg-laden and have considerably re- 
stricted flight. A female might be highly 
likely to oviposit in, or very near, the par- 
ticular area where she was a larva. The 
sampled sex ratios of Sialis iola were sig- 
nificantly male-biased in the ecotone but 
not in the marsh in 1998, 1999, and both 
years combined (Fig. 3, Table 2). If there is 
a 1:1 sex ratio in new marsh adults, the eco- 
tone male-biased samples suggest that 
many males leave the marsh and search for 
females in the ecotone. Further because the 
marsh sex ratio is female-biased, on the ay- 
erage, each marsh male competed with few- 
er other males for mates. 

In conclusion, we report that DMP sial- 
ids flew from early April through early 
June, and their abundances markedly var- 
ied among traps and habitats. We obtained 
baseline data on DMP sialids with the view 
of monitoring and managing them in the 
future. Malaise traps are an excellent 
means to monitor them because the traps 
can be run 24 hours per day, and these 
traps can collect large samples of sialids 
for detailed analysis. Malaise traps often 
obtain rare species that might be missed by 
visual censusing and hand collecting and 
other collecting means. Our data indicate 
that it would be most efficient to trap sial- 
ids in the ecotone, if limited resources 
would not permit sampling other habitats 
as well. Sample sizes from traps of the 
same style, which are contemporaneously 
run in the same habitat, can markedly vary. 
In 1998-1999, one ecotone trap had 34% 
more sialids than the other, and one marsh 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


1998 


a TR at 
G 
x) 
tee 
® 
ok 
/) 
= 
i 
“”) 
1 2 3 4 5 
Sampling period 
100 >, 

Cue 1999 
> 80. 
xe) | 
o 60 - 
©. 
—) 
5 40 
6 
Pl 

0 zal 

1 2 3 4 
Sampling period 
Fig. 4. Male Sialis iola were more common earlier in the flight season than females. Black parts of the bars 


represent females; gray parts, males. 4a, The 1998 collecting periods are 1 (12—28 April), 2 (28 April-10 May), 
3 (10-17 May), 4 (17-26 May), and 5 (26 May-—6 June). The traps had 0 female and 0.1 male per day in period 


Nn 


trap caught 67% more than the other. 
Therefore, monitoring should use at least 
two traps. More traps per habitat would 
provide an even more complete picture of 
sialid abundances from year to year than 
two traps. To understand sialid annual pop- 
ulation fluctuations and species composi- 


. 4b, The 1999 periods are 1 (11-25 April), 2 (25 April-9 May), 3 (9-23 May), and 4 (23 May-—6 June). The 
traps had 0.4 female and 0.7 male per day in period 1. 


tion better, it would be worthwhile to mon- 
itor them for at least 10 years with Malaise 
traps, hand-collecting, and other appropri- 
ate means throughout the Preserve. The 
high abundance of sialids in the ecotone 
indicates the great value of this habitat 
with its woody plants for these animals. 


VOLUME 107, NUMBER 3 


699 


Table 2. The sex ratios of Sialis iola were significantly different from 1:1 in the ecotone and usually in the 
marsh based on Malaise-trap samples from Dyke Marsh Preserve, Virginia.* 


Year Habitat No. of Sialids % Females x Je 
1998-1999 All three 3,606 B32 395.9 <0.0001 
Ecotone 3,016 30.3 468.0 <0.0001 

Forest 2 50.0 = — 
Marsh 288 57.6 6.7 0.0095 
1998 All three 1,168 30.9 170.3 <(0.0001 
Ecotone 1,020 PiTES 207.5 <0.0001 

Forest 2 50.0 — — 
Marsh 146 54.8 1.3 0.2467 
1999 All three 2,138 $35}57/ 227.9 <0.0001 
Ecotone 1,996 31.8 265.5 <0.0001 

Forest 0) — — — 
Marsh 142 60.6 6.4 0.0118 


* P values are from Chi-square tests in which the expected sex ratios were 1:1 in a particular habitat within 
year. The forest samples were too small for Chi-square analysis. 


ACKNOWLEDGMENTS 


We thank the Friends of Dyke Marsh, the 
National Park Service, and the Washington 
Biologists’ Field Club for supporting our 
research. Robert O’Hanlon (Alexandria, 
VA) provided invaluable help with traps. R. 
Edward DeWalt and Donald A. M. Mackay 
made many helpful comments about a pre- 
liminary draft of this paper, and Rusan 
Chen helped with statistical analysis. 


LITERATURE CITED 


Arnold, D. and W. A. Drew. 1987. A preliminary sur- 
vey of the Megaloptera of Oklahoma. Proceedings 
of the Oklahoma Academy of Science 67: 23-26. 

Azam, K. M., and N. H. Anderson. 1969. Life history 
and habits of Sialis rotunda and Sialis californica 
in western Oregon. Annals of the Entomological 
Society of America 62: 549-558. 

Barrows, E. M. and D. S. Kjar. 2005. Biodiversity Da- 
tabase of the Washington, D.C., Area (BDWA). 
Website. http://biodiversity.georgetown.edu (5 
March 2005). 

Brigham, W. U. 1982. Megaloptera, pp. 7.1—7.12. /n 
Brigham, A. R., W. U. Brigham, and A. Gnilka, 
eds. Aquatic Insects and Oligochaetes of North 
and South Carolina. Midwest Aquatic Enterprises, 
Mahomet, Illinois, 837 pp. 

Canterbury, L. E. and S. E. Neff. 1980. Eggs of Sialis 
(Sialidae: Megaloptera) in eastern North America. 
Canadian Entomologist 112: 409-419. 

Flint, O. S. 1964. New species and new state records 
of Sialis (Neuroptera: Sialidae). Entomological 
News 75: 9-13. 


Johnston, D. W. 2000. The Dyke Marsh Preserve Eco- 
system. Virginia Journal of Science 51: 223-272. 

Murnane, A. 2004. Neuroptera. Ant Lion, Dobsonfly, 

Owlfly. Internet file. http:// 
www.discoverlife.org/nh/tx/Insecta/Neuroptera/ 
(March 8, 2004) 

Oxford English Dictionary. 1971. The Compact Edi- 
tion of the Oxford English Dictionary, Vol. Il. A— 
O. Oxford University Press, Oxford, U.K., 2,048 
PP- 

Preacher, K. J. 2003. Calculation for the Chi-Square 
Test. Internet file. http://www.unc.edu/~preacher/ 
chisq/chisq.htm (5 March 2005) 

Pritchard, G. and T. G. Leischner. 1973. The life his- 
tory and feeding habits of Sialis cornuta Ross in 


Lacewing, 


a series of abandoned beaver ponds (Insecta; Me- 
galoptera). Canadian Journal of Zoology 51: 121— 
SHE 

Ross, H. H. 1937. Studies of Nearctic aquatic insects. 
1. Nearctic alder flies of the genus Sialis (Megal- 
optera, Sialidae). Illinois Natural History Survey 
Bulletin 21: 57-99. 

Sokal, R. R. and E J. Rohlf. 1969. Biometry. W. H. 
Freman, San Francisco, California, 776 pp. 

SPSS, Inc. 2003. SPSS Advanced Model 12.0. SPSS, 
Inc., Chicago, Illinois, 497 pp. 

Tarter, D. C., W. D. Watkins, D. L. Ashley, and J. T. 
Goodwin. 1978. New state records and seasonal 
emergence patterns of alderflies east of the Rocky 
Mountains (Megaloptera: Sialidae). Entomologi- 
cal News 89: 231-234. 

Townes, H. 1972. A light-weight Malaise trap. Ento- 
mological News 83: 239-247. 

Whiting, M. FE 
(Megaloptera, Sialidae) in North America. Ento- 
mological News 102(1): 50-56. 


1991. A distributional study of Sialis 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 700-728 


A SYSTEMATIC REAPPRAISAL OF THE GENUS DIURAPHIS AIZENBERG 
(HEMIPTERA: APHIDIDAE) 


Gary L. MILLER, MANYA B. STOETZEL, AND ETHAN C. KANE 


Systematic Entomology Laboratory, Plant Sciences Institute, Agricultural Research Ser- 
vice, U.S. Department of Agriculture, Bldg. 005, BARC-West, Beltsville, MD 20705, 
U.S.A. (email: gmiller@sel.barc.usda.gov; mstoetze@sel.barc.usda.gov; eckane@sel. 
barc.usda.gov) 


Abstract.—Adult female apterae and alates of the genus Diuraphis Aizenberg are de- 
scribed and illustrated, and keys are provided for identification. Diuraphis elymophila G.- 
x. Zhang is considered a new synonym of Diuraphis frequens (Walker) and Diuraphis 
muehlei (Borner) is considered a new synonym of Diuraphis noxia (Kurdjumov). A phy- 
logenetic analysis suggests that while the clade that contains Diuraphis noxia + Diuraphis 
mexicana is monophyletic, the previously recognized subgenus Holcaphis is paraphyletic. 


Key Words: aphid, Diuraphis, Holcaphis 


The genus Diuraphis was proposed in 
1935 with Brachycolus noxius Mordvilko 
[= Diuraphis noxia (Kurdjumov)] as its 
type species (Aizenberg 1935). As currently 
defined (Remaudiére and Remaudiére 
1997), there are 11 species in the genus re- 
ferable to two generally recognized subgen- 
era. The subgenus Diuraphis sensu stricto 
contains three species: D. mexicana 
(McVicar Baker), D. muehlei (Borner) and 
D. noxia (Kurjumov). The subgenus Hol- 
caphis Hille Ris Lambers (1939) comprises 
the remaining species and includes: D. 
agropyronophaga G.-x. Zhang, D. agrosti- 
dis (Muddathir), D. bromicola (Hille Ris 
Lambers), D. calamagrostis (Ossiannils- 
son), D. elymophila G.-x. Zhang, D. fre- 
quens (Walker), D. holci (Hille Ris Lamb- 
ers), and D. tritici (Gillette). Historically, 
Diuraphis sensu stricto differs from Hol- 
caphis by the presence of a supracaudal 
process on the eighth abdominal tergite and 
usually the presence of marginal tubercles 
on abdominal segments II-VI (Heie 1992). 
Although the two subgenera of Diuraphis 


are generally recognized (Eastop and Hille 
Ris Lambers 1976, Remaudieére and Re- 
maudiére 1997), their monophyly has not 
been tested. 

Diuraphis was not well known until the 
early 20" century when outbreaks of D. 
noxia (the Russian wheat aphid) and D. tri- 
tici (the western wheat aphid) in Russia and 
the western United States, respectively, 
brought attention to the destructiveness of 
these aphids on wheat. By the late 1970's, 
attention was once again focused on Diura- 
phis, especially D. noxia. Substantial range 
extension of D. noxia was first documented 
in South Africa in 1978 (Diirr 1983) and in 
the United States in 1986 (Stoetzel 1987). 
This aphid spread quickly and now has 
been recorded throughout much of the 
wheat growing regions of the world. In the 
United States, its damage to wheat and bar- 
ley was extensive and resulted in heavy 
crop losses in some fields (Stoetzel 1987). 
By 1993, D. noxia was a pest in 16 western 
states and caused cumulative losses esti- 
mated at $500—900 million dollars (Bernal 
et al. 1993, Morrison and Peairs 1998). 


VOLUME 107, NUMBER 3 


Because D. noxia is recognized as an 
economically important pest, much of the 
literature on Diuraphis has concentrated on 
this species. However, Zhang et al. (1991) 
gave a key to Diuraphis and discussed phy- 
logenetic relationships. They also included 
the descriptions of D. agropyronophaga 
and D. elymophila. Kovalev et al. (1991) 
provided a key to the Diuraphis apterous 
viviparous females and reviewed the Rus- 
sian literature. Descriptions, keys to apterae 
and alata, and illustrations of Diuraphis of 
Fennoscandia and Denmark were provided 
by Heie (1992). Halbert et al. (1992) also 
included keys to North American Diura- 
phis. 

Apterae and alatae of the genus Brachy- 
colus Buckton closely resemble Diuraphis, 
the main difference is the position of the 
cornicle (Heie 1992). In Brachycolus and 
most other genera of aphids, the cornicle is 
on the posterior portion of abdominal ter- 
gite V, whereas in Diuraphis the cornicle is 
on the posterior portion of abdominal ter- 
gite VI (Heie 1992). However, neither the 
relationship between Diuraphis and Bra- 
chycolus in a phylogenetic context nor the 
monophyly of the subgenera of Diuraphis 
have been examined. 

The objectives of this paper are to: (1) 
redescribe, illustrate, and present keys for 
the identification of apterae and alatae of 
Diuraphis species; and (2) test the hypoth- 
esis that Diuraphis is monophyletic and 
comprised of two distinct subgroups (i.e., 
Diuraphis (sensu stricto) and Holcaphis). 


MATERIALS AND METHODS 


Synoptic descriptions are taken from 
original descriptions, types, and identified 
material from the Aphidoidea collection of: 
Muséum National D’Histoire Naturelle 
(MNHN), Paris, France; National Museum 
of Natural History (USNM), Beltsville, 
MD, U.S.A.; Canadian National Collection 
of Insects (CNCI), Ottawa, Canada; The 
Natural History Museum, London, U.K. 
(BMNH); Institute of Zoology Academia 
Sinica (IZAS), Beijing, People’s Republic 


701 


of China, and University of Rostock Insect 
Collection (URIC), Sagerheide, Germany. 
Measurements are presented in millimeters 
as minimum and maximum ranges of rep- 
resentative specimens. 

Morphological terms and _ structures 
adapted from Stoetzel et al. (1999) are used 
in this work. Those terms are listed below, 
and equivalent terms that may be found in 
other literature are listed in parentheses for 
reference: terminal process (= unguis, pro- 
cessus terminalis); secondary sensoria (= 
secondary rhinaria); cornicle (= siphuncu- 
lus); fundatrix/fundatrices (= stem moth- 
er(s)); aptera/apterae (= wingless vivipa- 
rous female(s)); alata/alatae (= winged vi- 
viparous female(s)); and ovipara/oviparae 
(= egg-laying female(s)). 

The information under Specimens Ex- 
amined is organized to conserve space. Ab- 
breviations for fundatrices, apterae, alatae, 
Oviparae, apterous males, alate males, and 
immatures are listed as: fund.; ap.; al.; ov.; 
ap. ; al. d; and imm. respectively. If a col- 
lection was made at the same locality, but 
on a different date as a previously listed 
collection, duplicated information is not re- 
peated. For example, the documentation 
provided for a particular locality may be re- 
corded as: TEXAS: Big Bend, VII-11- 
1978, on Bromus unioloides |= Bromus ca- 
tharticus], 1. M. Miller coll., (2 al.) USNM; 
VIII-29—1957, XI-21—1957, IV-14—1978, 
on Bromus sp., (15 ap. on 15 sl.) USNM. 
In this hypothetical example, the second 
collection was also found at Big Bend, even 
though “Big Bend” 
When specimens are mounted on a single 
slide (sl.), it is not written as such but is 
assumed. Bracketed ([]) text represents sup- 
plemental information by the present au- 
thors for clarification purposes or refers to 
the original collection data of laboratory 
reared specimens. Collection data that are 
the same except for collection date are sim- 
ply listed sequentially. Host plants listed in 
the Specimens Examined sections are sum- 


was not repeated. 


marized in Table 2. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Data matrix used in the cladistic analysis. 

234567890 1234567890 1245678 
B. cerastit 1000110110 0000011011 0010011 
B. cucubali 0000000210 001011001 0010001 
B. stellariae 0010111000 QL AL AL aL ALO) (O(O}aE LL IL aL 
B. asparagi 1010121100 IOLLLOLUOD 1000010 
D. agropyronophaga 0010020310 (ONO) aL ALL aN aL a0 aL aL 1011001 
D. agrostidis 0001021001 ILOILO iL LO i LO 1000011 
D. bromicola 0011120011 LA OLO DAL i LOLLLLO 
D. calamagrostis 0000121201 1001110010 AL O)(0)1L (== 
D. frequens 1001020001 ILO MOOI aL al 1000100 
D. holci 0001010211 al) LAL Oat WO 1010000 
D. mexicana Hal A LAOS OAL LOOLLA LUO WA 1110000 
D. noxia LLAZLAO LOA 1101101001 1111000 
D. tritici 1002120300 MLO AAA 1010100 


Phylogenetic analysis.—Phylogenetic 
analyses of Diuraphis and selected out- 
groups were conducted to test the mono- 
phyly of Diuraphis and infer relationships 
within the genus to test the validity of cur- 
rently recognized subgenera of Diuraphis 
(Diuraphis sensu stricto and Holcaphis). 
Nine species (Diuraphis agropyronopha 
G.-x. Zhang, Diuraphis agrostidis (Mud- 
dathir), Diuraphis bromicola (Hille Ris 
Lambers), Diuraphis calamagrostis (Os- 
siannilsson), Diuraphis frequens (Walker), 
Diuraphis holci (Hille Ris Lambers), Di- 
uraphis mexicana (McVicar Baker), Diura- 
phis noxia (Kurdjumov), and Diuraphis tri- 
tici (Gillette)) are included in the present 
analysis. Examination of type material of 
Diuraphis muehlei (Borner) and Diuraphis 
elymophila Zhang revealed that those spe- 
cies are junior synonyms of D. noxia and 
D. frequens, respectively. 

Brachycolus stellariae (Hardy), Brachy- 
colus cerastii (Kaltenbach), Brachycolus 
cucubali (Passerini), and Brachycorynella 
asparagi (Mordvilko) are included as out- 
groups to provide a context in which to test 
the proposed monophyly of Diuraphis sen- 
su lato. Diuraphis is suspected to be closely 
related to Brachycolus and Brachycorynel- 
la. Members of these three currently rec- 
ognized genera were referable to Brachy- 
colus (Shaposhnikov 1964). Heie (1992) 


also noted Brachycorynella was similar to 
Brachycolus. 

Characters: Thirty morphological char- 
acters were examined. Two characters 
proved to be autapomorphic and were ex- 
cluded from the final data set as parsimony 
uninformative. Of the remaining 28 char- 
acters, 25 were coded as binary while three 
were treated as non-additive multistate 
characters. Unknown or indeterminable 
character states were coded as missing data. 
Character descriptions are outlined below 
and the final data matrix is presented in Ta- 
ble 1. 


Apterous viviparous female 
(excluding the fundatrix) 


Head 

1. Tips of median dorsal head setae: 

pointed or tapered (QO); blunt (1). 
Antennae 

2. Base of scape with slight posterior-lat- 
eral protuberance: absent (0); present 
(1). 

3. Antennal tubercle shape: undeveloped 
or flat (0); moderately developed or 
slightly raised (1). 

4. Antennal segment I: entirely pigmented 
(O); pigmented medially (1); pale (2). 

5. Antennal segment II venter: usually 
smooth (0); usually with some imbri- 
cations (1). 


VOLUME 107, NUMBER 3 


6. Antennal segment III: entirely pig- 
mented except for base (0); pigmenta- 
tion reduced to base or apex (1); pale 
(2). 

7. Antennal segment III: imbricated 
throughout (0); partly imbricated (1). 

8. Length of antennal segment ITV com- 
pared to length of segment V: segment 
IV usually longer than segment V (0); 
segment IV usually shorter than seg- 
ment V (1); length of segment IV sub- 
equal to length of segment V (2): 
length of segment IV variable when 
compared to segment V (3). 

9. Length of antennal segment III com- 
pared to length of antennal segment IV 
+V: shorter (0); longer (1). 

Mouthparts 

10. Ultimate rostral segment accessory se- 
tae: present (0); absent (1). 

11. Length of base of antennal segment VI 
compared to ultimate rostral segment: 
usually shorter or subequal (0); longer 
()). 

12. Number of setae anterolateral to post- 
clypeus: 2—3 (0); 1 (1). 

Thorax 

13. Lateral prothoracic tubercles: present at 
least sometimes (0); absent (1). 

14. Protibiae: not uniformly colored (0); 
uniformly colored (1). 

15. Metafemur: not uniformly colored (0); 
uniformly colored (1). 

Abdomen 

16. Large polygonal dorsal abdominal re- 
ticulation: present (0); absent (1). 

17. Intersegmental sclerites: present (0); 
absent (1). 

18. Lateral abdominal tubercles: present at 
least sometimes (0); absent (1). 

19. Apical flange of cornicle: present (0); 
absent (1). 

20. Cornicle with associated basal sclerite: 
present (0); absent (1). 

21. Cornicle position: anterior to stigmal 
pore VI (0); level or posterior to stig- 
mal pore VI (1). 

22. Dorsum of abdominal segment VIII: 


703 


usually not raised medially (0); raised 
medially (1). 
. Cauda: apically rounded (0); apically 
pointed (1). 
24. Mid-ventral caudal spicules: individu- 
ally separate (0); connected (1) 
25. Length of cauda compared to length of 
hind tarsus II: longer (0); shorter or ap- 
proximately equal (1). 


N 
eS) 


Alate viviparous females 

Antennae 

26. Antennal segment IV secondary sen- 
soria: usually present (0); usually ab- 
Sent) 

27. Length of antennal segments TV + V 
compared to antennal segment III: lon- 
ger (0); shorter or subequal (1). 


All phylogenetic analyses were per- 
formed using PAUP* (Swofford 2001). 
Maximum parsimony (MP) analyses were 
conducted using branch-and-bound search- 
es, and bootstrap analyses involved 1,000 
replicates of branch-and-bound searching. 
In the initial MP analyses, all characters 
were treated as unordered and as having 
equal weights. In subsequent analyses, the 
successive approximations approach to 
character weighting (SACW) was used to 
select the most cladistically reliable topol- 
ogy (Farris 1969). For the SACW analysis, 
characters were weighted based on the re- 
scaled consistency index. 


PHYLOGENETIC RESULTS AND DISCUSSION 

MP analysis generated 16 equally most 
parsimonious topologies all with a tree 
length of 83, consistency index (CI) of 
0.37, retention index (RI) of 0.43, and re- 
scaled consistency index (RC) of 0.16. 
Three iterations of SACW with characters 
weighted based on the rescaled consistency 
index resulted in a single most parsimoni- 
ous topology (Fig. 1) with a tree length of 
14.28, Cl of 0.63, RI of 0.75, and RC of 
0.47. Bootstrap analysis of the unweighted 
data set produced relatively low support 
values, with only two nodes recovered in 
greater than 50% of the replicates. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


L= 14.28 
Cl= 0.63 
RI= 0.75 
RC= 0.47 


62 


Fig. 1. 


. calamagrostis 
. holci 

. agrostidis 

. frequens 

. tritici 

. bromicola 

70 . mexicana 
noxia 

. asparagi 

. agropyronophaga 


. Stellariae 


7» © wm © S&S © & & SS & 


. cucubali 


B. cerastii 


Maximum parsimony topology resulting from three iteration of successive weighting and branch- 


and-bound analysis. Node labels indicate bootstrap proportions obtained from an analysis of the unweighted 
matrix. L = Length, CJ = Consistency Index, RJ = Retention Index, RC = Rescaled Consistency Index. 


The most parsimonious tree presented in 
Fig. | conflicts with current taxonomic con- 
cepts of Diuraphis in that the genus is not 
recovered as a well-supported monophylet- 
ic group. The recovery of B. asparagi with- 
in the Diuraphis clade is problematic, but 
not entirely unprecedented. B. asparagi 
shares many similarities with Diuraphis 
species and has been grouped with other 
species of Diuraphis is previous studies 
(e.g., Shaposhnikov 1964). The relative po- 
sition of the cornicle with respect to stigmal 
pore VI (character 21), although lacking 
strong bootstrap support, is potentially syn- 
apomorphic for these taxa. 

Regarding proposed subgeneric divisions 
of Diuraphis, the data examined in this 
study provides support for Diuraphis sensu 
stricto (D. mexicana + D. noxia) which 
was recovered with bootstrap support of 


70%. This clade is united by a slight pos- 
terior-lateral protuberance on the base of 
the scape and a medially raised dorsal sur- 
face of abdominal segment VIII. The 
grouping of D. mexicana + D. noxia carries 
interesting biogeographic implications since 
D. mexicana and D. noxia are of Nearctic 
and Palearctic origin, respectively. 

The remaining Diuraphis species, how- 
ever, were not recovered as a monophyletic 
sister-group to the D. mexicana + D. noxia 
clade and thus, it would be inappropriate to 
treat this as a subgeneric division of Diura- 
phis sensu lato. Therefore, although there is 
support for Diuraphis sensu stricto, we con- 
clude that it is not instructive to recognize 
the proposed subgenus Holcaphis (e.g., 
Eastop and Hille Ris Lambers 1976; Re- 
maudiére and Remaudiére 1997), since it 
most likely represents a non-monophyletic 


VOLUME 107, NUMBER 3 


grouping of the remaining Diuraphis spe- 
cies. 


DIAGNOSIS OF THE GENUS DIURAPHIS 


Diuraphis is characterized by an elongate 
body, relatively short antennae, antennal tu- 
bercles low or weakly developed, abdomi- 
nal dorsum usually without pigmented 
sclerites anterior to segment VI but some 
species have intersegmental abdominal 
sclerites, cornicles inconspicuous and usu- 
ally without an apical flange, and first tarsal 
segments in adults with 3—3—2 setae. Some 
species have abdominal tubercles, spinal 
supracaudal process present or absent, and 
dorsal setae occasionally spatulate. Most 
species produce wax. Diuraphis species 
generally are associated with leaves of var- 
ious cultivated and wild grasses (Poaceae). 


KEY TO APTERAE AND ALATAE DIURAPHIS 
(alatae of D. calamagrostis not included 
due to insufficient material) 


1. Abdominal segment VIII supracaudal process pre- 
sent as a well developed fingerlike projection (Fig. 
9D); cornicle with apical flange; lateral prothorac- 
ic tubercles and abdominal marginal tubercles pre- 
SEITE GN. GG oo cudhe Ener OTe, CRE, ch Deemer ea D. noxia 

— Abdominal segment VIII supracaudal process 

either absent, slightly raised, or present as a 
conical or triangular protuberance but not fin- 
gerlike; cornicle without apical flange; lateral 
prothoracic tubercles and marginal tubercles 


(ETS OF AOS noscdscocccs0vaaeadac 2 
2. Prothoracic tubercles or marginal tubercles pre- 

GE 6 o1d,a'c b 6 cuakewb alc) mo clo Ofbldin oO amo aisore 3} 
— Prothoracic tubercles or marginal tubercles ab- 

REE o odd wo Deel e ORE ONUn OL ROG CECE Some acacia 4 


3. Length of setae on antennal segment III ap- 
proximately % the diameter of the base; apterae 
with supracaudal process on abdominal seg- 
ment VIII present as a conical or triangular 
protuberance (Fig. 8D), abdomen without in- 
tersegmental sclerites, cauda triangular with 
pointed apex D. mexicana 

— Length of setae on antennal segment III sube- 
qual to the diameter of the base; apterae with- 
out supracaudal process on abdominal segment 
VIL, abdomen with intersegmental sclerites, 
cauda parallel sided with bluntly rounded apex 
Begskey D. calamagrostis (apterae only) (in part) 

4. Ultimate rostral segment with a pair of acces- 
SOGYISCLAG Hota e atid ehela dene! = eee te tools isos oh 5 


705 


— Ultimate rostral segment without accessory se- 


5. Length of antennal segment III shorter than an- 
tennal segments IV + VY; ultimate rostral seg- 
ment approximately 2 times as long as wide at 
base D. agropyronophaga 

— Length of antennal segment III subequal to 
longer than antennal segments [TV + V; ulti- 
mate rostral segment approximately 3 times as 
long asi widevatibaSeneisna sae D. tritici 

6. Abdomen with intersegmental sclerites, al- 
though sclerites may be extremely reduced in 
some alata; length of hind tarsus II shorter or 
subequal to length of cauda............. 7 

— Abdomen without intersegmental sclerites; 
length of hind tarsus II longer than length of 
cauda 
7. Legs stout, e.g., greatest width of hind tibiae 
subequal or wider than the length of penulti- 
mate antennal segment; rostrum length sube- 
qual to width at base 


D. calamagrostis (apterae only) (in part) 

— Legs more slender, e.g., greatest width of hind 
tibiae less than the length of penultimate an- 
tennal segment; rostrum longer than width at 


DASE 2s ets ton as Sie aes ee een Oem eae 8 
8. Antennal segment III usually longer than seg- 

ment IV + V, occasionally subequal to seg- 

ment IV+V; cornicles short, approximately % 

to % the length of the cauda ........ D. holci 


— Antennal segment III usually shorter than seg- 
ment IV + V, occasionally subequal to seg- 
ment IV+V; cornicles very short, porelike, ap- 
proximately ,, to % the length of the cauda 
wc) ig ee iy SEER), SSH, SEES J. are ats D. agrostidis 

9. Cornicle unpigmented, porelike, Y%) to 9 the 
length of the cauda; dorsal sclerites on abdom- 
inal segment VII absent or reduced to a few 
scattered polygonally sclerotized areas 
Lshatateicies automat ote tog: gates meee am Rem eNTaw ame D. bromicola 

— Cornicle pigmented, short but elongate, , to 
the length of the cauda; dorsal sclerites on ab- 
dominal segment VII well developed, extend- 

INC MMearhy stOKS PLAGE eatin arsenal D. frequens 


Diuraphis agropyronophaga G.-x. Zhang 
(Fig. 2) 

Diuraphis (Holcaphis) agropyronophaga 
G.-x. Zhang, 1991:327; Zhang et al.1991: 
123; Remaudiére and Remaudiére 1997: 
9] 


Type material.—Aptera holotype, No. 
6324-1-1-2, on Agropyron sp., Nei Mongol 
Automous Region, Fregzhen County, 19- 
VI-1976, G.-x. Zhang and T.-s. Zhong coll., 
not seen. We have studied a single paratype 


706 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 2. 
host plant information represents a summary of those 
s plants listed in the Specimens Examined sections. Al- 
though some of the host data may represent aberrant 
hosts, they are included for reference purposes. When 
common names for various hosts were used, they are 
recorded as such on the list and the scientific name is 
added in parentheses for cross-reference. Botanical 
names listed in the collection data were checked 
against the Integrated Taxonomic Information System 
(Anonymous 2004a), The International Plant Names 
Index (Anonymous 2004b), and the National Genetic 
Resources Program, Germplasm Resources Informa- 
tion Network (Anonymous 2004c). 


Host plants of Diuraphis. The following 


Agrostis stolonifera 
Diuraphis agrostidis Muddathir 
Agropyron sp. 
Diuraphis agropyronophaga G.-x. Zhang 
Diuraphis noxia (Kurdjumov) 
Avena sp. 
Diuraphis tritici (Gillette) 
Barley (see Hordeum sp.) 


Bromus catharticus Vahl 
Diuraphis mexicana (McVicar Baker) 
Diuraphis noxia (Kurdjumov) 
Bromus carinatus Hook. & Arn. 
Diuraphis mexicana (McVicar Baker) 
Diuraphis noxia (Kurdjumov) 
Bromus inermis Leyss. 


Diuraphis bromicola (Hille Ris Lambers) 


Bromus marginatus Nees ex Steud. 


Diuraphis mexicana (McVicar Baker) 
Diuraphis tritici (Gillette) 


Bromus polyanthus Scribn. 
Diuraphis mexicana (McVicar Baker) 
Bromus tectorum L. 
Diuraphis noxia (Kurdjumov) 
Bromus sp. 
Diuraphis mexicana (McVicar Baker) 
Calamagrostis lanceolata , 
Diuraphis calmagrostis (Ossiannilsson) 
Calamagrostis purpurea 
Diuraphis calmagrostis (Ossiannilsson) 
Downy brome (see Bromus tectorum) 
Echinochloa crus-galli (L.) Beauv. 
Diuraphis frequens (Walker) 
Elymus dahuricus Turcz. ex Griseb. 


Diuraphis frequens (Walker) 


Table 2. Continued. 


Elymus glaucus Buckl. 

Diuraphis frequens (Walker) 
Elymus sp. 

Diuraphis tritici (Gillette) 

Elytrigia repens var. repens 

Diuraphis frequens (Walker) 
Holcus lanatus L. 

Diuraphis holci (Hille Ris Lambers) 
Holcus mollis L. 

Diuraphis holci (Hille Ris Lambers) 
Hordeum murinum L. 

Diuraphis noxia (Kurdjumoy) 
Hordeum vulgare L. 

Diuraphis holci (Hille Ris Lambers) 

Diuraphis noxia (Kurdjumoy) 
Hordeum sp. 

Diuraphis noxia (Kurdjumov) 
Mountain brome (see Bromus marginatus) 
Oats (see Avena sp.) 

Pascopyrum smithii (Rydb.) A. Love 

Diuraphis tritici (Gillette) 

Phleum pratense L. 

Diuraphis noxia (Kurdjumov) 
Phleum pratense ssp. nodosum (L.) Arcang. 

Diuraphis frequens (Walker) 

Diuraphis noxia (Kurdjumoy) 
Phleum sp. 

Diuraphis noxia (Kurdjumoy) 
Quack grass (see Elytrigia repens var. repens) 
Triticum aestivum L. 

Diuraphis noxia (Kurdjumov) 
Triticum sp. 

Diuraphis noxia (Kurdjumov) 

Diuraphis tritici (Gillette) 


Wheat (see Triticum sp.) 


slide deposited in IZAS with left label, 
**9921—I—1 Agropyron 90.VI.1” and right 
label, ““PARATYPES, Holcaphis agropy- 
ronophaga ZHANG 16-VI-1990 Ningxia 
China.’ Additional paratypes listed in 
Zhang et al. (1991). 

Field features——Aptera grayish white, 


VOLUME 107, NUMBER 3 


707 


9,3) 332 


Figs. 2—3. 
side, aptera venter of head and antennal segments I-II. B, Antenna of alata. C, Cornicle of aptera. D, Cauda of 
aptera. 3, D. agrostidis. A, Right side, aptera dorsum of head and antennal segments; left side, aptera venter of 
head and antennal segments I-II. B, Antenna of alata. C, Cornicle of aptera. D, Cauda of aptera. 


covered with white powder (Zhang et al. 
1991). 

Recognition characters.—Aptera: Body 
length 2.220—2.232; width through eyes, 
0.378. Antenna (Fig. 2A) shorter than body; 
segment III 0.150—0.216 long; IV 0.066— 
0.102 long; V 0.066—0.078 long; base of VI 


2, Diuraphis agropyronophaga. A, Right side, aptera dorsum of head and antennal segments; left 


0.084—0.108 long; terminal process, 0.084— 
0.132 long. Head sclerotized, smooth, with- 
out spinulation; longest dorsal head setae 
subequal to width of antennal segment III. 
Rostrum extending to mesocoxae; ultimate 
segment 0.108—0.114 long, approximately 2 
times as long as wide at base, subequal to 


708 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


hind tarsal segment II, with 2 accessory se- 
tae. Pronotum without marginal tubercles. 
Hind tibia 0.468—0.570 long; hind tarsus II 
0.114 long. Abdomen smooth with ventral 
surface spiculose, dorsal surface spiculose 
on segments VI-VIII, with sclerite on seg- 
ment VII-VIII; marginal abdominal tuber- 
cles and supracaudal process absent. Cor- 
nicle (Fig. 2B) pigmented, short, 0.030 
long, apical flange undeveloped; without 
associated basal sclerite. Cauda (Fig. 2C) 
0.138 long, elongate, triangular, with slight 
medial constriction, 6—7 lateral setae. 

Alata: Body length 1.416; width through 
eyes 0.390—0.342. Antenna (Fig. 2D) short- 
er than body; segment HI 0.246—0.228 long, 
with 6—7 secondary sensoria restricted to 
approximately half circumference of seg- 
ment; IV, 0.102 long, with 2—4 secondary 
sensoria; V, 0.090—0.096 long, without sec- 
ondary sensoria; base of VI, 0.102—0.114 
long; terminal process, 0.138 long. Dorsal 
head setae longer than width of antennal 
segment III. Rostral length and setae similar 
to apterous female, ultimate segment 
0.102—0.108 long with 2 accessory setae. 
Pronotum without marginal tubercles. Hind 
tibia 0.600—0.618 long; hind tarsus II 
0.126—0.132 long. Abdominal tergum with- 
out patches or bands, surface sculpturing 
similar to aptera, abdominal tergite VIII 
with 3—5 setae; abdominal marginal tuber- 
cles and supracaudal process absent. Cor- 
nicle short, smooth, 0.024 long, similar to 
aptera. Cauda 0.120—0.120 long, similar to 
aptera, with 6 lateral setae. 

Notes.—Although Zhang (1991) illus- 
trates D. agropyronophaga with a slightly 
raised area on abdominal tergite VIII, this 
structure was not discernable on the speci- 
mens we examined. 

Diuraphis agropyronophaga resembles 
D. tritici. However, the ultimate rostral seg- 
ment is shorter than D. tritici (approximate- 
ly 2 times as long as wide at the base versus 
approximately 3 times as long as wide at 
the base for D. tritici). Also, in D. agro- 
pyronophaga, the length of antennal seg- 
ment III is shorter than antennal segments 


IV + V whereas the length of antennal seg- 
ment III is subequal to longer than antennal 
segments IV + V in D. tritici. See also 
Notes section for D. tritici. 

There has been some confusion in the lit- 
erature concerning the publication date of 
D. agropyronophaga (Zhang 1991). The 
date was listed by Remaudiére and Remau- 
diére (1997) as 1990, the date that appears 
in the title of Zhang’s (1991) publication. 
However, this work was not published until 
May 1991 (as printed on the publication). 
A description of D. agropyronophaga was 
also listed as “sp. nov.” in Zhang et al. 
(1991), but, because the actual time of is- 
suance has not been determined, that work 
would be listed as the last day in 1991. 

Specimens examined.—CHINA: Ni- 
ngxia, ex Agropyron sp., IZAS: 16-VI-1990 
(2 ap. paratypes, 2 al. paratypes, 2 alatoid 
imm. labeled ““PARATYPES’’). 


Diuraphis agrostidis (Muddathir) 
(Fig. 3) 


Holcaphis agrostidis Muddathir 1965:477. 

Diuraphis (Holcaphis) agrostidis: Eastop 
and Hille Ris Lambers 1976:175; Remau- 
diére and Remaudiére 1997:91. 


Type material.—Holotype, morphotypes, 
and paratypes deposited in BMNH (Mud- 
dathir 1965); not seen. 

Field features.—Aptera yellow green 
with white powder; head dark, almost 
black; antennae, legs, and cauda dark (Heie 
19D): 

Recognition characters.—Aptera: Body 
length 1.470—2.100; width through eyes 
0.318—0.384. Antenna (Fig. 3A) shorter 
than body; segment III 0.090—0.162 long; 
IV 0.060—0.084 long; V 0.072—0.084 long; 
base of VI 0.066—0.084 long; terminal pro- 
cess 0.102—0.120 long. Head sclerotized, 
smooth, without spinulation; longest dorsal 
head setae longer than width of antennal 
segment III. Rostrum ending before meso- 
coxae; ultimate segment 0.066—0.084 long, 
approximately 1.5 times as long as wide at 
base, shorter than hind tarsal segment II, 


VOLUME 107, NUMBER 3 


without accessory setae. Pronotum without 
marginal tubercles. Hind tibia 0 .390—0.420 
long; hind tarsus II 0.108—0.120 long. Ab- 
domen smooth with ventral surface spicu- 
lose, dorsal surface spiculose on segments 
VI-VIII, with small pleural and interseg- 
mental sclerites, large sclerites on segments 
V-VIII; abdominal marginal tubercles and 
supracaudal process absent. Cornicle (Fig. 
3B) slightly pigmented, short, 0.018—0.024 
long, apical flange undeveloped; with as- 
sociated basal sclerite. Cauda (Fig. 3C) 
0.120—0.144 long, elongate, triangular with 
rounded apex, 6 lateral setae and usually 
one preapical seta. 

Alata: Body length 1.860—1.902; width 
through eyes 0.360—0.372. Antenna (Fig. 
3D) shorter than body; segment IIT 0.192— 
0.198 long, with 4—5 secondary sensoria re- 
stricted to approximately half circumfer- 
ence of segment; IV 0.078—0.102 long, with 
O—1 secondary sensoria; V 0.090—0.102 
long, without secondary sensoria; base of 
VI 0.102—0.108 long; terminal process 
0.192—0.210 long. Dorsal head setae sube- 
qual to width antennal segment III. Rostral 
length and setae similar to apterous female, 
ultimate segment 0.060—0.072 long without 
accessory setae. Pronotum without margin- 
al tubercles. Hind tibia 0.540—0.636 long; 
hind tarsus IT 0.108—0.114 long. Abdominal 
surface sculpturing and sclerotization simi- 
lar to aptera, abdominal tergite VIII with 3— 
5 setae; abdominal marginal tubercles and 
supracaudal process absent. Cornicle short, 
smooth, 0.018 long, similar to aptera. Cau- 
da 0.120 long, similar to aptera. 

Notes.—Diuraphis agrostidis most 
closely resembles D. holci. Apterae and ala- 
tae of D. agrostidis have intersegmental 
sclerites that are larger than the adjacent 
spiracle and associated sclerotized area, and 
antennal segment III is usually shorter than 
antennal segments IV + V. Conversely, ap- 
terae and alatae of D. holci have interseg- 
mental sclerites that are subequal or smaller 
than adjacent spiracle and associated scler- 
otized and antennal segment ILI is usually 
longer than antennal segments IV + V. The 


709 


host of D. agrostidis is Agrostis stolonifera 
L. (Muddathir 1965, Heie 1992) whereas 
the host of D. holci are Holcus spp. See also 
Notes section for D. bromicola, D. fre- 
quens, and D. holci. 

Specimens examined.—UNITED KING- 
DOM: SCOTLAND: Frazerburgh, ex grass, 
H.L.G.S. coll., BMNH: 13-VIII-1959 (8 
ap., 2 imm.). ENGLAND: Northumberland, 
Newcastle, King’s College, ex Agrostis sto- 
lonifera, K. Muddathir coll., BMNH: 8-?- 
1963 (2 al.). 


Diuraphis bromicola (Hille Ris Lambers) 
(Fig. 4) 


Holcaphis bromicola Hille Ris Lambers 
1959-291 

Diuraphis (Holcaphis) bromicola: Eastop 
and Hille Ris Lambers 1976:175; Remau- 
diere and Remaudieére 1997:91. 


Type material—Two cotype slides seen. 
One slide with left label ““Holcaphis brom- 
icola nov. spec cotypes Det. D.H.R.L.” and 
right label ““N. Germany Pl. Bromus iner- 
mis oc.) Leipzie (Date (Oct toaom lace 
Miihle BM1984-340 ““(BMNH). Another 
slide with left label ““Holcaphis bromicola 
nov. spec cotypes Det. D.H.R.L.” and right 
label ““N. Germany Pl. Bromus inermis 
Loc. Leipzig Date 20-VI-1957 Leg. Mihle 
BM1984-340 “(BMNH). Cotypes of D. 
bromicola were originally deposited in the 
collection of Hill Ris Lambers (1959) but 
are presently in BMNH. 

Field features.—Oviparous female very 
light green with grey wax-powder; alata 
similar to ovipara but darker with less pow- 
der, head and thorax black (Hille Ris Lamb- 
ers) 1959): 

Recognition characters.—Aptfera: Body 
length 1.950—2.100; width through eyes 
0.348—0.360. Antenna (Fig. 4A) shorter 
than body; segment III 0.204—0.246 long; 
IV 0.096—0.108 long; V 0.090—0.102 long; 
base of VI 0.096—0.108 long: terminal pro- 
cess, 0.096—0.126 long. Head sclerotized, 
smooth, without spinulation; longest dorsal 
head setae shorter than width of antennal 


710 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 4—5. 4, Diuraphis bromicola. A, Right side, aptera dorsum of head and antennal segments; left side, 
aptera venter of head and antennal segments I-II. B, Antenna of alata. C, Cornicle of aptera. D, Cauda of aptera. 
5, D. calamagrostis. A, Right side, aptera dorsum of head and antennal segments; left side, aptera venter of 
head and antennal segments [-II. B, Cornicle of aptera. C, Cauda of aptera. 


segment III. Rostrum ending before meso- 
coxae; ultimate segment 0.072 long, ap- 
proximately 1.5 times as long as wide at 
base, shorter than hind tarsal segment II, 
without accessory setae. Pronotum without 
marginal tubercles. Hind tibia 0 .552—0.630 
long; hind tarsus II 0.132—0.144 long. Ab- 
domen smooth with ventral surface spicu- 
lose, dorsal surface spiculose on segments 
VI-VIII, small sclerites on segment VII and 
large sclerite on VIII; abdominal marginal 
tubercles and supracaudal process absent. 
Cornicle (Fig. 4B) very short, porelike, 
0.006—0.012 long, unpigmented. Cauda 
(Fig. 4C) 0.108—0.114 long, elongate, tri- 


angular with rounded apex, 4—6 lateral se- 
tae. 

Alata: Body length 1.650—1.818; width 
through eyes 0.330—0.384. Antenna (Fig. 
4D) shorter than body; segment III 0.252— 
0.276 long, with 3—5 secondary sensoria re- 
stricted to approximately half circumfer- 
ence of segment; IV 0.138—0.162 long, with 
O—1(usually absent) secondary sensoria; V 
0.120—0.144 long, without secondary sen- 
soria; base of VI 0.108—0.126 long; termi- 
nal process 0.150—0.162 long. Dorsal head 
setae shorter than width antennal segment 
III. Rostral length and setae similar to ap- 
terous female, ultimate segment 0.060— 


VOLUME 107, NUMBER 3 


0.072 long without accessory setae. Pro- 
notum without marginal tubercles. Hind tib- 
ia 0.642—0.744 long; hind tarsus II 0.132— 
0.150 long. Abdominal surface sculpturing 
and sclerotization similar to aptera, abdom- 
inal tergite VIII with 4 setae; abdominal 
marginal tubercles and supracaudal process 
absent. Cornicle very short, porelike, 
0.006—0.012 long, unpigmented, similar to 
aptera. Cauda 0.102—0.114 long, similar to 
aptera. 

Notes.—The original species description 
of D. bromicola was based upon oviparae, 
alate viviparae, and apterous males (Hille 
Ris Lambers 1959). The present description 
of the apterous female is the first time this 
stage of D. bromicola has been described. 

Diuraphis bromicola most closely resem- 
bles D. agrostidis, D. frequens, and D. holci 
but can be distinguished by the presence of 
porelike, unpigmented cornicles. Diuraphis 
agrostidis, D. frequens, and D. holci all 
have pigmented cornicles. Aptera and alata 
of D. bromicola also can be distinguished 
from D. agrostidis and D. holci by the ab- 
sence of intersegmental sclerites hind tarsus 
II is longer than the cauda. Diuraphis 
agrostidis and D. holci both have interseg- 
mental sclerties and hind tarsus II is shorter 
than the cauda. Aptera and alata of D. 
bromicola can be distinguished from those 
of D. frequens by the reduced dorsal scler- 
ite on abdominal segment VII. In D. fre- 
quens this sclerite is well developed. See 
also Notes section for D. agrostidis, D. fre- 
quens, and D. holci. 

Specimens examined.—GERMANY: 
Leipzig, ex Bromus inermis, Mihle, 
BMNH: X-1956 (4 ov. labeled “‘cotypes’’); 
20-VI-1957 (2 al. labeled “‘cotypes’”’ ); Leip- 
zig, ex Bromus inermis BMNH: 12-VII- 
1959 (2 ap., 2 al.). RUSSIA: Kursk, ex Bro- 
mus inermis, Agarvonova, BMNH: VII- 
HOS OR @rale): 


Diuraphis calamagrostis (Ossiannilsson) 
(Fig. 5) 


Holcaphis calamagrostis Ossiannilsson 
1959-25. 


711 


Diuraphis (Holcaphis) calamagrostis: Eas- 
top and Hille Ris Lambers 1976:175; Re- 
maudiere and Remaudiére 1997:91. 


Type material.—Holotype and paratypes 
deposited in the collection of the Institute 
of Plant Pathology and Entomology, Upp- 
sala, Sweden (Ossiannilsson 1959); how- 
ever, we have studied a single paratype 
slide deposited in BMNH. The paratype 
slide examined has left label ‘“Holcaphis 
calamagrostis Ossiannilsson apterae para- 
types BM 1984-340 Det. Ossiannilsson”’; 
right label ““N. Sweden Pl. Calmagrostis 
purpurea Loc. Jalla Vaksala date 1-VII- 
1952 Leg. Ossiannilsson 3317” (BMNH). 

Field features.—Aptera pale yellow, 
head, antenna and legs fuscous; alata simi- 
lar to aptera but thorax entirely fuscous, ab- 
dominal segments II-VI occasionally with 
large dark pigmented marginal sclerites 
(Ossiannilsson 1959). 

Recognition characters.—Aptera: Body 
length 1.566—2.538; width through eyes 
0.270—0.420. Antenna (Fig. 5A) shorter 
than body, 5- or 6-segmented; on 5-seg- 
mented specimens, segment III 0.114— 
0.174 long; IV 0.030—0.072 long; base of 
V 0.060—0.090 long; terminal process 
0.078—0.126 long; on 6-segmented speci- 
mens, segment III 0.096—0.150 long; IV 
0.054—0.078 long; V 0.060—0.072 long; 
base of VI 0.066—0.084 long; terminal pro- 
cess 0.114—0.138 long. Head sclerotized, 
smooth, without spinulation; longest dorsal 
head setae subequal to width of antennal 
segment III. Rostrum extending just past 
procoxae; ultimate segment 0.054—0.072 
long, subequal to width at base, shorter than 
hind tarsal segment II, occasionally with a 
single accessory seta. Pronotum with or 
without marginal tubercles. Hind tibia 
0.288—0.510 long; hind tarsus Il 0.090— 
0.126 long. Abdomen smooth with ventral 
surface spiculose, dorsal surface spiculose 
on segments VI-VIII, with small pleural 
and intersegmental sclerites, large sclerites 
on segments VI-VIII and occasionally a 
small sclerite on segment V; marginal tu- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


bercles present or absent; supracaudal pro- 
cess absent. Cornicle (Fig. 5B) short, 
0.012—0.024 long, with associated basal 
sclerite. Cauda (Fig. 5C) 0.096—0.138 long, 
elongate, parallel-sided to slightly triangu- 
lar with rounded apex, 4—6 lateral setae and 
sometimes one preapical seta. 

Alata: Not seen. See Ossiannilsson 
(1959) and Heie (1992) for description. 

Notes.—The presence of abdominal mar- 
ginal tubercles in D. calamagrostis may be 
variable. Kovalev et al. (1991) listed these 
structures as present in D. calamagrosStis; 
conversely, Heie (1992) placed this species 
in the Holcaphis group defined in part by 
the ‘“‘normal absence of marginal tubercles 
from abd. segm. H-—VI.” Ossiannilsson’s 
(1959) description of the aptera indicates 
the presence of marginal abdominal tuber- 
cles. However, in his diagnosis section, he 
states “‘the normal presence of marginal tu- 
bercles on some of the abdominal seg- 
ments’’ as a diagnostic character. This sug- 
gests that the marginal tubercles may not 
always be present. Examination of a single 
slide containing four paratypes revealed 
three of the specimens did not exhibit mar- 
ginal abdominal tubercles and one of the 
specimens exhibited prominent marginal 
abdominal tubercles. The paratype that ex- 
hibited abdominal tubercles also had pro- 
notal tubercles. The remaining paratypes 
that did not have abdominal tubercles, like- 
wise did not have pronotal tubercles. In an 
additional slide of D. calamagrostis deter- 
mined by Ossiannilsson, all eight determin- 
able adults had abdominal marginal tuber- 
cles. Pronotal marginal tubercles were not 
visible in all specimens. 

Diuraphis calamagrostis may be con- 
fused with other species of Diuraphis with 
intersegmental abdominal sclerites (D. 
agrostidis and D. holci), however, the ros- 
trum of D. calamagrostis extends only just 
past the procoxae. In D. agrostidis and D. 
holci, the rostrum extends to the mesocox- 
ae. In addition, the ultimate segment in D. 
calamagrostis 1s subequal to the width at 


its base. In D. agrostidis and D. holci, the 
rostrum is longer than the width at its base. 

Specimens examined.—SWEDEN: Sol- 
na:, ex Calamagrostis lanceolata, Ossian- 
nilsson, BMNH: 28-VIII-1948 (8 ap., 3 
imm., | indeterminable); Jalla Vaksala,, ex 
Calamagrostis purpurea, Ossiannilsson 
BMNH: 1-VII-1952 (4 ap. labeled “‘para- 
types’’). 


Diuraphis frequens (Walker) 
(Fig. 6) 


Aphis frequens Walker 1848:2219. 

Brachycolus  korotnewi Mordvilko 1901: 
325; Hille Ris Lambers 1939:97 [synon- 
ymy with Holcaphis frequens (Walker)] 

Holcaphis frequens: Hille Ris Lambers 
ISBO7. 

Diuraphis (Holcaphis) frequens: Eastop 
and Hille Ris Lambers, 1976:176; Re- 
maudiére and Remaudiére 1997:91. 

Diuraphis (Holcaphis) elymophila G.-x. 
Zhang 1991:327; Zhang et al., 1991:125. 
New synonymy. 


Type material.—The type depository for 
Aphis frequens was recorded as BMNH 
(Hille Ris Lambers 1939). The type speci- 
mens were not seen. Walker’s (1848) orig- 
inal description is based on oviparae and 
apterous males. Type material for Brachy- 
colus korotnewi is unknown. Type deposi- 
tory for Diuraphis elymophila is IZAS, ho- 
lotype and five paratypes seen. 

Field features.—Aptera geen with dark 
green head; antennae, legs and cauda black 
fuscous (Hille Ris Lambers 1939), cornicles 
brown (Heie 1992), covered with gray wax- 
powder (Hille Ris Lambers 1939). Ovipara 
dark green, somewhat glaucous, mottled 
with yellow, powdered with white; eyes 
dark red; antenna black, yellow at base; 
legs dull yellow, tarsi and tips of tibiae 
black (Walker 1848). Adult male wingless, 
darker than ovipara; antenna black (Walker 
1848). 

Recognition characters.—Aptera: Body 
length 1.560—2.070; width through eyes 
0.342-—0.378. Antenna (Fig. 6A) shorter 


VOLUME 107, NUMBER 3 


7M} 


Figs. 6-7. 
aptera venter of head and antennal segments I—II. B, Antenna of alata. C, Cornicle of aptera. D, Cauda of aptera. 
7, D. holci. A, Right side, aptera dorsum of head and antennal segments; left side, aptera venter of head and 
antennal segments I-II. B, Antenna of alata. C, Cornicle of aptera. D, Cauda of aptera. 


than body, 6-segmented; segment III 0.108— 
0.192 long; IV 0.072—0.114 long; V 0.066— 
0.090 long; base of VI 0.066—0.102 long; 
terminal process 0.090—0.120 long. Head 
sclerotized, smooth, with faint reticulate 
spinulation; longest dorsal head setae less 
than width of antennal segment III. Ros- 
trum extending to mesocoxae; ultimate seg- 
ment 0.066—0.084 long, approximately 


6, Diuraphis frequens. A, Right side, aptera dorsum of head and antennal segments; left side, 


1.36—1.66 times as long as side at base, 
shorter than hind tarsal segment II, acces- 
sory setae absent. Pronotum without mar- 
ginal tubercles. Hind tibia 0.420—0.600 
long; hind tarsus Il 0.114—0.156 long. Ab- 
domen smooth with ventral surface spicu- 
lose, dorsal surface spiculose on segments 
VI-VIII, pleural and intersegmental scler- 
ites absent, large sclerites on segments VI— 


714 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Vill; segment VIII sometimes with slight 
supracaudal process; abdominal marginal 
tubercles absent. Cornicle (Fig. 6B) equi- 
distant from spiracular openings on abdom- 
inal segments VI and VII, pigmented, short, 
0.025—0.031 long, apical flange undevel- 
oped; without associated basal sclerite. 
Cauda (Fig. 6C) 0.090—0.126 long, elon- 
gate, triangular with rounded apex, 4—6 lat- 
eral setae. 

Alata: Body length 1.800—2.130; width 
through eyes 0.360—0.372. Antenna (Fig. 
6C) shorter than body; segment II 0.186— 
0.270 long, with 4—6 secondary sensoria re- 
stricted to approximately half circumfer- 
ence of segment; IV 0.138—0.180 long, with 
O-—2 secondary sensoria; V 0.096—0.126 
long, without secondary sensoria; base of 
VI 0.096—0.114 long; terminal process 
0.168—0.198 long. Dorsal head setae shorter 
than width antennal segment III. Rostral 
length and setae similar to apterous female, 
ultimate segment 0.078—0.084 long without 
accessory setae. Hind tibia 0.660—0.732 
long; hind tarsus II 0.120—0.160 long. Ab- 
dominal surface sculpturing and sclerotiza- 
tion similar to aptera, abdominal tergite 
VIII with 4 setae; segment VIII sometimes 
with slight supracaudal process; lateral ab- 
dominal tubercles absent. Cornicle short, 
0.031 long, similar to aptera. Cauda 0.090— 
0.126 long, similar to aptera. 

Notes.—After examination of the type 
series, descriptions, and illustrations of D. 
elymophila Zhang et al. (1991), we have 
concluded that D. elymophila is a junior 
synonym of D. frequens. Zhang et al. 
(1991) believed D. elymophila differed 
from D. frequens by the length of the ter- 
minal process as compared to the base. 
They stated that the terminal process is 1.5 
times as long as the base in D. elymophila 
which distinguishes it from D. frequens, 
and this character was used for species sep- 
aration in a key (Zhang et al. 1991). The 
use of this character is suspect for two rea- 
sons. First, Zhang et al. (1991) stated in a 
subsequent couplet that contained D. fre- 
quens, that the terminal process was 1.66 


times the length of the base. Secondly, ex- 
amination of D. frequens specimens from 
the BMNH and USNM revealed that the 
length of the terminal process as compared 
to the base ranged from 1.1 to 1.7 for ap- 
terae and 1.4 to 2.0 for alatae. Heie (1992) 
recorded the length of the terminal process 
as 1.05—1.50 times the base for apterae and 
up to 1.75 times the base for alatae. Al- 
though the original description of D. fre- 
quens records Artemisia maritima L. (As- 
teraceae) as the host (Walker 1848), D. fre- 
quens has been recorded from other hosts 
including Agropyron repens | = Elytrigia 
repens var. repens (L.) Desv. ex B.D. Jack- 
son] (Heie 1992 and BMNH and USNM 
slide data) and Elymus glaucus Buckl. 
(USNM slide data). Diuraphis frequens 
previously has been recorded as distributed 
in Mongolia (Heie 1992) and is reported in 
the current work as occurring on another 
species of Elymus (E. dahuricus Turcz. ex 
Griseb.). 

As with D. agropyronophaga, there has 
also been confusion concerning the publi- 
cation date for D. elymophila (Zhang 1991). 
Remaudiére and Remaudiére (1997) listed 
1990 as the date for the description but it 
should be regarded as 1991. See also dis- 
cussion of D. agropyronophaga for details. 

Diuraphis frequens is most similar to D. 
agrostidis, D. bromicola, and D. holci. Ap- 
tera and alata of D. frequens are distin- 
guished from D. agrostidis by the absence 
of intersegmental sclerites, the presence of 
a short but elongate cornicle that is approx- 
imately equidistant between spiracles VI 
and VII, and the absence of a dorsal sclerite 
on segment VI. In D. agrostidis, aptera and 
alata exhibit intersegmental sclerites, the 
cornicle is porelike and situated closer to 
spiracle VI than VII, and a dorsal sclerite 
is present on segment VI. Aptera and alata 
of D. frequens are distinguished from D. 
bromicola by a short but sclerotized corni- 
cle and the presence of a large dorsal scler- 
ite on segment VII. In D. bromicola, the 
cornicle is porelike and unsclerotized and a 
large dorsal sclerite on segment VII is ab- 


VOLUME 107, NUMBER 3 


sent. Aptera and alata of D. frequens are 
distinguished from D. holci by the absence 
of intersegmental sclerites and antennal 
segment III is usually shorter than antennal 
segments IV + V. Conversely, D. holci ap- 
tera and alata exhibit intersegmental scler- 
ites and antennal segment III is usually lon- 
ger than antennal segments IV + V. See 
also Notes section for D. agrostidis, D. 
bromicola, and D. holci. 

Specimens examined.—AUSTRIA: Vi- 
enna, ex grass, R. & L. Burkhart coll., 
USNM: 27-V-1991 (1 ap., 2 al.). CANA- 
DA: MANITOBA: St. Pierre, ex Agropyron 
repens {|= Elytrigia repens var. repens], C. 
C. Gill coll., BMNH: 25-VIII-1965 (1 ap.); 
Montreal, ex Triticum repens [= Elytrigia 
repens var. repens], D.H.R.L. coll., BMNH: 
21-VIII-1956 (6 ap. on 2 sl.); NEW 
BRUNSWICK: Fredericton, ex Phleum 
pratense (?) and Echinochloa crus-galli, J. 
B. Adams coll., BMNH: 21-VIII-1964 (7 
ap. on 2 sl.). CHINA: Nei Mongol, ex Ely- 
mus dahuricus, 1ZAS: 27-VU-1984 (4 ap., 
2 al.on 2 sl., one slide labeled ““HOLO- 
TYPE” and “PARATYPES’’) ITALY: near 
Udine, ex grass, R.&L. Burkhart coll., 
USNM: 1-VII-1991 (2 ap., 1 al.). NETH- 
ERLANDS: Bennekom, ex Holcus (?), 
H.L.G.S. coll., BMNH: 20-VI-1949 (3 ap., 
1 al.); Wageningen, ex grass, D. Gonzalez 
coll., USNM: V-1990 (9 ap., 19 imm.). 
UNITED KINGDOM: ENGLAND, Tyne 
& Wear, Fatfield, BMNH: 31-VII-1948 (4 
ap.) ;? Essex, Harlow Hill, BMNH: 28- 
VIII-1948 (4 ap.); Herts., Rothamsted, G. 
D. Heathcote coll., BMNH: 8-VI-1965 (2 
al.); Middlesex, Enfield, ex grass blade, J. 
H. Martin, BMNH: 23-IX-1989 (6 ap.) 
UNITED STATES: COLORADO, Gunni- 
son Co., Litthke Gunnison Cr., W. Elk Wil- 
derness, ex Elymus sp., R. Hammon coll., 
USNM: 2-VIII-1990 (15 ap., 2 al., 27 imm. 
on 19 sl.); Throughline Trailhead, West Elk 
Wilderness, ex Elymus glaucus, R. Ham- 
mon and FE Judson coll., USNM: 11-VIII- 
1992 (2 ap., 4 imm.); Little Gunnison 
Creek, ex Elymus glaucus, F M. Judson 
coll., USNM: 16-IX-1990 (8 ap. on 3 sl.); 


715 


Routt Co., Rabbit Ears Pass, Ferndale Pic- 
nic Ground, ex Elymus sp., EF M. Judson 
coll., USNM: 12-VIII-1990 (4 ap., 5 al., 1 
imm. on 6 sl.); NORTH DAKOTA, Fargo, 
ex suction trap, USNM: 29-VII-1966 (1 
al.); Oregon, EK F Hasbrouck coll., USNM: 
11-VI-1953 (1 al.); WASHINGTON, Quin- 
cy, ex grass, B. J. Landis, USNM: 10-VI- 
1959 (1 al.); WYOMING, Fremont Co., ex 
quackgrass [ = Elytrigia repens var. re- 
pens], C. Wilbert coll., USNM: 29-VIII- 
1986 (14 ap., 6 imm. on 6 sl.). UNKNOWN 
LOCATIONS: ex Agropyron repens {| = 
Elytrigia repens var. repens|, BMNH: 1-IX- 
1968 (3 ap., 1 imm.); “‘Yellow pan Trap’’, 
A. Frowd coll., USNM: 26-VI-1975 (1 al.); 
“yellow-trap’, R. Sigvald coll., USNM: 
1976 (1 al.); “trapped, USNM: no data (1 
alae 


Diuraphis holci (Hille Ris Lambers) 
(Fig. 7) 


Aphis holci Hardy 1850:531; Hille Ris 
Lambers, 1956:229. Nomen nudum. 

Holcaphis holci Hille Ris Lambers 1939: 
97; Hille Ris Lambers 1956:229. 

Diuraphis (Holcaphis) holci: Eastop and 
Hille Ris Lambers, 1976:176; Heie 1992: 
104; Remaudiére and Remaudiére 1997: 
91. 


Type material.—One slide labeled cotype 
seen. Slide with left label “‘Holcaphis holci 
HR. sexuales  cotypes’ Det?) DIE-RE 
and right label ““N. Fd.-pl. Holcus lanatus 
Loc. Bennekom Date 3-XI-43 Leg. 
D.H.R.L.”” (BMNH). 

Field features.—Aptera green with dark 
green head and extremities; “covered with 
whitish grey waxpowder” (Hille Ris Lamb- 
ers 1939), cornicles and cauda brownish 
(Here 1992): 

Recognition characters.—Aptera: Body 
length 1.860—2.160; width through eyes 
0.378—0.444. Antenna (Fig. 7A) shorter 
than body; segment III 0.150—0.204 long; 
IV 0.072—0.102 long; V 0.072—0.102 long; 
base of VI 0.084—0.102 long; terminal pro- 
cess 0.102—0.120 long. Head sclerotized, 


716 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


smooth, with faint reticulate spinulation; 
longest dorsal head setae longer than width 
of antennal segment III. Rostrum extending 
prior to mesocoxae; ultimate segment 
0.066—0.084 long, approximately 1.4—1.8 
times as long as wide at base, shorter than 
hind tarsal segment II, without accessory 
setae. Pronotum without marginal tubercles. 
Hind tibia 0.528—0.660 long; hind tarsus II 
0.108—0.138 long. Abdomen smooth with 
ventral surface spiculose, dorsal surface 
spiculose on segments VI-—VIII, usually 
with small pleural and intersegmental scler- 
ites, intersegmental sclerites sometimes 
greatly reduced, large dorsal sclerites usu- 
ally present on segments VI-VIII; abdomi- 
nal marginal tubercles and supracaudal pro- 
cess absent. Cornicle (Fig. 7B) short, 
0.036—0.054 long. Cauda (Fig. 7C) 0.126— 
0.144 long, elongate, triangular with round- 
ed apex, 4—6 lateral setae and 0-2 preapical 
setae. 

Alata: Body length 1.500—1.590; width 
through eyes 0.333—0.372. Antenna (Fig. 
7D) shorter than body; segment III 0.198— 
0.216 long, with 4—5 secondary sensoria re- 
stricted to approximately half circumfer- 
ence of segment; IV 0.078—0.096 long, with 
O-—1 secondary sensoria; V 0.090—0.102 
long, without secondary sensoria; base of 
VI 0.102—0.120 long; terminal process 
0.180—0.210 long. Dorsal head setae longer 
than width antennal segment III. Rostral 
length and setae similar to apterous female, 
ultimate segment 0.066—0.078 long, occa- 
sionally with a single accessory setae. Hind 
tibia 0.600—0.642 long; hind tarsus II 
0.120—0.126 long. Abdominal surface 
sculpturing and sclerotization similar to ap- 
tera, abdominal tergite VIII with 3—5 setae; 
lateral abdominal tubercles and supracaudal 
process absent. Cornicle short, smooth, 
0.034—0.049 long, similar to aptera. Cauda 
0.120—0.132 long, similar to aptera. 

Notes.—Diuraphis holci was first de- 
scribed as Holcaphis holci (Hardy) in 1939 
by Hille Ris Lambers. Later, Hille Ris 
Lambers (1956) noted that Aphis holci Har- 
dy was actually a nomen nudum and 


‘“‘should be quoted H. holci H.R.L., 1947.” 
However, Hille Ris Lambers’s citation of 
the year “1947” work was probably in er- 
ror. The citation of any 1947 work men- 
tioned by Hille Ris Lambers was not listed 
in the literature section in Hille Ris Lamb- 
ers (1956). Examination of all of Hille Ris 
Lambers publications for 1947 did yield the 
description of Schizaphis holci Hille Ris 
Lambers (1947) that year. However, this 
species is not a synonym of D. holci and 
Hille Ris Lambers’s (1956) reference to 
1947 should be considered an error. Thus, 
the correct date for making the species de- 
scription available is 1939. 

Diuraphis holci is most similar to D. 
agrostidis, D. bromicola, and D. frequens. 
Aptera and alata of D. holci may be distin- 
guished from D. agrostidis by intersegmen- 
tal sclerites that are nearly equal or slightly 
smaller than the sclerotized area of the ab- 
dominal spiracles, and the presence of lon- 
ger cornicles. In D. agrostidis, intersegmen- 
tal sclerites are larger than the sclerotized 
area of the abdominal spiracles, and the 
cornicles are very small, nearly porelike. 
Aptera and alata of D. holci may be distin- 
guished from D. bromicola and D. frequens 
by dorsal head setae that are longer than the 
widest portion of antennal segment III. The 
dorsal head setae in both the aptera and ala- 
ta of D. bromicola and D. frequens are 
shorter than the greatest width of antennal 
segment III. In addition, aptera and alata of 
D. holci may be distinguished from D. 
bromicola by the length of hind tarsus II 
longer than the cauda, and the presence of 
pigmented cornicles. Aptera and alata of D. 
bromicola have a hind tarsus II that is 
shorter than the cauda, and the cornicles are 
unpigmented. The aptera and alata of D. 
holci may be distinguished from those of 
D. frequens by the length of hind tarsus II 
as compared to the caudal length and the 
position of the cornicle relative to adjacent 
spiracles. In D. holci, hind tarsus II is short- 
er than the length of the cauda and cornicle 
is closer to abdominal spiracle VI than VII. 
In D. frequens, hind tarsus II is longer than 


VOLUME 107, NUMBER 3 


the cauda and the cornicle is equidistant be- 
tween abdominal spiracle VI and VII. See 
also the Notes section for D. agrostidis, D. 
bromicola, and D. frequens. 

Specimens examined.—NETHER- 
LANDS: Bennekom, ex Holcus lanatus, 
A.G. Robinson & H.R.L. coll., USNM:17- 
VIII-1970 (1 ap., 1 nymph). UNITED KIN- 
DOM: ENGLAND, Berkshire, Silwood 
Park, ex grass, J. Packham via S. McNeill 
coll., BMNH: 11-VII-1980 (4 al.) ; Cum- 
berland, ex Holcus mollis, EH. Jacobs coll., 
BMNH: 21-VII-1943 (3 ap., 9 imm.); 
Northumberland, Ciosforth Park, ex Holcus 
mollis, K. Muddathir coll., BMNH: 12-VII- 
1965 (3 al.); Welton, ex Hordeum vulgare, 
BMNH: 3-VII-1972 (2 ap., 1 nymph); Sur- 
rey, Wisley Gardens, ex? Triticum repens, 
H.L.G.S. coll., BMNH: 29-X-1954 (3 ap.); 
Richmond Park, ex Holcus, V.EE. coll., 
BMNH: 19-VI-1972 (3 ap.); Kew Gardens, 
ex Holcus lanatus, V.EE. coll., BMNH: 9- 
IX-1962 (5 ap.). 


Diuraphis mexicana (McVicar Baker) 
(Fig. 8) 


Cuernavaca mexicana McVicar Baker 
1934: 210. 

Diuraphis mexicana: Aizenberg 1956:154. 

Bracycolus nodulus Richards 1959: 251. 

Diuraphis nodulus: Eastop and Hille Ris 
Lambers 1976:175; Pefia-Martinez 1981: 
178 [synonomy with Diuraphis mexicana 
(McVicar Baker)]| 

Diuraphis (Diuraphis) mexicana: Eastop 
and Hille Ris Lambers, 1976:175; Re- 
maudiére, and Remaudiere 1997:91. 


Type material.—Eighteen slides labeled 
““Cotype”’ deposited in USNM and listed 
below. 

Field features.—Not recorded. 

Recognition characters.—Aprtera (Fig. 
8A): Body length 1.620—1.860; width 
through eyes 0.372-0.414. Antenna (Fig. 
8B) shorter than body; segment HI 0.132— 


0.234 long; IV 0.078—0.126 long; base of 


V 0.096—0.108 long; terminal process 
0.102-0.126 long. Head _ sclerotized, 


TAY) 


smooth, with faint reticulate spinulation; 
dorsal head setae blunt-tipped, longest dor- 
sal head setae shorter than width of anten- 
nal segment III. Rostrum extending to me- 
socoxae; approximately 1.4—1.7 times as 
long as wide at base, ultimate segment 
0.066—0.072 long, without accessory setae 
and shorter than hind tarsal segment II. Pro- 
thorax with marginal tubercles, occasional- 
ly only one present. Hind tibia 0.486—0.648 
jong; hind tarsus II 0.108—0.138 long. Ab- 
domen (Fig. 8C) smooth with ventral sur- 
face spiculose, dorsal setae blunt-tipped, 
short; dorsal surface spiculose on segments 
VI-VII, with sclerite on segments VII— 
VIII; slight supracaudal process (Fig. 8D) 
on segment VIII, deltoid shaped, process 
usually with a pair of lateral setae and a 
basal pair of setae; abdominal marginal tu- 
bercles (Fig. 8E) usually present, occasion- 
ally absent. Cornicle (Fig. 8F) short, 0.030— 
0.042 long, without apical flange. Cauda 
(Fig. 8G) 0.108—0.144 long, elongate, tri- 
angular, usually with 4 lateral setae and O— 
2 preapical setae. 

Alata (Fig. 8H): Body length 1.380— 
1.578; width through eyes 0.270—0.284. 
Antenna (Fig. 81) shorter than body; seg- 
ment III 0.252—0.294 long, with 4—7 sec- 
ondary sensoria distributed in a straight 
row; IV 0.150—0.174 long, with O-—2 sec- 
ondary sensoria; V 0.132—0.162 long, with- 
out secondary sensoria; base of VI 0.108— 
0.132 long; terminal process 0.132—0.156 
long. Head setae and spinulation similar to 
apterous female. Rostral length and setae 
similar to apterous female, ultimate seg- 
ment 0.060—0.072 long. Prothorax with lat- 
eral marginal tubercles, occasionally only 
one present. Mesosternum with rows of mi- 
nute spinules; hind tibia 0.558—0.768 long; 
hind tarsus IT 0.120—0.138 long. Wing veins 
with fuscous highlighting. Abdominal ter- 
gum without patches or bands, surface 
sculpturing similar to aptera, supracaudal 
process on segment VIII sometimes re- 
duced, abdominal tergite VIII with 3—5 se- 
tae; abdominal marginal tubercles usually 
present, occasionally absent. Cornicle short, 


/i8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Se 
eas 


Weep 


Fig. 8. Diuraphis mexicana. A, Aptera ventral and dorsal habitus. B, Right side, aptera dorsum of head and 
antennal segments; left side, aptera venter of head and antennal segments I-II. C, Aptera lateral abdominal 
habitus. D, Supracaudal process. E, Abdominal tubercle. K Cornicle of aptera. G, Cauda of aptera. H, Alata 
dorsal and ventral habitus. I, Antenna of alata. J, Alata lateral abdominal habitus. 


VOLUME 107, NUMBER 3 


0.024—0.042 long, similar to aptera. Cauda 
0.120—0.132 long, similar to aptera, usually 
with 4 lateral setae and O—2 preapical setae. 
Notes.—Diuraphis mexicana most close- 
ly resembles D. noxia, however, the supra- 
caudal process of the aptera of D. mexicana 
is more deltoid-shaped. In D. noxia, this 
process is more fingerlike. In addition, the 
cornicle of D. mexicana does not have an 
apical flange whereas that of D. noxia does. 
See also Notes section of D. noxia. 
Specimens examined.—CANADA: 
BRITISH COLUMBIA, Summerland, ex 
orchard grasses, D. P. Pielou coll., CNCI: 
6-IX-1955 (2 ov., 8 al., | imm. on 11 sl. 
labeled “‘Holotype’? and “Paratype” of 
Brachycolus nodulus); Summerland, ex 
downy brome [= Bromus tectorum], M. B. 
Stoetzel coll., USNM: 5-IX-1990, 7-Ix- 
1990 (56 ap., 22 imm. on 34 sl.). MEXICO: 
Colonia Andahuae, Ciudad de Mexico, ex 
Bromus proximus genuinus [?},USNM: 16- 
VI-1934 (10 ap., 21 al., 34 imm. on 18 sl. 
labeled “‘cotype’’); Saltillo, Coahnila., ex 
Bromus unioloides [= Bromus catharticus], 
R. V. Carapia coll., USNM: 20-I-1988 (10 
ap., 10 al. on 10 sl.); D.E, ENCB-IPN, ex 
Bromus sp., R. Pena coll., USNM: 14-II- 
1991, 1-IIJ-1991 (42 ap., 23 al., 19 imm. on 
27 sl.); Xochimilco, ex Bromus catharticus, 
Pena and Stoetzel coll., USNM: 3-IV- 
1991,4-IV-1991, 5-IV-1991, 6-IV-1991 (59 
ape 2 al-e93 umm: on Sil sl): El Batan; 
CIMMYT- Lab colony, ex Bromus carina- 
tus, M. B. Stoetzel coll., USNM: 5-IV-1991 
(4 ap., 2 al., 4 imm. on 2 sl.) UNITED 
STATES: COLORADO, Rio Blanco Co., 
UCEPC-Meeker, ex Bromus marginatus, R. 
Hammon coll., USNM: 31-V-1990, 14-VI- 
1990, 12-IX-1991, 8-X-1991(63 ov., 82 ap., 
Meal so sm. of 75) sl.)> Ft. Collis, CO 
State Univ., ex Bromus marginatus, W. 
Meyer coll., USNM: 23-XII-1991 (6 al., 3 
imm. on 4 sl.); NEW MEXICO, Cimarron, 
R.S. Ranch, ex Bromus polyanthus, USNM: 
[no date] (31 ov., 1 ap., 5 imm. on 11 sl.); 
Meeker, ex mountain brome [= Bromus 
marginatus], R. W. Hammon coll., USNM: 


719 


8-14-X-1992 (30 ap., 5 ov., 6 ap. 6, 5 imm. 
ony 19s15): 


Diuraphis noxia (Kurdjumov) 
(Fig. 9) 


Brachycolus noxius Kurdjumov 1913:13. 

Diuraphis noxius: Aizenberg 1935:157. 

Cavahyalopterus graminarium Mimeur 
1942:67. 

Brachycolus miihlei Boérner 1950: 9. New 
synonymy. 

Cavahyalopterus noxius: Bodenheimer and 
Swirski 1957:287. 

Diuraphis noxia: Eastop and Hille Ris 
Lambers 1976:175; Diirr 1983:81; Ko- 
valev et. al. 1991:425; Remaudiére and 
Remaudiére 1997:91. 


Nomenclatural notes.—For nearly 80 
years there was some confusion as to the 
author of D. noxia. Kovalev et al. (1991) 
provided clarification on the authorship of 
this species. Kurdjumov (1913) not only 
gave information of D. noxia life history 
and morphological characters but also de- 
veloped a key for separating this species 
from other species of grain-damaging 
aphids. Mordvilko in Kurdjumovy’s (1913) 
work proposed the new epithet, Brachyco- 
lus noxius Mordvilko, but never published 
its description. 

Type material.—Types of D. noxia not 
seen. Slide of C. graminarium labeled 
‘““paratype’’ (MNHN) and slide of B. mueh- 
lei labeled *““Typen”’ (URIC) seen, listed be- 
low. 

Field features.—Aptera green (Durr 
1983), pale yellow green, or greygreen, 
wax-powdered (Heie 1992); eyes reddish 
(Diirr 1983). Alata head dark, thorax with 
dark spots; abdomen pale green, not prui- 
nose (Heie 1992). 

Recognition characters.—Aptera (Fig. 
9A): Body length 1.344—2.490; width 
through eyes, 0.330—0.450. Antenna (Fig. 
9B) shorter than body, 6-segmented; seg- 
ment III 0.096—0.192 long; IV 0.054—0.120 
long; V 0.066—0.114 long; base of VI 
0.060—0.102 long; terminal process 0.120— 


20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 9. Diuraphis noxia. A, Aptera ventral and dorsal habitus. B, Right side, aptera dorsum of head and 
antennal segments; left side, aptera venter of head and antennal segments I-IJ. C, Aptera lateral abdominal 
habitus. D, Supracaudal process. E, Abdominal tubercle. EK Cornicle of aptera. G, Cauda of aptera. H, Alata 
dorsal and ventral habitus. I, Antenna of alata. J, Alata lateral abdominal habitus. 


VOLUME 107, NUMBER 3 


0.180 long. Head sclerotized, smooth, oc- 
casionally with faint spinulation; dorsal 
head setae blunt-tipped, longest dorsal head 
setae shorter than width of antennal seg- 
ment III. Rostrum extending to mesocoxae; 
ultimate segment 0.072—0.090 long, ap- 
proximately 2 times as long as wide at base, 
without accessory setae and shorter than 
hind tarsal segment IJ. Prothorax with mar- 
ginal tubercles, occasionally only one pres- 
ent. Hind tibia 0.402—0.678 long; hind tar- 
sus II 0.090—0.138 long. Abdomen (Fig. 
9C) smooth with ventral surface spiculose, 
dorsal setae blunt-tipped, short; dorsal sur- 
face spiculose on segments VII—VIUII, with 
sclerite on segments VI—VIII; supracaudal 
process (Fig. 9D) on segment VIII, finger- 
like, process usually with a pair of lateral 
setae and a basal pair of setae; abdominal 
marginal tubercles (Fig. 9E) present. Cor- 
nicle (Fig. 9F) short, 0.042—0.048 long. 
Cauda (Fig. 9G) 0.090—0.180 long, elon- 
gate, triangular, with 4—6 lateral setae and 
Q—2 preapical setae. 

Alata (Fig. 9H): Body length 1.332- 
1.980; width through eyes 0 .300—0.420. 
Antenna (Fig. 91) shorter than body, 6-seg- 
mented; segment III 0.126—0.198 long, with 
3-7 secondary sensoria distributed in a 
straight row; IV 0.078—0.132 long, with 1— 
3 secondary sensoria; V 0.072—0.132 long, 
without secondary sensoria; base of VI 
0.072—0.102 long; terminal process, 0.162— 
0.216 long. Head setae and spinulation sim- 
ilar to apterous female. Rostral length and 
setae similar to apterous female, ultimate 
segment 0.066—0.078 long. Prothorax with 
marginal tubercles, occasionally only one 
present. Hind tibia 0.540—0.750 long; hind 
tarsus IT 0.102—0.132 long. Wing veins with 
fuscous highlighting. Abdomen (Fig. 9J) 
with lateral sclerites, tergum with sclerites 
on segments VII—-VIII, surface sculpturing 
similar to aptera, abdominal tergite VIII 
with 8—10 setae including those of supra- 
caudal process, supracaudal process on seg- 
ment VIII more reduced than aptera; mar- 
ginal abdominal tubercles present. Cornicle, 
0.036—0.054 long, similar to aptera. Cauda 


721 


0.114—0.156 long, similar to aptera, usually 
with 4—6 lateral setae and | preapical seta. 

Notes.—We place D. muehlei as a junior 
synonym of D. noxia. In Borner’s (1950) 
brief original description of D. muehlei, he 
noted that it was similar to D. noxia but had 
a shorter supracaudal process on the eighth 
abdominal segment (%4 to % the length of 
the cauda versus D. noxia with a supracau- 
dal process of % to % the length of the cau- 
da). He also stated that D. muehlei caused 
leaf rolling in Phleum pratense. Differences 
in the ratio between the supracaudal process 
have been confirmed by subsequent work- 
ers since BOrner’s original description (e.g., 
Shaposhnikov 1964, Kovalev et al. 1991, 
Zhang et al. 1991, Heie 1992). Examination 
of Borner’s types of D. muehlei revealed 
that the slide labeled ““Typen”’ contained 
Oviparae, apterous males, and immatures. 
While males were mentioned in the descrip- 
tion, there was no statement concerning the 
stage of the adult females. Bérner’s use of 
the supracaudal-process-to-caudal-length 
character for species elevation is suspect. 
The length of the supracaudal process and 
the length of the cauda of this series is with- 
in his published range for D. muehlei. Al- 
though the supracaudal process of D. noxia 
is often half as long as or more than the 
caudal length, a large series of oviparae in 
the USNM collected from a laboratory col- 
ony revealed some specimens that exhibited 
a short supracaudal-process-to-caudal ratio 
within the range described by Borner for D. 
muehlei. Examination of the supracaudal 
processes for alatae and apterae from other 
laboratory colonies of D. noxia also pro- 
duced individuals with a short supracaudal- 
process-to-caudal ratio. 

Thomas Thieme (personal communica- 
tion) adds important information to the D. 
muehlei story. According to Dr. Thieme, in 
1944 Borner originally received specimens 
of D. muehlei from Miihle who kept the 
aphids in culture on Phleum pratense. Later, 
Borner received additional material and he 
also kept the aphids in culture. These 
aphids were slide mounted (also in 1944) 


(22 


then described as D. muehlei and are rep- 
resented on Boérner’s ““Typen”’ slide. Sub- 
sequent workers have listed P. pratense as 
the sole host of D. muehlei (Shaposhnikov 
1964, Kovalev et al. 1991, Zhang et al. 
1991, Heie 1992). Phleum pratense has also 
been recorded as a host for D. noxia (Ko- 
valev et al. 1991, Heie 1992). 

Specimens examined.—AFGHANI- 
STAN: Paghman, ex wheat, E.J. Hambleton 
coll., USNM: 21-VI-1961 (3 al.); Corizi- 
Mir, ex alfalfa, E.J. Hambleton coll., 
USNM: 22-VI-1961 (1 al.). ALGERIA: Be- 
lezma (Constantine), ex Hordeum sativum 
[= Hordeum vulgare], Frezal coll., MNHN: 
V-1949 (3 ap., 2 al. on 2 sl.); Kenchela, 
Laporte coll., MNHN: 19-III-1951 (2 al.). 
CANADA: BRITISH COLUMBIA, Cres- 
ton, ex Phleum pratense, H. Gerber coll., 
USNM: 23-XI-1988 (1 ap.); Abbotsford, ex 
air suction trap, C. K. Chan coll., USNM: 
8-VI-1992 (1 al.); Vancouver, ex Agropy- 
ron, C. K. Chan coll., USNM: 21-IX-1992 
(1 ap.,l al. on 2 sl.); 14-I-1993 (2 ap.). 
CHILE: Santiago, M. Zekene coll., USNM: 
23-XI-1987 (2 ap., 18 al. on 10 sl.); San- 
tiago, Crianza laboratory, M. Zerene coll., 
USNM: 2-V-1988 (22 ap., 4 al. on 14 sl.); 
Llai Llay, ex wheat, D. Reed coll., USNM: 
12-XI-1990 (7 ap., 5 imm. on 8 sl.). IRAN: 
Karadj, ex Gramineae, G. Remaudiere coll., 
MNHN: 12-VII-1955 (1 al.); Mashad (?), 
ex Hordeum sp., G. Remaudiére coll., 
MNHN: 28-IX-1955 (2 ap.); Saadatabad, ex 
Triticum sp., G. Remaudiére coll., MNHN: 
4-V-1959 (4 ap.); 40 km E. Mashad, ex 
Hordeum sp., G. Remaudiere coll., MNHN: 
13-V-1966 (4 ap., 2 al. on 2 sl.); Neysha- 
pour, 97 km. s.w. from Mashad, ex wheat, 
D. Gonzalez coll., USNM: 12-V-1990 (23 
ap., 3 imm. on 11 sl.). ITALY: Brunico, ex 
Phleum sp., BMNH: 12-VI-1965 (2 al.). 
FRANCE: Perpignan (P.O.), ex Hordeum 
murinum, G. Remaudieére coll., USNM: III- 
1971 (2 ap., 2 al.); Behoust, EPL lab colony 
[origins: France, Antibes, 6-VI-1989, ex 
wheat, K. Hopper and G. Mercadier coll.; 
France SE, 764—1,150 m altitude, 7-VI- 5— 
6-VII 1989, G. Mercadier and G. Gruber 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


coll.; Jorden, DISI, 13-V-1989, ex wheat, 
K. S. Pike and L. Tanigoshi coll.; Syria, 
Aleppo, V-1989, ex wheat, R. Miller coll.; 
Turkey, Beypazari, 110 km w. Ankara, 14— 
19 VI 1988, ex barley, T. Paprawski and EF 
Gruber coll.; Moldavia, Kischenev and vi- 
cinity, 28 V- 2 VI 1989, ex wheat and bar- 
ley, T. Paprawski and E Gruber coll.; Unit- 
ed States, South Dakota, Brookings, VIII- 
1988, ex wheat, N. Elliott coll.], ex barley 
or wheat, M. B. Stoetzel coll., USNM: 7- 
IX-1989 (47 ap., 35 al., 8 imm. on 24 sl.); 
10-VIII-1989 (72 ap., 89 imm. on 30 sl1.); 
11-VITI-1989 (102 ap., 1 al., 139 imm. on 
36 sl.); unknown collection date (86 ap., 33 
al., 2 imm. on 58 sl.). GERMANY: ex 
Phleum pratense, Mihlei/Borner coll., 
URIC: 14-IV-1944 (6 ov., 1 ap. 6, 5 imm. 
on | sl. labeled ““Typen’’); Leipzig, BMNH: 
27-V1I-1959, ex Phleum pratense, (2 ap., 1 
al., 2imm.). LIBYA: Tripolitania, ex wheat, 
Hambleton coll., USNM: 6-H-1959 (2 ap., 
1 al., 1 immature on 2 sl.); 10-IV-1959 (2 
ap., 2 al. on 2 sl.); Tripoli, Azizia, ex to- 
matoes, Hambleton coll., USNM: 4-V-1960 
(3 al.); Bentmaya, Fezz-an, ex wheat, R. L. 
Linkfield coll., USNM: 5-IV-1962 (2 ap., 1 
al., 1 immature on 2 sl.). MEXICO: Saltillo, 
Coah, ex Bromus unioloides |= Bromus ca- 
tharticus|, R. V. Carapia coll., USNM: 20- 
I-1988 (14 ap., 20 al. on 12 sl.); El Baton, 
CIMYT lab colony, ex Bromus carinatus, 
USNM: 5-IV-1991 (8 ap., 1 al., 2 imm. on 
3 sl.);.Xochimilco, ex Bromus catharticus, 
Pefia and Stoetzel coll., USNM: 4-IV-1991 
(17 ap., 1 al., 2 imm. on 8 sl.); 5-IV-1991 
(1 ap., 1 al.); 6-I[V-1991 (8 ap., 3 al., 3 imm. 
on 5 sl.) MOLDAVIA: S. Halbert coll., 
USNM: 6-XI-1989 (1 ap., 4 al., 38 ov., 3 
ap.d, 1 imm. on 42 sl.). MONGOLIA: Al- 
tay, D. Gonzales coll., USNM: 10-VI-1991 
(18 ap., 29 imm. on 12 sl.); Wuqia, D. Gon- 
zales coll., USNM: 3-VII-1991 (8 ap., 16 
imm. on 5 sl.). MOROCCO: Rabat, ex Trit- 
icum, Bourleau coll., MNHN: 19-I-1939 (1 
ap., 1 al. on 2 sl., labeled “‘paratype’’). 
PAKISTAN: Quetta 1,800 m, ex Triticum 
sp., Inayatulla coll., MNHN: 18-III-1988 (1 
ap., 1 al.). PEOPLES REPUBLIC OF CHI- 


VOLUME 107, NUMBER 3 


NA: Tacheng, ex wheat, D. Gonzalez coll., 
USNM: 28-V-1990 (14 ap., 1 al., 10 imm. 
on 8 sl.); Yining, ex wheat, D. Gonzalez 
coll., USNM: 1-VI-1990 (13 ap., 1 al., 17 
imm. on 8 sl.). SOUTH AFRICA: E du Toit 
coll., USNM: 20-VI-1988 (11 ap., 23 al., 
11 imm. on 9 sl.); Orange Free State, Beth- 
lehem, small grain center, EK du Toit coll., 
USNM: XI-1988 (23 ap., 2 imm. on 6 sl.). 
TURKEY: Isparta-Egridir, ex Triticum, 
Kan., Kanort., and Yesil coll., USNM: 9-V- 
1961 (2 ap., 2 al., 2 imm. on 9 sl.); Kara- 
man, ex Hordeum sativum [|= Hordeum 
vulgare], Remaudieére coll., MNHN: 15-VI- 
1966 (1 al.); (?), ex Triticum sp., Remau- 
diere coll., MNHN: 14-VI-1966 (2 ap.); 
Porsuk, ex wheat, M. Elmali coll., USNM: 
10-VII-1990 (14 ap., 4 al. on 17 sl.); Bug- 
day, ex wheat, M. Elmali coll., USNM: 13- 
VII-1990 (5 ap., 4 al. on 10 sl.). UNITED 
KINGDOM: (?)Lincs., Sleaford, ex Phleum 
pratense, W.H. Golightly coll., BMNH: 15- 
VI-1976, 21-VI-1976 (4 ap., 1 al., 3 imm. 
on 2 sl.); Shaffords Bridge nr. St. Albans, 
ex Phleum nodosum [= Phleum pratense 
ssp. nodosum], R.N.B.P coll., BMNH: 12- 
VIII-1956 (5 ap.). 

UNITED STATES: ARIZONA, Pinal 
Co., Maricopa, ex wheat, D. Fullerton coll., 
USNM: 17-II-1987 (20 ap., 2 al., 15 imm. 
on 8 sl.); COLORADO, Baca Co., Spring- 
field, ex Triticum aestivum, FE B. Peairs 
coll., USNM: 18-IV-1986 (8 ap., 1 al. and 
2 imm. on 3 sl.); Waverly Co., Piedmont 
Farms, ex barley, USNM: 17-VIII-1990 (5 
ap., 1 al. on 2 sl.); Rio Blanco Co., UCEPC- 
Meeker, ex wheat, R. Hammon coll., 
USNM: 8-X-1991 (7 ap., 1 imm. on 4 sl.); 
Ft. Collins, E. Prospect, ex downy brome 
[= Bromus tectorum], USNM: 17-VI-1990 
(2 ap., 1 imm.); Ft. Collins [origin Weld 
Co., Colorado, 1988], ex wheat, USNM: 
16-VII-1990 (13 ap., 6 al., 2 imm. on 7 sl.); 
Meeker, ex wheat, Hammon and Stoetzel 
coll., USNM: 18-VII-1990 (9 ap., 5 al. on 
5 sl.); San Miguel Co., 3 mi w of Egnar, R. 
W. Hammon coll., USNM: 24-X-1995 (1 
ap., 10 imm. on 3 sl.); San Miguel Co., | 
mi s of Egnar, ex wheat, R. W. Hammon 


723 


coll., USNM: 8-XI-1995 (1 ap.); IDAHO, 
Parma [origin Caldwell, ID], Univ. of Ida- 
ho, SW ID R/E Center, M.- g. Feng coll., 
USNM: 27-VI-1987 (2 ap., 2 al. on 4 sl.); 
KANSAS, Stanton Co., ex wheat, P. E. 
Sloderbeck coll., USNM: 5-IV-1986 (2 ap., 
2 imm.); NEW MEXICO, Eddy Co., Carls- 
bad, ex wheat, D. Liesner coll., USNM: 11- 
IV-1986 (1 ap., 1 al., 3 imm.); DeBaco Co., 
Ft. Summer, W. Houghton coll., USNM: 
29-IV-1986 (2 ap., 2 imm.); OKLAHOMA, 
Cimarron Co., Boise City, ex wheat, B. 
Massey and S. Coppack coll., USNM: 9- 
IV-1986 (1 ap., 7 al., 18 imm. on 6 sl.); 
Stillwater [origin Texas, ex wheat, 26-III- 
1986], lab colony, ex winter wheat, R. L. 
Burton coll., USDA: 3-VII-1986 (6 ap., 7 
al., 3 imm. on 7 sl.); Stillwater, lab colony, 
ex winter wheat, R. L. Burton coll., USNM: 
19-VI-1987 (39 ap., 40 al., 48 imm. on 40 
sl.); Jackson Co., nr. Altus, ex wheat, M. 
Karner coll., USNM: 29-ITJ-1988 (5 ap., 2 
imm. on 3 sl.); Stillwater [origin Silverton, 
texas]; 76x barley,nMia "Bs Stoctzelscolle 
USNM: 17-IV-1989 (51 ap., 56 al. on 21 
sl.); Stillwater [origin Hays, Kansas], ex 
barley, M. B. Stoetzel coll., USNM: 20-IV- 
1989 (25 ap., 2 al. on 6 sl.); Stillwater [or- 
igin Ft. Collins, Colorado], ex barley, M.B. 
Stoetzel coll., USNM: 20-IV-1989 (51 ap., 
10 al. on 12 sl.) Stillwater [origin Moscow, 
Idaho]; ‘ex “barley, MB: Stoetzel “coll:; 
USNM: 21-IV-1989 (25 ap., 20 al., 6 imm. 
on 11 sl.); OREGON, Umatilla Co., 4 mi. 
w. Echo, ex wheat, Reed and Stoetzel coll., 
USNM: 18-VI-1990 (17 ap., 18 al., 3 imm. 
on 11 sl.); Hermiston, M. B. Stoetzel coll., 
USNM: 19-VI-1990 (3 ap., 6 al., | imm. on 
3 sl.) SOUTH DAKOTA, Jone Co., nr. 
Murdo, M. E. Gray coll., USNM: 10-VI- 
1986 (1 ap., | al., | imm.). TEXAS, Bailey 
Co., 18 mi SW of Muleshoe, ex wheat, E. 
Leach coll., USNM: 17-III-1986 (2 ap. on 
2 sl.); Swisher Co., ex wheat, L. Bush coll., 
USNM: 20-III-1986 (3 ap., | al., 3 imm. on 
2 sl.); Lubbock Co., 6 mi N Lubbock, ex 
wheat, W. P. Morrison coll., USNM: 25-LI- 
1986 (15 ap., 1 al., 24 imm. on 11 sl.); 
Palmer Co., 5 mi S of Farwell, ex wheat, 


724 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 10. Diuraphis tritici. A, Right side, aptera dorsum of head and antennal segments; left side, aptera 
venter of head and antennal segments III. B, Antenna of alata. C, Cornicle of aptera. D, Cauda of aptera. 


W. R. Morrison coll., USNM: 26-III-1986 
(2 ap., 3 al. on 3 sl.); Tom Green Co., ex 
wheat, T. Fuchs coll., USNM: IV-1986 (2 
ap., 2 imm.); Bushland, ex wheat, Rekle 
coll., USNM: 28-IV-1986 (15 ap., 1 al., 11 
imm. on 10 sl.); Lubbock, ex wheat, G. J. 
Poterka coll., USNM: 27-II-1989 (7 ap., 4 
imm. on 3 sl.); Abernathy, W 1 mile on TX 
597 and S 1 mile on Fm. Rd. 154, ex Bro- 
mus willdenowii [= Bromus catharticus], 
M. B. Stoetzel coll., USNM: 24-IV-1989 (1 
ap., 5 imm. on 2 sl.); Lubbock, Lubbock 
Airport—U.S. 87, ex Bromus tectorum, M. 
B. Stoetzel coll., USNM: 24-IV-1989 (1 
al.); Lubbock Co., U.S. 87 and TX 1294, 
ex Bromus willdenowii [= Bromus cathar- 
ticus|, M. B. Stoetzel coll., USNM: 24-IV- 
1989 (3 ap., 15 imm. on 3 sl.); Terry Co., 
Meadow, ex. Triticum aestivum, M. B. Sto- 
etzel coll., USNM: 25-IV-1989 (7 ap. on 2 
sl.); Swisher Co., 5 miles W Kress, Texas 
on TX 145, ex Triticum aestivum, M. B. 
Stoetzel coll., USNM: 26-IV-1989 (19 ap., 
1 al., 5 imm. on 7 sl.); Bailey Co., ex barley 
in greenhouse, G. J. Puterka coll., USNM: 
15-I-1990 (3 ap. on 4 sl.). WYOMING, 
Laramie Co., Archer Station, ex winter 
wheat, C. Burkhart, J. Edwards, L. Bennett 
coll., USNM: 2-VII-1986 (3 ap., 7 al., 8 


imm. on 5 sl.). YEMEN: Sanaa, ex wheat, 
D. M. Tuttle coll., USNM: 28-X-1977 (2 
ap., 14 al. on 4 sl.). 


Diuraphis tritici (Gillette) 
(Fig. 10) 


Brachycolus tritici Gillette 1911:441. 

Diuraphis (Holcaphis) tritici: Eastop and 
Hille Ris Lambers 1976:176; Remaudiére 
and Remaudiere 1997:91. 


Type material.—Since Gillette (1911) 
did not designate a holotype and maps 
drawn on the original series of slides are in 
error, we have selected a lectotype for clar- 
ification and to ensure stabilization of no- 
menclature. From the syntypes we have se- 
lected an aptera (at the 6 o’clock position) 
as the lectotype and have placed a label on 
the back of the slide with a map of the po- 
sition of the specimen and the label: Diura- 
phis tritici (Gillette) LECTOTYPE & PAR- 
ALECTOTYPES. The original left label 
and illustrated map incorrectly drawn on 
the front of the slide states ““Brachycolus 
tritici, n. sp. Types Al. viv. Apt. viv. C. P. 
Gillette”; top label ““U.S. Nat. Mus. Cat No 
41934”; right label “‘on Agropyron glauc- 
um 5—24—-11 Ft. Collins, Colo. Coll. L. C. 


VOLUME 107, NUMBER 3 


Bragg Colo. Agr. Exp. Sta. Ac. 5—24-11 
mt’d in xyl.balsam’’? (USNM). The slide 
also contains 2 paralectotype alatae and | 
paralectotype aptera. In addition, a single 
paralectoype slide containing 2 apterae and 
13 imm. with original left label and illus- 
trated map incorrectly drawn states “‘Bra- 
chycolus tritici, n. sp. Types 26 others = 
paratypes C. P. Gillette’’; top label “U.S. 
Nat. Mus. Cat No 41934”; right label ‘“‘on 
wheat 10—12-’08 Ft. Collins, Colo. coll. L. 
C. Bragg Colo. Agr. Exp. Sta. Ac. 10—12-— 
08 (mt’d in xyl. balsam) (USNM). This par- 
alectoype slide has a different date (“‘10— 
12-’08”) than that recorded in Gillette’s 
(1911) description (October 10, 1908). In 
addition, there are no males present on the 
slide. However, it is possible that the dis- 
crepancy in dates may have been a typo- 
graphical error. Despite these inaccuracies, 
we believe this slide is one of the original 
syntypes. A third slide labeled ‘‘CO- 
TYPE” in the USNM collection was col- 
lected in 1915, after the publication of the 
original description of D. tritici, and should 
not be considered a syntype. 

Field features.—Aptera: covered with 
fine white powder, body pale green to pale 
yellow, cornicle color similar to body or 
dusky, cauda and legs dusky. Alata: body 
also covered with powder, cauda black; 
head, thorax, and most of antennae black, 
abdomen light green, cornicles yellow or 
brown (Gillette 1911). 

Recognition characters.—Aptera: Body 
length 1.950—2.346; width through eyes, 
0.372—0.462 Head sclerotized, smooth, with 
faint reticulate spinulation; longest dorsal 
head setae shorter than width of antennal 
segment III; tips of dorsal head setae blunt. 
Base of scape without poterior-lateral pro- 
tuberance; sculpturing of anterior margin of 
antennal segment III differs from that of 
posterior margin; Antenna (Fig. 1OA) short- 
er than body; segment III 0.120—0.228 
long; IV 0.066—0.120 long; V 0.072—0.120 
long; base of VI 0.072—0.096 long; terminal 
process, 0.090—0.156 long. Rostrum ex- 
tending to meso-coxae; ultimate segment 


I25 


0.126—0.144 long, approximately 3 times as 
long as wide at base, subequal to hind tarsal 
segment II, with 2 accessory setae. Prono- 
tum without marginal tubercles. Hind tibia 
0.468—0.630 long; hind tarsus II 0.120-— 
0.150 long. Abdomen with faint fine retic- 
ulation on dorsum, surface of segments VI— 
VUI with additional spicules, pleural and 
intersegmental sclerites absent; ventral sur- 
face spiculose; abdominal marginal tuber- 
cles and supracaudal process absent. Cor- 
nicle (Fig. 10B) short, 0.012—0.036 long, 
apical flange undeveloped; associated basal 
sclerite absent. Cauda (Fig. 10C) 0.102— 
0.156 long, elongate, triangular with round- 
ed apex, 4—6 lateral setae and 2 preapical 
setae. 

Alata: Body length 1.590—1.980; width 
through eyes 0.318—0.408. Head spinula- 
tion less distinct and dorsal setae slightly 
shorter in comparison to apterous female. 
Antenna (Fig. 10D) shorter than body; seg- 
ment III of uniform color, 0.162—0.222 
long, with 4—7 secondary sensoria; IV 
0.096—0.120 long, with 1—3 secondary sen- 
soria; V 0.090—0.114 long, with O—I sec- 
ondary sensoria; base of VI 0.090—0.102 
long; terminal process 0.132—0.180 long. 
Rostral length and setae similar to apterous 
female, ultimate segment 0.120—0.144 long, 
with accessory setae. Pronotum without 
marginal tubercles. Hind tibia 0.552—0.696 
long; hind tarsus II 0.126—0.150 long, im- 
bricae usually without spinules. Abdominal 
surface sculpturing and sclerotization simi- 
lar to aptera, abdominal marginal tubercles 
and supracaudal process absent. Cornicle 
similar to aptera; 0.012—0.030 long. Cauda 
0.108—0.144 long, tapered with slight me- 
dial constriction. 

Notes.—Diuraphis tritici can be distin- 
guished from all other species of Diuraphis 
by its long ultimate rostral segment that has 
a single pair of accessory setae. It most 
closely resembles D. agropyronophaga, 
however, in D. tritici the ultimate rostral 
segment is approximately 3 times as long 
as wide at the base and the length of anten- 
nal segment III is subequal to longer than 


726 


antennal segments IV + V. In D. agropy- 
ronophaga, the ultimate rostral segment is 
approximately 2 times as long as wide at 
the base and the length of antennal segment 
III is shorter than antennal segments [V + 
V. See also Notes section for D. agropy- 
ronophaga. 

Specimens examined.—CANADA: Sas- 
katchewan, Swift Current, ex wheat, C. C. 
Gill, USNM: 8-VIII-1967 (1 ap.). UNITED 
STATES: COLORADO, Ft. Collins, ex 
grass, L.C.B., USNM: 31-V-1915 (2 al. on 
2 sl.); Ft. Collins, Colo. Exp. Stn., ex grass, 
Bragg, USNM: 1-VI-1915 (6 al.); Meeker, 
UCEPC lab colony, ex Bromus marginatus, 
R. Hammon, USNM: 2-V-1990 (22 ap. on 
9 sl.); Rio Blanco Co., UCEPC-Meeker, ex 
Mt. Brome ‘“‘Teton, R. Hammon, USNM: 
12-IX-1991 (3 ap.); Meeker, ex mountain 
brome [= Bromus marginatus], R. W. Ham- 
mon, USNM: 8-14-X-1992 (10 ap., 10 
imm. on 20 sl.); KANSAS, Stanton Co., ex 
western wheat, K. O. Bell, USNM: 16-XI- 
1989 (9 ap., 17 imm. on 7 sl.); Jewell Co., 
ex wheat, K. O. Bell, USNM: 13-VI-1990 
(5 ap.,l imm. on 3 sl.); MINNESOTA, ex 
Elymus, USNM: 11-VII-1903 (6 ap., 14 
imm.), 13-VII-1903 (9 ap., 4 al., 10 imm.); 
MONTANA, Judith Basin, ex wheat, S. J. 
Snow, USNM: 1-VI-1915 (lal., 1 imm.); 
Moccasin, ex wheat, J. R. Parker, USNM: 
10-VI-1915 (6 al.); Bozeman, ex wheat, S. 
Lajeunesse, USNM: spring 1986 (2 al.); 
NEW MEXICO, Curry Co., Clovis, ex 
wheat, G. L. Nielson, USNM: 18-VI-1959 
(3 ap.); Los Lunas, ex western wheat grass 
[= Pascopyrum smithii], F Quinones, 
USNM: 18-V-1977 (2 ap., 2 al., 6 imm. on 
3 sl.); Valencia Co., Grants, ex wheat, J. 
Fitzgerald, USNM: 9-V-1986 (3 ap. on 2 
sl.); Clovis, ex wheat, M. Garrett, USNM: 
ANEMITIOS (SS 2yOo5 Jil min, On dS) sil.) 
OKLAHOMA, Sentinel, ex wheat, E. 
Cleveland, USNM: 16-V-1974 (10 ap., 3 
al., 5 imm. on 4 sl.); Stillwater, USDA-ARS 
lab colony [origin Boseman, Montana, ex 
wheat, S. Lajeunesse, spring 1986], G. Pu- 
terka, USNM: 13-II-1992 (12 ap., 12 al., on 
12 sl.); SOUTH DAKOTA, Stanley Co., ex 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Agropyron cristatum and wheat, G. B. Or- 
lob, USNM: VIII-1922 (3 ap.); TEXAS, 
Floydada, ex oats [= Avena sp.], USNM: 
7-V-1922 (1 ap., | al.); Amarillo, ex wheat, 
N. E. Daniels, USNM: 14-III-1952 (5 ap.); 
WASHINGTON, Everson, ex grass, Chris- 
tenson, USNM: 1-III-1941 (4 ap.); WYO- 
MING, Platte Co., ex winter wheat, L. Ben- 
nett, C. Burkhardt, J. Edwards, USMN: 18- 
VI-1986 (5 ap., 7 al., 14 imm. on 5 sl.); 
Wheatland, ex wheat, E.G.L., USNM: no 
date (2 ap., 60 imm.); Ft. Collins [origin 
Weld Co., Colorado, 1988], ex wheat, 
USNM: 16-VH-1990 (13 ap., 6 al., 2 imm. 
on 7 sl.). 


CONCLUSIONS 


This generic evaluation has led to the 
synonymy of two species of Diuraphis (D. 
elymophila = D. frequens and D. muehlei 
= D. noxia). The current phylogenetic anal- 
ysis also suggests that while the monophly 
of the subgenus Diuraphis (sensu stricto) 
(D. noxia + D. mexicana) is supported, the 
currently recognized subgenus Holcaphis 
(i.e., Eastop and Hille Ris Lambers 1976; 
Remaudiere and Remaudiére 1997) is par- 
aphyletic. However, these findings also im- 
ply that the monophyly of Diuraphis (sensu 
lato) is also suspect since B. asparagi was 
included within the clade that contained Di- 
uraphis. Although it was not the purpose of 
this paper to determine the monophyly of 
Brachycolus or Brachycorynella, additional 
studies with these groups are encouraged to 
determine relationships among these two 
genera and Diuraphis. 


ACKNOWLEDGMENTS 


We thank A. Rung and R. Ochoa for their 
input and insight with the phylogenetic 
analysis and to J. Brown and D. Miller for 
manuscript review (USDA, Systematic En- 
tomology Laboratory). We are grateful to E. 
Maw (Agriculture Canada, Ottawa) for a re- 
view of an early draft of the manuscript and 
A. Jensen (Moses Lake, WA) for construc- 
tive discussions on the subject matter and 
his in-depth manuscript review. We are es- 


VOLUME 107, NUMBER 3 


pecially grateful to V. Blackman (BMNH), 
R. Foottit (CNCI), G. Remaudiere (MNHN) 
and T. Thieme (URIC), G.-x. Zhang (IZAS) 
for their assistance, correspondence, and 
loan of specimens. Special thanks are ex- 
tended to two anonymous reviewers for ex- 
cellent suggestions and helpful comments. 


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PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 729-730 


NOTE 


Scoloposcelis pulchella pulchella (Zetterstedt, 1838) in North America 
(Hemiptera: Heteroptera: Anthocoridae) 


The genus Scoloposcelis Fieber is Hol- 
arctic with four species in the Old World 
and two species in the New World. Two 
specimens from Alaska have been identified 
by me as Scoloposcelis pulchella pulchella 
(Zetterstedt, 1838), a Palearctic species 
whose range extends from western Europe 
to Far-Eastern Russia (Péricart 1972, 1996). 
The presence of these specimens in Alaska 
represents the first record for northwestern 
Nearctic Region. Label data are as follows: 
Alaska, Fairbanks, June 26, 1938 G.P. En- 
gelhardt (1 2); and Alaska, E.side Charley 
R., | mi. up from mouth, Circle, Alaska, 15 
Aug 1974. Ed Holsten. in large standing Pi- 
cea glauca (1 @). 

This species was reported from Quebec, 
Canada, as a port-of-entry interception 
(Maw et al. 2000). Scudder (1997), in his 
treatment of the Hemiptera: Heteroptera of 
the Yukon, listed only Scoloposcelis flavi- 
cornis Reuter, described from Texas in 1884 
and reported to be widespread in southern 
Canada and the continental United States 
(Kelton 1978, Henry 1988, Lattin 2000). 
Kelton provided an enlarged electron mi- 
crograph of the ostiola area (fig. 23) show- 
ing a canal quite different from the Alaska 
specimens—a canal whose appearance 
compares very well with this structure pro- 
vided by Péricart (1972) (fig. 152b). Kel- 
ton’s (1978) illustration shows a rather wide 
canal slightly curved anteriorly with a 
rounded apex while Péricart’s illustration 
shows an elongate, narrow canal gently 
curved anteriorly and gradually tapering to 
a sharp apex—near the anterior angle of the 
metapleuron, but not touching. The illustra- 
tion of a male from Spain (Péricart 1972) 
resembles the two Alaska specimens. This 
general appearance occurs in some North 
American species and is quite similar to the 


illustration of the adult of S. flavicornis 
Reuter (Kelton 1978). Reuter (1884: 152) 
described the ostiolar canal of the Texas 
type as “‘apice angularite arcuata bassinque 
versus longius producta pronotum lateribus 
versus basin late sinuatis.”’ Blatchley (1926: 
645) stated that the ‘“‘osteolar channel in our 
species long, angularly curved, not reaching 
base of metasternal plate.’ Both descrip- 
tions differ from Kelton’s (1978) illustra- 
tion of the canal. 

A synonymical review of Scoloposcelis 
pulchella is given below for North Ameri- 
can workers. 


Scoloposcelis pulchella pulchella 
(Zetterstedt, 1838) 


Anthocoris pulchella Zetterstedt 1838: 235. 
Lithuania. 

Scoloposcelis pulchella: Reuter 1884: 152, 
153. Sweden, Finland, France. 

Anthocoris nigrescens Harada 1929: 58. Ja- 
pan. 

Scoloposcelis nigrescens: Hiura 1960: 51. 
Japan. 

Scoloposcelis: Carayon 1972: 341. 

Scoloposcelis pulchella pulchella: Péricart 
1972: 279=283; Elov 1976: 78; 81; 
Zheng and Bu 1990: 27; Zheng and Bu 
1991: 126; Heidger 1995: 541-547; Per- 
icart 1996: 135. Europe, Asia (western 
Siberia, east to Eastern Siberia, Far East- 
ern Siberia); Maw et al. 2000: 102. 


This is another example of a western Pa- 
laearctic species whose range now is known 
to extend to Alaska. This occurrence on ei- 
ther side of the Behring Straight is also 
known for some other species of Hemip- 


tera: Heteroptera (e.g., Acalypta cooleyi 
Drake 1917: 213. Arizona, California, 


Montana, Oregon, Asia [Froeschner 1988: 
709], [Péricart and Golub 1996: 7. Ka- 


730 


zakhstan, China, Mongolia, Russia (East Si- 
beria, Far East)]. Heidger (1995) described 
the biology and ecology of the species. A 
review of North American species of Scol- 
oposcelis is needed. 

Acknowledgments.—My sincere thanks 
to L. Parks for careful attention to the man- 
uscript, T. Lewis for assistance in locating 
critical publications, and to a careful re- 
viewer. 


LITERATURE CITED 


Blatchley, W. S. 1926. Heteroptera or true bugs of 
Eastern North America, with especial reference to 
the faunas of Indiana and Florida. Nature Publ. 
Co., Indianapolis, 1116 pp. 

Carayon, J. 1972. Caractéres systématiques et classi- 
fication des Anthocoridae (Hemipt.). Annales de 
la Société Entomologique de France (N.S.) 8: 
309-349. 

Drake, C. J. 1917. New and noteworthy Tingidae from 
the United States. Ohio Journal of Science 17: 
213-216. 

Elov, E. S. 1976. Bugs of the family Anthocoridae 
(Heteroptera) in Soviet Central Asia and Kazakh- 
stan. Entomologiskoe Obozrenie 60: 369-378 
(English translation, 1976, Entomological Review 
55: 74-81). 

Froeschner, R. C. 1988. Family Tingidae Laporte, 
1807, pp. 708-733. In Henry, T. J. and R. C. 
Froeschner, eds. Catalog of the Heteroptera, or 
True Bugs of Canada and the Continental United 
States. E.J. Brill, Leiden, 958 pp. 

Harada, M. 1929. Ecological study on the scolytid bee- 
tles injurous to Picea jezoensis. Ezomatsu Kisei 
Kikuimushi-rui no setaiteki Kenkyu. I-vi, 1-18 (in 
Japanese). 

Heidger, C. 1995. Zur Biologie und Okologie von 
Scoloposcelis pulchella Zetterstedt (Heteroptera: 
Anthocoridae). Mitteilungen der Deutschen Ge- 
sellschaft fiir Allgemeine und Angewandte Ento- 
mologie 9: 541-547. 

Henry, T. J. 1988. Family Anthocoridae Fieber, 1837, 
pp. 12—28. In Henry, T. J. and R. C. Froeschner, 
eds. Catalog of the Heteroptera, or True Bugs, of 
Canada and the Continental United States. E.J. 
Brill, Leiden, 958 pp. 

Hiura, I. 1960. Contributions to the knowledge of An- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


thocoridae from Japan and its adjacent territories 
(Hemiptera—Heteroptera): 2. Bulletin of the Osa- 
ka Museum of Natural History 12: 43-55. 

Kelton, L. A. 1978. The insects and arachnids of Can- 
ada. Part 4. The Anthocoridae of Canada and 
Alaska. Canada Department of Agriculture, Re- 
search Branch. Publication 1639, 101 pp. 

Lattin, J. D. 2000. Minute pirate bugs (Anthocoridae), 
pp. 607—637. In Schaefer, C. W. and A. R. Panizzi, 
eds. Heteroptera of Economic Importance. CRC 
Press, Boca Raton, Florida, 828 pp. 

Maw, H. E. L., R. G. Foottit, K. G. A. Hamilton, and 
G. G. E. Scudder. 2000. Checklist of the Hemip- 
tera of Canada and Alaska. NRC—-CNRC, NRC 
Research Press, Ottawa, 220 pp. 

Péricart, T. J. 1972. Hémiptéres Anthocoridae: Cimic- 
idae et Microphysidae de |’ Ouest-paléarctique. In 
Faune de |’Europe et du Bassin Méditerranéen. 
Masson, Paris. 7: i-iv, 1—404. 

. 1996. Anthocoridae, pp. 108-140. Jn Auke- 
ma, B. and C. Rieger, eds. Catalog of Hemiptera 
of the Palaearctic Region, Vol. 2. Cimicomorpha 
1. The Netherlands Entomological Society, Am- 
sterdam, 361 pp. 

Péricart, T. J. and V. B. Golub. 1996. Super family 
Tingoidea Laporte, 1832, pp. 3-78. in Aukema, 
B. and C. Rieger, eds. Catalogue of the Heterop- 
tera of the Palaearctic Region, Vol. 2. Cimico- 
morpha I. Netherlands Entomological Society, 
Amsterdam. i—xiv + 361 pp. 

Reuter, O. M. 1884. Monographia Anthocoridarum Or- 
bis terrestris. Helsingforsiae, 204 pp. (also pub- 
lished in Acta Societatis Scientiarum Fennicae 14: 
(1885): 555-758. 

Scudder, G. G. E. 1997. True bugs (Heteroptera) of the 
Yukon, pp. 241-336. In Danks, H. V. and J. A. 
Downes, eds. Insects of the Yukon, Biological 
Survey of Canada (Terrestrial Arthropods), Otta- 
wa, 1,034 pp. 

Zetterstedt, J. W. 1838. Insecta Lapponica descripta. 
Ordo III. Lipsiae. Hemiptera, pp. 257-314. 

Zheng, L. Y. and W. J. Bu. 1990. A list of Anthocor- 
idae from China. Contributions from the Tianjin 
Natural History Museum 7: 23—27 (in Chinese). 

. 1991. Records of anthocorid bugs (Hemip- 

tera) new to China. Acta Zootaxonomica Sinica 

16(1): 126 (in Chinese). 


John D. Lattin, Department of Botany 
and Plant Pathology, Oregon State Univer- 
sity, Corvallis, OR 97331-2902, U.S.A. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 731-732 


NOTE 


Distributional Notes on Bertamyia notata (Loew), Including the First Report of the 
Family Platypezidae from the Caribbean (Diptera: Platypezidae) 


Kessel (1970) erected the genus Berta- 
myia, differentiating it from Agathomyia 
Verrall primarily by its lack of a postsutural 
supra-alar seta; the position of crossvein r- 
m, which is slightly more distal than in 
Agathomyia; and differences in larval struc- 
ture. Bertamyia also differs from known 
Agathomyia in forming a puparial cocoon 
within the fungal host (Kessel et al. 1973). 
The genus contains two species, the type 
species, B. notata (Loew), found widely in 
the Nearctic and Neotropical regions, and 
B. umacibise (Kessel and Clopton) from 
South Africa. Bertamyia notata has been 
reared from the bracket fungus 7Tyromyces 
chioneus (Fr.) P. Karst. (Kessel 1957, Chan- 
dler 2001) in North America. Adults, like 
those of many other platypezids, can be 
found running actively on the upper surfac- 
es of broad leaves. 

Another feature of Bertamyia that is well 
known, but has not been used as a diag- 
nostic feature, is the presence of well-de- 
fined, pale maculations with a light metallic 
bluish or greenish sheen on the dorsum of 
the thorax and on the abdominal tergites of 
both sexes as well as on the frons of the 
female. Only a few other platypezids in the 
genus Agathomyia Verrall (e.g., Agatho- 
myia collini Verrall) have silvery markings 
or a silvery sheen to the thorax (Chandler, 
personal communication). In B. notata, 
these markings are variable which led ear- 
lier authors to describe several species that 
subsequently have been synonymized (Kes- 
sel 1957, Kessel and Pearce 1966). In some 
specimens, these markings appear faint, 
which may be due in part to age of the 
specimen or greasing after preservation. On 
the other hand, the male genitalia have not 
been examined across the geographic range 
of the species, and it is possible that future 
work will reveal additional species. 


Bertamyia notata is widespread in the 
Nearctic Region, ranging from Alaska to 
Northwest Territories and Quebec, south to 
California and Georgia (Kessel and Pearce 
1966). Kessel and Pearce (1966) recorded 
the species from Mexico, Argentina, Brazil, 
and Paraguay in the Neotropical Region. 

I recently collected a single female of B. 
notata in Puerto Rico and a series from the 
Dominican Republic, which prompted me 
to examine material in the collection of the 
National Museum of Natural History, 
Smithsonian Institution (USNM). Addition- 
al distribution records of interest came to 
light. A specimen from Chiapas, Mexico, 
extends the range well east of Oaxaca, the 
southernmost previous record from Mexico 
(Kessel and Pearce 1966). Kessel and 
Pearce (1966) noted that they had not seen 
specimens from Central America. Material 
in the USNM collection from Costa Rica 
and specimens that I collected from Panama 
document the occurrence of B. notata there. 
The USNM collection also has a small se- 
ries from Madre de Dios Department, Peru, 
well west and north of previous South 
American records, and material that I re- 
cently collected in Bolivia. The collection 
also contains specimens from Jamaica, 
which together with my collections from 
Puerto Rico and the Dominican Republic, 
are the first records for the Caribbean and 
suggest a widespread occurrence in the re- 
gion. In fact, the family Platypezidae has 
not previously been recorded from the Ca- 
ribbean (Chandler, personal communica- 
tion), although the USNM collection con- 
tains specimens of an unidentified species 
of Microsania from Dominica, so that pla- 
typezid genus occurs in the Caribbean as 
well. The specific localities of material ex- 
amined are given below. Bertamyia notata 


732 


is a small, inconspicuous fly that is easily 
overlooked. The new records presented 
here provide a strong indication that the 
species occurs throughout much of the Neo- 
tropical Region. 

Adults of B. notata have habits similar to 
many other platypezids. They can be found 
on horizontal surfaces of large leaves in 
semi-moist to wet broadleaf forests, some- 
times in dappled sunlight. They usually are 
seen walking quickly in an erratic fashion 
in a small area on the leaf. 

Material examined (north to south). 
PUERTO RICO: 1 &, Caribbean National 
Forest, Road 186 at Quebrada Grande, 450 
meters, 18°18'24"N, 65°50'00"W, 17 June 
2003, N. E. Woodley. DOMINICAN RE- 
PUBLIC: 4 6, 3 @, Pedernales Province, 
Parque Nacional Sierra de Baoruco, Las 
Abejas, 18°09.011'N, 71°37.342’, 1,150 
meters, 11 July 2004, N. E. Woodley. JA- 
MAICA: 4 6, Runaway Bay, February 
1969, W. W. Wirth, “stream bed”. MEXI- 
CO: 1 d, Chiapas, Finca Prusia, 33 km S 
of Jaltenango, 1,000 meters, 12 May 1985, 
W. N. Mathis. COSTA RICA: 7 @, La Sui- 
za, various dates (April 1922, July 1926, 
August 1926, September 1926), Pablo 
Schild; 1 @, Turrialba, November 1922, 
Pablo Schild; 1 @, Turrialba, 15-19 July 
1965, P. J. Spangler. PANAMA: 1| 2, Bam- 
bito River, 30 August 1952, E S. Blanton; 
1 3, 1 2, Canal Zone, Barro Colorado Is- 
land, 10 June 1978, N. E. Woodley; 6 d, 1 
2, Canal Zone, Chiva Chiva Road, 8 air km 
north of Fort Clayton, 25 July 1978, N. E. 
Woodley. PERU: 2 d, 1 2, Madre de Dios, 
Manu, Rio Manu, Pakitza, 250 meters, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


12°07'S, 70°58'W, 9-23 September 1988, 
Amnon Friedberg; 1 ¢, Madre de Dios, 
Manu, Erika, near Salvacion, 550 meters, 
5—6 September 1988, Amnon Friedberg; 3 
6, Madre de Dios, Manu, Rio Manu, Cocha 
Salvador, 240 meters, 14 September 1988, 
Amnon Friedberg. BOLIVIA: 4 @, Santa 
Cruz Department, Ichilo Province, Hotel 
Flora y Fauna, 4—6 km SSE Buena Vista, 
17°29.95'S, 63°N33.15'W, 400-500 meters, 
6-8 November 2003, N. E. Woodley. 

I am very grateful to Peter Chandler 
(Burnham, Slough, England), for providing 
information and reading the manuscript, 
and John Brown and Allen Norrbom (both 
of the Systematic Entomology Laboratory, 
USDA) for their review of the manuscript. 


LITERATURE CITED 


Chandler, P. J. 2001. The flat-footed flies (Diptera: 
Opetiidae and Platypezidae) of Europe. Fauna En- 
tomologica Scandinavica 36: 1—276. 

Kessel, E. L. 1957. Distribution and variation in Aga- 
thomyia notata (Loew) (Diptera: Platypezidae). 
The Wasmann Journal of Biology 15(1): 69-80. 

. 1970. Bertamyia, a new genus of Platypezin- 
inae (Diptera: Platypezidae). The Wasmann Jour- 
nal of Biology 28(2): 185-190. 

Kessel, E. L., M. E. Buegler and P. M. Keyes. 1973. 
A survey of the known larvae and puparia of Pla- 
typezidae, with a key to ten genera based on im- 
mature stages (Diptera). The Wasmann Journal of 
Biology 31(2): 233-261. 

Kessel, E. L. and M. J. Pearce. 1966. The genus Aga- 
thomyia in the Americas south of the United 
States (Diptera: Platypezidae). The Wasmann 
Journal of Biology 24(2): 261-278. 


Norman E. Woodley, Systematic Ento- 
mology Laboratory, PSI, ARS, USDA, % 
Smithsonian Institution NHB-168, P.O. Box 
37012, Washington, DC 20013-7012. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, pp. 733-734 


NOTE 


A New Synonym and New Thailand Records of Cincticostella femorata (Tshernova) 
(Ephemeroptera: Ephemerellidae) 


Ishiwata (2003) and Jacobus and Mc- 
Cafferty (2003) recently contributed revi- 
sions to the mayfly genus Cincticostella A\- 
len (Ephemerellidae: Ephemerellinae) 
(McCafferty and Wang 2000). The Oriental 
fauna of the genus is relatively poorly 
known. One such Oriental species, C. fe- 
morata (Tshernova), is known only from 
the holotype, a larva collected in June 1968 
from the Red River (Song Koi), Bak Tkhai, 
Vietnam (Tshernova 1972: figs. Sa—1). Gose 
(1969: figs. 23—37) described a similar spe- 
cies based on two larvae collected from 
Chanta Buri, Thailand on June 20, 1961. 
Gose (1969) did not provide a formal name 
for this species, but Allen (1975) subse- 
quently provided the name C. boja Allen. 
Cincticostella boja also is known only from 
the type material. The two species are dif- 
ferentiated from one another by the pres- 
ence or absence of small occipital spines on 
the head; the number of denticles on the 
claws; the number of paired, submedian, 
tergal spines on the abdomen; and the rel- 
ative density of hairlike setae on the seg- 
ments of the caudal filaments. 

We recently examined long series of ben- 
thic macroinvertebrate collections taken as 
part of an extensive inventory of the may- 
flies and other aquatic insects of Thailand 
(e.g., Sites et al. 2001, Parnrong et al. 
2002). Certain sampling sites in northern 
Thailand were visited monthly for one year. 
Examination of a series of specimens from 
one of these repeatedly sampled sites, and 
additional material, indicated to us that C. 
femorata is morphologically variable with 
a mixture of individuals and instars corre- 
sponding to the original morphological 
characterizations of either C. femorata or C. 
boja. Intermediate forms also are repre- 
sented in the samples we examined. There- 


fore, we recognize a new synonym for C. 
femorata |Cincticostella femorata (Tsher- 
nova, 1972) = C. boja (Allen, 1975), new 
synonym]. 

Specifically, these specimens demon- 
Strate variation in the development of oc- 
cipital spines, including an individual with 
no such spines. The number of denticles on 
each claw varies from two to four; one in- 
dividual has two denticles on one claw and 
four on another. Paired, medial, tergal 
spines are present on abdominal segments 
1-10, 2-10, or 3-10. Hairlike setae are pres- 
ent laterally on the caudal filaments, but the 
distribution of these setae varies from 
sparse to dense. 

Cincticostella femorata is one of the 
most striking species of Ephemerellinae. Its 
head is recessed in a greatly expanded pro- 
thorax, and its femora are very broad with 
serrate margins. These characters combine 
to give the species a very dorsoventrally 
flattened appearance. The species also has 
hairlike setae densely situated along the lat- 
eral margins of the abdominal sterna and 
the posterior margin of sternum 9, reminis- 
cent of certain species of the genus Dru- 
nella Needham (Jacobus and McCafferty 
2004). 

Little is known about the biology of C. 
femorata, but at least in Doi Inthanon Na- 
tional Park, larvae were collected consis- 
tently from leaf packs near a waterfall. 
Alate stages have yet to be associated with 
the larva. 

Material examined.—THAILAND: 
Chiang Mai Prov., Doi Inthanon National 
Park: creek at twin pagodas, 18°33’N, 
98°28’E, 2,119 m, 1-V-2003, L-492, UMC 
& CMU teams, one larva; Siriphum Water- 
fall, 18°32’N, 98°31’'E, 1,460 m elev., 14-1- 
2003, 17-1i-2003, 15-i1i-2003, CMU team, 


734 


leaf pack, three larvae. Mae Hong Son 
Prov., Namtok Maw Pang, 19°22’N, 
98°22'E, 850 m elev., 19-11-2002, L-305, 
Sites, Vitheepradit, Kirawanich, one larva. 

This material presently is deposited in 
the Enns Entomology Museum; however, 
some specimens will be deposited in Thai- 
land with the National Science Museum, 
Pathum Thani, and the Royal Forestry De- 
partment, Bangkok. Photographs of the lo- 
calities for two of the collections (identified 
as L-492 and L-305), in which this species 
was collected, are available in a Locality 
Image Database via a link from the Internet 
site of the Enns Entomology Museum, Uni- 
_versity of Missouri-Columbia. 

Acknowledgments.—We thank Chawee- 
wan Hutacharern, Royal Forestry Depart- 
ment; Jariya Chanpaisaeng, Kasetsart Uni- 
versity; and Porntip Chantaramongkol, 
Chiang Mai University, for their kind assis- 
tance in obtaining permission from the Na- 
tional Research Council Thailand and Roy- 
al Forestry Department to make these col- 
lections. We thank Akekawat Vitheepradit 
(University of Missouri), and Penkhae 
Thamsenanupap, Chirayu Naewong, and 
Pensri Bunlue (CMU Team, Chiang Mai 
University) for assistance with field work. 
Support for RWS was provided in part by 
NSF project number DEB-0103144 and by 
MU project number PSSLO232. This ma- 
terial is based, in part, upon work supported 
under a National Science Foundation Grad- 
uate Research Fellowship to LMJ. 


LITERATURE CITED 


Allen, R. K. 1975. Ephemerella (Cincticostella): A re- 
vision of the nymphal stages (Ephemeroptera: 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Ephemerellidae). 
16-22. 

Gose, K. 1969. Mayflies (Ephemeroptera) from Thai- 
land. Nature and Life Science in Southeast Asia 
6: 125-138. 

Ishiwata, S. 2003. A revision of the genus Cincticos- 


Pan-Pacific Entomologist 51: 


tella (Insecta: Ephemeroptera: Ephemerellidae) 
from Japan. Species Diversity 8: 311—346. 

Jacobus, L. M. and W. P. McCafferty. 2003. Revision 
to the genus Crinitella (Ephemeroptera: Ephem- 
erellidae). Journal of the New York Entomological 
Society 111: 48—50. 

. 2004. Revisionary contributions to the genus 
Drunella (Ephemeroptera: Ephemerellidae). Jour- 
nal of the New York Entomological Society 112: 
127-147. 

McCafferty, W. P. and T.-Q. Wang. 2000. Phylogenetic 
systematics of the major lineages of pannote may- 
flies (Ephemeroptera: Pannota). Transactions of 


the American Entomological Society 126: 9-101. 

Parnrong, S., M. Buathong, and R. W. Sites. 2002. 
New records of Behningiidae, Potamanthidae, and 
Prosopistomatidae (Ephemeroptera) from Thai- 
land. ScienceAsia 28: 407—409. 

Sites, R. W., T.-Q. Wang, S. Permkam, and M. D. Hub- 
bard. 2001. The mayfly genera (Ephemeroptera) 
of southern Thailand. Natural History Bulletin of 
the Siam Society 49: 243-268. 

Tshernova, O. A. 1972. Some new Asiatic species of 
mayflies (Ephemeroptera, Heptageniidae, Ephem- 
erellidae). Entomologiskoe Obozrenie 51: 604— 
614. 


Luke M. Jacobus, W. P. McCafferty, and 
Robert W. Sites, (LMJ,WPM) Department 
of Entomology, Purdue University, West 
Lafayette, IN 47907, U.S.A. (e-mail: 
luke-jacobus @ entm.purdue.edu); (RWS) 
Enns Entomology Museum, Department of 
Entomology, University of Missouri, Co- 
lumbia, MO 65211, U.S.A. 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, p. 735 


NOTE 


Froeschneropsidea, A Replacement Name for the Preoccupied Genus Froeschnerisca 
(Hemiptera: Heteroptera: Miridae: Deraeocorinae: Hyaliodini) 


The hyaliodine genus Froeschnerisca 
was proposed by Henry and Ferreira (2003) 
in honor of the late Richard C. Froeschner 
to accommodate Hyaliodomiris andinus 
Carvalho, a species known only from Bo- 
livia and Peru (Carvalho 1953) and subse- 
quently transferred to the genus Hyaliodo- 
coris Knight (Carvalho 1957). We have dis- 
covered that Froeschnerisca Henry and 
Ferreira is preoccupied by Froeschnerisca 
Coscar6n, 1997 (Heteroptera: Reduviidae). 
As a result, we propose the replacement 
name Froeschneropsidea, new name, for 
the primary junior homonym Froeschner- 
isca Henry and Ferriera, 2003. Froeschner- 
opsidea andinus (Carvalho), the type of the 
genus and only included species, is a new 
combination. 

Acknowledgments.—We thank Manuel 
Baena (Departamento de Biologia y Geo- 
logia, I.E.S. Trassierra c/o Avda. Arroyo del 
Moro s/n14011 Cordoba, Spain) and Dim- 
itri Forrero (Department of Entomology, 
Cornell University, Ithaca, New York) for 
kindly bringing the above case of homon- 
ymy to our attention. 


LITERATURE CITED 


Carvalho, J. C. M. 1953. Neotropical Miridae, LX: 
New species of Hyaliodes Reuter and Hyaliodo- 
coris Carvalho (Hemiptera). Revista Brasileira de 
Biologia 13: 113-119. 

. 1957. Catalogue of the Miridae of the world. 
Part I. Cylapinae, Deraeocorinae, Bryocorinae. 
Arquivos do Museu Nacional, Rio de Janeiro 
44(1): 1-158. 

Coscar6n, M. C. 1997 (1996). Froeschnerisca nom. 
nov. for the junior homonym Froeschneriella 
Coscar6n (Heteroptera: Reduviidae). Physis, Secc. 
C. 54(126—127): 49. 

Henry, T. J. and P. S. E Ferreira. 2003. Three new 
genera and three new species of Neotropical Hy- 
aliodini (Hemiptera: Heteroptera: Miridae: Der- 
aeocorinae), with new combinations and new syn- 
onymy. Journal of the New York Entomological 
Society 111: 96-119. 


Thomas J. Henry and Paulo S. Fiuza Fer- 
reira. (TJH) Systematic Entomology Labora- 
tory, Plant Sciences Institute, Agricultural Re- 
search Service, U.S. Department of Agricul- 
ture, % P.O. Box 37012, National Museum of 
Natural History, Smithsonian Institution, 
Washington, DC 20013-7012 U. S. A. (e-mail: 
thenry@ sel.barc.usda.gov), (PSFF), Departa- 
mento de Biologia Animal, Universidade Fed- 
eral de Vigosa, Vigosa, Minas Gerais, 36571- 
000 Brazil (e-mail: pfiuza@utv.br). 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, p. 736 


BoOoK REVIEW 


Catalogue of Ceutorhynchinae of the 
World, with a Key to Genera. By Enzo 
Colonnelli. 2004. 124 pp. Euros 80+ Vat, 
~$100. Argania Edito, Barcelona, Spain. 
ISBN 84-931847-6-4, e-mail: Argania@ 
entomopraxis.com 


This catalog of Ceutorhynchinae of the 
world is a must for weevil workers. This is 
a slim, attractive volume with large double- 
columned pages that accurately catalogs 
1,316 species of ceutorhynchine weevils 
and all relevant bibliography (another 4,600 
entries). It includes a historical outline, sec- 
tions on nomenclature, distribution, and 
ecology of these weevils, and a section en- 
titled taxonomic remarks, which proposes 
new taxonomic groups. A major feature of 
this book is a key to the genera of the 
world. 

This book has several nice features. 
These include summary lists of taxa, host- 
plant families and genera, abbreviations of 
depositories and taxonomists names, and 
new combinations and synonymies. I most 
liked the three lists at the beginning, which 
outline the number of species per tribe and 
genus and the number of genera in each 
tribe. Another attractive feature is the inclu- 
sion of information on ecology after the en- 
try for each species, known host plants, and 
data on countries where each is found. This 
information, coupled with the list of plant 
families and host genera on page 9, will 
help collectors. 

There are also pleasing sections describ- 


ing in detail the proposal of a new tribe, the 
Hypohypurini, and the rationales for des- 
ignations of neotypes and lectotypes. I also 
liked the separate lists of new placements, 
changed combinations, new synonymies, 
changes of rank, revised statuses, and se- 
lections of original spellings. 

The 18-page key to genera is a major 
section of this book, but, unfortunately, it 
is not illustrated. That, and the fact that all 
the genera of the world are treated together, 
makes it a little difficult to use. In the in- 
troduction to the key, the author does fore- 
cast that some closely related taxa will not 
key out close to one another because of the 
choice of characters used. 

In this work, 16 new genera are pro- 
posed. These are diagnosed and described 
only by their positions and descriptions in 
the key. I find this unsatisfactory, given that 
one has to read the entire key in order to 
understand the author’s concept of a given 
genus. This, however, is the only major flaw 
in this otherwise very fine book. The author 
lists the etymologies for these genera to- 
gether at the beginning of the catalog. 

I recommend this catalog to all weevil 
workers and persons interested in the seri- 
ous study of Ceutorhynchinae. For the 
price, it is a wonderful investment. 


Catherine N. Duckett, Research Associ- 
ate, Department of Entomology, National 
Museum of Natural History, Smithsonian 
Institution, MRC 105, P.O. Box 37012, 
Washington, DC 20013-7012, U.S.A. (e- 
mail: duckettc @ si.edu) 


PROC. ENTOMOL. SOC. WASH. 
107(3), 2005, p. 737 


BOOK REVIEW 


Identification and Geographical Distri- 
bution of the Mosquitoes of North Amer- 
ica, North of Mexico. [Second Edition. ] 
Richard EF Darsie Jr. and Ronald A. Ward. 
2005. $70.00. University Press of Flori- 
da, Gainesville 12611-207 ISBN 0-8130- 
2784-5. 


Because of their importance as pests and 
disease vectors, mosquitoes are among the 
best known of all insects, yet new facts 
about them are being discovered each year. 
During the past 24 years several species 
have been added to the North American 
fauna, bringing the total to 174 species in 
14 genera and 29 subgenera. The mono- 
graph by Carpenter and LaCasse (1955) 
was indispensable for the identification of 
mosquitoes until the publication of the first 
edition of the present book in 1981 (Darsie 
and Ward 1981). This second edition au- 
thoritatively brings the subject up to date in 
a handsome, beautifully illustrated book. 

The main sections of the book present the 
following topics: a systematic index that in- 
cludes a list of taxa and discussions of var- 
ious name changes; morphology of adult fe- 
male mosquitoes; keys to the adult female 


mosquitoes; morphology of fourth-instar 
mosquito larvae; keys to the fourth-instar 
larvae; and geographical distribution of the 
Culicidae of North America, north of Mex- 
ico. The key couplets to both females and 
fourth-instar larvae are fully illustrated with 
nearby drawings by Chien C. Chang and 
Taina Litwack that greatly facilitate use of 
the keys. The 130 pages of the distribution 
section feature distribution maps for each 
species as well as tables listing the species 
for each state or province. The selected bib- 
liography lists 815 publications. 

This book is now the essential source for 
the identification of North American mos- 
quitoes. It is a very important milestone in 
the field of Culicidology. 


LITERATURE CITED 


Carpenter, S. J. and W. J. LaCasse. 1955. Mosquitoes 
of North America (North of Mexico). Berkeley, 
University of California Press, Berkeley, 360 pp., 
127 pls. : 

Darsie, R. F, Jr. and R. A. Ward. 1981. Identification 
and geographical distribution of the mosquitoes of 
North America, north of Mexico. Mosquito Sys- 
tematics Supplement 1: 1—313. 


William E. Bickley, 3/24 Gracefield Rd., 
Apt. 101, Silver Spring, MD 20904, U.S.A. 


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CONTENTS 
(Continued from front cover) 
KIMSEY, LYNN S.—Revision of the northern South American tiphiid genus Merithynnus 
Kaimsey:11991' Giymenoeptera: Tiphidaesmnwaninae)) 0402... se te ah ieee leita 


LaBONTE, J. R., A. D. MUDGE, and K. J. R. JOHNSON—Nonindigenous woodboring 
Coleoptera (Cerambycidae, Curculionidae: Scolytinae) new to Oregon and Washington, 
1999-2002: Consequences of the intracontinental movement of raw wood products and solid 
wood packing materials Wer neki imme aeeneee MG ik. (2 3 C3 UES RS SL 0 Oe nee a ae 


McCAFFERTY, W. P. and L. SUN—Mystaxiops: A new genus of small minnow mayflies 
(Ephemexoptera? Baetidas) tromibapua New Guinea), Seb tay ious ce eee rae a eater are ete re 


MILLER, GARY L., MANYA B. STOETZEL, and ETHAN C. KANE—A systematic reappraisal 
of the genus Diuraphis Auzenbers (Hemiptera; Aphididae) 227 es 2e-).2-ue) see ee ers 


PINTO, C. MIGUEL and GUILLERMO L. CLAPS—First record of Cuterebra almeidai 
(Guimaraes and Carrera) from Argentina, new host records for Cuterebra apicalis Guérin- 
Méneville, and list of Cuterebra (Diptera: Oestridae) in the collection of the Instituto 
Pundacion; Miguel eMloniucumaney Amo enicitic es mye ce a) baiela a2 one iy eee erat annem eal er 


PINTO, JOHN D.—Descriptions of additional New World Trichogrammatidae (Hymenoptera): 
The genus Nicolavespa and a new species of Haeckeliania ....... 0.06. ccc eee cece een eens 


PLAKIDAS, JOHN D.—A new species of Porricondyla (Diptera: Cecidomyiidae) from south- 
WeSterm Pennsylvania yaoi U ane wiih One nt ieee meee OU he I cee hele a RON aR A 


POINAR, GEORGE, JR.—Culex malariager, n. sp. (Diptera: Culicidae) from Dominican amber: 
The first fossil mosquito vector of Plasmodium ................ 2c c ccc cece cence cece eee ees 


ROBBINS, ROBERT K.—Phylogenetic relationships among the species of Panthiades Hiibner 
(by caenidae Phechina|y Burmaeiman) cee see eae ess 2 Lite aN 0b, ted i eal Se CC LEP 


RUEDA, LEOPOLDO M., RICHARD C. WILKERSON, and CONG LI—Anopheles (Anopheles) 
lesteri Baisas and Hu (Diptera: Culicidae): Neotype designation and description ............ 


SMITH, DAVID R. and IAN C. STOCKS—A new trigonalid wasp (Hymenoptera: Trigonalidae) 
from eastern North Anmnenica 1 ssgessen yaciaat sae tits ee ae Va Meh HEN Cried coc MP ARR a Uae 


TAUBER, CATHERINE A., GILBERTO S. ALBUQUERQUE, and MAURICE J. TAUBER— 
Characteristics of the Loyola Navas male (Neuroptera: Chrysopidae: Apochrysinae) ........ 


YAMAN, MUSTAFA, IRFAN ASLAN, ONDER CALMASUR, and FIKRETTIN SAHIN—Two 
bacterial pathogens of Helicoverpa armigera (Hiibner) (Lepidoptera: Noctuidae) ............ 


NOTES 


HENRY, THOMAS J. and PAULO S. FIUZA FERRERIA—Froeschneropsidea, a replacement 
name for the preoccupied genus Froeschnerisca (Hemiptera: Heteroptera: Miridae: 
Deracocorimaes Hy altos) 8) ea ke eee SN pac Malema eT a eae UR se acer a a 


JACOBUS, LUKE M., W. P- McCAFFERTY, and ROBERT W. SITES—A new synonym and new 
Thailand records of Cincticostella femorata (Tshernova) (Ephemeroptera: Ephemerellidae) 


LATTIN, JOHN D.—Scoloposcelis pulchella pulchella (Zetterstedt, 1838) in North America 
(Hemuptera:;-Heteroptera;Anthoconidac) a esse aaa eens ke ee en ele en 


WOODLEY, NORMAN E.— Distributional notes on Bertamyia notata (Loew), including the first 
report of the family Platypezidae from the Caribbean (Diptera: Platypezidae) .............. 


BOOK REVIEWS 


BICKLEY, WILLIAM E.—/dentification and Geographical Distribution of the Mosquitoes of 
North America, North of Mexico {Second Edition], by Richard F. Darsie, Jr. and Ronald A. 
DE hie Ieee RS Bed ie An Sey) OU ARR. TAU PIe ORIN hom RE any a ete ter, AS eRe ren meter OEE Ly. 


DUCKETT, CATHERINE N.—Catalogue of Ceutorhynchinae of the World, with a Key to Genera, 
by Bnzoi Colonel wat lie Neer Muar stitial elas < Ala) aca ARR Ears Te ko JT aD RCs A 


576 


554 


536 


700 


D172 


627 


652 


548 


501 


604 


530 


543 


623 


T35 


133 


729 


731 


niin 


| 
} 


VOL. 107 OCTOBER 2005 NO. 4 


Ql (ISSN 0013-8797) 
24x PROCEEDINGS 


ca 1 
of the 


ENTOMOLOGICAL SOCIETY 
of WASHINGTON 


PUBLISHED 
QUARTERLY 


BRAILOVSKY, H.—A new species of Dalader Amyot and Serville, with a key to the Malaysian 
species (Hemiptera: Heteroptera: Coreidae: Coreinae: Daladerini) ...............02--.00000- 883 
BURCKHARDT, DANIEL, PAUL HANSON, and LUIS MADRIGAL—Diclidophlebia lucens, 
n. sp. (Hemiptera: Psyllidae) from Costa Rica, a potential control agent of Miconia calvescens 
(MACIANIOITIALACC AG) MIN AGW AULy tists sacle cits aie canal tssioy « sleiere boeta fete: abobe.c clave ld n’vin,vis, ova wat piel acetaivin ate ¢ 741 
BURNS, JOHN M. and DANIEL H. JANZEN—What’s in a name? Lepidoptera: Hesperiidae: 
Pyrginae: Telemiades Hiibner 1819 [Pyrdalus Mabille 1903]: New combinations Telemiades 
corbulo (Stoll) and Telemiades oiclus (Mabille)—and more ..............00ceeceeeeece eee eees 770 
DARSIE, RICHARD F., JR.—Key to the pupae of the mosquitoes (Diptera: Culicidae) of Florida 892 


DERR, DAWN P. and JERRY L. COOK—Morphology of the antenna of Caenocholax fenyesi 
Pierce (Strepsiptera: Myrmecolacidae) based on scanning electron microscopy ...........-.. 762 


GANDHI, KAMAL J. K., DANIEL W. GILMORE, GEORGE E. BALL, RALPH W. HOLZEN- 


THAL, STEVEN A. KATOVICH, JESSICA J. KOEHLE, KIRK J. LARSEN, WILLIAM J. 
MATTSON, and STEVEN J. SEYBOLD—A review of ground beetle species (Coleoptera: 


Carabidae) of Minnesota, United States: New records and range extensions ................ 917 
HALL, JASON P. W. and KEITH R. WILLMOTT—A new species of Paiwarria (Lepidoptera: 
BY CAPRIOde EP IMDACHAL) MOM WESLEIIECIAMOL ,/atd po sje se ials «init einielaie ribie soles ae cle smeasinn view © ase 960 


HARBACH, RALPH E., RAMPA RATTANARITHIKUL, and BRUCE A. HARRISON— 
Baimaia, a new subgenus for Anopheles kyondawensis Abraham, a unique crabhole-breeding 


PAA ARGS PATOL CARTCL INU ASIA! <0. ceive sah ht« k Mianivd al cele ae Wolof tsetse a's s'oinid foie elie slp mend acine A 750 
JOSE, JOSEPHINE, YOSHIMI HIROSE, and JEFFREY Y. HONDA—Two new species of 

Trichogramma (Hymenoptera: Trichogrammatidae) from the Ryukyu Islands, Japan ........ 782 
KINKOROVA, JUDITA—Notes on the fruit flies (Diptera: Tephritidae) of California .......... 851 
KONDRATIEFF, B. C., R. F KIRCHNER, and DAVID LENAT— Two new species of Haploperla 

Navas (Plecoptera: Chloroperlidae) from North Carolina, U.S.A. ............2.e eee ce cree ees 859 


MAIER, CHRIS T.—First records of alien insects in Connecticut (Orthoptera: Tettigoniidae; 
Coleoptera: Buprestidae, Chrysomelidae; Diptera: Rhagionidae, Tephritidae; Hymenoptera: 
Mae SOMME Eso a Ge Ieee 2c Mee Di ale a dyes Ub ais'ele aitgds Zhen bao Waetab Seles bo wn asin e's 947 


(Continued on back cover) 


THE 


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PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 741-749 


DICLIDOPHLEBIA LUCENS, N. SP. (HEMIPTERA: PSYLLIDAE) FROM 
COSTA RICA, A POTENTIAL CONTROL AGENT OF MICONIA 
CALVESCENS (MELASTOMATACEAE) IN HAWAII 


DANIEL BURCKHARDT, PAUL HANSON, AND LUIS MADRIGAL 


(DB) Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Switzerland (e- 
mail daniel.burckhardt @ unibas.ch); (PH) Escuela de Biologia, Universidad de Costa Rica, 
San Pedro, San Jose, Costa Rica (e-mail phanson @biologia.ucr.ac.cr); (LM) same address 
(e-mail lama 752003 @ yahoo.com) 


Abstract.—A new species of Diclidophlebia (Psyllidae: Paurocephalinae), D. lucens, 
is described from Miconia calvescens (Melastomataceae) in Costa Rica. This plant is an 
invasive weed in various Pacific islands and the new species described here represents a 
potential biological control agent. The egg, fifth instar larva, and adults are illustrated, 
the former being the first illustration of an egg for any species in the genus. Preliminary 


observations on the life cycle of this species are also provided. 


Key Words: 
tropics 


Most of the approximately 3,000 de- 
scribed jumping plant-louse species have 
very narrow host plant ranges within the 
Dicotyledones. Some psylloids develop on 
crop, forest or ornamental plants where they 
can become important pests (Burckhardt 
1994). Other species are associated with 
weeds and constitute potentially useful con- 
trol agents. Examples are Heteropsylla spi- 
nulosa Muddiman et al. in Australia and 
New Guinea for the control of Mimosa di- 
plotricha C.W. Wright ex Sauvalle (= in- 
visa Martius) (Muddiman et al. 1992, Swar- 
brick 1997), Prosopidopsylla flava Burck- 
hardt in Australia for the control of mes- 
quite (Prosopis spp.) (Van Klinken 2000), 
and Boreioglycaspis melaleucae Moore in 
Florida for the control of Melaleuca quin- 
quenervia (Cav.) S.T. Blake (Wineriter et al. 
2003). Here we describe a new species of 
Diclidophlebia which is being studied as a 
potential biological control agent of Micon- 
ia calvescens Schrank and Mart ex DC. 
(Melastomataceae). 


taxonomy, new species, Paurocephalinae, biological control of weeds, neo- 


The pantropical genus Diclidophlebia 
Crawford has been redefined within the 
subfamily Paurocephalinae (Psyllidae) by 
Burckhardt and Mifsud (2003) to include 
24 described species associated with spe- 
cies of seven different host plant families. 
Prior to the revision of Burckhardt and Mif- 
sud (2003) Diclidophlebia species were as- 
signed to six different genera, reflecting the 
large variation encountered in the forewing 
shape, pattern and venation as well as in the 
male and female terminalia. Of the 12 de- 
scribed New World species, five are known 
to develop on Melastomataceae: D. fava 
(Brown and Hodkinson) and D. longitar- 
sata (Brown and Hodkinson) on Miconia 
argentea (Sw.) DC. (both from Panama), D. 
paucipunctata (Brown and Hodkinson) and 
D. tuxtlaensis (Conconi) on Conostegia xal- 
apensis (Bonpl.) D. Don (from Panama and 
Mexico, respectively), and D. heterotrichi 
(Caldwell and Martorell) from Heterotri- 
chum cymosum (J.W. Wendl. ex Spreng) 


742 


Urb. in Puerto Rico. Of the remaining New 
World species, hosts are known for four, 
and these are associated with Sterculiaceae, 
Tiliaceae, and Ulmaceae. Old World species 
of Diclidophlebia are associated with Eu- 
phorbiaceae, Malvaceae, Rhamnaceae, Ster- 
culiaceae, and Tiliaceae. 

The new species of Diclidophlebia de- 
scribed below feeds on Miconia calvescens, 
a plant that is native to the Neotropical Re- 
gion that was introduced as an ornamental 
plant into Tahiti and subsequently into Ha- 
waii. Because of its ability to grow quickly 
and to form dense stands, thereby creating 
deep shade that many native species cannot 
tolerate, M. calvescens is among the most 
threatening introduced plants on many Pa- 
cific islands (Meyer 1996). Since its intro- 
duction in 1937, this plant has taken over 
about two thirds of the land surface of Ta- 
hiti and directly threatens nearly half the 
island’s endemic plant species (Meyer and 
Florence 1996). There is currently serious 
concern that this invasive weed could have 
similar effects in Hawaii (Medeiros et al. 
1997). Herbicide spraying and hand remov- 
al are the principal control measures (Co- 
nant et al. 1996), but these methods are lim- 
ited by the prolific production of very 
small, bird-dispersed seeds and the longev- 
ity (up to eight years) of the seed bank 
(Meyer 1998). Thus biological control is an 
important component of the long-term plan 
for controlling populations of M. calvescens 
on Pacific islands. 


Diclidophlebia lucens Burckhardt, 
Hanson, and Madrigal, new species 
(Figs. 1-16) 


Diagnosis.—Adult: Body color bright 
orange reddish; forewing coloration uni- 
formly yellowish orange. Forewing oblong 
oval; median third of vein Rs and vein M 
straight and subparallel; surface spinules 
forming cellular pattern. Metatibia weakly 
expanded apically with a crown of sclero- 
tised apical spurs. Male proctiger globular 
(Fig. 5), parameres broadly lamellar (Fig. 
6), aedeagus 2-segmented with a large bi- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


lobed ventral process in the middle of the 
distal segment (Fig. 7), apex tubular. Fe- 
male terminalia cuneate, short, pointed api- 
cally (Fig. 8), circumanal ring cruciform. 

Fifth instar larva: Antenna 10-segment- 
ed. Forewing bud relatively large, with 4— 
6 marginal sectasetae. Legs moderately 
long, tarsal arolium only slightly longer 
than claws. Caudal plate angular posterior- 
ly; area of extra pore fields extended, sep- 
arated into two curved rows of distinct oval 
patches. Sectasetae pointed. Caudal plate 
laterally near fore margin with 3—4 secta- 
setae on either side, and near the circumanal 
ring with 3+3 sectasetae. 

Description.—Adult: Coloration: Bright 
orange reddish (male more reddish than fe- 
males), eyes grey. Antenna dirty yellowish, 
apices of segments 4, 6 and 8, and entire 
segments 9 and 10 brown. Ventral face of 
head and abdomen bright orange. Legs and 
terminalia yellowish. Forewing transparent, 
yellowish orange, apex of vein Cu,, brown; 
hindwing transparent, whitish. 

Structure: Head (Fig. 1) weakly inclined 
from longitudinal body axis, about as wide 
as mesoscutum. Vertex trapezoidal, surface 
finely sculptured with microscopic setae; 
median suture fully developed. Eyes subg]- 
obular. Genae evenly rounded, with a pair 
of long setae on either side of frons. Frons 
forming large triangular sclerite. Antenna 
(Fig. 2) 10-segmented, with a single, large 
subapical rhinarium on each of segments 4, 
6, 8 and 9; margin of rhinaria bearing long 
spines proximally and short ones distally; 
terminal setae distinctly longer than seg- 
ment 10. Clypeus flattened, pyriform. Tho- 
rax weakly arched, with fine microsculpture 
and microscopic setosity; mesoscutellum 
swollen, metascutellum with small subacute 
tubercle. Forewing (Fig. 3) oblong oval, 
widest in middle; pterostigma ending be- 
yond middle of vein Rs; vein Rs relatively 
straight in median third, curved in a 45° an- 
gle towards fore margin apically; vein M 
straight, subparallel to basal two-thirds of 
vein Rs; veins M,,, and M,,, relatively 
short; vein Cu,, weakly curved, moderately 


VOLUME 107, NUMBER 4 743 


aN 


eqn} pua 
peZi}OJa|OsS 


paramere 


S14 
> SAK 


ARs 


subgenital plate 


Figs. 1-8. Diclidophlebia lucens. 1, Head, dorsal view. 2, Antenna. 3, Forewing. 4, Portion of cell Rs 
indicating cellular pattern of surface spinules. 5, Male terminalia, in profile. 6, Paramere, inner face. 7, Distal 
portion of aedeagus. 8, Female terminalia, in profile. 


744 
long; surface spinules leaving spinule-free 
stripes along veins, absent from basal half 
of cell c+sc, forming a hexagonal pattern 
(Fig. 4). Hindwing slightly shorter than 
forewing, with indistinctly grouped costal 
setae; vein M+Cu, developed. Metacoxa 
with large, horn-shaped, subacute meracan- 
thus; metatibia long, slender, weakly ex- 
panded apically, bearing an anteriorly and 
posteriorly open crown of 8—9 sclerotised 
apical spurs which are laterally slightly 
larger than anteriorly. Abdominal tergites 
with a tubercular bump in the middle. Male 
terminalia (Fig. 5) with tubular proctiger; 
subgenital plate subglobular. Paramere 
shortly lamellar, anterior margin weakly 
curved, posterior margin angular subapical- 
ly, outer and inner face (Fig. 6) covered in 
long setae, ending in sclerotised tooth. Ae- 
deagus 2-segmented, distal portion (Fig. 7) 
with a large bilobed ventral process in the 
middle, apex tubular; sclerotised end tube 
of ductus ejaculatorius long and almost 
straight. Female terminalia (Fig. 8) cuneate, 
short; dorsal margin of proctiger concave, 
apex pointed; subgenital plate shorter than 
proctiger, abruptly narrowed in apical third, 
pointed; circumanal ring cruciform. 

Measurements in mm and ratios (3 6, 3 
2): head width (HW) 0.33—0.38; Antenna 
length (AL) 0.36—-0.38; forewing length 
(WL) 0.74—0.94; male proctiger length 
(MP) 0.10—0.12; paramere length 0.09; 
length of distal portion of aedeagus 0.08— 
0.10; female proctiger length 0.28—0.30; 
AL/HW 1.00—1.15; antennal segment 3/an- 
tennal segment 4 length ratio 1.50—3.00; 
WL/HW 2.24—2.57; WL/forewing width ra- 
tio; rostrum length/HW 0.30—0.36; metati- 
bia length/HW 0.68—0.78; MP/HW 0.29— 
0.36; FP/HW 0.80-—0.84; FP/circumanal 
ring length ratio 2.24—2.69; FP/female sub- 
genital plate length ratio 1.75—1.81. 

Fifth instar larva: Coloration: Orange; 
cephalothoracic sclerite orange brownish. 
Antenna, legs and wing-buds yellowish. 
Caudal plate orange, brownish laterally. 

Structure (Fig. 9): Body elongate, sparse- 
ly covered in microscopic rod and normal 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


setae. Antenna 10-segmented with a rhinar- 
ium on each of segments 4, 6, 8 and 9, and 
following numbers of pointed sectasetae on 
each segment from | to 10: 1 (0), 2 (1-2), 
35) 4 @)5' (©), 6 G=2); 7 O-=) sO> 
1), 9 (0), 10 (O). Dorsal thoracic sclerites 
small. Forewing bud moderately large with 
4—6 marginal pointed sectasetae; hindwing 
pad with 1—2 marginal pointed sectasetae. 
Legs relatively long with pointed sectasetae 
on tibiae; tarsal arolium slightly longer than 
claws. Caudal plate angular posteriorly with 
3—4 pointed sectasetae laterally near ante- 
rior margin on either side and 3+3 pointed 
sectasetae dorsally near circumanal ring. 
Extra pore fields extended consisting of two 
curved rows of distinct oval patches on ei- 
ther side of caudal plate. 

Measurements in mm and ratios (4 lar- 
vae): body length (BL) 0.73—0.77; antenna 
length (AL) 0.34—0.38; AL/forewing pad 
length ratio 1.29—1.38; body breath/BL 
0.70—0.80; caudal plate breadth/length ratio 
1.82—2.16; cicumanal ring breadth/caudal 
plate breadth ratio 0.18—0.20. 

Egg: Coloration: Pale yellowish orange 
when young, becoming dark when mature; 
empty chorion blackish after larva ecloses. 

Structure (Fig. 10): Elongate ovoid, with 
a basal pedicel and an apical filament; about 
0.1 mm wide by 0.25 mm long (excluding 
pedicel and apical filament). Apical fila- 
ment very long (about as long as egg itself), 
and curved upward and backward above 
egg. 

Type materialHolotype ¢: Costa Rica, 
Cartago, Sabanilla de Tucurrique, 800 m, 
x1.2000, Miconia calvescens (P. Hanson); 
collection in the field 20.xi.2000, samples 
from the greenhouse plant x-x11.2000 (The 
Naturhistorisches Museum, Basel, dry 
mounted). Paratypes: 23 6, 71 2, 31 larvae 
(Natural History Museum, London: 10 6, 
6 2 dry mounted; 3 d, 3 &, 8 larvae, slide 
mounted; 50 @, 10 fifth instar larvae in 
95% _ ethanol—Naturhistorisches Museum, 
Basel: 4 6, 6 2, 4 larvae, dry mounted; 3 
3,3 @, 9 larvae, slide mounted—Muséum 
d’ Histoire Naturelle, Genéve: 1 d, 1 @, dry 


VOLUME 107, NUMBER 4 745 


0.50 mm ae 


10 


Figs. 9-10. Diclidophlebia lucens. 9, Fifth instar larva, left dorsal, right ventral face; with details of forewing 


—_— ee 


pad, extra pore fields and apex of hind leg. 10, Egg. 


746 


mounted—National Museum of Natural 
History, Washington, DC, collection in 
USDA Beltsville; MD} cri, oe dry. 
mounted—Muséum National d’ Histoire 
Naturelle, Paris: 1 3d, 1 ¢, dry mounted), 
same data as holotype. 

Relationships.—Diclidophlebia lucens 
forms a morphologically homogeneous, 
probably monophyletic, group with other 
Melastomataceae inhabiting Diclidophlebia 
species, viz. D. fava, D. longitarsata (both 
on Miconia argentea), D. paucipunctata, D. 
tuxtlaensis (both on Conostegia xalapensis, 
the latter also on Miconia sp.), and D. het- 
erotrichi (on Heterotrichum cymosum). The 
group is defined within Diclidophlebia by 
the oblong-oval forewing with partially 
subparallel veins Rs and M, the hexagonal 
pattern of the surface spinules, the tubular 
male proctiger, the short, broadly lamellar 
paramere with long setae on the outer and 
inner face, the short, cuneate female ter- 
minalia with short suddenly narrowed sub- 
genital plate, and the cruciform circumanal 
ring. D. lucens shares with D. fava and D. 
longitarsata, both associated with Miconia, 
the presence of a ventral process on the dis- 
tal portion of the aedeagus. D. lucens dif- 
fers from the last two species in the lack of 
a dark forewing pattern and details in the 
male and female terminalia. 

The last instar larvae of Diclidophlebia 
have not been treated monographically. 
There are several isolated species descrip- 
tions from which it is difficult to discern the 
taxonomic significance of characters. For 
the present work we examined material of 
following species: D. dahli (Riibsaamen), 
D. eastopi Vondraéek, D. excetrodendri (Li 
and Yang), D. fava, D. fremontiae (Klyver) 
and D. xuani Messi. In addition, we used 
published descriptions of D. longitarsata, 
D. menoni (Mathur), D. nebulosa (Brown 
and Hodkinson), D. paucipunctata and D. 
tuxtlaensis. Diclidophlebia dahli, D. easto- 
pi, D. excetrodendri, D. menoni and D. 
xuani have 9-segmented antennae and the 
tarsal arolium is much larger than the claws, 
in contrast to D. lucens, which has 10-seg- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mented antennae and a tarsal arolium which 
is only slightly longer than the claws. The 
latter character is similar in D. fremontiae 
which has, however, 9-segmented antennae. 
D. nebulosa differs in the truncate sectase- 
tae which are pointed in D. lucens. D. fava 
differs from D. lucens in the shorter legs, 
the higher number of lateral sectasetae on 
the forewing buds (about 7—8) and the 
smaller extra pore fields on the caudal plate. 
From D. longitarsata, D. paucipunctata and 
D. tuxtlatensis, D. lucens differs in the 
smaller dimensions; e.g., antenna shorter 
than 0.4 mm in D. /ucens and longer in the 
other three species. 

Etymology.—From the Latin verb lucere 
meaning to be bright, to shine, referring to 
the bright orange color. 


BIOLOGY 


Miconia calvescens is very scarce in 
Costa Rica, occurring in just a few isolated 
locations on the Caribbean slope, below 
1,000 meters altitude (usually between 700 
and 1,000 m), generally in sites receiving 
substantial sunlight, and often on steep 
slopes. Although individuals of D. lucens 
are small-sized psyllids, they can be readily 
detected in the field by their waxy secre- 
tions (appearing as small cottony masses, 
Figs. 11-13). Thus far, populations of D. 
lucens have been found on M. calvescens 
in six of the eight sites sampled. Studies of 
host-plant range are currently in progress, 
but preliminary results suggest that D. lu- 
cens does not feed on plants other than Me- 
lastomataceae, and even within this plant 
family it is probably restricted to a narrow 
range of species. 

All life stages are found on the host 
plant, primarily on the terminal buds (in- 
cluding both leaf and flower buds), and ex- 
panding young leaves. Females oviposit on 
the youngest leaves or in open buds, and 
rarely on the outer surface of closed buds. 
An egg may be laid on either the upper or 
lower surface of the leaf, often into the 
small space between a vein and the leaf 
lamina. 


VOLUME 107, NUMBER 4 


747 


Figs. 11-16. Diclidophlebia lucens (photographs with Nikon Coolpix). 11, Infestation showing wax fila- 
ments. 12, Infestation of flower buds. 13, Damage to young leaves. 14, Psyllids, wax, spherical excrement on 
leaf. 15, Male and female mating. 16, Female on purple underside of leaf. 


Upon eclosion, the first instar larvae be- 
gin to feed on buds and barely expanded 
leaves, and almost immediately begin to 
produce long waxy filaments under which 
they become concealed. All five larval in- 
stars produce these filaments and when 
populations are high these cottony masses 
of wax can become quite extensive. The 


larvae also produce spherical globules of 


liquid excrement, about one millimeter in 
diameter, which are apparently coated with 
wax, since the globules are whitish in color 
and maintain their spherical shape (Fig. 14). 
The psyllids do not appear to be regularly 
tended by ants, although more field obser- 
vations are required. Thus far, no parasit- 
oids have been reared from D. lucens; al- 
though various parasitoids are known from 


748 


Psylloidea in Costa Rica, none have yet 
been reared from psyllids on Melastomata- 
ceae (Noyes and Hanson 1996). 

Duration of the various life stages ap- 
pears to be similar to that reported for D. 
tuxtlaensis in Mexico (Conconi 1972). In 
the latter species the duration of the stages 
are: eggs, 3 days; first instar larvae, 3—5 
days; second instar larvae, 4—5 days; third 
instar larvae, 5—7 days; fourth instar larvae, 
4-5 days, fifth instar larvae, 5—6 days. 
Thus, the time from oviposition to eclosion 
of the next generation of adults ranges from 
about 24 to 31 days, depending upon the 
temperature. Conconi (1972) reported that 
wing buds first appear in the second instar 
larvae and that each successive instar be- 
comes slightly more active, except for late 
fifth instars which become inactive. The 
same author also reported that adult lon- 
gevity varied from 30 to 38 days. While 
males and females of D. lucens are readily 
distinguishable as adults (males being 
smaller, brighter red, and with enlarged 
genitalia, Figs. 15—16), no sexual differenc- 
es have been found for distinguishing the 
larvae (nor were any found by Conconi 
1972). Adults are generally quite inactive 
except when disturbed. 

The insect is easily reared on potted 
plants under greenhouse conditions. Ongo- 
ing studies will determine the effects of D. 
lucens on M. calvescens. Preliminary ob- 
servations suggest that by feeding near the 
apical meristem, this species potentially re- 
tards growth and reproduction. Moreover, 
high populations appear to result in pre- 
mature dehiscence of infested leaves. 


ACKNOWLEDGMENTS 


We thank Pablo Allen and Eduardo Cha- 
con for their help in the field, David Mifsud 
for preparing Figs. 1-9, Kenji Nishida for 
the photographs (Figs. 11-16), David Hol- 
lis and Tracy Johnson for their comments 
on the manuscript, and Edgar Rojas and 
Clifford Smith for making the Miconia pro- 
ject possible. We also acknowledge eco- 
nomic support from the state of Hawaii, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Department of Land and Natural Resources, 
USGS Biological Resources Division and 
the National Park Service, via the Univer- 
sity of Hawaii Cooperative Studies Unit, 
and from USDA Forest Service Internation- 
al Programs. 


LITERATURE CITED 


Burckhardt, D. 1994. Psylloid pests of temperate and 
subtropical crop and ornamental plants (Hemip- 
tera, Psylloidea): a review. Trends in Agricultural 
Sciences, Entomology 2: 173-186. 

Burckhardt, D. and D. Mifsud. 2003. Jumping plant- 
lice of the Paurocephalinae (Insecta, Hemiptera, 
Psylloidea): systematics and phylogeny. Contri- 
butions to Natural History, Bern 2: 3-34. 

Conant, P., A. C. Medeiros and L. L. Loope. 1996. A 
multiagency containment program for Miconia 
(Miconia calvescnes), an invasive tree in Hawai- 
ian rainforests, pp. 249-254. In Luken, J. O. and 
J. W. Thieret, eds. Assessment and Management 
of Plant Invasions. Springer-Verlag, New York. 

Conconi, J. R. E. de 1972. Descripcién y biologia de 
Paurocephala tuxtlaensis sp. nov. (Homoptera 
Psyllidae) de la regié6n de Los Tuxtlas en Vera- 
cruz, México. Anale del Instituto de Biologia, 
Universidad Nacional Aut6noma de México 
43(1): 51-66. 

Medeiros, A. C., L. L. Loope, P. Conant, and S. 
McElvaney. 1997. Status, ecology, and manage- 
ment of the invasive plant, Miconia calvescens 
DC (Melastomataceae) in the Hawaiian Islands. 
Bishop Museum Occasional Papers 48: 23-36. 

Meyer, J.-Y. 1996. Status of Miconia calvescens (Me- 
lastomataceae), a dominant invasive tree in the 
Society Islands (French Polynesia). Pacific Sci- 
ence 50: 66-76. 

. 1998. Observations on the reproductive biol- 
ogy of Miconia calvescens DC (Melastomata- 
ceae), an alien invasive tree on the Island of Tahiti 
(South Pacific Ocean). Biotropica 30: 609—624. 

Meyer, J.-Y. and J. Florence. 1996. Tahiti’s native flora 
endangered by the invasion of Miconia calves- 
cens. Journal of Biogeography 23: 775-781. 

Muddiman, S. B., I. D. Hodkinson, and D. Hollis. 
1992. Legume-feeding psyllids of the genus Het- 
eropsylla (Homoptera: Psylloidea). Bulletin of 
Entomological Research 82: 73-117. 

Noyes, J. S. and P. Hanson. 1996. Encyrtidae (Hyme- 
noptera: Chalcidoidea) of Costa Rica: the genera 
and species associated with jumping plant-lice 
(Homoptera: Psylloidea). Bulletin of The Natural 
History Museum, Entomology Series 65: 105— 
164. 

Swarbrick, J. T. 1997. Weeds of the Pacific Islands. 
Technical paper No. 209, South Pacific Commis- 
sion, Noumea, New Caledonia. 124 p. 


VOLUME 107, NUMBER 4 


Van Klinken, R. D. 2000. Host-specificity constrains 
evolutionary host change in the psyllid Prosopi- 
dopsylla flava. Ecological Entomology 25: 413— 
422. 


749 


Wineriter, S. A., S. E. Halbert, and J. P. Cuda. 2003. 
Boreioglycaspis melaleucae Moore (Insecta: He- 
miptera: Psyllidae). (access code http://crea- 
tures.ifas.ufl.edu/beneficial/b_melaleucae.htm) 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 750-761 


BAIMAIA, A NEW SUBGENUS FOR ANOPHELES KYONDAWENSIS 
ABRAHAM, A UNIQUE CRABHOLE-BREEDING ANOPHELINE IN 
SOUTHEASTERN ASIA 


RALPH E. HARBACH, RAMPA RATTANARITHIKUL, AND BRUCE A. HARRISION 


(REH) Department of Entomology, The Natural History Museum, Cromwell Road, 
London SW7 5BD, U.K. (email: rharbach@nhm.ac.uk); (RR) Museum of World Insects, 
72 Nimamhemin 13, Huay-Kaeo Road, Chiangmai 50200, Thailand (email: insects_ 
museum @hotmail.com); (BAH) Public Health Pest Management, North Carolina Depart- 
ment of Environment and Natural Resources, 585 Waughtown Street, Winston-Salem, NC 
27107 (bruce.harrison@ncmail.net) 


Abstract.—Baimaia, n. subg., is introduced as a new subgenus of Anopheles for the 
unusual crabhole species, An. kyondawensis Abraham, in Southeast Asia. A diagnosis of 
the subgenus is provided that features unique anatomical characters of the adult, larval, 
and pupal stages of the type species. The larva of An. kyondawensis is redescribed and 
the previously unknown adult female, adult male, and pupa are described in detail. The 
affinities of Baimaia and An. kyondawensis are discussed in terms of their position in the 


phylogeny of Anophelinae, and their bionomics and distribution are reviewed. 


Key Words: 


Abraham (1947) described Anopheles 
(Anopheles) kyondawensis from larvae 
found in shallow ground pools along 
streams near the village of Kyondaw 
(Moulmein Township, Mon State) in south- 
ern Myanmar. The species was not encoun- 
tered again until 1966 when a single larva 
was found in a crabhole at Ban Pha Man 
located near the Laos border in Nan Prov- 
ince of Thailand (Harrison and Scanlon 
1975). Harrison and Scanlon (1975) sug- 
gested that the larvae collected by Abraham 
in Myanmar may have been swept out of 
freshwater crabholes by high water because 
they were found in association with larvae 
he identified as An. (Cellia) leucosphyrus 
Do6nitz [probably An. baimaii Sallum and 
Peyton (recently described in Sallum et al. 
2005) since it is the dominant species of the 
Leucosphyrus Group in Myanmar], which 
“normally occur in small temporary ground 


Culicidae, Anophelinae, taxonomy, mosquito 


pools that follow flooding or rains.” In 
1979, another larva was collected from a 
stream in Huai Kop, Sai Yok District of 
Kanchanaburi Province in western Thailand 
(Harrison et al. 1991). Likewise, it is likely 
that this larva may have been dislodged 
from a crabhole because the margin of the 
stream where it was captured was lined 
with these habitats at water level. Oo et al. 
(2004) found one larva of An. kyondawen- 
sis in a shaded pool along a stream in the 
vicinity of Innwaing near the type locality 
during collections made in Myanmar be- 
tween May 1998 and March 2000. These 
authors did not indicate whether the pool 
was associated with crabholes. Material ex- 
amined during the present study, however, 
confirms that An. kyondawensis does in fact 
breed in burrows created by freshwater 
crabs. This material consists of larvae, and 
adults reared from larvae and pupae col- 


VOLUME 107, NUMBER 4 


lected from crabholes in Ban Tham Sua, 
Tak Province, located in northwestern Thai- 
land. 

Reid and Knight (1961) included An. 
kyondawensis in the Culiciformis Group of 
subgenus Anopheles based on the reduced 
setae 5,6,7-C, and other setae of the larval 
head capsule. Harrison and Scanlon (1975) 
considered this placement tentative until the 
adult and pupal stages were known. Fol- 
lowing the discovery of the previously un- 
known adult and pupal stages, we initiated 
studies of this species, and after having 
considered all of the unique features in the 
adult, larval, and pupal stages noted below, 
we concluded that An. kyondawensis does 
not belong in any currently recognized spe- 
cies group of Anopheles. 

The traditional classification of subfami- 
ly Anophelinae included three genera: 
Anopheles Meigen, Bironella Theobald, 
and Chagasia Cruz. The phylogenetic re- 
lationships of these genera, based on a cla- 
distic analysis of morphological data (Har- 
bach and Kitching 1998), reflect the intui- 
tive hypothesis (Ross 1951) that Anophel- 
inae is a monophyletic clade comprised of 
Chagasia in a sister-group relationship to 
Bironella + Anopheles. Molecular phylog- 
enies inferred from nuclear and mitochon- 
drial gene sequences also support this hy- 
pothesis of relationships (Besansky and 
Fahey 1997; Foley et al. 1998; Krzywinski 
et al. 200la, b), but more recent studies 
based on both morphological (Sallum et al. 
2000, Harbach and Kitching 2005) and mo- 
lecular data (Sallum et al. 2002) suggest 
that Anopheles is a paraphyletic assemblage 
relative to Bironella. In the absence of sup- 
port for the generic status of Bironella, Sal- 
lum et al. (2000) formally synonymized 
this taxon with Anopheles s.s. This synon- 
ymy, however, is not supported (see below) 
by the later studies of Sallum et al. (2002) 
and Harbach and Kitching (2005), which 
indicate that Bironella should be regarded 
as a subgenus of Anopheles. 

Taking account of independent lines of 
evidence, especially sequence data for the 


751 


slowly evolving single-copy nuclear white 
gene, Krzywinski and Besansky (2003) hy- 
pothesized that Bironella diverged from the 
main lineage of Anopheles following the 
earlier separation of Chagasia. Although 
this hypothesis is not supported by the mor- 
phological and molecular phylogenetic 
studies of Sallum et al. (2000 and 2002, re- 
spectively), it is not inconsistent with the 
results of the more recent cladistic analysis 
of Harbach and Kitching (2005), one aim 
of which was to investigate the phyloge- 
netic position of An. kyondawensis. This 
latter analysis placed An. kyondawensis as 
sister to Bironella + all other Anopheles, 
with Chagasia as sister to these three taxa. 
Although this arrangement of taxa, ex- 
pressed parenthetically as Chagasia + (An. 
kyondawensis + (Bironella + other Anoph- 
eles)), raises questions concerning the bio- 
geography of anophelines, support for this 
set of relationships (assessed using Bremer 
and relative Bremer support) is strong and 
indicates that both An. kyondawensis and 
Bironella are independent lineages relative 
to the rest of Anophelinae (see Harbach and 
Kitching (2005) for a full assessment of re- 
lationships and character support). These 
results agree with the suggestion by Sallum 
et al. (2002) that ““Bironella may be plau- 
sibly regarded as a subgenus of Anophe- 
les’, and imply in accordance with appli- 
cation of the principle of equivalent rank 
(Hennig 1966) that An. kyondawensis 
should also be afforded subgeneric rank. 
Hence, a new subgenus is proposed herein 
for this species. 


MATERIALS AND METHODS 


This study is based on a small number of 
larvae, and adults reared from larvae and/ 
or pupae collected from crabholes (as in- 
dicated above), and the holotype larva of 
An. kyondawensis deposited in The Natural 
History Museum (NHM), London (see Ma- 
terial examined following the species de- 
scription). Because the medium in which 
the holotype was mounted on a microscope 
slide had turned black with age, the speci- 


2 


men was removed following the procedures 
of Brown and De Boise (2005) and re- 
mounted in Euparal on the same slide. The 
head and dissected mouthparts of the larva 
were mounted under a separate coverslip. 
Following stereoscopic examination, the 
head of the only available female was re- 
moved, cleared in 5% NaOH for 2 h at 
50°C, and mounted, with the mouthparts 
separated from the head capsule, in euparal 
on a microscope slide for more detailed 
study. The genitalia of 2 available males 
were also dissected and likewise cleared 
and mounted on individual microscope 
slides. Pinned adults were examined under 
simulated natural light; dissections, larvae, 
and larval and pupal exuviae were studied 
with differential interference contrast op- 
tics. Measurements and counts were made 
from all available specimens. Numbers in 
parentheses represent modes, when appar- 
ent, of the reported ranges. The anatomical 
terminology and abbreviations used in the 
descriptions and illustrations, respectively, 
follow Harbach and Knight (1980, 1982). 
The symbols 2, 5, Le, Pe, and L used in 
the literature summary and material exam- 
ined sections for An. kyondawensis repre- 
sent female, male, larval exuviae, pupal ex- 
uviae, and fourth-instar larva, respectively. 
An asterisk (*) after one of these symbols 
in the literature summary section indicates 
at least part of the life stage was illustrated 
in the publication cited. 


TAXONOMIC TREATMENT 


Anopheles subgenus Baimaia, Harbach, 
Rattanarithikul, and Harrison, new 
subgenus 


Type species.—Anopheles kyondawensis 
Abraham, 1947. 

Diagnosis.—Baimaia appears to be relat- 
ed to subgenus Anopheles Meigen, espe- 
cially the Aitkenii, Alongensis, and Culici- 
formis Groups, but differs principally in 
features of the male genitalia, which are 
unique within genus Anopheles. The gono- 
coxite does not have differentiated parabas- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


al and inner setae, and the gonostylus is 
flattened and mitten-shaped distally and 
lacks a gonostylar claw. Males also have 
uniquely developed maxillary palpi, which 
are straight and very nearly cylindrical with 
palpomeres 4 and 5 barely swollen and only 
slightly flattened. Adults lack thoracic scal- 
ing, and females have a dense covering of 
long sensilla between the antennal whorls 
that depart a fuzzy appearance to the anten- 
nae. The immature stages are found in crab- 
holes, or in pools after having been washed 
out of these habitats. Pupae have a trumpet 
that appears undifferentiated and interme- 
diate between angusticorn and laticorn, spi- 
racular scars of abdominal segments H—VII 
and seta 9-II-VIII borne ventrally, seta 1- 
III plumose, and a long fringe of spicules 
on the inner and outer margins of the pad- 
dle. Larvae have strongly inwardly curved 
antennae, setae 5,6,7-C reduced, long, sin- 
gle, compressed or flattened, somewhat lan- 
ceolate setae on the thorax and abdomen, 
and setae 6-[V—VI as long as setae 6-J-—III. 
Baimaia is monobasic: see in the System- 
atics section following the description of 
An. kyondawensis for discussion. 

Etymology.—Baimaia is a patronymic 
honoring Prof. Visut Baimai of Mahidol 
University, Bangkok, for his many impor- 
tant contributions to our knowledge of the 
cytogenetics and systematics of Anopheles 
mosquitoes in southeastern Asia. We have 
chosen to Latinize Visut’s surname by add- 
ing the feminine suffix “*-a”’ rather than the 
masculine “*-us’’ because Baimadia is more 
euphonious and easier to pronounce (Bi-mi- 
0). The three-letter abbreviation Bmi. is rec- 
ommended for this subgenus. 


Anopheles (Baimaia) kyondawensis 
Abraham, 1947 
(Fig. 1) 


Anopheles (Anopheles) kyondawensis Abra- 
ham, 1947 (L*); Delphin and Rao 1957 
(L key); Reid and Knight 1961 (classifi- 
cation); Reid 1968 (L key); Kyi 1971 
(distribution); Rattanarithikul and Harri- 
son 1973 (L* key); Harrison and Scanlon 


VOLUME 107, NUMBER 4 153 


ES 


N 


St: ie Zale 
Ss 2, i 


25 -"\ 6 
Jee 7) y “ot 
1 7 Boe 
es. IN ‘ 
uw 5 a 


sev 


sternal 


° 


2 SS 
| ane SS 


tergal 1 


7. Froward 


Fig. 1. Pupa and male genitalia of Anopheles (Baimaia) kyondawensis. A,B, Pupa: (A) left side of cepha- 
lothorax, dorsal to right; (B) dorsal (left) and ventral (right) aspects of metathorax and abdomen. C, Male 
genitalia, aspects as indicated. Ae = aedeagus; Cl = claspette; CT = cephalothorax; Ge = gonocoxite; GL = 
genital lobe; Gs = gonostylus; Pa = paddle; T = trumpet; I-VIII = abdominal segments I-VIII; 0-14 = setal 


numbers for specified areas, e.g., seta 1-III. Scales in mm. 


754 


1975 (L*, distribution, taxonomy); Har- 
rison et al. 1991 (collection record); Har- 
bach 1994, 2004 (classification); Oo et al. 
2004 (collection record). 


Female.—Head: Vertex with sparse 
long, dark, narrow, truncate erect scales me- 
dially (centrally), abruptly graded into 
equally long attenuate setae laterally and 
anteriorly, latter indistinguishable from oc- 
ular setae that are removed from eye mar- 
gin, decumbent scales absent; frontal tuft 
comprised of few slender piliform scales 
and 4 long forward projecting setae, scales 
and setae slightly paler (more golden) than 
those of vertex. Clypeus without scales. 
Antenna length about 1.5 mm; pedicel bare, 
yellowish; flagellum with dense covering of 
relatively long anteriorly curved setalike 
sensilla between antennal whorls, flagello- 
mere | with a few inconspicuous narrow 
scales on dorsomesal surface among most 
proximal setae and sensilla, other flagello- 
meres without scales. Proboscis long and 
slender, entirely dark-scaled, labella paler 
than prementum, with 6 long and few short- 
er basal setae that fan out from ventral and 
lateral surfaces, length about 2.35 mm, 
1.15 length of forefemur, slightly longer 
than maxillary palpus (about 1.1). Max- 
illary palpus long and slender (palpomere 5 
very slightly swollen), length about 2.15 
mm, entirely dark-scaled, palpomeres | and 
2 with semierect scales, palpomere 1 with 
1 long and several shorter setae, palpomere 
2 with few short inconspicuous setae on 
proximal half, palpomeres 4 and 5 with 
rather widely spaced, short, inconspicuous, 
forward-projecting setae on ventrolateral 
margin. Cibarial armature absent. Thorax: 
Without scales; integument light brown. 
Scutum with prominent dark brown setae 
on all areas (anterior promontory, acrosti- 
chal, dorsocentral, lateral prescutal, fossal, 
antealar, supraalar, and prescutellar areas); 
parascutellar seta equally prominent. Scu- 
tellum evenly rounded with many long and 
few short dark brown setae. Mesopostno- 
tum bare. Antepronotum with row of dark 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


brown setae along anterior margin. Post- 
pronotum with 1—3 inconspicuous setae on 
extreme posterodorsal margin (could be 
confused for prespiracular setae). Pleura 
with dark setae as follows: 1 upper proe- 
pisternal, 1 prealar, 1 upper mesokatepister- 
nal, 2 lower mesokatepisternal, 3 or 4 upper 
mesepimeral, no lower mesepimeral. Wing: 
Length 3.25 mm; entirely dark-scaled; hu- 
meral crossvein and alula without scales; 
cell M, about 0.55 length of cell R,; cell M, 
about 0.6 length of vein M,,;. Dorsal sur- 
face with elongate semierect scales on R,, 
R,,;, R>, proximal 0.5 of R;, M,,,, and dis- 
tally on 1A, remaining veins with decum- 
bent spatulate scales; ventral surface with 
similar scales on middle of R,, R,,; and dis- 
tally on CuA and 1A. Halter: Pedicei 
mainly pale, scabellum and capitellum dark, 
capitellum dark-scaled. Legs: Slender; cox- 
ae without scales, prominent dark brown se- 
tae on anterior surface of forecoxa, outer 
surface of midcoxa and posterior surface of 
hindcoxa; trochanters with dark scales and 
short dark setae on ventral surfaces; femora, 
tibiae, and tarsi entirely dark-scaled; fore- 
femur length about 2.05 mm, 0.87 length of 
proboscis. Abdomen: Integument dark 
brown dorsally, lighter ventrally; without 
scales; with long brown setae. 
Male.—Like female except antennal 
whorls with more numerous and much lon- 
ger setae, long setalike sensilla only present 
on 2 terminal flagellomeres, which are dis- 
proportionately long compared to the other 
flagellomeres, apical flagellomere about 
0.75 length of penultimate flagellomere, 
which is 4X longer than more proximal fla- 
gellomeres. Maxillary palpus slightly short- 
er than proboscis; palpomeres 4 and 5 near- 
ly cylindrical, very slightly expanded. Same 
numbers of pleural setae as in female ex- 
cept 2—4 upper mesepimeral setae present. 
Genitalia (Fig. 1C): Tergum IX without se- 
tae; sternum IX small, not fused to tergum. 
Gonocoxite relatively short and stout, with- 
out scales, setae on dorsolateral, lateral, and 
ventrolateral areas strongly developed, 
more distal ones longer than gonocoxite 


VOLUME 107, NUMBER 4 755 
Table 1. Range of numbers of branches for pupal setae of Anopheles (Baimaia) kyondawensis. 
Seta Cephalothorax gbdomine) Peemens Paddle 
No. cT I iit I IV V VI VI IX P 
0) — — ] ] 1 1 | ] _ — 
1 2,3(2) ~300 14-21 26—36(28) 1-4(4) 1 | 1 ] l 
2 2,3(2) 5-9 3=5(5) 3,4(4) 1,2(2) 1,2(2) 1,2(2) 2 — 2 
3 2 1,2(1) 3—9(7) 5-11 8-11 1S) 5,6 (3 (6) = 
4 2,3(2) 2,3(3) 1—3(3) 2,3(2) 1—3(2) PP3(6)) | — — 
5 2-6(2) 3—6(5) 2,3(2) 7—11(8) 1,2(1) ] l a —_ 
6 2 2-4 1—3(2) ] 1 1 I NA) 9 == = 
7 2,3(3) 1 1—3(1) 14 1—3(2) 1,3(3) ] 13) = 
8 1,2(1) — m 2,3(2) 1—3(2) 1,2(2) D3; SAG) = 
9 14(2) 1 1 ] 1 1 l | — a 
10 4—6(4) — — 24 | I ] 1-3(2) — _— 
11 5-7(6) — — 1 1 i ] 1 = — 
12 4—6(5) = = = = = = = = = 
14 _ — — it ] 1 l | - — 


and gonostylus, setae on inner and parabas- 
al areas not developed or distinct from other 
prominent setae on dorsomesal surface; 
gonostylus short, curved, laterally flattened, 
and expanded in distal half, expanded por- 
tion with thumblike projection at approxi- 
mately middle of lateral margin and rela- 
tively sparse covering of minute setae on 
ventral surface, dorsal surface with sparse 
row of similar setae that extends onto prox- 
imal portion; gonostylar claw absent. Ae- 
deagus jong and slender, leaflets absent; 
claspette a single undivided lobe, bearing a 
close-set apical row of 6 (apparent) similar, 
straight setae. 

Pupa (Fig. 1A, B).—Character and po- 
sitions of setae as illustrated; numbers of 
branches in Table 1. Cephalothorax: Even- 
ly and lightly to moderately pigmented. 
Seta 4-CT significantly longer that 5-CT, 
approaching length of 6-CT. Trumpet: Un- 
differentiated and intermediate between an- 
gusticorn and laticorn; without tracheoid 
area; pinna without fold opposite cleft (1 of 
3 available specimens with an unnatural 
fold due to mounting), rim thin and uni- 
form. Abdomen: Evenly and moderately 
pigmented; length 2.9—3.0 mm. Spiracular 
scars of segments II—VII borne ventrally 
near middle of lateral margins. Seta 9-II— 
VIII inserted ventral to caudolateral corner 


of segments, 9-II—III short, peglike, 9-[V— 
VII progressively longer and more sharply 
pointed on succeeding posterior segments, 
9-VII ventral and inserted close to 7-VII 
(but when mounted, see Fig. 1B, appears to 
be near posterolateral corner), 9-VIII long, 
with well-developed branches (14—23); 1- 
III plumose, with 26—36 branches, 1;5-IV— 
VII equally strongly developed, 1.4—1.6x 
length of tergum, all single except 1-IV 
with 1—4(4) branches arising well beyond 
base; 3-III anterior to 1-III; 7-IV—VII, and 
sometimes 7-III, inserted on fold line, 7-V— 
VII at posterior margin of sternum. Genital 
lobe: Length 0.30 mm in male; 0.18 mm 
in female. Paddle: Lightly pigmented; 
asymmetrical, outer part broadest in basal 
half, inner part broadest in distal half; 
length about 0.75 mm, width about 0.50 
mm, index about 1.5; refractile border 
about 0.25 paddle length; long dense spic- 
ules on both inner and outer margins, about 
0.5 length of seta 1-Pa. Seta 1-Pa relatively 
long, single, curved; 2-Pa double, about 0.8 
length of 1-Pa. 

Larva, fourth-instar—As illustrated by 
Rattanarithikul and Harrison (1973) and 
Harrison and Scanlon (1975) except where 
noted below; ranges of setal branching in 
Table 2. Head: Slightly wider than long, 
length 0.67 mm, width 0.70—0.82 mm; 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


756 


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VOLUME 107, NUMBER 4 


more or less evenly pigmented, collar and 
most posterior part of frontal ecdysial line 
darkly tanned. Seta 1-C long, attenuate; 2- 
C single, simple, very long, length 0.3—0.4 
mm, arising very close to its mate, their al- 
veolar sockets more or less confluent; 3-C 
short and rather stout, length 0.20—0.25 that 
of 2-C; 4-C single, slender, simple, about 
length of 3-C, inserted far forward of 5,6- 
C, more widely separated from its mate 
than setae 3-C; 5,6,7-C very short, 
branched from base; 8,9-C short, single, 
simple; 11-C well developed, about as long 
as antenna, plumose. Antenna: Cylindrical, 
markedly curved inward, with relatively 
few spicules on proximal half of mesal sur- 
face; length 0.25—0.28 mm. Seta 1-A very 
short, length about diameter of antenna at 
point of insertion on proximal 0.17 of an- 
tenna, with 2—5(3) branches from middle of 
stem; 4-A with 4—10 branches, slightly lon- 
ger that 2,3-A. Thorax: Integument hya- 
line, smooth. Seta 1-P without setal support 
plate, with branches arising from short 
stem; 2-P about 5X length of 1-P, borne on 
small setal support plate; 3-P single, simple 
and slightly flattened; 9,10,12-P,M and 
9,10-T long, single, simple, common sup- 
port plates of these setae with very short 
spine; 12-T short, normally single and 
slightly flattened (characterized as “‘bifid 
distally” by Harrison and Scanlon 1975); 
13-T long, single, simple and slightly flat- 
tened. Abdomen: Integument hyaline, 
smooth; tergal plates small, less than 0.25 
width of segments, segments IV—VII, and 
sometime III, each with small median ac- 
cessory tergal plate. Setae 0-II-VIUI and 14- 
I-VI (14-II-VII incorrectly shown on 
posterior margins of segments II-VI in il- 
lustration of Harrison and Scanlon 1975) 
minute, single; 1-I minute, usually single, 
occasionally double, 1-II with slender un- 
pigmented lanceolate leaflets, 1-III—VII ful- 
ly palmate, leaflets broad, flat and pig- 
mented, with distinct shoulders and apical 
filaments; 2-IV—VI, 3-I-VI and 4-VI,VII 
long, single and flattened, 3-I1V long, flat- 
tened and bifid or trifid distally; seta 5 very 


/ 


small on segment I and progressively larger 
on segments II to VII, 5-I usually double, 
5-11 triple and 5-IV—VII increasing in 
turn on average from 3 or 4 branches to 6 
or 7 branches; 6-III as long as 6-I,I, nor- 
mally with 8 or 9 short widely spaced 
branches mainly on proximal half (not 
‘flattened’? as indicated by Harrison and 
Scanlon 1975), 6-IV-VI as long as 6-II, 
simple, 6-VII very small with 2 or 3 
branches; 10-I and 13-VII occasionally 
double, otherwise these setae and 10-V,VI, 
1i-H, 12-I,VI,VU and 13-V single, simple 
and flattened; 13-I-IV,VI small, branched 
from short basal stem. Pecten plate with 
14—16 subequal spines, each with strong 
denticles arising from basal half on dorsal 
side. Saddle moderately pigmented, length 
0.27—0.28 mm; seta 1-X single, simple and 
slightly flattened, about 1.3 saddle length, 
inserted close to margin of saddle (incor- 
rectly illustrated on edge of saddle by Har- 
rison and Scanlon 1975). 
Systematics.—The number and positions 
of specialized setae on the gonocoxites of 
the male genitalia have served as the pri- 
mary basis for the subgeneric classification 
of Anopheles since the pioneering work of 
Christophers (1915). Disregarding a few 
apparent departures from the usual condi- 
tion in subgenus Anopheles, the arrange- 
ment and characteristics of these setae are 
constant within the subgenera. In compari- 
son with the development and positions of 
these setae in the six traditionally recog- 
nized subgenera of Anopheles (see Reid 
1968: fig. 26), it is obvious that An. kyon- 
dawensis (Fig. 1C) does not fit within any 
of these groups. Furthermore, the notion 
that the specialized gonocoxal setae of 
Anopheles evolved from ordinary setae of 
the gonocoxite is supported by the relative- 
ly unspecialized condition of apparently ho- 
mologous setae in this species. This, as well 
as the thin, very nearly cylindrical maxil- 
lary palpi of males, with the two apical pal- 
pomeres only slightly more swollen than 
those of Chagasia, reinforces the hypothe- 
sis that the ancestral stock of Anopheles 


758 


gave rise to An. kyondawensis and a lineage 
from which all other Anopheles evolved. 
Disregarding the unique features of the 
male genitalia, An. kyondawensis exhibits a 
number of characters in common with spe- 
cies of the Alongensis and Culiciformis 
groups of subgenus Anopheles. The adults 
are small, unadorned insects and the larvae 
have cranial setae 5, 6, and 7 strongly re- 
duced. The Alongensis Group includes two 
species (An. alongensis Venhuis and An. 
cucphuongensis Vu, Nguyen, Tran, and 
Nguyen) and the Culiciformis Group in- 
cludes three species (An. culiciformis Cog- 
ill, An. sintoni Puri, and An. sintonoides 
Ho) (Harbach 2004) in the Oriental Region. 
Larval habitats utilized by members of the 
Alongensis Group (known only from Viet- 
nam) include rock holes in a limestone cave 
and small holes in limestone outcrops in 
forested hills (An. alongensis and An. cuc- 
phuongensis, respectively). These habitats 
share a generic likeness to the shaded, cryp- 
tic crabholes occupied by larvae of An. 
kyondawensis. Larvae of the Culiciformis 
Group are found primarily in treeholes and 
other plant containers. Adult females of An. 
kyondawensis are easily distinguished from 
those of the Alongensis and Culiciformis 
Groups by the dense covering of long sen- 
silla between the antennal whorls and the 
presence of minute setae on the postero- 
dorsal margin of the postpronotum. The in- 
wardly curved antennae of An. kyondaw- 
ensis are found otherwise only in An. cuc- 
phuongensis (specimens were unavailable 
to determine whether this character also oc- 
curs in the closely related An. alongensis). 
The flattened single thoracic and abdominal 
setae on the larva of An. kyondawensis are 
unique, and the pupa is distinguished by the 
plumose condition of seta 1-III and the ven- 
tral placement of seta 9-II-VIII. Also, the 
pupal trumpet of An. kyondawensis is un- 
differentiated and intermediate between the 
angusticorn- and laticorn-shaped trumpets 
that Reid and Knight (1961) recognized and 
used to divide subgenus Anopheles into two 
taxonomic sections, the Angusticorn and 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Laticorn Sections. The trumpet of An. kyon- 
dawensis has a fairly long meatus, a shal- 
low meatal cleft, a pinna that is widely 
flared, and an uncertain axis. In fact, it is 
very similar to other primitive species with 
generalized trumpets, e.g., An. sintonoides 
(see Harrison and Scanlon 1975) and An. 
sintoni (see Tewari and Hiriyan 1992) of 
the Culiciformis Group. 

Harbach and Kitching (2005) recently in- 
cluded An. kyondawensis in a phylogenetic 
study of Anophelinae based on morpholog- 
ical characters. Their findings revealed that 
An. kyondawensis is the earliest taxon de- 
rived from an ancestor that gave rise to all 
other Anopheles. A search of the matrix 
containing 167 characters that occur in spe- 
cies of Anopheles that were used in that 
study revealed 19 characters (numbers 2, 4, 
NO; IS, 23, 22!) 4, a), Oss 10; 19, YZ, G3; 
96, 97, 101, 107, 141, and 156) that occur 
in species of Anopheles that retain apparent 
ancestral (plesiomorphic) characters. Only 
two of these 19 characters occur in more 
derived species of subgenus Cellia (number 
24 in An. funestus Giles and 96 in An. ci- 
nereus Theobald and An. superpictus Gras- 
S1), aS Well as species in subgenus Anoph- 
eles. Of the other 17 characters, two (92, 
93) are unique to An. kyondawensis, and the 
remaining 15, plus the two that occur in the 
three species of Cellia, were otherwise 
found only in Chagasia and Anopheles 
(species in subgenera Anopheles, Bironella, 
Kerteszia, Lophopodomyia, and Stetho- 
myia). The shared plesiomorphic characters 
that occur in An. kyondawensis and other 
species are listed in Table 3. 

Bionomics.—Most of what is known 
about the bionomics of An. kyondawensis is 
noted in the introduction. The immature 
stages apparently inhabit crabholes associ- 
ated with streams or streamlets and are oc- 
casionally washed out of these habitats. 
Larvae have been found only in crabholes 
and small pools along the sides of streams 
shaded by forest in hilly and mountainous 
areas. Species found in association with An. 
kyondawensis in crabholes include Culex 


VOLUME 107, NUMBER 4 759 


Table 3. Shared plesiomorphic characters of Anopheles (Biamaia) kyondawensis that occur in various species 
in other subgenera of Anopheles (Anopheles, Bironella, Lophopodomyia, and Stethomyia), and Chagasia (char- 
acter numbers taken from Harbach and Kitching 2005). These characters, which are exhibited by taxa that occur 
in nearly every zoogeographical region, corroborate the more basal position that An. kyondawensis occupies in 


the phylogeny of Anopheles, and justify Baimaia as a new subgenus. 


No. Shared Characters Character Nos. 


Species Zoogeographic Region 
An (Ano.) aitkenii James Oriental 
An. (Ano.) algeriensis Theobald Palaearctic 
An. (Ano.) asiaticus Leicester Oriental 
An. (Ano.) atropos Dyar and Knab Nearctic 
An. (Ano.) corethroides Theobald Australasian 
An. (Ano.) cucphuongensis Vu, 

Nguyen, Tran and Nquyen Oriental 
An. (Ano.) implexus Theobald Afrotropical 
An. (Ano.) interruptus Puri Oriental 
An. (Ano.) judithae Zavortink Nearctic 
An. (Ano.) sintonoides Ho Oriental 
An. (Bir.) confusa Bonne-Wepster Australasian 
An. (Bir.) gracilis Theobald Australasian 
Kw5S0Bir.) hollandi Taylor Australasian 
An (Ker.) bambusicolus Komp Neotropical 
An. (Ker.) cruzii Dyar and Knab Neotropical 
An. (Lph.) oiketorakras Osorno- 

Mesa Neotropical 
An. (Ste.) nimbus (Theobald) Neotropical 
An. (Ste.) kompi Edwards Neotropical 
Ch. fajardi (Lutz) Neotropical 


2,4,15 

AMS) 

79 

15 
2,4,107,141 


Ree N W 


15 

2 

79 

15,76,101 

4,15,76 
2,4,10,15,24,76,97,101 
10,15,24,68,97,101 
15,24,64,65,68,97,101 
76 

76 


RS Re YO COW WRK ee 


97 

2,24,65,96,107 
2,24,65,96,107 
2,10,23,64,65,141,156 


NNN 


(Lophoceraomyia) bengalensis Barraud, 
Cx. (Lop.) minor (Leicester), Cx. (Lop.) spi- 
culosus Bram and Rattanarithikul, Urano- 
taenia (Pseudoficalbia) abdita Peyton, Ur. 
(Pfc.) koli Peyton and Klein, Ur. (Pfc.) 
stricklandi Barraud, Ur. (Uranotaenia) 
macfarlanei Edwards, and unidentified spe- 
cies of Verrallina (Harrison and Scanlon 
1975; collection records for the specimens 
listed below). Larvae collected with An. 
kyondawensis in stream pools include An. 
(Anopheles) bengalensis Puri, An. (Ano.) 
insulaeflorum (Swellengrebel and Swellen- 
grebel de Graaf), and members of the An. 
(Cellia) dirus complex (Abraham 1947, Oo 
et al. 2004). Adults of this species have 
never been collected in the wild, and noth- 
ing is known about their biology or behav- 
i0r. 

Distribution.—Anopheles kyondawensis 
has been found only in places on either side 
of the Thai-Myanmar border located rough- 


ly between 14 and 17° north (Kanchanaburi 
and Tak Provinces, Thailand; Mon State, 
Myanmar), and at one other place (approx- 
imately 18°50'’N 100°50’E) nearly 400 km 
northeast of these localities near the Thai- 
Laos border in Nan Province, Thailand. As 
noted by Harrison and Scanlon (1975), this 
suggests that An. kyondawensis is more 
widely distributed in forested hilly and 
mountainous areas of mainland Southeast 
Asia. 

Material examined.—Eleven specimens: 
Dl ie. SSRe wand 452 incindine 
the holotype. Holotype L, MYANMAR: 
Mon State, Kyondaw [currently Kyondo] 
(16°36'N 98°04’E), small shallow pool 
along side of stream, 2 Dec 1943 (NHM). 
Other specimens, | d Pe (1—518-18), THAI- 
LAND: Tak Province, Ban Tham Sua 
(16°41'N 98°41’'E), crabhole in shallow 
stream in forest, 8 Jun 90 (Rampa); 1° Pe 
(1-520-105), 1¢éLePe (1-520-13), 2L (1- 


760 


520), Ban Tham Sua (16°41'N 98°42'B), 
crabhole in streamlet in forest, 8 Jun 90 
(Chamnong). 


ACKNOWLEDGMENTS 


We are grateful to Theresa Howard (The 
Natural History Museum, London) for pre- 
paring figure 1, and Jim Pecor (Walter Reed 
Biosystematics Unit, Smithsonian Institu- 
tion, Washington, DC) for the loan of spec- 
imens critical to this study. 


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Besansky, N. J. and G. T. Fahey. 1997. Utility of the 
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Harbach, R. E. and I. J. Kitching. 1998. Phylogeny and 
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PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 762—769 


MORPHOLOGY OF THE ANTENNA OF CAENOCHOLAX FENYESI PIERCE 
(STREPSIPTERA: MYRMECOLACIDAE) BASED ON SCANNING 
ELECTRON MICROSCOPY 


DAWN P. DERR AND JERRY L. COOK 


Department of Biological Sciences, Sam Houston State University, Huntsville, TX, 
77341-2116, U.S.A. (e-mail: bio_jlc@shsu.edu) 


Abstract.—The antenna of Caenocholax fenyesi Pierce includes two types of sensilla. 
Sensilla chaetica are located at the distal ends of segments three and four. These sparsely 
abundant sensilla are thought to be tactile sensory structures. Sensilla coeloconica are 
found abundantly on segments three through seven, including the flabellum of segment 
three. These sensilla coeloconica are chemoreceptors that are presumably used as phero- 
mone receptors used in location of the female, which remains an endoparasite. The mor- 
phology of the antenna of C. fenyesi is shown using scanning electron microscopy. 


Key Words: 


Caenochoax fenyesi, a member of the 
strepsipteran family Myrmecolacidae, was 
first described by Pierce (1909). The de- 
scription was based on four males collected 
in Cordoba, Veracruz, Mexico by Dr. A. 
Fenyes. No females were collected, and the 
host species was not identified. Subsequent- 
ly, both sexes have been collected (Kathir- 
ithamby and Johnston 2003). Adult male C. 
fenyesi are free-living, while adult females 
are permanent endoparasites. Although 
most Strepsiptera are host specific, C. fen- 
yesi, as well as other myrmecolacids, are 
distinctive in that females parasitize mem- 
bers of Orthopteroidea whereas males par- 
asitize members of the family Formicidae 
(Hymenoptera) (Kathirithamby 1989). 
Caenocholax fenyesi appears to be either an 
atypical generalist among the Strepsiptera 
or a species complex, with males known to 
parasitize a variety of ant species (Cook et 
al. 2004). The adult male C. fenyesi life- 
span is limited to a few hours (Cook 1996) 
and in this time it must locate the endopar- 
asitic female to successfully mate. Caeno- 


sensilla, antenna, morphology, Caenocholax fenyesi 


choax fenyesi has a widespread distribution 
ranging from Argentina to the southern 
United States (Cook et al 1997, Kathiri- 
thamby and Hughs 2002). 

Pierce (1909), in his original description, 
and Bohart (1941), in a review of the order, 
described the adult male Caenochoax fen- 
yesi antennae as being seven-jointed or seg- 
mented. Given that scanning electron mi- 
croscopy (SEM) was not available, the de- 
scriptions are general and do not give any 
detail of cuticular sensilla of the antennae. 
In fact, little has been reported concerning 
the strepsipteran antennae, except for the 
morphological study by Kinzelbach (1971). 
Kinzelbach reported that strepsipterans 
have two types of antennal sensilla, sensilla 
trichodea and sensilla basiconica. Addition- 
ally, Strepsiptera have a sensory depression, 
named Hofeneder’s organ, on the fourth an- 
tennal segment, except in the families Elen- 
chidae and Bohartilidae where it is on the 
third segment (Hofeneder 1910, Kinzelbach 
1971, Kathirithamby 1989). 

Based on the work of Snodgrass (1935) 


VOLUME 107, NUMBER 4 


and Schneider (1964), Zachurak (1985) 
compiled a listing of ten categories or types 
of insect sensilla. By definition sensilla are 
organs that possess “‘structural and func- 
tional mechanisms needed for accepting 
stimuli, generating a nerve impulse mes- 
sage and conducting this message to an ap- 
propriate receiving cell” (Zachurak 1985). 
Sensilla are found on several areas of the 
body, namely antennae, mouthparts, legs 
and wings, genitalia and anal cerci, as well 
as others; and come in many forms, to in- 
clude hairs, bristles, spines, pegs, cones, 
plates and scales. In investigating sensilla, 
researchers often attempt to attribute some 
function to a given structure—chemorecep- 
tor, thermoreceptor, and hygroreceptors— 
just to list a few. To understand both the 
morphology and function of sensilla, it is 
important to have an understanding of the 
life history of the organism. 

Early studies of antennal sensilla yielded 
information on the number of types of sen- 
silla on the flagella of numerous insects at 
both the adult (male and female) and some 
instar stages. For example, adult Heliothis 
zea (Lepidoptera) and Stomoxys calcitrans 
(Diptera) have four distinct types while 
Apis mellifera (Hymenoptera) have ten sen- 
silla types (Zachurak 1985). Recent work 
using the scanning electron microscope has 
allowed for detailed study of the cuticular 
sensilla of Diptera (De Freitas Fernandes et. 
al. 2002), Hemiptera (Liang 2001) and Co- 
leoptera (Merivee et. al. 2000, 2002) that 
includes not only typology but also size, 
number and distribution patterns. 

Gross morphology of the antennae of C. 
fenyesi is described by Pierce (1909) as 
“seven jointed; the first two joints trans- 
verse, cylindrical, cupped; third joint trans- 
verse, cupped, but produced outwardly be- 
neath in a long flabellum, which is almost 
as long as the metathorax; fourth joint 
transverse, cylindrical; fifth elongate five 
sixth as long as the width of the head; sixth 
seven-tenths as long as the fifth and slightly 
surpassed by the flabellum of the third; sev- 
enth four-fifth as long as the fifth.” This 


763 


description was accompanied by a figure, 
which we reproduce below to compare with 
the antennae of our study. Kathirithamby 
and Johnston (1992) provided a SEM of the 
head that included a portion of the anten- 
nae, but provided no antennal description. 
The objective of this study was to identify 
the different morphological types and lo- 
cation of cuticular sensilla on the antennae 
of adult male Caenochoax fenyesi. 


MATERIALS AND METHODS 


Preserved specimens of adult male Caen- 
ocholax fenyesi from the collection at Sam 
Houston State University were studied us- 
ing a scanning electron microscope. Male 
Strepsipteran specimens were obtained 
from parasitized individuals in colonies of 
Solenopsis invicta Buren collected from 
Brazos County, Texas. 

Specimens preserved in 100% ethanol 
were air dried for 15 to 30 seconds on filter 
paper, mounted on aluminum SEM stubs, 
and coated with gold using a Cressington 
108 sputter coater. Micrographs were taken 
using a VEGA TS 5130SB scanning elec- 
tron microscope (Sam Houston State Uni- 
versity, Huntsville, Texas) at 15 kV. 

Morphological terminology used for 
classification is based on Schneider (1964) 
and Zacharuk (1985). 


RESULTS AND DISCUSSION 


The antenna drawn by Pierce (1909) il- 
lustrated specimens from the type locality 
in Cordoba, Mexico. Pierce drew an anten- 
na similar in form to the antennae of our 
study, but lacking any detail of the sensory 
structures. Figure | shows a comparison of 
Pierce’s antenna with an antenna from the 
Texas specimens used for our study. The 
dimensions and shape of antennae were 
consistent to Fig. 1B in all specimens we 
viewed. 

Non-sensory, multi-cellular processes of 
the antennal cuticle are found on the surface 
of all segments of the antennae, but are ex- 
cluded from the areas where sensilla are lo- 
cated. These are platelike extensions ar- 


764 


Fig. 1. 
at 453; scale bar = 100 pm. 


ranged in alternating rows and terminating 
with a slender spine or hairlike structure. 
We also found these extensions in conjunc- 
tion with sensory structures, the cuticle sur- 
rounds the sensilla and may provide protec- 
tion. These are the only processes found on 
the first and second basal segments of the 
antenna (Fig. 2). 

Two types of sense organs were identi- 
fied—sensilla chaetica (ch) and sensilla 
coeloconica (co). Sensilla chaetica were 
found on segments three and four, while 
sensilla coeloconica were located on seg- 
ments three through seven. 

Sensilla chaetica are sensory bristles or 
spines usually set into a socket. They are 
generally thought to be tactile, but some 
have been identified as chemoreceptors. In- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Antenna of Caenocholax fenyesi. 1A, Antenna redrawn after Pierce 1909. 1B, Antenna SEM taken 


nervation is by one or more neurons (Za- 
charuk 1985). In Caenochoax fenyesi these 
sensilla are sparsely located along the distal 
third of segment three (on the margin near 
the attachment to segment four) and in a 
similar location on segment four (Fig. 3). 
The extended portion, the flabellum, of seg- 
ment three does not contain any chaetica 
sensilla. The bristles range from 7.3 to 11.5 
wm from the socket insertion and taper 
slightly from base to distal end (Fig. 4). As 
these sensilla are usually considered tactile, 
the location may suggest that they are used 
in confirming the positioning of the long 
flagellar segment. Chaetica sensors have 
been identified on the flagella of Lepidop- 
tera, Orthoptera, Blattaria, Hemiptera, Co- 
leoptera, Siphonaptera, and Diptera 


VOLUME 107, NUMBER 4 


SS cat 


Figs 2-3. Antennal segments of Caenocholax fenyesi. 2, Basal antennal segments (segments I & ID). 3, 
Segments III & IV. Scale bar = 10 pm. 


766 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 4-5. Sensilla of Caenocholax fenyesi antennal segments. 4, Sensilla chaetica on antennal segment III. 
5, Sensilla coeloconica surrounded by non-sensory structures. Scale bar = 5 pm. 


VOLUME 107, NUMBER 4 


Figs. 6-7. 


Male Caenocholax fenyesi. 6, Sensilla coeloconica on the three terminal antennal segments. 


Segment VII is at the bottom right; Scale bar = 50 pm. 7, Ventral view of the head of Caenocholax fenyesi. 


Scale bar = 200 pm. 


(Schneider, 1964). These structures are pre- 
sumably the same as those identified in oth- 
er Strepsiptera by Kinzelbach (1971) but la- 
beled as sensilla trichodea. The difference 
between these two sensilla is that sensilla 
chaetica are set in a socket while sensilla 
trichodea are freely moveable on a basal 
membrane with variable basal insertion on 


the cuticle (Zacharuk 1985). The sensilla on 
the antenna of C. fenyesi are clearly emerg- 
ing from sockets and bristlelike; and are 
therefore sensilla chaetica. Our study had 


the luxury of seeing these structures with 
the electron microscope, making it easier to 
distinguish between sensilla chaetica and 


sensilla trichodea. However, some strepsip- 


768 


terans may indeed have sensilla trichodea, 
or a combination of both of these sensilla. 
More studies using an electron microscope 
to examine other strepsipteran species is 
needed to elucidate which of these struc- 
tures are present on strepsipteran antennae. 

Sensilla coeloconica are thin-walled sen- 
sory cones on the floor of depressions or 
pits in the cuticle. The pegs are innervated 
by four or five neurons forming a bundle 
that terminates at the tip of the peg. Coe- 
loconic organs are most often reported to 
be thermo-, chemo-, or hygroreceptors (Za- 
charuk 1985). In Caenochoax fenyesi the 
pegs appear to be smooth, ball-like struc- 
tures nestled in deep depressions that are 
surrounded by numerous non-sensory, mul- 
ti-cellular processes (Fig. 5). The pegs 
range from 2.5 to 6.5 wm. Smaller pegs 2.5 
to 4.2 wm are located on segment four, 
while pegs from 4.5 to 6.5 wm can be found 
on the elongated portion of segment three 
as well as segments five, six and seven (Fig. 
6). Where coeloconica pegs are found, they 
are evenly distributed. Sensilla coeloconica 
have been described on antennae of Hemip- 
tera (Liang 2001), Diptera (De Freitas Fer- 
nandes et. al. 2002), Coleoptera (Merivee 
et. al. 2000, 2002), and Orthoptera (Schnei- 
der 1964). The life history of the adult male 
Caenochoax fenyesi suggests that chemo- 
receptivity plays an important role in locat- 
ing a female with which to mate. Caeno- 
cholax fenyesi males only live a few hours, 
in which time they must locate an endopar- 
asitic adult female, if they are going to suc- 
cessfully mate (Cook 1996). The large 
number of coeloconica sensilla on the an- 
tennae and their natural history may rein- 
force the proposition that these sense or- 
gans are chemoreceptors. The distribution 
of the coeloconica sensilla on the head of 
the male C. fenyesi could facilitate detec- 
tion and location of pheromones given off 
by the female (Fig. 7). 


ACKNOWLEDGMENTS 


We thank Brandon Lowery for help in 
assembling the plates and Sam Houston 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


State University for access to the scanning 
electron microscope. 


LITERATURE CITED 


Bohart, R. M. 1941. Revision of the Strepsiptera with 
special reference to the species of North America. 
University of California Publications in Entomol- 
ogy 7(6): 91-160. 

Cook, J. L. 1996. A study of the relationship between 
Caenocholax fenyesi Pierce (Strepsiptera, Myr- 
mecolacidae) and the red imported fire ant, Sole- 
nopsis invicta Buren (Hymenoptera, Formicidae). 
Ph.D. Dissertation, Texas A&M University, Col- 
lege Station, Texas. 

Cook, J. L., J. S. Johnston, R. E. Gold, and S. B. Vin- 
son. 1997. Distribution of Caenocholax fenyesi 
(Strepsiptera: Myrmecolacidae) and the habitats 
most likely to contain its stylopized host, Sole- 
nopsis invicta (Hymenoptera: Formicidae). Envi- 
ronmental Entomology 26: 1258-1262. 

Cook, J. L., L. A. Calcaterra, and L. Nufiez. 2004. First 
record of Caenocholax fenyesi (Strepsiptera: Myr- 
mecolacidae) parasitizing Solenopsis invicta (Hy- 
menoptera: Formicidae) in Argentina with a dis- 
cussion on its distribution and host range. Ento- 
mological News 115(2): 61-66. 

De Freitas Fernandes, F, P. M. Linardi, and H. Chiar- 
ini-Garcia. 2002. Morphology of the antenna of 
Dermatobia hominis (Diptera: Cuterebridae) 
based on scanning electron microscopy. Journal of 
Medical Entomology 39(1): 36-43. 

Hofeneder, K. 1910. Mengenilla n.g. chobautii n.sp. 
Eine neue Stsepsiptere aus Nordafrika. Berichte 
des Naturwissenschaftlich-Medizinischen Bereins 
in Innsbruck 32: 33-58, 2 pl. 

Kathirithamby, J. 1989. Review of the order Strepsip- 
tera. Systematic Entomology 14: 41-92. 

Kathirithamby, J. and D. P. Hughes. 2002. Caenocho- 
lax fenyesi (Strepsiptera: Myrmecolacidae) para- 
sitic in Camponotus planatus (Hymenoptera: For- 
micidae) in Mexico: Is this the original host? An- 
nals of the Entomological Society of America 
95(5): 558-563. 

Kathirithamby, J. and J. S. Johnston. 1992. Stylopi- 
zation of Solenopsis invicta (Hymenoptera: For- 
micidae) by Caenocholax fenyesi (Strepsiptera: 
Myrmecolacidae) in Texas. Annals of the Ento- 
mological Society of America 85: 293-297. 

. 2003. The discovery after 94 years of the elu- 
sive female of a myrmecolacid (Strepsiptera), and 
the cryptic species of Caenocholax fenyesi Pierce 
senso lato. Proceedings of the Royal Society, Lon- 
don Series B (Supplement), Biology Letters, pub- 
lished online 17 September 2003 (DOI 10.1098/ 
rsbl.2003.0078). 

Kinzelbach, R. K. 1971. Morphologische Befunde an 
Facherfliiglern und ihre phylogenetische Bedeu- 


VOLUME 107, NUMBER 4 


tung (Insecta: Strepsiptera). Zoologica 41 (119: 1. 
und 2. Halfte): 1-256. 

Liang, A. 2001. Scanning electron microscopy of am- 
tennal sense organs in Prosapia bicincta (Say) 
(Hemiptera: Cercopidae). Journal of Entomologi- 
cal Science 36(4): 335-341. 

Merivee, E., A. Ploomi, M. Rahi, J. Bresciani, H. P. 
Ravn, A. Luik, and V. Sammelselg. 2002. Anten- 
nal sensilla of the ground beetle Bembidion pro- 
perans Steph. (Coleoptera, Carabidea). Micron 
33: 429-440. 

Merivee, E., A. Ploomi, M. Rahi, A. Luik, and V. Sam- 
melselg. 2000. Antennal sensilla of the ground 
beetle Bembidion lampros Hbst (Coleoptera, Car- 
abidae). Acta Zoologica 81: 339-350. 


769 


Pierce, W.D. 1909. A monographic revision of the 
twisted winged insects comprising the order Strep- 
siptera Kirby. United States National Museum 
Bulletin 66: 1—232. 

Schneider, D. 1964. Insect antennae. Annual Review 
of Entomology 9: 103-122. 

Snodgrass, R. E. 1935. Principles of Insect Morpkol- 
ogy. McGraw-Hill Book Company, Inc., 
York and London, pp. 510-549. 

Zacharuk, R.Y. 1985. Chapter 1, Antennae and Sen- 
silla, pp. 1-69. In Kerkut, G. A. and L. I. Gilbert, 
eds. Comparative Insect Physiology, Biochemis- 
try, and Pharmacology, Vol. 6. Pergamon Press, 
Oxford. 


New 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 770-781 


WHAT?’S IN A NAME? LEPIDOPTERA: HESPERIIDAE: PYRGINAE: 
TELEMIADES HUBNER 1819 [PYRDALUS MABILLE 1903]: 
NEW COMBINATIONS TELEMIADES CORBULO (STOLL) 

AND TELEMIADES OICLUS (MABILLE)—AND MORE 


JOHN M. BURNS AND DANIEL H. JANZEN 


(JMB) Department of Entomology, National Museum of Natural History, Smithsonian 
Institution, PO. Box 37012, MRC 127, room E-515, Washington, DC 20013-7012, USA 
(e-mail: burnsj@si.edu); (DHJ) Department of Biology, University of Pennsylvania, Phil- 
adelphia, PA 19104, USA (e-mail: djanzen @sas.upenn.edu) 


Abstract.—Both the Central American skipper butterfly Achlyodes oiclus Mabille and 
the South American Pyrdalus corbulo (Stoll) belong in Telemiades. Pyrdalus becomes a 
junior synonym of Telemiades. Pyrdalus corbulo cora Evans, which is really a species 
(not a subspecies), is a new synonym of Telemiades oiclus, new combination. Though 
differing sharply in wingshape and color pattern, T. oiclus and Telemiades corbulo, new 
combination, share a distinctive male secondary sex character and are, in both sexes, 
genitalically similar to each other and to T. nicomedes (Moschler). Grewn caterpillars of 
T. oiclus and T. nicomedes resemble each other (and suggest slugs). DNA barcoding 
further supports the relationship of these species. With its brown-forewing/brown-and- 
yellow-hindwing adult color pattern, T. oiclus superficially resembles 13 other species of 
skippers reared in the Area de Conservacion Guanacaste (ACG) of northwestern Costa 
Rica. Of these presumably mimetic species, one is raised from reduction to subspecific 
rank, and two are raised from synonymy, to gain reinstated status: 7. gallius (Mabille), 
T. chrysorrhoea (Godman and Salvin), and Eracon lachesis (Dyar). The pupa of T. oiclus 
shares distinctive features with the pupae of other species of Telemiades. All eight species 
of Telemiades reared in the ACG feed only on leaves of plants in the family Fabaceae. 
Six eat various species of Jnga and, in a relatively few cases, species in three other 
mimosoid genera, whereas 7. oiclus and T. nicomedes each use two species in one pap- 
ilionoid genus—Dioclea and Machaerium, respectively. 


Key Words: nomenclature, genitalia (male and female), secondary sex character, facies, 
mimicry, pupa, caterpillar, foodplants, Fabaceae, Dioclea, Machaerium, 


Inga, Area de Conservacion Guanacaste, Costa Rica 


Our grasp of evolution and its biodiver- 
sity products improves with better scientific 
names. 

More than a century ago, the Central 
American species of skipper butterfly that is 
central to this paper was described, named, 
and misplaced in an older genus by a worker 
who, 14 years later, proposed a new and ap- 


propriate genus—but only for a different- 
looking South American skipper species. 
Half a century after that, another worker 
misapplied the specific name of the Central 
American species and described and named 
that taxon again—but placed it (as a subspe- 
cies of the South American species) in the 
appropriate genus. Details follow. 


VOLUME 107, NUMBER 4 


Mabille (1889: 25, fig. 3) described Ach- 
lyodes oiclus from a single female from 
“Chiriqui’” (today a Pacific province in 
western Panama) with a cursory printed ac- 
count of superficial appearance and a crude 
black-and-white cut of dorsal and ventral 
wing surfaces. Godman and Salvin (1895: 
WOl, Z; so SLO, WOl. Bs le wo, mss, ZO, Zl) 
briefly redescribed the type specimen of A. 
oiclus and captured its odd facies in excel- 
lent color paintings of dorsal and ventral 
wing surfaces, thus making it relatively 
easy to identify, both then and now. Mabille 
(1903) described the genus Pyrdalus for 
one South American species, Pyrdalus cor- 
bulo (Stoll). This skipper was originally de- 
scribed (from Surinam) in the genus Papilio 
in 1781 and was transferred, a century 
thereafter, to Celaenorrhinus. 

Evans (1953: 80) treated what he called 
“oiclus Mabille’ as a taxon in the genus 
Bolla. Pointing out that Evans, in so doing, 
had to be dealing with something other than 
Achlyodes oiclus, Steinhauser (1989) re- 
stored that combination and applied it to a 
female from Turrialba, Costa Rica, in the 
Allyn Museum of Entomology. Because 
Evans’s concept of oiclus was wrong, he 
failed to recognize a male and female of 
this species (from Honduras) in the British 
Museum (Natural History) and described 
them anew as Pyrdalus corbulo cora Evans 
(1953: 32). Here his genus-level insight was 
good; but, as in many other cases, he 
lumped at the species level (Burns and Jan- 
zen 2001: 19-20). What he described is re- 
ally a distinct species, not a subspecies of 
P. corbulo. Pyrdalus cora Evans, new sta- 
tus, is a new synonym of Pyrdalus oiclus 
(Mabille), new combination. 

But Pyrdalus, which groups two closely 
related species of skippers, does not warrant 
generic rank. In characterizing Pyrdalus, 
Evans (1953: 31) wrote, “‘Genitalia, palpi 
and antennae as in Telemiades. ¢ 
up[per]f[orewing] with a costal fold... ,” 
which also applies to most species of Te- 
lemiades. It turns out that Pyrdalus is a 
small subdivision of the sizeable, diverse, 


771 


and taxonomically challenging neotropical 
genus Telemiades. 

From data in three of the foregoing no- 
menclatural efforts, we know that Telemi- 
ades oiclus, new combination, ranges, at 
the very least, from Honduras to Panama. 
Judging from our experience in the Area de 
Conservacion Guanacaste (ACG) in north- 
western Costa Rica (Burns and Janzen 
2001, Janzen and Hallwachs 2005), 7. oi- 
clus is a rainforest skipper, occurring from 
about 520 to 320 m on Atlantic (Caribbean) 
foothills of the Cordillera de Guanacaste 
and feeding, as a caterpillar, on mature 
leaves of two large rainforest vines, Dioclea 
malacocarpa Ducke and D. wilsonii Standl. 
(Fabaceae). It may well go lower in this 
general region because its foodplants do. 
However, the bioinventory has yet to reach 
the ACG lowest limit (200 m) and will not 
exceed that limit. A series of 21 adults 
(reared from 36 wild-caught caterpillars) all 
beseeching an informative name catalyzed 
this study. 


MATERIALS 


Except for those marked AMNH (which 
are in the American Museum of Natural 
History, New York, NY, USA), adult spec- 
imens examined of each of the two species 
here moved to Telemiades are in the Na- 
tional Museum of Natural History 
(USNM), Smithsonian Institution, Wash- 
ington, DC, USA; and this depository does 
not appear in the following specimen lists. 
These start with numbers of adult males and 
females examined, then (in parenthesis) 
numbers of male and female genitalia com- 
pared. Wherever applicable, the lists spec- 
ify Janzen and Hallwachs ACG rearing 
voucher codes and adult eclosion dates, and 
(in parenthesis) genitalia dissection codes 
(those of Burns begin with X-). The rearing 
codes give, in tripartite-and-hyphenated 
form, the last two digits of the year that the 
caterpillar was collected, SRNP, and a num- 
ber unique for that year. SRNP stems from 
Santa Rosa National Park, which is now a 
part of the ACG. 


772 


‘ ai 
Cae Bele 


Figs. 1-3. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Male genitalia of Telemiades oiclus from the ACG, Costa Rica (00O-SRNP-21533, X-5061); scale 


= 1.0 mm. 1, Tegumen, uncus, and gnathos in dorsal view. 2, Complete genitalia (minus right valva), with 
everted vesica, in left lateral view. 3, Cornuti at distal end of vesica in another view. 


Information on rearing of caterpillars and 
on handling and storage of genitalia may be 
found in Burns and Janzen (2001) and Jan- 
zen and Hallwachs (2005) and in Burns 
(1997), respectively. 

Telemiades oiclus.—12 6, 13 2 (6 46,6 
2). COSTA RICA: Area de Conservaci6n 
Guanacaste: Sector Pitilla, Sector San Cris- 
tobal, and Rincon Rainforest, 9 ¢, 12 2 as 
follows: 99-SRNP-5157, 27 May 1999, ° 
(X-5810); OO-SRNP-11806, 15 Aug 2000, 
3; 00-SRNP-21533, 31 Oct 2000, 3 (X- 
5061); OO-SRNP-22202, 7 Jan 2001, 2 (X- 
5062); 01-SRNP-2923, 4 Sep 2001, ?; 01- 
SRNP-2924, 17 Sep 2001, 2 (X-5808); 
01-SRNP-2947, 8 Sep 2001, 6 (X-5069); 
01-SRNP-3300, 9 Oct 2001, 6; O01-SRNP- 
3727, 8 Nov 2001, ¢; 01-SRNP-3782, 23 
Oct 2001, 2 (X-5070); 01-SRNP-25089, 14 
Jan 2002, 3 (X-5809); 02-SRNP-2271, 26 
Apr 2002, 2; 02-SRNP-19803, 21 Dec 
2002, d (X-5807); 03-SRNP-6806, 12 Aug 


2003, 2 (X-6043); 03-SRNP-7308, 6 Aug 
2003, 2; 03-SRNP-7387, 17 Aug 2003, 6; 
03-SRNP-9990, 30 Nov 2003, ¢; 03- 
SRNP-12536.1, 3 Oct 2003, ¢; 03-SRNP- 
12588.1, 19 Oct 2003, 2; 03-SRNP-20865, 
3 Oct 2003, 2; 04-SRNP-30865, 26 Apr 
2004, 2. Guapiles, May, 1 3 (X-5071). 
Turrialba, 27 Mar 1969, 1 6, V. King. 
PANAMA: Canal Zone, La Pita, 11 Jun 
1963, 1 ¢ (S. S. Nicolay genitalia dissec- 
tion H359), G. B. Small. Cerro Campana, 
1500 ft (455 m), 10 Jan 1964, 1 2 (K- 
5072), G. B. Small. 

Telemiades corbulo, new combina- 
Ui) 6, 42 SG Ss 3 LY). WwIRAVAILE 
Pard, Obidos, no date, 1 3 (E. L. Bell gen- 
italia dissection G645), 1 2, AMNH. Ron- 
dénia, vic. Cacaulandia, 10°32'S, 62°48'W, 
160-350 m, 22 Oct 1991, 1 6 (K-5077), J. 
MacDonald. ECUADOR: Napo, Limonco- 
cha, Rio Napo, 240 m, 9 Feb 1971, 1 6 G. 
S. Nicolay genitalia dissection H466), S. S. 


VOLUME 107, NUMBER 4 


773 


Figs. 4-6. 


& S. Nicolay; Puerto Napo, 540 m, 7 Nov 
1988, 1 ¢ (X-5075), 1 2 (X-5076), S. S. 
Nicolay, 9 Nov 1988, 1 6, D. H. Ahren- 
holz. FRENCH GUIANA: Cayenne, Dec 
1903, | 2 (X-6045). St. Laurent, no date, 
1 6, AMNH. Vidal, 4°52’N, 52°18'W, O- 
S0imea/ Novel 9838; 4D: Jy Hanvey;: Ei 
Nov 1988, 1 2 (X-5074), D. J. Harvey; 18 
Nov 1988, 2 6 (X-5073), D. J. Harvey. 
PERU: Loreto, Iquitos, 21 Feb 1932, 1 6, 
AMNH. 


DISCUSSION 


Genitalia.—Telemiades oiclus and T. 
corbulo are united (figuratively) by their 
genitalia, which, in each sex, are clearly 
variations on the same theme (cf. Figs. 1— 
10). However, despite many basic similari- 
ties, the intricate male genitalia differ in un- 
mistakable ways. These include (a) the mid- 
dorsal, distal end of the uncus—in dorsal 
view, convex (Fig. 1) in 7. oiclus vs. con- 
cave (Fig. 4) in 7. corbulo; (b) the middor- 


Male genitalia of Telemiades corbulo from Puerto Napo, 540 m, Napo, Ecuador, 7 Nov 1988, 
S. S. Nicolay (X-5075); scale = 1.0 mm. 4, Tegumen, uncus, and gnathos in dorsal view. 5, Complete genitalia 
(minus right valva), with everted vesica, in left lateral view. 6, Cornuti at distal end of vesica in another view. 


sal center of the uncus—with a small dor- 
sally-directed cone (Figs. 1—2) in 7. oiclus 
vs. no such protuberance (Figs. 4—5) in 7. 
corbulo; (c) the dentate, upcurved, distal 
end of the valva—in lateral view, with two 
or more major points (Fig. 2) in 7. oiclus 
vs. one very major point (Fig. 5) in T. cor- 
bulo; and (d) a short secondary sac off the 
main everted vesica—with 0, 1 (Fig. 2), 2, 
or 3 spines in 7. oiclus vs. a row of spines 
(Fig. 5) in 7. corbulo. The simpler female 
genitalia differ most notably in the width of 
the midventral U-shaped notch in the pos- 
terior margin of the lamella postvaginalis— 
narrow (Fig. 7) in 7. oiclus vs. wide (Fig. 
9) in T. corbulo. Having been caught in co- 
pulo, the male and female of 7. corbulo 
whose genitalia appear in Figs. 4—6 and 
Figs. 9-10 were literally united by them. 
Previously, the male genitalia of 7. cor- 
bulo have been adequately, albeit inexactly, 
figured by Williams and Bell (1933: pl. IV, 
fig. 3) and by Evans (1953: pl. 28, fig. 


774 


E[.]7). Although Williams and Bell (1933: 
83) claimed that “‘the left [valva] is similar 
to the right but the apex is more rounded 
externally,” there is no sign of such asym- 
metry in the 7. corbulo dissections at hand. 
Nevertheless, two species of Telemiades 
described by Evans do have distinctly 
asymmetric valvae (Evans 1953: pl. 27, 
figs. E.6.7.trenda, E.6.9.sila), as do a few 
other described and undescribed species of 
Telemiades. Mielke (1993: 614, 616, fig. 
22) illustrated the female genitalia of what 
he variously called the holotype/lectotype 
of Pythonides insulsus Draudt, which is 
considered one of the synonyms of 7. cor- 
bulo (the other is Eudamus praestes Hew- 
itson). Noting that the female genitalia do 
not agree with those of Pyrdalus corbulo 
whereas the rest of the type specimen ex- 
ternally looks like P. c. corbulo from south- 
ern Brazil, Mielke (1993: 614) allowed that 
the abdomen may be that of another spe- 
cies. It must be, because Mielke’s fig. 22 
does not look like Figs. 9—10 (either that, 
or Draudt’s insulsus is not a synonym of T. 
corbulo). 

Where in Telemiades does Pyrdalus fit? 
It clearly relates to T. nicomedes (MO6s- 
chler). This species—originally described 
from Colombia, and now thought to range 
from at least adjacent northeastern Peru and 
northern Brazil to Mexico—may be a spe- 
cies complex. To date, in the ACG, only 2 
3,2 @ of this possible complex have been 
reared. The ACG sample is too small, and 
comparative material of diverse geographic 
origin is too limited, to resolve the matter 
at this time (but the four ACG specimens 
themselves appear to be a single species). 
So the name 7. nicomedes is used here in 
the sense of “7. nicomedes or an unde- 
scribed species quite near it.” 

The intricate male genitalia of T. oiclus 
and T. corbulo (Figs. 1-6) are remarkably 
similar to those of T. nicomedes. In T. ni- 
comedes, both the valvae and the cornuti 
resemble those of 7. corbulo, while the 
middorsal, distal end of the uncus (which is 
squared to rather rounded) approaches that 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


of 7. oiclus. Notwithstanding the close 
overall resemblance, 7. nicomedes differs in 
various small ways. In all three species (and 
in many other species of Telemiades, as 
well), two pairs of lateral prongs project 
backward from the posterior end of the teg- 
umen. In all three species, the more dorsal 
prong is pointed at its tip, while the more 
ventral prong is rounded (Figs. 1—2, 4—5). 
In both 7. oiclus and T. corbulo, these 
prongs go straight backward; and the 
round-tipped prong extends well back 
(Figs. 1-2, 4—5)—at times, almost as far as 
does the pointed one (Figs. 1—2). But in 7. 
nicomedes, the ventral, round-tipped prong 
extends back only as far as the base of its 
dorsal, pointed counterpart; and that point- 
ed prong curves perceptibly dorsad instead 
of going essentially straight backward. Fur- 
thermore, the anterior end of the aedeagus 
bends to the left in 7. nicomedes but re- 
mains straight in 7. oiclus and T. corbulo. 

Likewise, the simpler female genitalia of 
T. oiclus and T. corbulo (Figs. 7-10) are 
very like those of 7. nicomedes. A notable 
difference is the more anterior position, in 
T. nicomedes, of a narrow, C-shaped “‘ring”’ 
of sclerotization around the posterior end of 
the ductus bursae, posterior to the entrance 
of the sperm duct. Anterior to that, in all 
three species, the long, narrow, mostly to 
wholly membranous ductus bursae increas- 
es in diameter and becomes sclerotized be- 
fore uniting with the membranous corpus 
bursae. 

DNA barcoding (a la Hebert et al. 2004) 
of some 360+ species of reared ACG hes- 
peruds independently indicates that 7. oi- 
clus and T. nicomedes are closely related, 
but distinct, species—and that they are 
more distinct from the other species of Te- 
lemiades, but cluster with them (Hajibabaei, 
Hebert, Burns, Janzen, and Hallwachs, un- 
published). 

Male secondary sex character.—Telemi- 
ades oiclus and T. corbulo share, in addi- 
tion to the costal fold, a male secondary sex 
character not otherwise evident in the genus 
Telemiades. On the dorsal hindwing, a tuft 


VOLUME 107, NUMBER 4 TUS 


Figs. 7-8. Female genitalia, ovipositor lobes, and part of ductus seminalis of Telemiades oiclus from the 
ACG, Costa Rica (O0-SRNP-22202, X-5062); scale = 1.0 mm. 7, Ventral view. 8, Right lateral view. 


776 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 9-10. Female genitalia, ovipositor lobes, and part of ductus seminalis of Telemiades corbulo from 
Puerto Napo, 540 m, Napo, Ecuador, 7 Nov 1988, S. S. Nicolay (X-5076); scale = 1.0 mm. 9, Ventral view. 
10, Right lateral view. 


VOLUME 107, NUMBER 4 


of long hairlike scales begins at the base of 
space 7 and projects distally, along space 7, 
halfway to the outer margin; surrounding 
wing scales, in space 8 and the proximal 
three-quarters of space 7 (plus, in T. oiclus, 
the proximal end of space 6), appear gray- 
ish and shiny (Figs. 19—20). On the adja- 
cent ventral forewing, scales with a simi- 
larly grayish, shiny look—in at least the 
proximal three-fifths of spaces la and Ib 
and the lower, proximal end of space 2— 
surround an elongate, oval androconial 
patch near the proximal end of the lower 
half of space 1b (Figs. 12, 16). In T. cor- 
bulo the grayish forewing scales in space 
la extend nearly to the outer margin (Fig. 
16). Both the dorsal hindwing tuft (Fig. 20) 
and the ventral forewing androconial patch 
(Fig. 12) that is associated with it are some- 
what orangy tan in 7. oiclus. In T. corbulo 
the tuft is mostly brown (Fig. 19), the an- 
droconial patch is cream (Fig. 16), and 
there is another creamy patch on the dorsal 
hindwing directly beneath the tuft. 

Facies.—Telemiades oiclus is a brown 
skipper in which the tornal half of the ven- 
tral hindwing is yellow (Figs. 11—14). The 
yellow is clearer and a little more extensive 
in females (Fig. 14) than it is in males (Fig. 
12). (However, the one Panamanian male 
examined has no yellow at all.) In sharp 
contrast, 7. corbulo is a brown skipper that 
weakly reflects a bluish/purplish color from 
the dorsal surface of both pairs of wings 
(Figs. 15-18). 

Telemiades oiclus and T. corbulo differ 
greatly in wingshape. Telemiades corbulo 
displays the usual hesperiid sexual dimor- 
phism in which the wings of females (Figs. 
17-18) are broader and rounder than those 
of their mates (Figs. 15-16). But T. oiclus, 
like one of four new species of Venada 
(Burns and Janzen 2005), suppresses this 
dimorphism so that the wings of males are 
as broad and round as those of females 
(Figs. 11-14). The difference in shape be- 
tween males of 7. oiclus and males of 7. 
corbulo is especially pronounced in the 
hindwing, which is relatively long and nar- 


777 


row in T. corbulo (cf. Figs. 19-20). The 
degree of sexual dimorphism in wingshape 
shown by 7. corbulo is representative of the 
genus Telemiades. Note the similarity of 
male wingshape in T. corbulo (Figs. 15—16) 
and 7. nicomedes (Figs. 23-24). 

Mimicry.—A color pattern essentially 
similar to that of 7. oiclus has evolved, 
more or less independently, in 13 other spe- 
cies of skippers reared in the ACG (an as- 
terisk marks those whose hindwing yellow 
is dorsal as well as ventral): the pyrgines 
*Typhedanus ampyx (Godman and Salvin), 
*Astraptes phalaecus (Godman and Sal- 
vin), A. chiriquensis (Staudinger), *A. an- 
aphus annetta Evans, Ocyba calathana cal- 
anus (Godman and Salvin), *Telemiades 
gallius (Mabille), reinstated status, *T7. 
chrysorrhoea (Godman and Salvin), rein- 
stated status, Eracon lachesis (Dyar), re- 
instated status, Aethilla lavochrea Butler, 
and Achlyodes busirus heros Ehrmann; and 
the hesperiines *Tromba xanthura (God- 
man), Damas immacula Nicolay (females 
only), and Aroma henricus (Staudinger). 
Images of these presumably mimetic hes- 
periids appear in Janzen and Hallwachs 
(2005). 

Tangential comments on the three species 
with reinstated status are in order. Evans 
(1953: 25) erroneously treated Telemiades 
gallius as a subspecies of 7. centrites (Hew- 
itson) and went on to synonymize T7. 
chrysorrhoea with T. centrites gallius. 
Judging from his comparative figures of the 
male valvae (Evans 1953: pl. 27, figs. 
E.6.4.centrites/gallius), T. gallius must be 
specifically distinct from 7. centrites. Fur- 
thermore, 7. gallius and T. chrysorrhoea 
are separate (but sister) species. Their gen- 
italia differ subtly, in both sexes, as do their 
facies: the hindwing yellow of T. gallius is 
orange (and a little less extensive) in T. 
chrysorrhoea; and all 139 reared ACG 
specimens of 7. chrysorrhoea lack the three 
small, hyaline, subapical white spots in 
spaces 6, 7, and 8 of the forewing that mark 
T. gallius. In the ACG, T. chrysorrhoea is 
a common cloud forest species (800—1400 


778 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 11-18. Adults, in dorsal (left) and ventral (right) views, of two species moved to Telemiades. 11—14, 
T. oiclus, reared in the ACG, Costa Rica. 11-12, 6 (1.8), 01-SRNP-3727. 13-14, 2 (X1.5), 03-SRNP-9990. 
15-18, T. corbulo, wild-caught at Vidal, 0-50 m, 4°52'N, 52°18’W, French Guiana, by D. J. Harvey. 15-16, 3 
(X1.6), 7 Nov 1988. 17-18, 2 (X1.6), 11 Nov 1988 (X-5074). 


VOLUME 107, NUMBER 4 


Figs. 19-26. Various features and stages of Telemiades species that (except for 7. fides) are closely related. 


19-20, Dorsal hindwings showing different overall shapes, plus tufts of a secondary sex character, in 7. corbulo 
(left) and T. oiclus (right). 21, Pupa 01-SRNP-2441 of 7. oiclus in ventral view. 22, Last instar caterpillar 01- 
SRNP-2441 of T. oiclus. 23-24, Adult 03-SRNP-21802 (X-5784) of 7. nicomedes in dorsal (left, X1.1) and 
ventral (right, 1.2) views. 25, Pupa 03-SRNP-10917 of 7. fides in dorsal view. 26, Last instar caterpillar 03- 
SRNP-27835 of 7. nicomedes. 


780 


m elevation), while 7. gallius is a rare, mid- 
elevation, rainforest species (600—900 m). 
DNA barcoding cleanly separates these two 
species (Hajibabaei, Hebert, Burns, Janzen, 
and Hallwachs, unpublished). 

Never having seen specimens of Ebrietas 
lachesis Dyar, Evans (1953: 66) hesitantly 
(and erroneously) placed this taxon in his 
new genus Morvina, as a subspecies of M. 
falisca (Hewitson). Mielke (2004) lists 
Ebrietas lachesis as a synonym of Eracon 
paulinus (Stoll). Eracon lachesis and E. 
paulinus are close—but distinguishable— 
species in what may be a species complex. 
Unlike E. lachesis, E. paulinus, in dorsal 
view, is proximally gray on both pairs of 
wings and is, on the distal hindwing, so 
blackish brown that the dark bands there are 
obscured, especially in males. Ventrally, 
males of E. paulinus do not express as 
much hindwing yellow as do males of E. 
lachesis. Male genitalia differ slightly: near 
the anterodorsal corner of the valva, a nar- 
row strip of fine dentation running along 
most of the dorsal edge of the valva bends 
sharply ventrad onto the inner surface of the 
valva, and extends farther ventrad in E. 
paulinus than in E. lachesis. Dissection and 
close comparison of male genitalia included 
the holotype of /achesis (from Teapa, Ta- 
basco, Mexico) in the USNM. Eracon lach- 
esis 18 the larger species. It ranges from 
southern Mexico to Panama, whereas E. 
paulinus is mainly South American. Nev- 
ertheless, the two species were taken to- 
gether, in 1969 and again in 1970, by S. S. 
Nicolay at about 455 m on Santa Rita 
Ridge, east of Colon, in Colon province, 
Panama—a locality pinpointed (as Cerro 
Santa Rita) in Ridgely (1976: 359). So far, 
40 3, 29 @ of E. lachesis have been reared 
in the ACG. 

Pupa.—The pupa of T. oiclus (Fig. 21) 
shares distinctive features with the pupae of 
other species of Telemiades, including T. fi- 
des Bell (Fig. 25). These features include a 
short, fat, conspicuous cone that projects 
straight forward from an interocular posi- 
tion at the anterior end of the head; a swol- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


len “‘midsection”’ (at the level of the ante- 
rior end of the abdomen); and, posterior to 
that, an abdomen that tapers sharply to the 
cremaster. All Telemiades pupae are shiny 
amber to amber brown, lacking a white 
waxy layer and markings (see images in 
Janzen and Hallwachs 2005). 

Caterpillar—Despite having dorsolateral, 
longitudinal, brown stripes peculiar to it, 
the last instar of T. oiclus (Fig. 22) resem- 
bles that of 7. nicomedes (Fig. 26). Both 
are light to medium brownish and heavily 
dotted dorsally with fine, glistening white 
spots that give an overall impression of a 
moist slug. In this connection, the caterpil- 
lar of 7. oiclus is especially plump and 
squat. Caterpillars of both species crawl 
with a slow, somewhat gliding motion. 
Moreover, if prodded, they hold their 
ground (like a slug), instead of crawling 
quickly away or turning to bite the molester. 
Penultimate and ultimate instar 7. oiclus 
have been found resting exposed on the 
dorsal surface of a foodplant leaf, rather 
than in a silk-and-leaf shelter like most 
skippers. Last instar 7. nicomedes make 
only a halfhearted attempt at a shelter, 
which may be just one leaf lightly tacked 
with silk to the leaf below. 

Foodplants.—So far, eight species of Te- 
lemiades have been reared in the ACG 
(from a total of 2068 Telemiades caterpil- 
lars). All eight feed strictly on plants in the 
Fabaceae, and six feed mainly or exclusive- 
ly on mature leaves of various species of 
the mimosoid genus /nga. Different species 
of Telemiades commonly share the same 
species of Inga. For example, five species 
of Telemiades eat I. oerstediana Benth. ex 
Seem. and J. sapindoides Willd., and four 
eat J. punctata Willd. Some of these /nga- 
eaters occasionally attack the mimosoid 
genera Pithecellobium, Zygia, and Cojoba. 
Each of two species of Telemiades reared 
in large numbers has been found on as 
many as 18 mimosoid legumes. Moss 
(1949) found three species of Telemiades in 
Amazonian Brazil feeding on /nga, and two 
of those on Pithecellobium, as well. 


VOLUME 107, NUMBER 4 


In contrast, 7. oiclus specializes on the 
papilionoid legume Dioclea, eating mature 
leaves of both D. malacocarpa and D. wil- 
sonii. Likewise, the rare T. nicomedes uses 
the papilionoid genus Machaerium, eating 
mature leaves of both M. acuminatum 
Kunth and M. seemannii Benth. 


ACKNOWLEDGMENTS 


We thank Donald J. Harvey, Patricia 
Gentil, Elizabeth A. Klafter, and Richard 
G. Robbins for dissecting 105 Telemiades 
and 38 Eracon genitalia, and Young T. 
Sohn for drawing four of the former; Fred- 
erick H. Rindge for access to material in 
his care at AMNH; W. Hallwachs, R. Mor- 
aga, G. Sihezar, G. Pereira, L. Rios, M. Pe- 
reira, O. Espinosa, E. Cantillano, M. Perei- 
ra, R. Franco, J. Perez, H. Ramirez, EK Cha- 
varria, M. M. Chavarria, C. Moraga, P. 
Rios, C. Cano, D. Garcia, E Quesada, E. 
Araya, M. Carmona, and M. Rios for cat- 
erpillar hunting and husbandry; R. Espino- 
sa, A. Guadamuz, and N. Zamora for iden- 
tifying foodplants; and two anonymous re- 
viewers for comments. This study was sup- 
ported most recently by NSF grants 
(D.H.J.) DEB 97-05072 and DEB 00- 
72730, the administration of the ACG, and 
a grant (J.M.B.) from the National Museum 
of Natural History Small Grants Program. 


LITERATURE CITED 


Burns, J. M. 1997. Presidential address 1996: On the 
beauties, uses, variation, and handling of genitalia. 
Journal of the Lepidopterists’ Society 51: 1-8. 

Burns, J. M. and D. H. Janzen. 2001. Biodiversity of 
pyrrhopygine skipper butterflies (Hesperiidae) in 
the Area de Conservaci6n Guanacaste, Costa 
Rica. Journal of the Lepidopterists’ Society 55: 
15-43. 

. 2005. Pan-neotropical genus Venada (Hesper- 

iidae: Pyrginae) is not monotypic: four new spe- 


781 


cies occur on one volcano in the Area de Conser- 
vacion Guanacaste, Costa Rica. Journal of the 
Lepidopterists’ Society 59: 19-34. 

Evans, W. H. 1953. A catalogue of the American Hes- 
periidae indicating the classification and nomen- 
clature adopted in the British Museum (Natural 
History). Part III. Pyrginae. Section 2. British Mu- 
seum, London. 246 pp., pls. 26—53. 

Godman, FD. and O. Salvin. 1879-1901. Biologia 
Centrali-Americana; Insecta; Lepidoptera-Rhopal- 
ocera. Vol. 2, 782 pp. Vol. 3, 113 pls. 

Hebert, DINE EB Ee Eenton Jee Vi Burns.eD)» Ei 
Janzen, and W. Hallwachs. 2004. Ten species in 
one: DNA barcoding reveals cryptic species in the 
neotropical skipper butterfly Astraptes fulgerator. 
Proceedings of the National Academy of Sciences 
of the United States of America 101: 14812— 
14817. 

Janzen, D. H. and W. Hallwachs. 2005. Event-based 
database of caterpillars, their host plants, and their 
parasitoids in the Area de Conservaci6n Guana- 
caste, northwestern Costa Rica. (http://janzen.sas. 
upenn.edu). 

Mabille, P. 1889. Diagnoses de lépidoptéres nouveaux. 
Le Naturaliste (2)3(45): 25. 

. 1903. Lepidoptera, Rhopalocera, Fam. Hes- 
peridae. Genera Insectorum 17a: 1—78. 

Mielke, O. H. H. 1993. Sobre os tipos de Hesperiidae 
(Lepidoptera) neotropicais descritos por M. 
Draudt. Revista brasileira de Entomologia 37: 
611-638. : 

. 2004. Hesperioidea, 95. Hesperiidae, pp. 25— 
86. In Lamas, G., ed. Checklist: Part 4A, Hesper- 
i0idea—Papilionoidea. /n Heppner, J. B., ed. Atlas 
of Neotropical Lepidoptera. Association for Trop- 
ical Lepidoptera, Scientific Publishers, Gaines- 
ville, Florida, xxxvi + 439 pp. 

Moss, A. M. 1949. Biological notes on some “Hes- 
periidae” of Para and the Amazon (Lep. Rhop.). 
Acta Zoologica Lilloana 7: 27—79, pls. I-V. 

Ridgely, R. S. 1976. A guide to the birds of Panama. 
Princeton University Press, Princeton, New Jer- 
sey, xv + 394 pp. 

Steinhauser, S. R. 1989. Taxonomic notes and descrip- 
tions of new taxa in the neotropical Hesperiidae. 
Part I. Pyrginae. Bulletin of the Allyn Museum 
127: 70 pp. 

Williams, R. C., Jr. and E. L. Bell. 1933. Studies in 
the American Hesperioidea. Paper I. (Lepidop- 
tera). Transactions of the American Entomological 
Society 59: 69-84, pl. IV. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 782-788 


TWO NEW SPECIES OF TRICHOGRAMMA (HYMENOPTERA: 
TRICHOGRAMMATIDAE) FROM THE RYUKYU ISLANDS, JAPAN 


JOSEPHINE JOSE, YOSHIMI HIROSE, AND JEFFREY Y. HONDA 


(JJ) Center for Biodiversity, Department of Biological Sciences, San Jose State Uni- 
versity, One Washington Square, San Jose, CA 95192-0100, U.S.A. Present address: De- 
partment of Entomology, 320 Morrill Hall, University of Illinois, 505 S. Goodwin Ave., 
Urbana, IL 61801, U.S.A.; (YH) Institute of Biological Control, Faculty of Agriculture, 
Kyushu University, Fukuoka 812-8581, Japan. Present Address: 349 Asano, Munakata 
811-3415, Japan; (JYH) Center for Biodiversity, Department of Biological Sciences, San 
Jose State University, One Washington Square, San Jose, CA 95192-0100, U.S.A. and 
Institute of Biological Control, Faculty of Agriculture, Kyushu University, Fukuok 812- 
8581, Japan (e-mail: jhonda@email.sjsu.edu) 


Abstract.—Two new species of Trichogramma Westwood from the Ryukyu Islands 
were collected in green pepper fields using trap cards containing Ephestia kuehniella 
Zeller (Lepidoptera: Pyralidae) eggs and from parasitized Plutella xylostella (Linnaeus) 
(Lepidoptera: Yponomeutidae) eggs. ITS-2 sequences and scanning electron micrographs 
were obtained for both Trichogramma species. Herein we describe Trichogramma cul- 


tellus n. sp., and T. umerus, n. sp. 


Key Words: 


Species of Trichogramma Westwood 
(Hymenoptera: Trichogrammatidae) in the 
Ryukyu Islands are poorly known. Ishi 
(1938, 1941) first recorded 7. chilonis Ishii 
as an egg parasitoid of Tetramoera (= Eu- 
cosma) schistaceana (Snellen) (Lepidop- 
tera: Totricidae) from Okinawa. Since then, 
there have been no records of additional 
species of Trichogramma from the Ryukyu 
Islands. In 1997, members of the Shimane 
Agricultural Experiment Station collected 
Trichogramma from trap Ephestia kuehn- 
iella Zeller (Lepidoptera: Pyralidae) egg 
cards and from parasitized Plutella xylos- 
tella (Linnaeus) (Lepidoptera: Yponomeu- 
tidae) eggs in Ishigaki and Iriomote Islands, 
respectively. Unique ITS-2 DNA sequences 
and microscopic examination indicated two 
taxa that have not been previously de- 
scribed. Herein we describe two new spe- 
cies from the Ryukyu Islands, Japan. 


Trichogramma, new species, Ryukyu Islands 


MATERIALS AND METHODS 


Species descriptions and specimens pre- 
pared for scanning electron microscopy 
(SEM) and DNA analysis are based on 
methods described previously (Taylor et al., 
in press). Species descriptions and mea- 
surements follow the anatomical terminol- 
ogy, morphological measurements, and ra- 
tios (relative dimensions) used in Pinto 
(1999). All types are deposited in the En- 
tomological Laboratory, Faculty of Agri- 
culture, Kyushu University (ELKU). 


Trichogramma cultellus Jose, Hirose, 
and Honda, new species 
(Figs. 1-6) 


Diagnosis.—Trichogramma cultellus is 
similar to the following species: 7. thalense 
Pinto and Oatman, 7. parkeri Nagarkatti, T. 
pintoi Voegele, and T. elegantum Sorokina. 


VOLUME 107, NUMBER 4 783 


Figs. 1-3. Trichogramma cultellus, male. 1, Dorsal view of genital capsule; scale bar = 50 pm. 2, Ventral 


view of genital capsule (IVP partially broken in preparation); scale bar = 50 jum. 3, Ventral view of apical 
distance (IVP partially broken in preparation); scale bar = 30 pm. 


784 


The female ovipositor length is much short- 
er in these species measuring 0.79, 0.86— 
0.95, 0.84, and 0.95 the hind tibia length 
for T. thalense, T. parkeri, T. pintoi, and T. 
elegantum, respectively, vs. 1.04—1.07 in 7. 
cultellus. The genital capsule is generally 
broader in 7. thalense with a width to 
length ratio greater than 0.30. Ventral pro- 
cesses are also positioned at the base of the 
intervolsellar process (IVP). In contrast, T. 
cultellus has a genital capsule width to 
length ratio of about 0.30 and ventral pro- 
cesses located distinctly anterior to the base 
of the IVP. Flagella setae are much longer 
in T. cultellus measuring over 3.5 X the fla- 
gellum width vs. 7. pintoi (2.56%), T. par- 
keri (2.85 X), and T. elegantum (2.2). The 
posterior extension of the dorsal lamina is 
much narrower in 7. cultellus and T. par- 
keri than in T. pintoi. However, the dorsal 
lamina is only 1.5X as long as wide in T. 
cultellus but over twice as long as wide in 
T. parkeri. 

Description.—Quantitative data taken 
from 7 males with a hind tibial length 
(HTL) of 0.14—0.16 mm and 2 females with 
HTL = 0.14—0.15 mm unless otherwise in- 
dicated. Color recorded from dried speci- 
mens wrapped in tissue paper. Both sexes 
with extensive brown suffusion on meta- 
soma and a dull yellow to tan mesosoma. 

Forewing relatively narrow, 0.25 + 0.01 
mm wide (n = 5); width (FWW) 0.52 + 
0.02 length (FWL) (n = 5); setation rela- 
tively sparse, 7-13 (n = 4) setae between 
4th and 5th setal tracks; longest fringe setae 
0.13 + 0.03 (n = 5) FWW, approximately 
1.5—2.1 hind tibial width (HTW). Hind 
wing with O (n = 2) and 3—5 (n = 2) setae 
in anterior and posterior tracks, respective- 
ly, the latter tract occupying 35-36% dis- 
tance from hamuli to wing apex. Scutellum 
with anterior pair of setae 91-95% (n = 2) 
length of posterior pair. 

Male: Flagellum length (FL) 0.16—0.19 
mm, approximately 2.4 longer than scape, 
FL 6.57 + 0.63X greater than flagellum 
width (FW), FL 1.22 + 0.11% greater than 
HTL; flagelliform setae long, tapering to- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ward apex, longest flagelliform setae 3.53 
+ 0.57X greater than FW; without unsock- 
eted setae; basiconic peg sensilla (BPS) rel- 
atively small, slightly inflated, formula 2-2- 
2-1-1-1 (n = 6); terminal placoid sensilla 
(PLS) with apical 0.23—0.34 extending be- 
yond flagellum apex. 

Genital capsule (GC) relatively narrow, 
0.30 + 0.01 as wide as long; apical distance 
(AD) 0.19 + 0.01 entire genital length 
(GL); apical width (AW) 0.68 + 0.04 gen- 
ital width (GW); dorsal aperture length 
(DAL) 0.49 + 0.02 GL; dorsal lamina 
(DLA) arising in anterior half of GC with- 
out approaching GC sides, slender posterior 
DLA extension narrows and sharply tapers 
toward apex occupying O-—0.42 AD (see 
Variation), width at level of intervolsellar 
process (IVP) less than that of aedeagus; 
DLA 1.68 + 0.34 (see Variation) as long as 
wide and 0.47 + 0.04 GL; IVP short, sub- 
triangular, occupying 0.11—0.17 AD; vol- 
sellae (VS) relatively straight, occupying 
0.42—0.61 AD; ventral ridge (VR) narrow 
and abruptly widening anteriorly, occupy- 
ing 0.45—0.58 basal distance (BD); ventral 
processes (VP) not obviously protuberant, 
slightly laterally displaced to VR and po- 
sitioned distinctly anterior to 1VP. Aedeagus 
length (AL) ca. equal to GL, 0.76 + 0.07 
(n = 6) HTL; apodemes occupying ca. 0.58 
AL. 

Female: Antennal funicle with 1 BPS on 
first funicular segment (F1) and 2 on second 
(F2). Ovipositor length (OL) 1.04—1.07 
that of HTL. 

Types.—Holotype 6: JAPAN. Okinawa 
Prefecture: Iriomote Island; 4 November 
1997; E. kuehniella trap host; Y. Narai and 
N. Itagaki. Paratypes: 6 d, 12 including 
allotype 2°, same data as holotype. 

Etymology.—Derived from the Latin 
word cultellus meaning small knife, refer- 
ring to the knife like appearance of the pos- 
terior extension of the DLA, used as a noun 
in apposition. 

Variation.—The posterior extension of 
the DLA usually occupies 0.22—0.42 of the 


VOLUME 107, NUMBER 4 


Figs. 4-6. 
Hindwing. Scale bars = 0.05 pm. 


AD, although in one of the paratypes it fails 
to reach the level of the AD. 

Remarks.—Trichogramma cultellus keys 
to couplet 23’ in Pinto’s (1999) key to the 
North American Trichogramma but is sep- 
arated from 7. parkeri by characters dis- 
cussed in the Diagnosis section. 

DNA sequence.—The 431 bp ITS-2 
DNA sequence has been deposited in the 
NCBI Database with the following acces- 
sion number: AY518693. We compared this 
sequence with sequences of species includ- 
ed in the Parkeri section to which T. cul- 
tellus belongs. A 581 bp 7. pintoi (acces- 
sion AY 182757) and a 556 bp T. bourara- 
chae Pintureau and Babault sequence (ac- 
cession AF043626) was found to be only 
39% and 46% similar to 7. cultellus. 


Trichogramma umerus Jose, Hirose, and 
Honda, new species 
(Figs.7—13) 


Diagnosis.—Trichogramma umerus most 
closely resembles T. pretiosum Riley and T. 
minutum Riley. Females of the latter species 
have a longer ovipositor, measuring 0.99 
and 1.14 the hind tibial length, respec- 


Trichogramma cultellus. 4, Dorsal and ventral view of male genital capsule. 5, Forewing. 6, 


tively, vs. 0.85 for T. umerus. The T. min- 
utum male is most easily differentiated from 
T. umerus in that the latter has longer fla- 
gellar setae, a shorter ventral ridge, and a 
lesser extension of the terminal placoid sen- 
silla on the flagellum. From T. pretiosum, 
T. umerus is most easily distinguished by 
having broader shoulders on the dorsal lam- 
ina. The dorsal lamina width to length ra- 
tios measure 1.38 and 1.67 for 7. wmerus 
and 7. pretiosum, respectively. 

Description.—Quantitative data from 8 
males with a hind tibial length (HTL) of 
0.13—0.16 mm and 3 females with HTL = 
0.14—0.17 mm, unless otherwise indicated. 
Both sexes apparently yellow with brown 
suffusion on mesosoma and an entirely 
brown metasoma. 

Forewing narrow, 0.21 + 0.02 (n = 7) 
mm wide; width (FWW) 0.47 = 0.03 (n = 
7) length (FWL); 9-21 setae between 4th 
and 5th setal tracts (n = 5); longest fringe 
setae 0.19 + 0.02 (n = 7) FWW, ca. 2.38X 
greater than hind tibial width (HTW) (n = 
7). Hind wing with |I—3 and 4-8 setae in 
anterior and posterior tracts, respectively, 
latter tract occupying approximately 52% of 


786 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 7-9. Trichogramma umerus, male. 7, Dorsal view of genital capsule; scale bar = 50 um. 8, Ventral 
view of genital capsule; scale bar = 50 pm. 9, Ventral view of apical distance; scale bar = 20 wm. 


VOLUME 107, NUMBER 4 


ee 


Figs. 10-13. 
Antenna. 13, Hindwing. Scale bars = 0.05 pm. 


le 


distance from hamuli to wing apex. Scutel- 
lum with anterior pair of setae measuring 
86-91% posterior pair (n = 2). 

Male: Flagellum length (FL) 0.15—0.18 
mm, 1.92—2.42x< longer than scape, FL 
6.85 + 0.44 greater than flagellum width 
(FW), FL 1.16 + 0.09 HTL; flagelliform 
setae long, 3.0 + 0.12 greater than FW; 
without unsocketed setae; basiconic peg 
sensilla (BPS) relatively small, slightly in- 
flated, formula 1(2)-2-2-0-1-1, terminal 
placoid sensilla (PLS) with apical 0.09— 
0.16 (n = 6) extending beyond flagellum 
apex. 

Genital capsule (GC) 0.33 + 0.01 as 
wide as long; sides not constricted at level 
of intervolsellar process (IVP); parameres 
(PM) relatively straight, slightly convergent 
at apex; apical distance (AD) measuring 


787 


13 


Trichogramma umerus. 10, Dorsal and ventral view of male genital capsule. 11, Forewing. 12, 


0.26 + 0.01 entire genital length (GL); api- 
cal width (AW) 0.60 + 0.03 genital width 
(GW); dorsal aperture length (DAL) 0.56 = 
0.02 GL; dorsal lamina (DLA) originating 
posterior to middle of GL, moderately 
notched at base with moderately developed 
shoulders, not reaching sides of GC; sides 
of DLA gradually narrowing posteriorly to 
form a sublinguiform posterior extension, 
distinctly acuminate at apex, with width ap- 
proximately equal to that of aedeagus at in- 
tervolsellar process (IVP) level; DLA 1.38 
+ 0.05 as long as wide 0.39 + 0.02 GL, 
occupying 0.42—0.57 AD; volsellae (VS) 
slightly bowed occupying 0.42—0.57 AD; 
IVP elongate, narrowing apically toward 
pointed apex, occupying 0.30—0.43 AD; 


tuberant, positioned immediately basal to 


788 


IVP; VR occupying 0.24—0.34 basal dis- 
tance (BD). Aedeagus length (AL) approx- 
imately equal to GL, attaining 0.77 + 0.05 
HTL; apodemes about 0.49 AL. 

Female: Antenna with | and 2 BPS on 
first funicular segment (Fl) and second fu- 
nicular segment (F2), respectively. Ovipos- 
itor length (OL) 0.82—0.85 HTL. 

Types.—Holotype d6: JAPAN. Okinawa 
Prefecture: Ishigaki Island; 13 March 1997; 
P. xylostella; Y. Narai and N. Kajitani. 
Paratypes: 7 6 and 3 @ including allotype 
2, same data as holotype. 

Etymology.—Derived from the Latin 
word umerus meaning shoulder, referring to 
the distinct shoulders of the DLA present 
in the type series, used as a noun in appo- 
sition. 

Remarks.—Trichogramma umerus keys 
to couplet 74’ in Pinto’s (1999) key to the 
North American Trichogramma which ter- 
minates with 7. pretiosum. These species 
are remarkably similar and even share the 
characteristic of having a poorly sclerotized 
posterior extension of the dorsal lamina. 
However, 7. umerus has a much different 
ITS-2 sequence (see below) and these spe- 
cies can be separated by characters dis- 
cussed in the Diagnosis section. 

DNA sequence.—The 526 bp ITS-2 
DNA sequence for 7. umerus has been de- 
posited in the NCBI Database (accession 
AY518694). We performed pair-wise com- 
parisons with both 7. minutum (NCBI 
#AF408658) and JT. pretiosum (NCBI 
#AF082822) to determine their sequence 
compatibility. The 7. minutum 420 bp se- 
quence was 59% compatible to that of T. 
umerus while 400 bp T. pretiosum sequence 
was 53% compatible. Interestingly, in T. 
minutum and T, pretiosum, there was a high 
degree of compatibility at the very begin- 
ning and middle of the sequence alignment 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


although a few extra tandem repeats were 
found in the 7. umerus sequence. In this 
section there appeared to be higher com- 
patibility in 7. minutum than in T. pretios- 
um as more base pairs were shared. How- 
ever, the final one-third sequence alignment 
could not be matched in either 7. minutum 
or T. pretiosum as there was a large exten- 
sion of ca. 100 base pairs in T. umerus. 


ACKNOWLEDGMENTS 


We thank John Pinto and Gary Platner 
for their suggestions and comments. We 
also thank Fujio Aoyama for providing 771- 
chogramma specimens reared in his labo- 
ratory at the Shimane Agricultural Experi- 
ment Station. Laurel Taylor provided pho- 
tomicrographs. This work was_ partially 
funded through a JSPS fellowship award to 
JYH. 


LITERATURE CITED 


Ishii, T. 1938. On the species of Trichogramma and 
their ecology in Japan. Oyo-Dobutusugaku-Zasshi 
10: 139-141 [in Japanese]. 

. 1941. The species of Trichogramma in Japan 
with descriptions of two new species. Kontyda 14: 
169-176. 

NCBI handbook [Internet]. Bethesda (MD): National 
Library of Medicine (US), National Center for 
Biotechnology Information; 2002 Oct. Available 
from http://www.ncbi.nlm.nih.gov/entrez/query. 
fegi?db = Books 

Pinto, J. D. 1999 [1998]. Systematics of the North 
American species of Trichogramma Westwood 
(Hymenoptera: Trichogrammatidae). Memoirs of 
the Entomological Society of Washington No. 22, 
287 pp. 

Platner, G. R., R. K. Velten, M. Planoutene, and J. D. 
Pinto.1999. Slide-mounting techniques for Tricho- 
gramma (Trichogrammatidae) and other minute 
parasitic Hymenoptera. Entomological News 110: 
56-64. 

Taylor, L., N. Yashiro, Y. Hirose, and J. Y. Honda. In 
press. A new species of Trichogramma (Hyme- 
noptera: Trichogrammatidae) parasitic on Jvera 
auripes (Butler) (Lepidoptera: Lymantriidae) from 
Hokkaido, Japan. Annals of the Entomological 
Society of America. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 789-807 


THE GENUS OMMATIUS WIEDEMANN, DILATIPENNIS SPECIES GROUP 
(DIPTERA: ASILIDAE) 


AUBREY G. SCARBROUGH AND JESSICA E. COSTANTINO 


(AGS) Department of Biological Sciences, Towson University, 8000 York Road, Bal- 
timore, MD 21252 U.S.A. (e-mail: ascarbrough@towson.edu); (JEC) Center for Conser- 
vation Research and Training, University of Hawaii, 3050 Maile Way, Gilmore Hall #408, 
Honolulu, HI 96822 U.S.A. (e-mail: jecostantino@smcm.edu) 


Abstract.—Six species of the dilatipennis species group of Ommatius Wiedemann are 
reported. The species group is known only from the Oriental Region. Its members are 
most similar to the genus Merodontina Enderlein but lack the prominent ventrobasal digit 
on the hind femur of the male. The species group includes O. catus, n. sp., and O. 
insectatus, n. sp., from southeastern Asia, O. forticulus, n. sp., from Sarawak, O. picti- 
pennis Bigot from Malaysia, O. torulosus Becker from Taiwan, and O. dilatipennis Wulp 
from Indonesia. Ommatius dilatipennis Wulp, recently transferred to Merodontina, is re- 
assigned [revised combination] to Ommatius. Lectotypes are designated for O. dilati- 
pennis and O. torulosus. The dilatipennis species group is diagnosed. Illustrations of 


selected characters and a key to species are also included. 


Key Words: 


The Ommatiine fauna of the Oriental Re- 
gion is poorly known with about 125 spe- 
CleSminmsixe Seneras Otethe. lattenespecies: 
most (60%) belong to the genus Ommatius 
Wiedemann. Species from this region have 
been collected largely in the more populat- 
ed regions of China, India, Philippines, Sri 
Lanka, Taiwan, and Indonesia (Oldroyd 
1972, 1975; Tsaras and Artigas 1994; Jo- 
seph and Parui 1998; Scarbrough and Mar- 
ascia 1999; Scarbrough and Hill 2000a, b). 
Furthermore, no attempt has been made to 
group closely related species. The primary 
objective of this study is to characterize the 
dilatipennis species group of Ommiatius, 
which is most similar to Merodontina En- 
derlein, and to assign six species to the 
group. Ommatius pictipennis Bigot and O. 
torulosus (Becker) are redescribed. Omma- 
tius dilatipennis Wulp, transferred recently 
to Merodontina (Scarbrough and_ Hill 


Asilidae, Ommatius, dilatipennis species group, Oriental, new species 


2000a), is returned to Ommatius. Lecto- 
types are designated for O. dilatipennis and 
O. torulosus. The new species are described 
and illustrations of significant characters 
useful for identifying the species are in- 
cluded. A key separating the six species is 
also included. 


METHODS AND MATERIALS 


The descriptive terminology and acro- 
nyms of museums follow McAlpine (1981) 
and Arnett et al. (1993), respectively. Spec- 
imens for this study were obtained via loans 
from curators and/or studied during visits at 
the following museums: Ben Burgge, Zool- 
ogische Museum, Amsterdam (ZMAN); 
Neil Evenhuis, Bernice P. Bishop Museum, 
Honolulu (BPBM); David Grimaldi, Amer- 
ican Museum of Natural History, New York 
(AMNH); Darren Mann, Oxford University 
Museum of Natural History, Oxford, U.K. 


790 


(UMO); Frank Menzel, Deutsches Ento- 
mologisches Institut, Miincheberg, Germa- 
ny (DEI); E. J. van Neiukerken and C. van 
Achterberg, Nationaal Natuurhistorischen 
Museum (‘‘Naturalis’’?), Leiden (RMNH); 
Mark O’Brian, University of Michigan, 
Museum of Zoology, Ann Arbor, (UMMZ); 
Thomas Pape, Museum of Zoology, Uni- 
versity of Copenhagen (ZMUC); Gary 
Steck, Florida State Collection of Arthro- 
pods, Gainesville (FSCA); and F Christian 
Thompson, USDA, Systematic Entomology 
Laboratory, Washington, D.C. (USNM). 
Dissections of the terminalia and prepara- 
tion of illustrations follow Scarbrough and 
Marascia (1999) and Scarbrough (2002b). 
Descriptions of new species are composites 
of all specimens examined. In recording la- 
bel data of specimens, an ~ symbol denotes 
data present on a second label. Square 
brackets are used for additional information 
not present on labels. Only confirmed dis- 
tribution records are included under each 
species covered herein. Structures in illus- 
trations are labeled only for the first species 
and are not repeated for comparable struc- 
tures of the following species. Illustrations 
of the terminalia in dorsal and ventral po- 
sitions are accompanied by a single scale 
line. An asterisk (*) denotes a syntype. 


Ommatius Wiedemann, dilatipennis 
species group 


Diagnosis.—The dilatipennis species 
group is characterized by its similarity to 
Merodontina Enderlein, especially in the 
strong costal dilation being on the apical 
half of the wing and the acutely pointed R, 
cell (Figs. 1-7) in males and the long, slen- 
der body and legs which lack stout ventral 
bristles. It differs from Merodontina in the 
absence of a prominent, ventrobasal, digi- 
tate process on the hind femur of the male. 


KEY TO THE OMMATIUS DILATIPENNIS 
SPECIES GROUP 


1. Femora and tarsi entirely black to brown.... 2 
— Femora and tarsi partly yellow ........... 3 


2. Epandrium slender apically, apex acutely 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


pointed; hypandrium with only black vestiture, 
terminalia otherwise as in Figs. 29-33; Malay- 
SAMY Wad cocks amine Somes ee a pictipennis Bigot 
— Epandrium wide apically, podiform with a 
rounded base ventrally and an pointed toe dor- 
sally; hypandrium broadly rounded apically; 
Indonesia (Java, Sumatra), Philippines, Malay- 
sia (Sarawak, Sabah)........ dilatipennis Wulp 
3. Dorsal postocular bristles short, apex of longest 
just before posterior margin of compound eye; 
scutellar bristles yellow ................ 4 
— Dorsal postocular bristles long, apex of longest 
well before posterior margin of compound eye; 
scutellar bristle brown ................- 5 
4. Large, robust species (28.0 mm); postocular 
bristles yellow; wide angles of tergites 2—5 yel- 
low to reddish yellow; epandrium slender api- 
cally, apex round, terminalia as in Figs. 17-21; 
Malaysia (Sarawak) ......... forticulus, n. sp. 
— Smaller, slender species (13.4—17.8 mm); pos- 
tocular bristles usually brown; tergites 2—5 
brown, margins narrowly yellow; epandrium 
capitate apically, terminalia as in Figs. 34—40: 
ALWWATINIS SYS e hee) SEES CAReS SA Pe torulosus (Becker). 
5. Middle femur mostly yellow; terminalia as in 
Figs. 22-28; Thailand ....... insectatus, Nn. sp. 
— Middle femur mostly black, terminalia as in 
Figs. 8-16; Thailand, Laos, Vietnam 


catus, N. sp. 


TAXA 


Ommatius catus Scarbrough and 
Costantino, new species 
(Figs. 1-2, 8-16) 


Male.—Black body, femora extensively 
black. Body 17.5-18.9 mm; wing 16.0— 
16.4 mm. Head: Face yellow tomentose, 
vestiture abundant, mostly yellow; one or 
two brownish-yellow bristles present; 
FHWR 1.0:8.8. Palpus and proboscis white 
to yellowish setose. Antenna black setose; 
flagellum slightly longer than scape or ped- 
icel, about twice as long as wide. Frons 
dark brown tomentose, mostly yellow se- 
tose. Ocellar tubercle black setose, longest 
seta subequal to basal three antennal seg- 
ments combined; five or six black posto- 
cular bristles, longest with apex about half 
distance toward ocellar tubercle. 

Thorax: Brown tomentose dorsally, nar- 
row sides and posterior brownish yellow to 
yellowish gray; setae short, mostly black; 
three black dorsocentral and four lateral 


VOLUME 107, NUMBER 4 


4 


Figs. 1-7. 


791 


Right wing of species of the dilatipennis species group of Ommatius Wiedemann. 1, O. catus, 


male. 2, O. catus, female. 3, O. dilatipennis, male. 4, O. forticulus, male. 5, Ommatius insectatus, male. 6, O. 
pictipennis, male. 7, O. torulosus, male. Abbrevations for cells: r, = first radial, r, = fourth radial, m, = first 


medial, m, = third medial. 


bristles present, dorsocentral bristles thinner 
and shorter than lateral bristles. Scutellum 
dull yellowish gray tomentose, setae mixed 
yellow and black, mostly yellow; two mar- 
ginal bristles and preapical groove present. 
Pleuron yellow to yellowish-gray tomen- 
tose; setae and most bristles yellowish; thin, 


black anepimeral bristle present. Halter yel- 
low. 

Wing (Fig. 1): Surface brownish, ante- 
rior half darkest. Cell r, triangular, base 
wide, just beyond apex of cell d. Crossvein 
r-m short, well before middle of cell d. Cell 
m, with narrow base, about third as wide as 


192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


iy c ay 


WIL S*0 


12 0.5 mm 


Figs. 8-12. Male terminalia of Ommatius catus. 8, Left epandrium. 9-10, Dorsal and ventral views. 11, 
Gonostylus. 12, Aedeagus, lateral view. Abbrevations: Ce = cercus, Ep = epandrium, Hp = hypandrium, Ge 
= gonocoxite, Gs = gonostylus. 


VOLUME 107, NUMBER 4 


basal third of cell; weak preapical constric- 
tion present. Cell m, with CuA,+M, short, 
slightly shorter than or as long as r-m cross- 
vein; apical vein slightly arched, oblique 
near base of cell m,. 

Legs: Coxae and trochanters black; cox- 
ae gray tomentose with whitish to yellowish 
vestiture, stout bristles absent. Fore femur 
dorsally and mostly anteriorly black, oth- 
erwise yellow, ventral setae long, yellow. 
Middle and hind femora black except ex- 
treme narrow yellow base; most anterior 
bristles black; anteroventral bristles long, 
thin, and black. Hind femur widest medi- 
ally, ventroapical margin narrowed, slightly 
concave; mostly short black bristles present 
posteroventrally; several, long, thin bristles 
present on basal half. Tibiae mostly yellow, 
narrow apex of fore and middle tibiae and 
apical two-thirds of hind tibia black; bris- 
tles thin, mostly black; fore and middle tib- 
iae with yellow bristles laterally; hind tibia 
bent basally. Tarsi mostly black with black 
bristles; basal tarsomere of fore and middle 
tarsi mostly yellow, apex black. 

Abdomen: Black. Apex of each segment 
yellow; tomentum mostly yellowish gray 
with yellow setae; tergites brown tomentose 
dorsally with brown setae; tergites laterally 
with unusually long, abundant setae, setae 
yellow on basal five tergites, brown on api- 
cal three. 

Terminalia (Figs. 8-12): Brown setae 
basally, yellow apically. Epandrium with 
flared, yellow apex. Hypandrium strongly 
produced medially. 

Female.—Differs from male as follows. 
Body 14.1—19.1 mm; wing 13.1—16.0 mm. 
Head: Face with 5—6 brown bristles, some- 
times absent; face 1/9.8 as wide as head. 
Frons yellow setose. Anepimeral bristle 
yellow or brown. Wing (Fig. 2): Normal, 
without strong costal dilation, dark corru- 
gated anterior cells; cell r, wide basally, 
well beyond apex of cell d; mm crossvein 
at or just before middle of cell d; cell m, 
with sides diverging to wing margin. Legs: 
Femora color variable; fore and middle 
femora yellow except dorsum and narrow 


793 


anterodorsal stripe to entirely brown; hind 
femur with basal third yellow to entirely 
brown. Tibiae with bristles entirely brown; 
hind tibia with apical fourth to half brown. 
Basal tarsomere of hind tarsus sometimes 
yellowish brown basally. Abdomen: Ter- 
gites with only sparse thin bristles laterally. 
Terminalia (Figs. 13—16): Tergite 9 short, 
straplike, about one-sixth as long as cercus. 
Sternite 8 with several long stout bristles 
laterally. Genital fork with stout arms, 
mostly sclerotized; base and inner flange of 
sclerotized arms membranous. 

Type specimens.—Holotype d, VIET- 
NAM: Ap Hung-Lam, 21 km. NW of Di 
Linh [11°38’N 108°07’S], 1,100 m, 29.ix- 
5.x.1960, C. M. Yoshimoto, deposited in 
BPBM, Honolulu. Paratypes: VIETNAM: 
1 2, Fyan [11°52’60"N 108°12’0"E], 900— 
12000 hms Lae vii-9 svar Ol NE ReeSpencer 
(BPBM). LAOS: 1 6, Borikhane Prov., 
Pakkading, vi.1965, Native Collector ~ 
BISHOP MUSEUM (BPBM). THAI- 
LAND: 1 @, Chiengmai [= Chiang Mai], 
3,500—4,000 ft., v.3.69 ~ G. R. Ballmer 
(BPBM). 

Other specimens examined.—VIET- 
NAM: 1 @ [genitalia absent], Fyan 
[11°52'60"N_ 108°12'0"E], 900-1,000 m, 
11.vi1.-9.vili."61 ~ N. R. Spencer (BPBM). 
THAILAND: 12, Chiengmai [= Chiang 
Mai] Prov., Doi Suthep, 3,500—4,000 ft., 
3.vi.69 G. R. Ballmer (BPBM). 

Etymology.—Latin catus, adjective for 
‘crafty or skillful’, referring to the preda- 
tory behavior of asilids. 

Distribution.—Laos, Thailand, Vietnam. 

Remarks.—The largely black middle and 
hind femora, wing venation (Figs. 1—2), the 
brown apical two-thirds of the hind tibia, 
and terminalia (Figs. 8-16) distinguish O. 
catus from congeners. The basally curved 
hind tibiae and the presence of a flat, spur- 
like, apical bristle are similar to O. insec- 
tatus but it is distinguished from that spe- 
cies by the combined characters of the ter- 
minalia. 


794 


Ommatius dilatipennis Wulp, 
revised combination 
(Fig. 3) 


Ommatius dilatipennis Wulp, 1872: 261. 
Lectotype ¢, [RMNH]. Type locality: 
Indonesia, Java. Hull 1962: 436; Oldroyd 
1975: 132, catalog. 

Merodontina dilatipennis: Scarbrough and 
Hill 2000a: 93, combination. 


Specimens examined.—INDONESIA: 
Java, 1 6, 1 2, Nederlands Indie, Java 
1,800'—2,400’, TJL[AJOENAN, SOEKA- 
NEGARA, eind okt. 1941, J.M.A.V. Gro- 
enedael (ZMAN); 2 ¢, Nederlands Indie, 
W. Java 1,800’, DJAMPANG TENGAH, 
18, 25.v.1939, J.M.A.V. Groenedael 
(ZMAN); 1 3, same label data except 
x.1941, J.M.A.V. Groenedael (ZMAN); 1 
6, E. Jacobson, Genoeng, Oengaran Java, 
xii.1909, Ommatius dilatipennis, Getekend 
(ZMAN); 4 2, Batoerrden, G. Slamat. Java, 
3, 14.vi.1928, FE C. Drescher (ZMAN); 3 ¢, 
2 @, Batoerrden, G. Slamat. Java, 
29.vii.1928, E C. Drescher (ZMAN); 1 6, 
Java, Jacobson (ZMAN); 1 ¢, Doeters V. 
Leerwon, Geenoeng Oengaran, Mei 1910, 
Ommatius dilatipennis, det. de Meijere 
(ZMAN); 1 3* [good condition, LECTO- 
TYPE], Diard [collector], Java, Ommatius 
dilatipennis Wulp, coll. EM.d.v. Wulp 
[round label] ~ syntype [red label] 
(RMNH); 1 d* [poor condition], Diard 
[collector], Java, Ommatius dilatipennis 
Wulp, coll. EM. Wulp [round label] ~ syn- 
type [red label] (RMNH); 1 ¢* [poor con- 
dition], Muller [collector], Java [round la- 
bel] ~ Ommatius dilatipennis, v. d. Wulp 
~ syntype [red label] (RMNH); | 6 * [poor 
condition], Muller [collector], Java [round 
label] ~ Ommatius dilatipennis, v. d. Wulp 
~ syntype [red label] (RMNH); 1 @° * [good 
condition], Blume [collector], Java [round 
label] ~ Ommatius dilatipennis v. d. Wulp 
~ syntype [red label] (RMNH); 1 @, W. 
Java, Djampang, Tengeh, 300-600 m, 
4.1.1940, coll. N. Groenendael ~ Ommatius 
dilatpennis v. d. W. [Wulp], det. H. Over- 
beck 1967 (RMNH); Sumatra: 1 2, 69/27, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Edw. Jacobson, Suban Ajam, Sum. 7.1916, 
Getekend (ZMAN); 1 2, Muller, Sumatra 
~ Ommatius dilatipennis v.d.w., coll. EM. 
v.d. Wulp (RMNH); 1 2, 69, 26 ~ Edw. 
Jacobson, Buban Ajam, Sum. 7.1916 ~ 
Ommatius dilatipennis v.d. Wulp, det. de 
Meijere (RMNH). MALAYSIA: Sabah, 1 
6, British North Borneo, Tawau [4°17'30"N 
117°54'14”E], Quoin Hill, Cocoa Res. Sta. 
malaise trap, 6.ix.1962, Y. Hirashima 
(BPBM); Sarawak, | 6, 1 2, Nauga Pela- 
gus nr. Kapit 180-585 m, 7—14.vii.’58, T. 
C. Maa (BPBM); 1 F [abdomen missing], 
Muller, Borneo ~ Ommatius dilatipennis 
v.d.w., Coll. EM.v.d. Wulp (RMNH); Sa- 
rawak, 1 6, 1 2, Bau District, Pangkalan 
Tabang, 300—450 m, 7—8.ix.’58, T. C. Maa 
(BPBM). PHILIPPINES: 1 d3, Mindanao, 
Lanao, Butig Mts., 24 km NE of Butig, 
1,080 m, 2,180 m, 21.vi.1958, in jungle, 
along stream, H. E. Milliron (BPBM); 1 2, 
Palawan, Tarumpitao Pt., 3.vi.1958, jungle 
clearing, H. E. Milliron (BPBM); | 2, Pa- 
lawan, Brooke’s Pt., Macagua, 75 m, 
7.iv.1962, M. Thompson (BPBM); | &, Pa- 
lawan, Mouth of Malabangan River, 
28.v.1958, jungle clearing, H. E. Milliron 
(BPBM); 1 &, Mindanao, Surigao, L. 
Mainit, 23.xi-1.xii.1959, C. H. Yoshimoto 
(BPBM); 1 ¢, Mindanao Is., Agusan del 
Norte, Cabadbaran, Belang-Balang Forest 
Road, 1.iv.1963 H. M. Cullamar (UMMZ). 

Remarks.—In addition to the characters 
in the key, the black setose palpus, entirely 
black femora and tarsi, wing venation (Fig. 
3), and combined characters of the termin- 
alia (see Figs. 4-15, Scarbrough and Hill 
2000a), especially the dorsally pointed foot- 
like apex of the epandrium, distinguish it 
from congeners. 

Scarbrough and Hill (2000a) assigned O. 
dilatipennis to the insula species group of 
Merodontina based on the similarity of the 
characters of the wings, legs, and termina- 
lia. Furthermore, the digitate process on the 
hind femur of males from Borneo and Sa- 
rawak was minute with a bristle at its apex. 
Thus, we assumed the digitate process was 
a variable character in that genus. However, 


VOLUME 107, NUMBER 4 


13 


0.5 mm 


15 


Figs. 13-16. 


16, Spermatheca. Abbrevations: Tg, = tergite 9, St, = 


further study of type material from Java and 
new material from Indonesia proved this as- 
sumption incorrect. Here we return O. di- 
latipennis to Ommatius and assign it to the 
dilatipennis species group. 

Box number 39 in RMNH has seven 


Female terminalia of Ommatius catus. 


795 


will €'°0 
wil 2'0 


13, Tergite 9 and circus. 14, Sternite 8. 15, Genital fork. 
sternite 8. 


specimens of O. dilatipennis of which five 
have syntype labels (*). All have some 
parts missing, varying degrees of [insect] 
damage, and their bodies are partly covered 
by fungus. To fix and stabilize the current 
concept of the name, the syntype male in 


796 


the best condition from Java is selected lec- 
totype. 


Ommatius forticulus Scarbrough and 
Costantino, new species 
(Figs. 4, 17—21) 


Male.—Measurements, body 28.0 mm; 
wing 18.7 mm. Brown. Head: Face, frons, 
and vertex yellow to yellowish-gray tomen- 
tose, facial vestiture whitish, bristles pale 
yellow, setae abundant and white; face nar- 
row, about % as wide as head. Frons yellow 
setose. Palpus mostly yellowish setose, 
brown setae dorsally. Proboscis black with 
whitish setae. Antenna mostly yellow se- 
tose, pedicel with brown setae ventrally; 
flagellum long and thin, 2.5 as long as 
wide, about as long as scape and pedicel 
combined. Ocellar tubercle brown setose, 
setae about as long as basal three antennal 
segments. Occiput with large brown tomen- 
tose spot dorsally, tomentum otherwise yel- 
lowish gray to gray, setae whitish, that dor- 
sally slightly pale yellowish; postocular 
bristles mostly yellow, proclinate with lon- 
gest about half distance toward ocellar tu- 
bercle. 

Thorax: Black. Mesonotum mostly 
brown tomentose dorsally, postpronotal 
spot, sides and prescutellum narrowly yel- 
low; vestiture sparse, mostly brown; short, 
scattered brown setae medially, sparse yel- 
lowish setae present laterally; bristles 
mixed brown and yellow, three lateral and 
two dorsocentral bristles yellow; three thin, 
short dorsocentral bristles present. Scutel- 
lum with yellowish-brown tomentum and 
three yellow marginal bristles; setae yellow. 
Pleuron yellow tomentose with sparse yel- 
low setae and bristles; anepimeral bristle 
present. Halter dull yellow, club dull 
brown. 

Wing (Fig. 4): Cell R, immediately be- 
hind costal dilation brown, apical half of 
wing only slightly brownish with brown 
microtrichia. Crossvein r-m well before 
middle of cell d. Base of m1 about half as 
wide as that at basal third, weak preapical 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


constriction present. Apex of cell m, just 
before apex of cell d. 

Legs: Coxae, trochanters, and femora 
blackish; coxae yellowish tomentose with 
mostly whitish to yellowish vestiture; fem- 
ora with mostly yellow vestiture, two or 
three bristles brown, bristles mostly long 
and thin ventrally. Hind femur ventrally 
without long digitate process basally. Fore 
and middle tibiae with mostly yellow bris- 
tles laterally. Tibiae yellow, narrow apex of 
middle tibia and apical third of hind tibia 
brownish to brown; setae yellow, mostly 
short, fore tibia with abundant, long, yellow 
setae and yellow bristles, latter sparse, bris- 
tles usually brown. Basal tarsomere of fore 
and middle tarsi mostly yellow; bristles 
blackish. 

Abdomen: Mostly black, wide triangles 
of tergites 2—6 laterally, apical margins of 
all tergites and apical margins of sternites 
1—4 brownish yellow to red; tomentum 
thin, mostly yellow with yellow vestiture; 
tergites with brown tomentum spot medi- 
ally. Tergites 6—8 with brown setae dorsal- 
ly; long yellow setae present on basal four 
segments, abundant on tergites laterally, 
scattered, sparse on sternites. 

Terminalia (Figs. 17-21): Epandrium 
brown, exceptionally narrow apically, 
slightly curved behind genital cavity, apex 
round. Gonostylus slender, slightly curved. 
Apical corners of gonocoxite stout. Hypan- 
drium narrowed, strongly produced apical- 
ly, apical margin with abundant yellow ves- 
titure apically. 

Female.—Unknown. 

Type material—Holotype 6, MALAY- 
SIA: Sarawak, N. W. Borneo, Kuching 
[1°32’N 110°20’E], coll. 18.v.03, Pres. 1908 
by the Sarawak Museum ~ Kuching 
DISAIN WNOrAO a, Wea, WOR =~. TOS. 
1447, deposited in UMO. 

Etymology.—Latin forticulus, an adjec- 
tive for ‘quite bold, rather brave’, referring 
to the flies hunting habits. 

Remarks.—Ommatius forticulus is easily 
recognized from congeners by the charac- 
ters in the key and the combined characters 


VOLUME 107, NUMBER 4 797 


21 (0.5mm 


Figs. 17-21. Male terminalia of Ommatius forticulus. 17, Lateral view. 18, Gonostylus. 19—20, Dorsal and 
ventral views. 21, Aedeagus, lateral view. 


798 


of the terminalia (Figs. 17—21). It is most 
similar to O. pictipennis in size, wing ve- 
nation, the entirely dark femora, and unusu- 
ally slender epandrium. The yellow basal 
tarsomere of the tarsi, the yellow setae of 
the hypandrium, and remaining characters 
of the terminalia distinguish it from O. pic- 
tipennis. 


Ommatius insectatus Scarbrough and 
Costantino, new species 
(Figs. 5, 22-28) 


Male.—Brown. Measurements, body 
19.0—20.0 mm, wing 14.2—16.0 mm. Head: 
Face, frons, and vertex yellow tomentose 
with yellow vestiture; face narrow, 1/8.9— 
1/9.6 as wide as head. Palpus and proboscis 
yellow to yellowish setose. Antenna and 
ocellar tubercle brown setose. Occiput with 
large brown tomentose spot dorsally, eye 
margined narrowly with yellowish gray to- 
mentum and yellowish setae, white ventral- 
ly with white setae. 

Thorax: Mesonotum mostly brown to- 
mentose dorsally, sides narrowly yellow, 
prescutellum with large yellowish-brown 
spot on each side of dorsocentrals; vestiture 
sparse, mostly brown; short, scattered 
brown setae medially, sparse yellowish se- 
tae present laterally; bristles brown, long 
and thin, only two dorsocentral bristles pre- 
sent posteriorly. Scutellum with yellowish- 
brown tomentum and two brown marginal 
bristles. Pleuron yellowish tomentose ante- 
riorly, white posteriorly and ventrally; setae 
and bristles sparse, yellowish to white; ane- 
pimeral bristle present. Halter pale yellow- 
ish white. 

Wing (Fig. 5): Anterior cells yellow to 
brownish yellow, r, darkest with strongest 
corrugations; apical half dense brown mi- 
crotrichose. Cell r, triangular, base wider 
than O. auricular, apex more horizontal, 
only slightly angled posteriorly at wing 
margin. Crossvein well before middle of 
cell d. Cell m, narrow basally, about 7 as 
wide as basal third. Cell m, strongly arched 
dorsally, CuA,+M, about as long as r-m. 

Legs: Coxae dark brown, whitish tomen- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


tose with whitish vestiture. Fore and middle 
femora mostly yellow, apex brown; fore fe- 
mur with narrow pale yellowish brown to 
brown streak on apical third to half anteri- 
orly; middle femur yellowish-brown to 
brown anteriorly, darkest dorsally. Hind fe- 
mur mostly brown, base yellow, transition 
from yellow to brown abrupt; ventrobasal 
digitate tubercle absent. Femoral setae 
mostly brown, generally thin and short; an- 
terior bristles mostly brown, stout and long; 
anteroventral bristles of middle femur thin, 
short, posteroventral bristles thicker, longer, 
and yellow; hind femur with mostly long, 
thin, brown bristles, one anteriorly and 
three or four on basal third pale yellow, 
posteroventral bristles short, thick, and nu- 
merous. Fore and middle tibiae with mostly 
yellow bristles laterally. Hind tibia with 
basal third slightly curved, apical half 
brown. Basal tarsomere of fore and middle 
tarsi mostly yellow; all tarsi ortherwise 
brown with blackish bristles. 

Abdomen: Brown, narrow apical margin 
of tergites 2—3 brownish yellow. Tomentum 
thin, mostly brown, gray to brownish-gray 
basally. Tergites 3-8 and sternites 6—8 
mostly to entirely brown setose; tergites 1— 
2, tergites 3-6 and sternites 1-6 yellowish 
to yellow setose. Apical corner of tergite 4 
with several, long, thin yellowish bristles, 
length as long as segment 4. 

Terminalia (Figs. 22—26): Cercus with 
narrow apex, apical tuft of yellow setae pre- 
sent ventrally. Epandrium strongly nar- 
rowed preapically, apex much wider, trun- 
cate and yellow; low flange present along 
dorsal margin, extending onto inner sur- 
face. Hypandrium produced medially, api- 
cal margin truncate with abundant long yel- 
low setae. 

Female.—Measurements, 16.6—19.0 mm, 
wing 15.2-15.4 mm. Head: Face 1/8.0—1/ 
9.3 as wide as head. Flagellum about as 
long as scape and pedicel combined. Tho- 
rax: Mesonotum with postpronotal spot, 
sides, and posteriorly brownish yellow to- 
mentose; three dorsocentral bristles present. 
Wing: Similar to catus (see Fig. 2). Brown- 


VOLUME 107, NUMBER 4 799 


wil G*0 


24 25 


26 0.5 mm 


Figs. 22-26. Male terminalia of Ommatius insectatus. 22, Lateral view. 23, Gonostylus. 24—25, Dorsal and 
ventral views. 26, Aedeagus, lateral view. 


800 


Ulu €°0 


27 


Figs. 27-28. 


ish, darkest anteriorly, dilation absent; 
crossvein r-m slightly before middle of cell 
d, and slightly longer than CuA,+M,,; cells 
r, apically and r, basally normal; cell m, 
narrow basally, gradually becoming wider 
apically, base about one-third as wide as 
cell at basal third. Fore and middle femora 
slightly brownish-yellow, apex and apical 
third anteriorly with narrow streak brown. 
Hind femur with basal third yellow; five or 
six long yellow posteroventral bristles pre- 
sent on basal half, short, stout brown bris- 
tles present only on apical third. Abdomen: 
Narrow apical margin of segments 1—4 yel- 
low, setae short, mostly brown dorsally, 
basal four or five tergites with sparse yel- 
low setae laterally; all tergites with at least 
brown setae medially, tergites 5-8 mostly 
to entirely brown setose; most sternites with 
only short, sparse yellow setae; sternites 6— 
8 brown setose; apical three tergites sparse- 
ly tomentose, blue reflections present. Ter- 
minalia (Figs. 27-28): Tergite 9 narrow, 
Straplike; sternite 8 simple, several long 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


wil 20 


28 


Female terminalia of Ommatius insectatus. 27, Genital fork. 28, Spermatheca. 


brown bristles present laterally (see Figs. 
13-14). Narrow with sclerotized arms, arms 
connected apically by membranous connec- 
tion; basally mostly membranous. Sperma- 
theca normal, basal bulb symmetrical. 
Type specimens.—Holotype ¢, THAI- 
LAND: Chieng [= Chiang] Mai Province, 
Doi Suthep N. P.: Konthathan, 6—700 m 
30.1x.1981, Zool. Museum Copenhagen leg, 
deposited in ZMUC. Paratypes: THAI- 
LAND: 2 2 Doi Suthep—Pui Natn. Park, 
Konthathan, Waterfalls area, 600 m, 20— 
27.x.1979, Zool. Museum Copenhagen leg. 
(ZMUC); 1 od, same data as holotype 
(ZMUC). THAILAND: 1 &, Chieng [= 
Chiang] Mai Province, Doi Inthanon N. P.: 
Huai Sai, Luang 10—1,100 m, 14.x.1981, 
Zool. Museum Copenhagen leg. (USNM); 
THAILAND: 1 6 Chieng [= Chiang] Mai 
Province, Doi Inthanon N. P.: Siripum 12— 
1,300 m 5.x.1981, Zool. Museum Copen- 
hagen leg. (ZMUC). THAILAND: 1 6, 
Chieng [= Chiang] Mai Province, Doi 


VOLUME 107, NUMBER 4 


Suthep 1,100 m 2.x.1981, Zool. Museum 
Copenhagen leg (USNM). 

Etymology.—Latin insectatus, ‘to attack 
or to pursue’, refers to the predaceous be- 
havior of the fly. 

Remarks.—Ommatius insectatus is most 
similar to O. catus but differs significantly 
in the combined characters of the terminalia 
(Figs. 22—28). Additionally, the mostly yel- 
low fore and middle femora, the brown api- 
cal half of the hind tibia and the sparse, 
long, thin yellow bristles on the apical cor- 
ner of abdominal tergite 4 further distin- 
guish it from O. catus. 


Ommatius pictipennis Bigot 
(Figs. 6, 29-33) 


Ommatius pictipennis Bigot, 1875: 246. 
Holotype 6d, type locality: Malaysia, 
Poulo-Pinang (UMO); Hull 1962: 436; 
Oldroyd 1975: 135. 


Redescription,—Male: Black. Body 
13.9—-24.7 mm; wing 10.0—16.2 mm. Head: 
Face and frons yellow tomentose. Facial 
vestiture mostly to entirely yellow, one 
brown bristle sometimes present; setae 
sparse dorsally, more abundant, longer ven- 
trally; bristles thin, only slightly thicker 
than other vestiture; face 1/7.8—1/9.4 as 
wide as head. Palpus mostly brown setose, 
yellow ventrobasally. Proboscis black, yel- 
lowish setae. Antenna, frons, and ocellar tu- 
bercle brown setose. Antenna brown; fla- 
gellum with narrow base yellow, 2.5X as 
long as wide, longer than either scape or 
pedicel, subequal in length to the two com- 
bined. Ocellar tubercle with longest seta as 
long as scape, pedicel and flagellum com- 
bined. Occiput mostly yellowish-gray to 
white tomentose with dense white setae, 
large brown tomentose spot and scattered 
brown setae present dorsally; 10—12 long, 
thin, brown postocular bristles present, lon- 
gest with apex just before or above ocellar 
tubercle. 

Thorax: Black. Mesonotum brown to- 
mentose dorsally, narrowly brownish-yel- 
low to yellow tomentose laterally and pos- 


801 


teriorly; setae and bristles sparse and black, 
yellow setae present laterally; bristles thin 
and short, four dorsocentral and seven to 
eight lateral bristles present. Scutellum 
brownish-yellow to yellow tomentose, 
mostly or entirely yellow setose, four or 
five thin marginal bristles present, two of 
these slightly thicker than others; preapical 
groove present. Pleuron yellow tomentose 
anteriorly, light yellowish gray posteriorly; 
vestiture sparse, mostly yellow, 10—12 
brown katatergal bristles and yellowish ane- 
pimeral bristle present. Halter stalk dull yel- 
low, knob reddish. 

Wing (Fig. 6): Apical margin strongly 
dilated anteriorly; brown on apical half, 
darkest with corrigations in cells immedi- 
ately behind dilation; dense microtrichia 
present beyond crossvein r-m, mostly bare 
basally. Cell r, triangular, unusually narrow 
basally, sides divergent on basal half, wid- 
est beyond with vein r, slightly angled pos- 
teriorly; base well beyond apex of cell d. 
Crossvein r-m short, just beyond basal third 
of d cell, about twice as long as CuA,+M,. 
Cell m, wide basally, weak preapical con- 
striction present. Apex of cell M, just be- 
fore base of m,. 

Legs: Coxae, trochanters, and femora 
black. Coxae yellowish tomentose with 
abundant yellowish to whitish vestiture. 
Femora with short, black setae anteriorly 
and dorsally, abundant, long, yellow setae 
posteriorly and ventrally; anterior bristles 
mostly black except one on hind femur ba- 
sally; ventral bristles thin and long, stoutest 
bristles basally. Hind femur with anteroy- 
entral bristles black, posteroventral bristles 
yellowish. Tibiae yellow with apex of each 
brown, apical half to third of hind tibia 
brown. Fore tibia with fringe of long yellow 
setae and four yellow bristles laterally; mid- 
dle tibiae with fringe of shorter setae, lateral 
bristles brown. Tarsi black with black bris- 
tles:; basal tarsomere of fore tarsus with 
fringe of long yellow setae. 

Abdomen: Black with narrow apical 
margin of most segments dull red to brown- 
ish yellow; tomentum mostly brown, dense 


802 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


33 90.5 mm 


Figs. 29-33. Male terminalia of Ommatius pictipennis. 29, Lateral view. 30, Gonostylus. 31—32, Dorsal and 
ventral views. 33, Aedeagus, lateral view. 


VOLUME 107, NUMBER 4 


and slightly grayish laterally on basal three 
or four tergites and sternites, thin on apical 
four or five tergites color with slight bluish 
reflections; tergites mostly black setose, 
long, yellowish to white setae present on 
tergites 1—4 laterally and basal 5 sternites; 
sternites 6—8 with long, erect, black setae. 

Terminalia (Figs. 29-33): Epandrium 
narrow and hooked on apical half, apex 
clawlike. Gonostylus (Fig. 30). Gonocoxite 
with enlarged, birdlike process laterally and 
median spinelike process medially. Hypan- 
drium strongly enlarged in lateral view, 
with abundant black setae and bristles. Ae- 
deagus narrow apically; pair of short, claw- 
like ventral processes present. 

Female: Unknown. 

Specimens examined.—MALAYSIA: 
Holotype d (UMO); 1 d, [Kuala Selangor 
3°23'22"N 101°17'06"E] Selangor Templar 
Park, 1.1x.1961, L. W. Quate (BPBM). 

Remarks.—Ommatius pictipennis is rec- 
ognized from congeners by the combined 
characters of the male terminalia (Figs. 29— 
33) especially the long, narrow, clawlike 
epandrium, shape of the gonostylus, lateral 
birdlike process of the gonocoxite, and 
abundant black vestiture of the hypandrium. 

The holotype male is in fair condition, 
missing only the left wing, right leg, and 
hind left tarsus. The body is partially cov- 
ered with fungus and debris, especially the 
terminalia. The data labels are as follows: 
1) circular label with red border and the 
word ‘holotype’; 2) white label with Om- 
matius pictipennis J. Bigot [in script], and 
the number ‘783 in Bigot Coll.’; and the 
UMO type label, Type Dip 289, Ommatius 
pictipennis Bigot, Hope Dept Oxford. The 
species was described from a single male 
specimen from Poulo-Pinang, Malaysia. 


Ommatius torulosus (Becker) 
(Figs. 7, 34—40) 


Ommatinus torulosus Becker 1925: 124. 
Lectotype 6, DEI. Type localities: Tai- 
wan: Kankau. 

Ommatius torulosus: Hull 1962: 436; Old- 
royd 1975: 135, catalog. 


803 


Redescription.—Male: Measurements, 
body 13.4—17.8 mm, wing 12.0—13.8 mm. 
Head: Brown, yellowish-gray tomentose 
with whitish to yellowish setae. Face and 
frons white setose, two to four brown facial 
bristles present. Palpus and proboscis yel- 
lowish to whitish setose. Antenna brown, 
brown setose; flagellum twice as long as 
wide, longer that either scape or pedicel, 
about two-thirds as long as combined 
length of scape and pedicel. Ocellar tuber- 
cle brown setose, longest setae as long as 
scape and pedicel. Occiput with large 
brown tomentose spot dorsally, yellowish 
laterally with yellow setae and gray ven- 
trally with white setae; post ocular bristles 
thin, mostly yellowish, longest with apex 
just beyond margin of eye. 

Thorax: Mesonotum dark brown, posta- 
lar callus and laterotergite dull yellowish 
brown; brown tomentose dorsally, narrow 
sides yellow, postalar corners and prescu- 
tellum brownish yellow; setae sparse, short, 
mostly brown, yellow setae laterally; three 
lateral and three dorsocentral bristles yel- 
low, anterior notopleural bristle brown. 
Scutellum brownish-yellow tomentose, two 
marginal bristles and scattered setae, yel- 
low; preapical groove present. Pleuron yel- 
lowish tomentose, sparse yellow setae and 
bristles present; anepimeral bristle setalike. 
Halter brownish yellow, stalk dull, some- 
what creamy. 

Wing (Fig. 7): Brownish on anterior api- 
cal half, cell r, darkest with most prominent 
corrugations behind dilation; dilation well 
beyond middle of wing; brown microtrichia 
most dense from r-m crossvein to apex of 
wing, sparse basally. Cell r, acutely point- 
ed, r, triangular with narrow base, sides 
abruptly diverge, wide apically. Crossvein 
rm well before middle of cell d, longer 
than CuA,+M,. Cell m, narrow basally, 
about third as wide as that at basal third, 
narrowly constricted preapically. Cell m, 
wide, apex angular; m-m long with cell dor- 
sally well before base of m). 

Legs: Coxae and trochanters brown, for- 
mer grayish tomentose with whitish setae 


804 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


—) 
N 
3 
3 


ww S*0 


37 


38 | 0.5 mm 


Figs. 34-38. Male terminalia of Ommatius torulosus. 34, Lateral view. 35, Aedeagus. 36—37, Dorsal and 


ventral views. 38, Aedeagus. 


and yellowish bristles. Fore and middle 
femora mostly yellow, slightly brownish 
yellow anteriorly, darkest dorsally and api- 
cally, mostly short brown setae present, 


much longer yellow setae ventrally, bristles 
mostly yellow and thin, one or two brown 
bristles present anteriorly. Hind femur 
brown, narrow base yellow, mostly long, 


VOLUME 107, NUMBER 4 


thin, yellow bristles present ventrally. Tib- 
iae mostly yellow, narrow apices of anterior 
two and apical third to half of posterior tib- 
ia brown; bristles mostly brown, two or 
three yellow bristles present on fore tibia 
laterally. Basal tarsomere of fore and mid- 
dle tibiae mostly yellow; remaining tarso- 
meres of all tarsi brown with brown bris- 
tles. 

Abdomen: Blackish, narrow apical mar- 
gin of segments light to dull brownish yel- 
low; tomentum sparse, mostly brown with 
brown setae; grayish to brownish-gray to- 
mentum on basal segments; long yellowish 
setae most abundant on tergites 1—3 or 4 
laterally, scattered and less abundant on 
sternites 1—4, generally sparse; sternites 6— 
8 entirely brown setose, mixed yellow and 
brown on sternite 5. 

Terminalia (Figs. 34—38): Brown, brown 
setose. Apex of epandrium capitate, rect- 
angular, about 3X higher than long, brown- 
ish yellow. Hypandrium broad apically with 
rounded apical corners, row of six Icng, 
slender, yellowish bristles present. 

Female: Differs from male as follows. 
Measurements, body 13.8—18.6 mm; wing 
12.0-14.6 mm. Head: Antenna brown se- 
tose; ocellar setae about as long as scape 
and pedicel combined. Thorax: Yellow. 
Wing: Costal margin straight, without a di- 
lation; apex of cell r, wide, not acutely 
pointed apically. Legs: More yellow than 
male, basal third of hind femur yellow. Tar- 
sal bristles entirely brown; hind tarsus with 
basal half of metatarsomere yellowish 
brown. Abdomen: Yellowish to yellowish 
brown tomentose with mostly yellow ves- 
titure; apical segments and narrow middle 
of basal tergites with brown vestiture. Ter- 
minalia (Figs. 39—40): Tergite 9 straplike. 
Sternite 9 simple, brown vestiture, abundant 
long thin bristles laterally. Genital fork with 
stout, brown arms, fork basaily membra- 
nous. Spermatheca carrot-shaped, base 
slightly bulbus, sides gradually narrowed 
apically, apex narrow, not acutely pointed, 
duct attached apically. 

Specimens examined.—TAIWAN: 1 d* 


805 


[abdomen absent], Fuhosho, Formosa _ H. 
Sauter O09, torulosus Becker [red label 
with word “Typus’] (AMNH); 1 2, Toa Tsui 
Kutsu (Formosa), H. Sauter v. 1914, C.E 
Baker Coll. 1927 (AMNH); 1 3, 1 2, TAL 
WAN: Taipei Co. [24°56’13"N 121°29' 
50"E], Maiyueyuan, 900 m, 16—17.v.1989, J. 
Heppner and H. Wang (FSCA); 1 6, TAI- 
WAN: Taichung Co., Chingshan [25°08'39"N 
121°43'18’E], 1,100 m, 8—11.v.1989, J. He- 
ppner and H. Wang (FSCA); 1 6, Toa Tsui 
Kutsu (Form.), H. Sauter v.1914 ~ torulosus 
~ C.E Baker collection 1927 (USNM); 1 °, 
TAIWAN: NanTowCo, SunMoonLake, 760 
m, June 20—25 1980, D.R. Davis, Forest 
(USNM); 1 ¢*, Fuhosho, Formosa, H. Sauter 
09 ~ torulosus Becker ~ [red label] Typus 
(USNM); 2 2, Toa Tsui Kutsu (Form.), H. 
Sauter v.1914 ~ CF Baker coll. (USNM); 1 
2, Kanshizei, Formosa, Sauter v.08 (DEI); 2 
9, Formosa, Hoozan [= Hozan 38°50’56"N 
125°75’03”E] 08-10, Sauter (DED; 1 ¢ [ab- 
dominal segments 4-terminalia absent] Ko- 
sempo, Formosa, Sauter v.12 (DED; 1 ¢ Ko- 
sempo, Formosa, Sauter v.12 (DEI); 2 d Toa 
Tsui Kutsu (Form.), H. Sauter v.1914 
(DED; 1 3, Toa Tsui Kutsu, Formosa, H. 
Sauter v.1914 (DED; | 6, Tappani, Formosa, 
H. Sauter (DEI); 2 6*, 12 *[abdomen ab- 
sent], Koshun [= Heng-Chun 22°2’15"N 
120°49'24”E], Formosa, Sauter 111.07 09 
(DEI); 1 d3*, Fuhosho, Formosa, H. Sau- 
ten 09 (DED Al 8G S38 Rankauwi(CKo- 
shun [= Heng-Chun 22°2'15’N_ 120°49' 
24"E]), Formosa, H. Sauter 1912 (DET); 1 d* 
[lectotype], Kankau (Formosa), H. Sauter 
vi.1912 (DEI). 

Remarks.—Ommatius torulosus is distin- 
guished by the characters in the key, the 
small, slender body, yellow vestiture of the 
mesonotum, and characters of the termina- 
lia (Figs. 34—40). 

Becker (1925) listed 25 males and 21 fe- 
males in the syntype series collected from 
Koshun, Kankau, Fuhosho, Tou-Tsui Kutsu, 
and dates of 1909 and 1912. Eleven males 
and five females present in DEI have red 
syntype labels. However, three of these fe- 
males [Formosa, Hoozan O8—10, Sauter or 


806 


39 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


wil €°0 
wil 20 


40 


Figs. 39-40. Female terminalia of Ommatius torulosus. 39, Genital fork. 40. Spermatheca. 


Kanshizei, Formosa, Sauter v.08] and seven 
males [Kosempo, Formosa, Sauter v.12, 
Toa Tsui Kutsu (Formosa) H. Sauter v. 
1914, Tappani?, Formosa, H. Sauter] have 
collecting locations and dates differing 
from that reported by Becker. Additionally, 
one male and three females in the AMNH 
and USNM have similar labels, often with 
red syntype labels [Toa Tsui Kutsu (For- 
mosa) H. Sauter v. 1914 C.E Baker Coll.]. 
Only the 8 specimens[*] in the AMNH, 
DEI, and USNM with label data that cor- 
respond to that reported by Becker are here 
considered a part of the original syntype se- 
ries. To fix and stabilize the current concept 
of the name, the syntype male with the label 
“Kankau (Formosa), H. Sauter vi.1912 
(DET) is selected lectotype. 


ACKNOWLEDGMENTS 


For the loan of specimens, we are in- 
debted to the curators listed in the methods 


section. We also thank Darren Mann and 
Frank Menzel for loans of the types of Om- 
matius pictipennis Bigot and O. torulosus 
(Becker), respectively. AGS thanks Darren 
Mann, David Grimaldi, EK Christian Thomp- 
son, E. J. van Neiukerken, C. van Achter- 
berg, Ben Burgge, and Gary Steck for hos- 
pitality and assistance during his visits to 
their institutions. JC acknowledges support 
from NSF-REU grant BBI 0097478. We 
also thank the anonymous referees for their 
helpful comments. 


LITERATURE CITED 


Arnett, R. H. Jr., G. A. Samuelson, and G. M. Nishida. 
1993. The Insect and Spider Collections of the 
World. 2™ Edition. E. J. Brill, Gainesville, Florida, 
220 pp. 

Becker, T. 1925. H. Sauter’s Formosa-Aushaute: Asi- 
linae III. (Diptera) Entomologische Mitteilungen 
14: 62-85, 123-139, 240-250. 

Bigot. 1875. Diptéres nouveaux ou peu connus. 4° par- 


VOLUME 107, NUMBER 4 


tie (1). V. Asilides exotique nouveaux. Annales de 
la Société Entomologique de France (5)5: 237— 
248. 

Hull, E M. 1962. Robber flies of the world. The genera 
of the family Asilidae. United States National Mu- 
seum Bulletin No. 224, 907 pp. 

Joseph, A. N. T. and P. Parui. 1998. The fauna of India 
and adjacent countries, Diptera (Asilidae). Part I. 
Zoological Survey of India, Calcutta, 278 pp. 

McAlpine, J. EF 1981. Morphology and terminology— 
Adults, pp. 9-63. In McAlpine, J. E, B. V. Peter- 
son, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, 
and D. M. Wood, eds. Manual of Nearctic Diptera. 
Vol. 1. Agriculture Canada. Monograph 27, 674 


Oldroyd, H. 1972. Robber flies (Diptera: Asilidae) of 
the Philippine Islands. Pacific Insects 14: 201— 
B3i- 

. 1975. Family Asilidae, pp. 99-156. Jn Delfin- 

do, M. D., and D. E. Hardy, eds. A Catalog of 


807 


Diptera of the Oriental Region. Volume 2. Uni- 
versity of Hawaii Press, Honolulu. 

Scarbrough, A. G. and H. N. Hill. 2000a. The Jnsula 
species group of Merodontina Enderlein (Diptera: 
Asilidae). Studia dipterologica 7: 93-108. 

Scarbrough, A. G. and H. N. Hill. 2000b. Ommatiine 
robber flies (Diptea: Asilidae) from Sri Lanka. 
Oriental Insects 34: 341—407. 

Scarbrough, A. G. and C. G. Marascia. 1999. Synopsis 
of the Oriental and Australian species of Emphy- 
somera Schiner (Diptera: Asilidae). Deutsche En- 
tomologische Zeitschrift 46:203—229. 

Tsacas, L. & J. Artigas. 1994. Le Genere Cophinopho- 
ra Hull, 1958 (Diptera: Asilidae), A repartition 
subcosmopolite inhabituelle. Annales de la Socié- 
té Entomoloque de France (Nouvelle Série) 30: 
447-479. 

Wulp, E M. van der. 1872. Bijdrage tot de Kennis der 
Asiliden van den Oost-Indischen Archipel. 
Tijdschrift voor Entomologie 15: 129-279. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 808-811 


ADDITIONAL HISTORIC RECORDS OF CICINDELA DORSALIS SAY AND 
CICINDELA PURITANA HORN (COLEOPTERA: CARABIDAE: CICINDELIND) 
FROM THE CHESAPEAKE BAY REGION, USA 


JONATHAN R. MAWDSLEY 


Department of Entomology, National Musuem of Natural History, Smithsonian Insti- 
tution, PO. Box 37012, MRC 187, Washington, DC 20013-7012 (e-mail: jonathan. 
mawdsley @nfwtf.org) 


Abstract.—Previously unpublished historic records are presented for two tiger beetle 
taxa of conservation concern from the Chesapeake Bay region, Cicindela dorsalis Say 
and C. puritana Horn. Cicindela dorsalis dorsalis was encountered regularly at Chesa- 
peake Bay sites by collectors during the 1940s and 1950s, when other populations of this 
subspecies were experiencing a severe decline. The historic presence of C. dorsalis media 
LeConte in the southern Chesapeake Bay is documented by specimens collected in 1882. 
Cicindela puritana has been collected regularly at sites in Calvert County, Maryland, 
since 1911, sometimes in large numbers. Both C. d. dorsalis and C. puritana are currently 


listed as ‘“‘Threatened”’ under the U.S. Endangered Species Act. 


Key Words: 
tion, endangered species 


The shores of the Chesapeake Bay are 
home to an unusual tiger beetle fauna 
(Boyd 1975, Glaser 1976) which includes 
two taxa currently listed as ““Threatened”’ 
under the U.S. Endangered Species Act 
(U.S. Fish and Wildlife Service 1990). My 
recent curatorial work in the collections of 
the Smithsonian Institution’s National Mu- 
seum of Natural History (NMNH) resulted 
in the discovery of additional historic spec- 
imens of both of these species which were 
collected at sites along the Chesapeake Bay. 

Specimens from museum collections 
have already provided information that is 
essential to the development of conserva- 
tion strategies for these tiger beetles. Be- 
sides documenting historic populations and 
population declines (Knisley et al. 1987; 
Hill and Knisley 1993, 1994), museum 
specimens have also provided insights into 
historic patterns of genetic variation in both 


Cicindela dorsalis, Cicindela puritana, tiger beetle, conservation, distribu- 


species (Vogler and DeSalle 1993; Vogler 
et al. 1993a, b). Consequently, I felt that a 
short paper summarizing these additional 
historic records would be appropriate and 
relevant to current conservation efforts for 
these beetles. 


Cicindela dorsalis Say 


Two subspecies of C. dorsalis occur on 
beaches along the Maryland and Virginia 
shores of the Chesapeake Bay, C. d. dor- 
salis Say and C. d. media LeConte (Knisley 
and Schultz 1997). The specimens listed be- 
low represent additional historic records of 
both subspecies from the Chesapeake Bay 
region. 


Cicindela dorsalis dorsalis Say 


This subspecies is currently listed as 
“Threatened”’ under the U.S. Endangered 
Species Act (U.S. Fish and Wildlife Service 


VOLUME 107, NUMBER 4 


1990). Once found in vast numbers on 
sandy beaches along the northeast coast of 
the United States, C. d. dorsalis experi- 
enced significant declines during the mid- 
twentieth century and is no longer found at 
most of its historic collecting localities (Sta- 
matov 1972, Knisley et al. 1987, Hill and 
Knisley 1994). 

Although C. d. dorsalis has disappeared 
from much of its former range, populations 
along the Maryland and Virginia shores of 
the Chesapeake Bay appear to be relatively 
robust (Hill and Knisley 1994). Indeed, sur- 
veys over the past thirty years have re- 
vealed many more populations of this spe- 
cies in the Chesapeake Bay region than had 
previously been known (Boyd and Rust 
1982, Knisley et al. 1987, Hill and Knisley 
1994, Knisley, personal communication). 

Cicindela d. dorsalis was first recorded 
from the Chesapeake Bay region by Boyd 
(1975), who reported recent collections 
from several localities in Calvert County, 
Maryland: Calvert Beach, at the base of 
Calvert Cliffs, Cliffs of Calvert, Cove 
Point, and Matoaka Beach. Glaser (1976, 
1984) reported this species from two addi- 
tional Calvert County localities, Camp Roo- 
sevelt and Flag Ponds. Boyd and Rust 
(1982) noted these localities and also men- 
tioned three older specimens collected on 
25.V1I.1941 in Mathews County, Virginia. 
Following extensive surveys in the 1980s 
and early 1990s, Hill and Knisley (1994) 
reported that adult beetles were found at 13 
sites in Calvert County, Maryland, and 55 
sites in Virginia. 

Other early collecting records of C. d. 
dorsalis from the Chesapeake Bay region 
have not been mentioned in the published 
literature. While curating the NMNH col- 
lection of these beetles, I found 30 speci- 
mens which had been collected prior to the 
publication of the first report of C. d. dor- 
salis from the Chesapeake Bay region. 

USA: Maryland: Calvert County: Breezy 
Point Beach, 14.VII.1950, O. L. Cartwright 
(3 6); Calvert Beach, 4.VII.1973, J. M. 
Sheppard (1 2); Cove Point, Chesapeake 


809 


Bay, 27.VIII.1972, J. M. Sheppard (4 6, 2 
2); Flag Pond, 3 miles south Kenwood 
Beach, 24.VI.1959, O. L. Cartwright (2 9), 
P. & P. Spangler (1 5); Kenwood Beach, 5 
miles south Prince Frederick, 5.VII.1950, 
GEV Nelsonr (igs 2) 29a MESS OsGe 
EH Nelson” (2) males): Elum y,Lomt 
24.VII.1949, O. L. Cartwright (1 2); Port 
Republic, 20.VII.1950, M. H. Hatch (1 @); 
Prince Frederick, 21.VII.1950, G. H. Dieke 
(2 5,1 2). St. Mary’s County: Point Look- 
out, on sandy beach, Chesapeake Bay, 
2I-N TNO AAS Gwe Bay Ofte Gleucinmn lame): 
1. VII.1944, G. B. Vogt (1 6, 2 @). 
Virginia: state label only (1 @). 

These specimens demonstrate that adults 
of C. d. dorsalis were regularly encountered 
by collectors at Chesapeake Bay beaches in 
the 1940s and 1950s, during a time when 
many other populations of this subspecies 
were experiencing severe declines (Stama- 
tov 1972, Knisley et al. 1987, Hill and 
Knisley 1994). Knisley (personal commu- 
nication) reports that C. d. dorsalis has been 
found at or near many of these same sites 
in his recent surveys. Two exceptions are 
the Breezy Point Beach, which has become 
a heavily used area and is therefore now 
unsuitable for beach tiger beetle popula- 
tions, and the Point Lookout site in St. 
Mary’s County, where Knisley failed to find 
C. d. dorsalis in recent years, indicating an- 
other possible extirpation. The continued 
presence of C. d. dorsalis at many of the 
same sites where it was found in the 1940s 
and 1950s suggests that there may be 
grounds for optimism regarding the long- 
term survival of this subspecies in the 
Chesapeake Bay region. 


Cicindela dorsalis media LeConte 


Cicindela dorsalis media LeConte has 
experienced declines in parts of its range 
and is currently listed as ““Endangered”’ by 
the state of Maryland (Yarbrough and Knis- 
ley 1994). Knisley and Schultz (1997) were 
the first to report this subspecies from the 
Chesapeake Bay. Five specimens of this 
subspecies in NMNH document its historic 


810 


presence in the southern Chesapeake Bay. 
Three of these specimens lack a precise 
year of collection but, given that they were 
collected by H. G. Hubbard and E. A. 
Schwarz, the specimens would have been 
collected some time prior to Hubbard’s 
death in 1899. 

Virginia: Hampton County, Fort Monroe, 
WAHL INSS2 CS. I Ly ilo, WU [limo syestel|, 
H. G. Hubbard and E. A. Schwarz (3 <G). 


Cicindela puritana Horn 


Since 1911, this species has been regu- 
larly collected at sites along the western 
shore of the Chesapeake Bay in Calvert 
County, Maryland (Hill and Knisley 1993). 
I recently discovered a wealth of additional 
material from these sites among the unsort- 
ed and unidentified tiger beetle specimens 
in NMNH. 

USA: Maryland: Calvert County: Cal- 
vert, 13.VII.1967, J. Stamatov (2 6, 2 @); 
Calvert Beach, 4.VH.1973, J. M. Sheppard 
(12 6, 24 @), 8.VII.1974, J. M. Sheppard 
(6 6, 3 2), VII.1974 (4 36, 4 @); Calvert 
Cliffs, VII.1973 (3 36, 2 2); Calvert Cliffs 
State Park, 7.VIII.1984, S. W. Gross (5 ¢ 
and | 2); Chesapeake Beach, 28.VI.1911, 
E. Shoemaker (2 6, 4 2); 21.VI.1912, E. 
Shoemaker (1 6); 22.VI.1912, E. Shoe- 
maker (1 6); 24.V1I.1914, E. Shoemaker (4 
Oy 5 Lye BOVIGIGA® (SOs WTS), 
MBN MNOBS, INS INieolesy > SO, 2 2), 
23.V1.1933 (6 6, 4 2), 9.V1.1934 (4 3, 6 
NiO SAA Nicolaya@® <n sues): 
22.V1.1934, O. L. Cartwright (3 6, 3 &), 
DANN NOB (2 5 3 2), ZAN MNES), 2X. INTC= 
olay (1 d), 1.VHI.1941, on sandy beach be- 
lon Clini Ge IB, Vost @ 6, 3 2), 
24.VII.1949, at foot of cliffs, G. B. Vogt (6 
3), 20.VI.1981, Gordon and Wilder (2 ¢); 
Dares Beach, I.VII.1934, A. Stone (1 d, 1 
2) 26.VI19495 He Dozier (aaa 2): 
Flag Pond, 24.V1.1959, O. L. Cartwright (3 
3d, 1 2); Matoaka Beach, Calvert Cliffs, 
IS WRIA, lek 12 ekonrel Cl c, 7 2)s Jeahuren 
Point, 24.V1I.1949, O. L. Cartwright (9 6, 
5 2), 28.VI.1950, O. L. Cartwright (6 6, 3 
2), 30. VII.1950, on beach, B. H. Dozier (1 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


6): 4aVMIISSIeeS. 1s Parhint G6) Sernce 
Frederick, 21.VII.1940 (2 ¢, 2 2); Scientist 
Cliffs, 28.VII.1979, W. E. Steiner (4 5, 1 @). 
Chesapeake Bay populations of this spe- 
cies have clearly been popular with tiger 
beetle collectors. It is interesting to note 
that most collections are of multiple indi- 
viduals and that large series of specimens 
have been collected on a single date (e.g., 
36 specimens collected on July 4, 1973, at 
Calvert Beach). Taken together, these two 
facts suggest that this species was probably 
abundant at certain times and localities. 


ACKNOWLEDGMENTS 


I thank Terry L. Erwin for sponsoring my 
continued work as a Research Associate at 
the National Museum of Natural History. C. 
Barry Knisley of Randolph Macon College 
and Michael A. Valenti of the Delaware 
Forest Service reviewed the manuscript and 
provided helpful comments and suggestions 
for its improvement. 


LITERATURE CITED 


Boyd, H. P. 1975. The overlapping ranges of Cicindela 
dorsalis dorsalis and C. d. media, with notes on 
the Calvert Cliffs Area, Maryland. Cicindela 7(3): 
55-59. 

Boyd, H. P. and R. W. Rust. 1982. Intraspecific and 
geographic variation in Cicindela dorsalis Say 
(Coleoptera: Cicindelidae). The Coleopterists Bul- 
letin 36(2): 221-239. 

Glaser, J. D. 1976. Cicindelids of Chesapeake Bay re- 
visited. Cicindela 8(1): 17—20. 

. 1984. The Cicindelidae (Coleoptera) of Mary- 
land. Maryland Entomologist 2(4): 65-76. 

Hill, J. M., and C. B. Knisley. 1993. Puritan tiger bee- 
tle (Cicindela puritana G. Horn) recovery plan. 
U.S. Fish and Wildlife Service, Hadley, Massa- 
chusetts, 39 pp. 

. 1994. Northeastern beach tiger beetle (Cicin- 
dela dorsalis dorsalis Say) recovery plan. U.S. 
Fish and Wildlife Service, Hadley, Massachusetts, 
45 pp. 

Knisley, C. B. and T. D. Schultz. 1997. The Biology 
of Tiger Beetles and a Guide to the Species of the 
South Atlantic States. Virginia Museum of Natural 
History, Martinsville, Virginia, viii + 210 pp. 

Knisley, C. B., J. I. Luebke, and D. R. Beatty. 1987. 
Natural history and population decline of the 
coastal tiger beetle, Cicindela dorsalis dorsalis 
Say (Coleoptera: Cicindelidae). Virginia Journal 
of Science 38: 93-303. 


VOLUME 107, NUMBER 4 


Stamatov, J. 1972. Cicindela dorsalis endangered on 
northern Atlantic coast. Cicindela 4: 8. 

U.S. Fish and Wildlife Service. 1990. Endangered and 
threatened wildlife and plants; determination of 
threatened status for the Puritan tiger beetle and 
the northeastern beach tiger beetle. Federal Reg- 
ister 55(152): 32088-32094. 

Vogler, A. P. and R. DeSalle. 1993. Phylogeographic 
patterns in coastal North American tiger beetles, 
Cicindela dorsalis inferred from mitochondrial 
DNA sequences. Evolution 47: 1192-1202 

Vogler, A. P., C. B. Knisley, S. B. Glueck, J. M. Hill, 
and R. DeSalle. 1993a. Using molecular and eco- 


811 


logical data to diagnose endangered populations 
of the Puritan Tiger Beetle, Cicindela puritana. 
Molecular Ecology 2: 375-383 

Vogler, A. P., R. DeSalle, T. Assmann, C. B. Knisley 
and T. D. Schultz. 1993b. Molecular population 
genetics of the endangered tiger beetle, Cicindela 
dorsalis (Coleoptera: Cicindelidae). Annals of the 
Entomological Society of America 86: 142-152 

Yarbrough, W. W. and C. B. Knisley. 1994. Distribu- 
tion and abundance of the coastal tiger beetle, Ci- 
cindela dorsalis media (Coleoptera: Cicindelidae), 
in South Carolina. Entomological News 105(4): 
189-194. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 812-819 


CONTRIBUTION TO THE TAXONOMY AND FAUNISTICS OF THE GENUS 
MEROPLEON DYAR (LEPIDOPTERA: NOCTUIDAE) 


JAN METLEVSKI 


Department of Entomology, Kansas State University, Manhattan, KS 66506-4004, 


U.S.A (e-mail: jmetlevs @oznet.ksu.edu) 


Abstract.—A new species of owlet moth, Meropleon linae (Lepidoptera: Noctuidae) 
from Kansas, is described and illustrated. Meropleon linae is distinguished from other 
species of Meropleon by forewing color and pattern, shape of the valva, and structure of 
the aedoeagus. Meropleon linae is most similar morphologically to M. cinnamicolor Fer- 
guson. New distribution records for M. ambifuscum (Newman) and the first record for M. 


titan Todd in Kansas are given. 


Key Words: 
Nearctic 


Meropleon Dyar, 1924, is a small, mor- 
phologically distinct genus endemic to east- 
ern North America. Members of this genus 
can be recognized using male and female 
genitalic characters. Males of Meropleon 
can be recognized by a simplified valva 
with no cucullus, corona or harpe, but with 
a short rounded and flattened costal process 
curved inside the genital capsule, and by 
the aedoeagus with several strong spinelike 
cornuti located near the base of the vesica 
or on the carina. Females of Meropleon can 
be recognized by the following combina- 
tion of genitalic characters: ovipositor 
short; papillae anales heavily sclerotized, 
distally dorsoventrally flattened and curved 
ventrad; gonapophyses with strong, thick- 
ened shaft, paddle-shaped, anteriors ones 
very short, several times shorter than pos- 
teriors; ostium bursae wide; ventral plate of 
ostium bursae a strongly sclerotized band 
broadly arched ventrad, broadly and strong- 
ly attached at its ends to ends of 8th tergite; 
ductus bursae weakly sclerotized, its length 
subequal to length of bursa copulatrix; bur- 
sa copulatrix membranous, with no signa, 


new species, new state record, distribution, Konza Prairie, Riley County, 


length of bursa copulatrix subequal to its 
width; appendix bursae very short, lobelike; 
the caudal edge of 7th sternite with a rim 
of short non-deciduous hairs. 

Five Meropleon species have been de- 
scribed (Ferguson 1982, Poole and Gentili 
1996) and one more species is described be- 
low. The known larvae feed internally in 
sedges and large grasses (Ferguson 1982). 
Species of the genus are not commonly 
caught using traditional lighting techniques, 
and are strictly associated with habitats 
where food plants grow, although in such 
places they can be numerous. Kansas is on 
the western edge of the range of this genus. 
Two species, M. ambifuscum (Newman, 
1948) and M. diversicolor (Morrison, 
1874), have been recorded from eastern 
Kansas (Ferguson 1982). 

One new species of Meropleon and a 
new Kansas state record were discovered 
during a survey of owlet moths at the Kon- 
za Prairie Biological Station, Riley County, 
Kansas. I take this opportunity to describe 
this new species and present new distribu- 
tion records for species of Meropleon in 
Kansas. 


VOLUME 107, NUMBER 4 


Fig. 1. 
lomere. A, length of short branch; B, length of long 
branch; C, width of flagellomere; D, length of flagel- 
lomere. 


Measure distances of an individual flagel- 


MATERIALS AND METHODS 


The majority of the material used in this 
work was obtained by night collecting at 
Konza Prairie Biological Station. Addition- 
al data were obtained from specimens in the 
Snow Entomological Collection at the Uni- 
versity of Kansas. Night collecting was 
conducted several times monthly from the 
beginning of July to the beginning of No- 
vember in 2003, and from the end of Feb- 
ruary to the middle of November in 2004. 
A white sheet and 175-watt mercury vapor 
light were used for night collecting. Col- 
lected specimens were spread and dried on 
spreading boards and examined with a Lei- 
ca MZ 9.5 stereomicroscope with 10 oc- 
ulars. Individual flagellomeres were mea- 
sured with an ocular micrometer in a Leica 
MZ APO stereomicroscope with 10 ocu- 
lars. Measurements as shown on Fig. | 
were taken viewing the right antenna in 
ventral view under high power (80%). 
Some specimens were dissected to study 
genitalic structures. Genitalia were photo- 
graphed with the use of a Leica MZ APO 
stereomicroscope and a Nikon DXM 1200 
digital camera. Terminology for genitalic 
structures follows Hacker (2004), and ter- 
minology for wing patterns follows Forbes 
(1954). Paratypes of Meropleon cinnami- 
color Ferguson, 1982, were examined. 

Abbreviations for institutions mentioned 
in the text are as follows: KPBS = Konza 


813 


Prairie Biological Station, Riley County, 
Kansas; KSEM = Snow Entomological 
Collection, University of Kansas, Lawr- 
ence; KSU = Kansas State University, 
Manhattan; KSU-MEPAR = Kansas State 
University Museum of Entomological Prai- 
rie Arthropod Research; USNM = National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 

The abbreviations AL, 20C, and THP 
that occur on some collecting data men- 
tioned in the text designate certain water- 
sheds at KPBS. 


Meropleon linae Metlevski, new species 
(Figs. 2, 4-11) 


Diagnosis.—The structure of the male 
and female genitalia clearly shows that this 
species belongs to the genus Meropleon. 
Meropleon linae can be differentiated from 
all other species of Meropleon using struc- 
tures of the male genitalia and the color and 
pattern of the wings. In the male genitalia 
of M. linae, the ventral margin of the valva 
is noticeably curved at the distal end of the 
sacculus and the aedoeagus has 5-7 ‘thick, 
heavily sclerotized spinelike cornuti (one 
near the base of the vesica and 4—6 on the 
ventral plate of the carina). In all other spe- 
cies of Meropleon, the ventral margin of the 
valva is not noticeably curved and the ae- 
doeagus has 2—4 cornuti (no more than one 
cornutus on the carina). The following fea- 
tures can be used to differentiate Meropleon 
linae from the most similar M. cinnamico- 
lor (Fig. 3): in M. linae the ground color of 
forewings is duskier than in M. cinnami- 
color; in M. linae, the forewings have no 
whitish scales, while in M. cinnamicolor 
there is a whitish area in the proximal part 
of forewing between veins R and Cu; the 
forewing pattern is much more reduced (its 
elements are almost completely absent) in 
M. linae than in M. cinnamicolor; in M. lin- 
ae the hindwings are whitish, compared to 
light brown in M. cinnamicolor. Meropleon 
linae can be easily differentiated from M. 
ambifuscum, M. diversicolor, M. titan Todd, 
1958, and M. cosmion Dyar, 1924, using the 


814 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 2-5. 
olina. 4—5, Holotype of M. linae. 4, Wing pattern. 5, Antenna. 


forewing pattern. In M. linae the forewings 
are unicolorous with an almost completely 
reduced wing pattern, while in M. ambifus- 
cum, M. diversicolor, and M. titan the fore- 
wings have the proximal half with dark 
brown black elements and a much lighter 
distal half, as well as well developed ele- 
ments of the wing pattern. In M. linae the 
forewings have no white elements, while in 
M. cosmion the white elements are well de- 
veloped and contrast strongly with the fore- 
wing’s ground color. 

Description.—Male: Head: Light red- 
dish brown, some brownish-gray hairs 
above eyes; vertex and upper part of front 
with tuft of flattened hairs, longest on ver- 
tex and shorter toward front; labial palpus 
gray, with longitudinal band of dark gray 
scales; antenna bipectinate, shaft covered 
with whitish scales, 61 flagellomeres, ratio 
length of short branch/length of long 


Meropleon linae and M. cinnamicolor. 2, Paratype of M. linae. 3, M. cinnamicolor, South Car- 


branch/thickness of flagellomere/length of 
flagellomere equal: 15th flagellomere—20/ 
29/13/13, 35th flagellomere—19/22/11/11. 
Thorax: Light reddish brown dorsally, 
covered with long flattened and slightly 
spatulate hairs; pale gray ventrally except 
brownish-gray area between head and fore- 
legs, covered with long simple hairs; fore- 
leg brownish gray, other legs pale gray. 
Forewing: Length 14-17 mm; reddish 
brown except for grayish shadow not con- 
trasting strongly with remainder of wing in 
postmedial area between inner margin and 
vein Cu2; markings almost completely ab- 
sent, postmedial line and reniform spot 
slightly lighter than ground color but hardly 
visible; veins A, Cu, and R and branches of 
veins Cu, R, and M with many brownish- 
gray scales, appearing dark gray; terminal 
line grayish brown; fringe slightly darker 
than ground color; underside gray, small 


VOLUME 107, NUMBER 4 


815 


8 


Figs. 6-9. 


9 


Male genitalia of Meropleon linae (holotype). 6, Genital capsula (posterior). 7, Right valva. 8, 


Aedoeagus with vesica everted left lateral view). 9, Aedoeagus with vesica and ventral plate of carina everted 


(right ventrolateral view). 


discal spot and area between vein A and 
inner margin whitish; fringe gray. 

Hindwing: Whitish with scattered gray 
scales in outer part; vein R and branches of 
veins R and M gray; discal spot light gray, 
barely visible; fringe whitish, light pinkish 
brown at apex; underside whitish, small 
dark gray discal spot, many scattered dark 
gray scales; area between vein Cu and inner 
margin with few dark gray scales; fringes 
white, gray at apex. 

Abdomen: Light gray; first segment with 
dorsal tuft of long slightly spatulate hairs. 

Male genitalia (Figs. 6—9): Uncus api- 
cally expanded and flattened dorsoventral- 
ly; sclerotized costa separates from dorsal 
edge of valva approximately at middle, ter- 
minated as a rounded flattened process 


turned inside the genital capsula; sacculus 
flattened dorsally, with setose area on dor- 
sal surface; clasper a longitudinal, almost 
straight, narrow, sclerotized bar extending 
from distal end of sacculus to ventral base 
of costal process, along and very close to 
ventral margin of valva; ventral margin of 
valva noticeably curved at distal end of sac- 
culus, slightly concave medially, slightly 
sclerotized distally from sacculus; ventral 
sclerotization of valva diverging from 
clasper at about one-third of clasper’s 
length before distal end of clasper; distal 
portion of valva from costal process to apex 
slightly sclerotized and setose apically; 
apex of valva rounded; distal portion of val- 
va equal to half length of valva’s dorsal 
edge; saccus somewhat produced, not ta- 


816 


pering to a point, abruptly narrowed in dis- 
tal half; aedoeagus tubular, slightly arcuate, 
extended moderately at distal end, with 
well-developed and strongly sclerotized 
ventral plate of carina; ventral plate of ca- 
rina with 4—6 thick, heavily sclerotized 
spines, arranged by size, largest located on 
rounded apex of ventral plate of carina, and 
remaining decreasing in size toward base of 
ventral plate (Fig. 9); everted vesica finely 
granulose, broad, short, angled slightly ven- 
trolaterally, then inflected ventrad, nar- 
rowed after inflection and prolonged by 
ductus ejaculatorius, with one short, round- 
ed, broad at base diverticulum located be- 
fore inflection; vesica with one thick, long, 
spinelike cornutus located near base of ve- 
sica, ventrolaterally on right side and point- 
ed dorso-cephalad (Fig. 8). 

Female: Coloration and markings similar 
to male; antenna filiform. 

Female genitalia (Figs. 10—11): Ovipos- 
itor short; papilla analis strong, heavily 
sclerotized, full length flattened ventrally, 
and in distal third dorsally, flat distal third 
curved ventrad, apex rounded, densely se- 
tose with short setae; apophysis posterioris 
strong, well sclerotized, paddle-shaped, lit- 
tle longer than papilla analis; eighth seg- 
ment strongly sclerotized, narrow, curved 
ventrad and cephalad, its ends broadly and 
strongly attached to ends of the ventral 
plate of ostium bursae; apophysis anterioris 
very short and wide; ostium bursae wide, 
ventral plate as strongly sclerotized narrow 
band, broadly arched ventrally, broadened 
at ends, dorsal plate moderately sclerotized, 
its caudal edge with small triangular shear 
medially; ductus bursae gelatinous-sclero- 
tized, dorsoventrally flattened in its caudal 
third, wide at ostium bursae, narrowed 
cephalad, abruptly asymmetrically expand- 
ed before meeting bursa copulatrix, with a 
mesal lobe on dorsal side; bursa copulatrix 
membranous, wide caudally, narrowed an- 
teriorly, as long as wide in its caudal part; 
appendix bursae very short, as a small lobe 
on the left side of caudal part of the bursa 
copulatrix; ductus seminalis extending from 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


the base of appendix bursae, on its right 
side close to the base of ductus bursae. 

Types.—Holotype: dg, “KANSAS, Riley 
County, KPBS, Nature trail area, 
39°06.28'N, 96°35.75'W, 22.X.2003, J. Me- 
tlevski,” ““KSU-MEPAR Genitalia Prep. by 
J. Metlevski No 161.” Deposited in KSU- 
MEPAR. Paratypes: 1 d, same as holotype, 
deposited in USNM; 2 6, “KANSAS, Ril- 
ey, County, INPBSa VAI 39,0652 ae 
96°35.70'W, 3.X.2004, J. Metlevski’; 1 6, 
“KANSAS, Riley County, KPBS, 
39°06.13'N, 96°36.35'W, 19.X.2004, J. Me- 
tlevski’’; 1 6, same label data as above ex- 
cept “21.X.2004,” ““KSU-MEPAR Geni- 
talia Prep. By J. Metlevski No 231,” de- 
posited in KSU-MEPAR; 1 <6, same label 
data as above except “24.X.2004,”’ “KSU- 
MEPAR Genitalia Prep. By J. Metlevski No 
232,” deposited in KSU-MEPAR; 1 2, 
“KANSAS, Riley County, KPBS, Main 
headquarters, 3.X.2004, J. Metlevski,”’ 
**“KSU-MEPAR Genitalia Prep. By J. Met- 
levski No 233,” deposited in KSU-ME- 
PAR. 

Biology.—Unknown. 

Discussion.—The habitat where speci- 
mens of M. linae were caught is a hilly tall- 
grass prairie with wooded areas along a 
creek and on some slopes. Other species of 
Meropleon fly in the autumn and are uni- 
voltine, and the collection data suggest that 
M. linae is the same. Meropleon linae is 
most similar morphologically to M. cin- 
namicolor. Both species are characterized 
by broadly bipectinate male antennae, fore- 
wings almost unicolorous reddish brown 
with wing pattern very much reduced, and 
abdominal tuft on the first abdominal ster- 
num only (in other species of Meropleon 
the tuft is present also on segments 3 and 
4). Meropleon linae and M. cinnamicolor 
also share the following characters of the 
male genitalia: an apically expanded and 
flattened uncus; an extended distal part of 
the valva, which is at least half as long as 
length of the valva’s dorsal edge (compared 
to less than half in other species of Mero- 
pleon),; and ventral sclerotization of the val- 


VOLUME 107, NUMBER 4 


817 


lit 


Figs. 10-12. Meropleon linae and M. titan. 10-11, Female genitalia of M. linae. 10, Ventral view. 11, Right 


lateral view. 12, M. titan. Kansas, Riley County, KPBS. 


va diverging from the clasper well before 
its distal end. From the point of morpho- 
logical similarity of M. cinnamicolor and 
M. linae the known distribution of both spe- 
cies is a challenge. Meropleon cinnamico- 
lor is known only from several areas in the 
coastal marshes of South Carolina and 
North Carolina and M. linae is found on the 
opposite side of the range of the genus. 


Etymology.—Meropleon linae is named 
in honor of my beloved wife Lina. 

Specimens of M. cinnamicolor exam- 
ined.—All are in USNM. Paratypes: 1 ¢, 
“Wedge Plantation, South Santee River, 
Charleston County, S[outh] C[arolina], 22 
November 1967, Douglas C. Ferguson,” 
“USNM Genitalia Slide by DCF No 
56474’; 1 2, same label data as above ex- 


818 


cept ““Noy.21.1967”; 1 2, “McClellanville, 
South Carolina 29458, 31 Oct. 1967, Light, 
R. B. Dominick, Charles R. Edwards”; 1 
2, ‘Wedge Plantation, 7 mi. NE Mc- 
Clellanville, Charleston County, S[outh] 
Car[olina], Nov.1.1967,” “Coll. by C. R. 
Edwards,’ ““USNM Genitalia Slide by 
DCF No 56488.”’ Other specimens: | 4, 
‘““Wedge Plantation, 7 mi. NE Mc- 
Clellanville, Charleston County, S[outh] 
Carlolmal SOceI9o7e > Coll by (Cy IR: 
Edwards,’’ ““USNM Genitalia Slide by 
DCF No 56473”; 1 6, South Carolina, 
Wedge Plantation, McClellanville, 28 No- 
vember 1970, D. C. Ferguson.” 


Meropleon ambifuscum (Newman) 


The distributional range of this species 
includes Connecticut (Ferguson 1982), 
Maryland (Metzler et al., in press), Ohio 
(Rings et al. 1992), Michigan (Ferguson 
1982), Wisconsin (Metzler et al., in press), 
Indiana (Metzler et al. in press), Illinois 
(Metzler et al., in press), Iowa (Metzler et 
al., in press), Missouri (Ferguson 1982), 
Nebraska, (Metzler et al. in press) Kansas 
(Ferguson 1982), Kentucky (Metzler et al. 
in press), Tennessee (Metzler et al., in 
press), Arkansas (Ferguson 1982), South 
Carolina (Ferguson 1982), Georgia (Met- 
zler et al., in press), Mississippi (Ferguson 
1982), and Louisiana (Metzler et al., in 
press). Meropleon ambifuscum was known 
in Kansas only from Douglas County (Fer- 
guson 1982) in the easternmost part of the 
state. New data on the distribution of M. 
ambifuscum in Kansas are given below. 
One specimen, labeled ““Topeka, Ks. Po- 
penoe.,’’ was found in KSEM, and nineteen 
specimens were collected at KPBS: 1 4, 
“Riley County, KPBS, 20C, 3.VIII.2003, J. 
Metlevski,”’ ““KSU-MEPAR Genitalia Prep. 
by J. Metlevski No 157” (KSU-MEPAR); 
2 3, “KPBS, Nature trail area, 39°06.28'N, 
96°35.75'W, 30.1X.2003, J. Metlevski,”’ 
“KSU-MEPAR Genitalia Prep. by J. Me- 
tlevski No 156,” ““KSU-MEPAR Genitalia 
Prep. by J. Metlevski No 158”? (KSU-ME- 
PAR); 6 3, same label data as above except 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


**19.X%.2003,”’ ““KSU-MEPAR Genitalia 
Preps By Jp Metleyska No [59% KesE 
MEPAR Genitalia Prep. By J. Metlevski No 
160”’ (KSU-MEPAR); 8 6, ““Riley County, 
KPBS, AL. S906 00N> 2 oGks 5-7. 
15.1X.2004, J. Metlevski’”’; 2 ¢, “Riley 
Gounty, KREBS] Ele So 206so7mNe 
96°33.80'W, 20.1X.2004, J. Metlevski.” 


Meropleon titan Todd 
(Fig. 12) 


This species is known from New Jersey 
(NatureServe 2004), Maryland (Stevenson 
1989, J. D. Glaser in litt.), Virginia 
(NatureServe 2004), North Carolina 
(NatureServe 2004), South Carolina (Fer- 
guson 1982), Georgia (J. K. Adams in litt.), 
Mississippi (Ferguson 1982), Missouri 
(Ferguson 1982), and north central Texas 
(Blanchard, 1973); it has not been reported 
from Kansas. Two males were collected at 
KPBS in October 2003. Both specimens 
have the following label data: KANSAS, 
Riley County, KPBS, Nature trail area, 
39°06.28'N, 96°35.75'W, 22.X.2003, J. Me- 
tlevski (KSU-MEPAR). Genitalia prepara- 
tion for one of them was made: KSU-ME- 
PAR Genitalia Prep. by J. Metlevski No 
162. Twelve more specimens were collected 
at KPBS in 2004: 1 d and 1 2, “KANSAS, 
Riley County, KPBS, AL, 39°06.12’'N, 
96°35.70'W, 26.1X.2004, J. Metlevski’’; 9 
6 and 1 2, same label data as above except 
*3.X.2004.”’ 


ACKNOWLEDGMENTS 


I am grateful to Gregory Zolnerowich 
(KSU) and Robert R. Kula (KSU) for help- 
ful comments on the manuscript and pre- 
paring pictures. I thank Michael G. Pogue 
(Systematic Entomology Laboratory, 
USDA [USNM]) and James S. Ashe 
(KSEM) for loans of specimens used in this 
study; Eric H. Metzler (Columbus, Ohio) 
for reviewing the manuscript and making 
helpful comments; James K. Adams (Dal- 
ton State College, Georgia), John D. Glaser 
(Maryland) and Dale E Schweitzer 
(NatureServe, Arlington, Virginia) for sup- 


VOLUME 107, NUMBER 4 


plying information on distribution of Mer- 
opleon titan; and Enid Cocke (KSU) for ed- 
iting the English text of the manuscript. I 
acknowledge the cooperation of Konza 
Prairie Biological Station. This article is 
Contribution No. 05-84-5 from the Kansas 
Agricultural Experiment Station (KAES) 
and was supported in part by KAES Hatch 
Project No. 583, Insect Systematics. 


LITERATURE CITED 


Dyar, H. G. 1924. A new noctuid from Louisiana. In- 
secutor Inscitiae Menstruus 12: 21-22. 

Blanchard, A. 1973. Records and iliustration of some 
interesting moths flying in Texas (Sphingidae, 
Ctenuchidae, Noctuidae, Notodontidae, Geometri- 
dae, Pyralidae, Cissidae). Journal of the Lepidop- 
terist’s Society 27(2): 103-109 

Ferguson, D. C. 1982. A revision of the genus Mero- 
pleon Dyar, with descriptions of a new species and 
subspecies (Lepidoptera: Noctuidae). Entomogra- 
phy 1: 223-225. 

Forbes, W. T. M. 1954. Lepidoptera of New York and 
neighboring states. Part III. Cornell University 


819 


Agricultural Experiment Station, Memoirs 329, 
433 pp. 

Hacker, H. H. 2004. Revision of the genus Caradrina 
Oscsenheimer, 1816, with notes on other genera 
of the tribus Caradrini (Lepidoptera, Noctuidae). 
Esperiana 10: 7—690. 

Metzler, E. H., J. A. Shuey, L. A. Ferge, P. Z. Gold- 
stein, R. A. Henderson. In press. Contributions to 
the Understanding of Tallgrass Prairie-Dependent 
Butterflies and Moths (Lepidoptera) and: their Bio- 
geography in the United States. Ohio Biological 
Survey, Columbus, Ohio. 

NatureServe. 2004. NatureServe Explorer: An online 
encyclopedia of life [web application]. Version 
3.0. NatureServe, Arlington, Virginia. Available 
http://www.natureserve.org/explorer (accessed: 
July 02, 2004). 

Poole, R. W. and P. Gentili, eds. 1996. Nomina Insecta 
Nearctica. Volume 3. Diptera, Lepidoptera, Si- 
phonaptera. Entomological Information Services, 
Rockville, Maryland, 1,143 pp. 

Rings, R. W., E. H. Metzler, EF J. Arnold, and D. H. 
Harris 1992. The Owlet Moths of Ohio. Bulletin 
of the Ohio Biological Survey, New Series 9, 223 
Pp. 

Stevenson, H. G. 1989. The Genus Meropleon Dyar 
(Lepidoptera: Noctuidae: Amphipyrinae) in Mary- 
land. Maryland Entomologist 3: 71—73. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 820-834 


REVIEW OF THE AULACIDAE (HYMENOPTERA) OF CHILE AND 
ADJACENT ARGENTINA 


DAvID R. SMITH 


Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- 
ment of Agriculture, % National Museum of Natural History, Smithsonian Institution, 


RO, Box BIOUZ, IMURE 
dsmith @sel.barc.usda.gov) 


168, Washington, 


DC 20013-7012, U:S.A. (e-mail: 


Abstract.—Five species of Aulacidae occur in Chile: Aulacus braconiformis (Kieffer), 
A. krahmeri Elgueta and Lanfranco, A. brevis, n. sp., Pristaulacus capitalis (Schletterer), 
and P. rubriventer (Philippi). Two of these, A, krahmeri and A. brevis, occur in adjacent 
Argentina. Aulacus brevis also is recorded from the Juan Fernandez Islands. A key to 
species is presented and diagnoses and new distribution records are given. Hosts are wood- 


boring beetles of the family Cerambycidae. 


Key Words: 
dez Islands 


Elgueta and Lanfranco (1994) revised the 
Chilean species of Aulacidae and included 
two species of Aulacus Jurine and two spe- 
cies of Pristaulacus Kieffer. During my 
study of New World Aulacidae, I have ex- 
amined about 90 additional specimens of 
aulacids from Chile and adjacent Argentina, 
and here I present distribution records ex- 
tending the distribution of some species, re- 
cord two species from southern Argentina 
for the first time, give the first records of 
Aulacidae from the Juan Fernandez Islands, 
and describe a new species of Aulacus. As 
Elgueta and Lanfranco (1994) have already 
provided detailed descriptions, only diag- 
noses of those species are given here. None 
of the species represented in Chile and 
southern Argentina occur elsewhere, and 
the fauna is rather sparse considering the 
estimated 100 species in the rest of the 
Neotropics. 

For generic synonymy and complete ref- 
erences to each species, see Smith (2001). 
Terminology follows Huber and Sharkey 
@i995)): 


parasitoids, new species, key, Cerambycidae, South America, Juan Fernan- 


Acronyms for collections from which 
specimens were borrowed are as follows: 
AEI, American Entomological Institute, 
Gainesville, FL; AMNH, American Muse- 
um of Natural History, New York, NY; 
CAS, California Academy of Sciences, San 
Francisco; CNC, Canadian National Collec- 
tion, Ottawa; FSCA, Florida State Collec- 
tion of Arthropods, Gainesville; PC, Pagli- 
ano Collection, Torino, Italy; USNM, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, DC. 


KEY TO GENERA AND SPECIES 


1. Tarsal claws simple or with minute tooth at 
base; head without occipital carina (Aulacus) 


AR CERELE Ean 5 ROME ed Ona Ors autso ot 6's) 6 D} 
— Tarsal claw comblike, with 3 or 4 inner teeth; 

head with complete occipital carina (Pristau- 

LAGUS) 2 BRS SARTO EK 8 rR ae eae 4 


2. Small, 6 mm or less in length; ovipositor 
length half length of forewing; forewing cell 
1M narrow, 3.5X longer than broad, and vein 
M+Cu mostly atrophied (Fig. 19) 
Seas, COO aire Rice WRC eee eae A. brevis, n. sp. 

— Large, 9 or more mm in length; ovipositor 
length longer than forewing length; forewing 


VOLUME 107, NUMBER 4 


cell 1M usually broad, about 2—3< longer than 
broad, and vein M+Cu complete (Figs. 4, 11) 
3. Wings hyaline to very lightly dusky, costal cell 
and stigma black (Fig. 11); antenna black with 
flagellomeres 5, 6, and part of 7 white; hind 
tarsus black with segments 2 and 3 white; mark 
on each side of vertex reddish brown; front of 
head evenly convex (Figs. 9-10) ... A. krahmeri 
— Wings black with spot under stigma and stigma 
yellowish (Fig. 4); antenna mostly yellow with 
apical 2—3 flagellomeres black; hind tarsus and 
head uniformly black; front of head projected 
into a shelflike protuberance above antennae 
(higs2=3) pee yer oi cliicne A. braconiformis 
4. Entirely black; wings entirely hyaline (Fig. 26) 


P. capitalis 
— Black with metasoma red; forewing hyaline 
with black spot at apex (Fig. 33) 


P. rubriventer 
Aulacus braconiformis (Kieffer) 
(Figs. 1-7) 


Neuraulacinus braconiformis Kieffer 1911: 
2M 

Aulacus braconiformis: Townes 1950: 
113.—Smith 2001: 269. 

Pristaulacus sp. 1: Barriga 1990: 57, 58. 


Diagnosis.—Length, about 15-16 mm. 
Antenna yellow with scape, pedicel and 
apical 2—3 flagellomeres black. Head, body, 
and legs black. Forewing darkly infuscated 
with yellow spot under stigma and stigma 
mostly yellow (Fig. 4). Frons angulate, pro- 
duced forward above antennae (Figs. 2—3); 
head dull with fine microsculpture (Figs. 1— 
3). Mesonotum smooth, subshining, with- 
out punctures or carinae. Notauli of meso- 
scutum meet posteriorly near transscutal ar- 
ticulation (Fig. 6). Upper surface of hind 
coxa rugose (Fig. 5); ovipositor guide not 
indicated; hind prefemur distinct. Oviposi- 
tor length 1.3 forewing length. Both sexes 
known. 

Distribution.—Recorded only from the 
Region Metropolitana de Santiago by El- 
gueta and Lanfranco (1994). 

New records.—CHILE: O’ HIGGINS: 
Rancagua, XII-85, Pérez Arce (4 2°, 
FSCA). SANTIAGO: La Obra, 1.70, Alfaro 
Col. (1 6, AMNH); Santiago, Rio Clarillo 
Nat. Pk., 2-XII-1988—3-I-1989, Malaise 


821 


trap (1 2, FSCA). Province unknown: Sal- 
to, I-10—1937, Dr. Reed, E. P. Reed collec- 
tion (1 2, CAS); Salto, XI-1922, Dr. Reed 
(1 6, USNM). 

Host.—Recorded as Pristaulacus sp.1 by 
Barriga (1990), who reared this species 
from Hephaestion lariosi (Bosq) (Ceram- 
bycidae). Adults have been collected from 
flowers of Maytenus boaria Molina (Celas- 
traceae) (Elguega and Lanfranco 1994). 

Note.—The holotye of N. braconiformis 
is in The Natural History Museum, London, 
UK. 


Aulacus krahmeri Elgueto and Lanfranco 
(Figs. 8-14) 


Aulacus krahmeri Elgueto and Lanfranco 
1994: 90.—Smith 2001: 272. 
Pristaulacus sp. 2: Barringa 1990: 58. 


Diagnosis.—Length, about 13 mm. An- 
tenna black with flagellomeres 5, 6, basal 
half of 7, and sometimes apical third of 4 
white. Head black with reddish-brown 
marks on each side of vertex. Mesosoma 
and metasoma black. Legs black with hind 
tarsal segments 2 and 3 white and 4 brown- 
ish. Wings hyaline to evenly dusky, only 
costal cell of forewing black; stigma black 
(Fig. 11). Head evenly rounded in front, 
dull with fine microsculpture (Figs. 8—10). 
Mesonotum with transverse carinae. Notau- 
li of mesoscutum not meeting behind, sep- 
arately meeting transscutal articulation (Fig. 
13). Upper surface of hind coxa with trans- 
verse carinae (Fig. 12); ovipositor guide at 
lower center on inner surface indicated by 
depression; hind prefemur distinct. Ovipos- 
itor length about 1.6 forewing length. 
Both sexes known. 

Distribution.—Recorded from VII Re- 
gion (Talca) to X Region (Valdivia) by El- 
gueta and Lanfranco (1994). Specimens I 
have seen fall within the known range and 
include western Argentina. 

New records.—ARGENTINA: NEU- 
QUEN: Lago Nothoé, 30.1.949, Coll. E 
Monros (1 ¢, AEI); S. Martin Andes, Pu- 
cara, Dec. ‘69, Schejovskor Coll. (2 &, 


822 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-5. 
surface of hind coxa. 


FSCA). CHILE: CURICO: El Coigo, Cord. 
Curico, Oct./Nov. “59, L. Pefia Guzman (1 
2, AEI). LINARES: Romehual, Cord. Par- 
ral, 5/10-XI-60, Pena (2 2, CNC); Fundo 
Malcho, Cord. Parral, 11-20-XI-1964, L. E. 
Pena (5 2, CNC). MALLECO: Sierra de 
Nahuelbuta, W. of Angol, 1,200 m, I-3—51, 
leg Ross & Michelbacher (6 @°, CAS): 
Cord. Lonquimay, I-5—1962, Luis Pefia, La 
Fusta (1 2, AEI). NUBLE: Las Trancas, 
1EGOO) my Da 13283 3 ePena (aS AED: 


Aulacus braconiformis. 1, Head, dorsal 


ASS SR Aree SSS ITS 


. 2, Head, lateral. 3, Head, front. 4, Forewing. 5, Dorsal 


TALCA: Vilches, XI-1990, Pérez Arce (4 
2, FSCA). Province unknown: Invernada, 
XII.70, Chillan, Luis Pena (1 2, AED). 

Hosts.—Recorded as Pristaulacus sp. 2 
by Barriga (1990), who reared this species 
from wood from which Callisphyris spp., 
Calydon submetallicus (Blanchard), and C. 
havrylenkoi Bosq. (Cerambycidae) also 
emerged. 

Note.—The holotype is in the Museo Na- 
cional de Historia Natural, Santiago, Chile. 


VOLUME 107, NUMBER 4 823 


» «es 
© *@% sue im 


Pee ‘a! 
- 


Figs. 6-7. Aulacus braconiformis. 6, Mesosoma, dorsal. 7, Mesosoma, lateral. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 8-12. Aulacus krahmeri. 8, Head, dorsal. 9, Head, lateral. 10, Head, front. 11, Forewing. 12, Dorsal 
surface of hind coxa. 


Aulacus brevis Smith, new species 
(Figs. 15—22) 


Diagnosis.—Length, 6 mm or less; black 
with legs yellow; malar space 0.3 eye 
height; ovipositor length about half wing 
length; forewing cell 1M narrow, 3.5 X lon- 
ger than broad and vein M-Cu atrophied ba- 
sally. 

Female.—Length, 5.0—6.0 mm; wing 
length, 4.0—5.0 mm; ovipositor length, 2.0— 
2.5 mm. Color: Antenna black with scape 
and pedicel reddish brown. Head_ black: 
mandible, except reddish apex, and mouth- 
parts yellow. Mesosoma black. Legs yellow 
brown, with dark brown hind femur, hind 
tibia, and hind tarsus. Metasoma black with 
various amounts of yellowish toward and 
increasing in size to apex. Wings hyaline; 
stigma black; veins brown. 

Head: Head from above narrowing be- 


hind eyes (Fig. 16). Shiny, pubescent, and 
uniformly punctate with punctures separat- 
ed by flat, shiny interspaces mostly greater 
than diameter of punctures (Figs. 16—18). 
In front view, eyes small and far apart, low- 
er interocular distance about 1.2X eye 
height. Malar space 0.3 eye height (Fig. 
18). Distance between lateral ocellus and 
eye subequal to distance between lateral 
ocelli. Distance between torulus and eye 
slightly longer than distance between toruli. 
Clypeus with small projecting tooth at cen- 
ter; about 3.0 broader than long. Lengths 
of scape, pedicel, and first 3 flagellomeres 
as 1.0: 0.6:1.0:1.6:1.4. 

Mesosoma: Notauli of mesoscutum not 
meeting behind, separately meeting transs- 
cutal articulation (Fig. 21). Mesoscutum 
and scutellum with transverse rugae (Fig. 
21). Propleuron shining with scattered 


VOLUME 107, NUMBER 4 


Figs. 13-14. Aulacus krahmeri. 13, Mesosoma, dorsal. 14, Mesosoma, lateral. 


826 


Fig. 15. 


punctures. Pleurae and propodeum mostly 
uniformly reticulate (Fig. 22) with unsculp- 
tured shining areas on upper and posterior 
pronotum and upper central area of meso- 
pleuron; posterior margin of mesopleuron 
scrobiculate. Forewing (Fig. 19) with cell 
IM small, narrow, 3.5 longer than broad; 
cell 2M long, 4.0X longer than broad; cell 
1M touching cell IRs, not separated by part 
of vein Rs+M; vein M-cu partially atro- 
phied toward Sc. Hind coxa dull, with 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


i 


Aulacus brevis, lateral view. Length, excluding ovipositor, 6.0 mm. 


punctures and rugae on upper surface (Fig. 
20); inner surface dull, with carinae, with- 
out ovipositor guide. Hind prefemur dis- 
tinct. Hind tibia 1.2 longer than hind fe- 
mur; hind basitarsus 1.2 longer than re- 
maining tarsal segments combined; lengths 
of hind tarsal segments as 1.0:0,4:0.2:0.1: 
OZ. 

Metasoma: About 1.2 longer than me- 
sosoma. Ovipositor length 0.5 wing 
length. 


VOLUME 107, NUMBER 4 


‘ 14.fe4 4 
j Hill, i w, 


827 


YW, 


Figs. 16-20. Aulacus brevis. 16, Head, dorsal. 17, Head, lateral. 18, Head, front. 19, Forewing. 20, Dorsal 


surface of hind coxa. 


Holotype.—Female, labeled ‘‘Pichina- 
huel, Cord. Nahuelbuta, Arauco, 1-X-1959, 
L Pefia’’ (CNC). 

Paratypes.—ARGENTINA: Neuquén, 
Pucana San Andes, XII-12-’68, Porter (1 2, 
FSCA). CHILE: Same data as for holotype 
except dates, 20-28-I-1959 (1 2, CNC), 1- 
10-I-1959 (1 2, CNC): Bio-Bio, El Aban- 
ice, XII-30-1950, leg Ross & Michelbacher 
(1 2, CAS); Acou. Zapallar, XII-15-1950, 
Ross & Michelbacher collectors (1 &, 
CAS); Cautin, Pucon (Peninsula), 10-XI-9- 
XII-1989, C. Porter, Malaiste trap (1 °, 
FSCA); Puetrihue, Osorno, I-20-67, Lionel 
Stange (1 2, AEI); Malleco Prov., 4 km W 
Victoria, 300 m, 26-31-x1i.76, S. Peck (1 &, 
AEI); Talca,Vilches, XII.’79, Luis Pena (1 
2, AEI); Valdivia, 30-IX-1988, leg. Krah- 


mer (6 @, PC); Chovellén, Maule, 
5 Xi 1953) L; EB. Penal( 2E€NG): El Coi- 
mo, C, Cimco, ENGI, IL, By deen (Se 
CNC): Fundo Malcho, Cord. Parral, XII- 
195i ple e eeena (202 CNC) samelexcept 
I-1958 (1 2, CNC); Alto de Vilches, 70 km 
E Talca, 5.XII.84—20.11.85, S. & J. Peck (1 
2, CNC); Curanipe, Maule, 4.XII.1953, L. 
E. Pefia (1 2, CNC); Tregualemu, Maule— 
Nuble; 7 Xdi953. ePenandiase ss CNG): 
Butomalal, Cord. Nahuelbuta, Arauco, 10- 
1959) 400mm: ba benders CNG): 
Icalma, 1,000 m, Cord. Lonquimay, 29-XII- 
58, L. E. Pema (1 2, CNC). J. Fernandez 
Isl., Robinson Crusoe Is., Plazoleta—E]l 
Punque, 23-29-I-1992, S. A. Marshall (4 2, 
CNC), same except without “El Punque”’ 
and 24-29-I-1992 (3 2, CNC); J. Fernandez 


828 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 21-22. Aulcus brevis. 21, Mesosoma, dorsal. 22, Mesosoma, lateral. 


VOLUME 107, NUMBER 4 


829 


Figs. 23-27. 


Isl., Robinson Crusoe Is., Bahia Cumber- 
land, 1-11-I-1993, S. A. Marshall, MT (5 
2, CNC), same except 3.1.1993, Malaise 
over creek (1 2, CNC). 

Distribution.—Argentina (Neuquén) and 
Chile from Aconcagua in the north, south 
to Osorno and the Juan Fernandez Islands. 

Etymology.—The species name reflects 
the small size, the smallest aulacid known 
from this region. 

Discussion.—The similar color, size, 
general habitus, short ovipositor, and sculp- 
turation place this species close to A. brev- 
icaudus (Cushman) and A. schiffi Smith 


Pristaulacus capitalis. 23, Head, dorsal. 24, Head, lateral. 25, Head, front. 26, Forewing. 27, 
Dorsal surface of hind coxa. 


from North America. In A. brevicaudus and 
A. schiffi, the malar space is long, half the 
length of the eye height; the lower intero- 
cular distance is 1.4 the eye height; the 
toruli are closer together than the distance 
between the eye and the torulus; the head 
is less punctate; the legs are entirely reddish 
brown; coarse transverse Carinae are present 
on the mesonotum; and cell 1M of the fore- 
wing is larger, only about two times longer 
than wide; and vein M-cu of the forewing 
is distinct. 

No other Neotropical aulacid species is 
comparable. The small size, small forewing 


830 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


4 ewe jar 


tnt Kegs 


Figs. 28-29. Pristaulacus capitalis. 28, Mesosoma, dorsal. 29, Mesosoma, lateral. 


VOLUME 107, NUMBER 4 


oO 
oS) 
— 


Figs. 30-34. Pristaulacus rubriventer. 30, Head, dorsal. 31, Head, lateral. 32, Head, front. 33, Forewing. 
34, Dorsal surface of hind coxa. 


cell 1M, atrophied vein M-cu, and short 
Ovipositor are unique to A. brevis. 
The biology and hosts are unknown. 


Pristaulacus capitalis (Schletterer) 
(Figs. 23—29) 


Aulacus capitalis Schletterer 1889: 520, fig. 
138. 

Pristaulacus capitalis: Kieffer 1902: 12.— 
Elgueta and Lanfranco 1994: 88.—Smith 
2001: 280. 


Diagnosis.—Length about 8—10 mm. En- 
tirely black. Wings hyaline, without black 
spots (Fig. 26). Head dull and finely rugose, 
with fine microsculpture, rugae denser and 
more prominent on frons (Figs. 23-25). 


Mesoscutum with notauli meeting at about 
transscutal articulation (Fig. 28). Upper sur- 
face of hind coxa with cross rugae (Fig. 
27); with ovipositor guide distinct on inner 
surface near apex. Hind prefemur distinct. 
Tarsal claws with 3 teeth and small basal 
lobe. Ovipositor length slightly shorter than 
forewing length. Male unknown. 
Distribution—Recorded only from the 
Region Metropolitana de Santiago by El- 
gueta and Lanfranco (1994). The following 
records extend the distribution from Co- 
quimbo in the north to Linares in the south. 
New records.—CHILE: COLCHAGUA: 
3 km N. Callejones, I-22-1967, E. I. 
Schlinger (1 2; CAS). COQUIMBO: El 


VOLUME 107, NUMBER 4 


Naranjo Tilama, Marzo 1968, Coll: J. Mo- 
lina (1 2, AMNH). LINARES: Fundo 
Malcho, I-1957, L. E. Pema (1 2°, CNC). 
MAULE: Forel Carrizahilo, 250 m, 30 
Janson eby 198i, Le Eo Pena (1 2, 
USNM). SANTIAGO: XII-21—47, L. Pena 
(1 2, AEI); Q. Sn. Ramé6n, I.10.75, Luis 
Pena (10 2, AEI); El Manzama, II-1983, 
900—1,000 m, L. Pefia (1 2, AEI); Rio 
Clarillo Nat. Pk., 23-I-23-II-1989, C. Por- 
ter & Ch. Gonzalez, Malaise trap (1 °, 
FSCA), same except 2-23-I-1989 (1 2, 
FSCA); Cordillera Reserve, Rio Clarillo, 
23-I-23-II-1989 (2 2, FSCA). Province 
unknown: Banos de Cariguenes, 2-II-1890, 
P. Herbst, E.P. Reed Collection (1 2; CAS; 
1 2 MCZ). 

Notes.—The biology and hosts are un- 
known. The holotype is in the Museum fiir 
Naturkunde der Humbolt-Universitat Ber- 
lin, Germany. 


Pristaulacus rubriventer (Philipp) 
(Figs. 30—36) 


Aulacostethus rubriventer Philippi 1873: 
202. 

Aulacus rubriventer: Schletterer 1889: 423. 

Pristaulacus rubriventer: Kieffer 1903: 
455.—Elgueta and Lanfranco 1994: 


87.—Smith 2001: 295. 


Diagnosis.—Length, about 13 mm. 
Head and mesosoma black except antennal 
flagellum, gena, and spot at extreme bases 
of femora reddish brown; metasoma red 
with black spot at base. Forewing hyaline 
with black spot at apex (Fig. 33). Head 
shining, gena with few, widely separated 
punctures; vertex and frons more densely 
punctate with punctures closer together 
than interspaces (Figs. 30-32). Notauli 
meeting behind near transscutal articula- 
tion (Fig. 35). Mesoscutum and scutellum 
with coarse transverse carinae (Fig. 35). 
Hind coxa shining, punctate with faint ca- 
rinae on apical half (Fig. 34); ovipositor 
guide on inner surface distinct, near apex 
of coxa. Hind prefemur distinct. Tarsal 
claws with 4 teeth and a very small basal 


833 


lobe. Ovipositor length subequal to or 
slightly longer than forewing length. Male 
unknown. 

Distribution.—Elgueta and Lanfranco 
(1994) recorded this species from the Re- 
gion Metropolitana de Santiago to VIII Re- 
gion (Nuble). I have seen four specimens 
from the Santiago area. 

New records.—CHILE: SANTIAGO: Q. 
San Ramon, I-10—75, Luis Pefa (2 8°, 
AEI); Lo Cafias, I-1980, Luis Pefia (1 9, 
AED; Q. SN. Ramon, Dec. 1974 (1 @, 
CNC). 

Notes.—Elgueto and Lanfranco (1994) 
stated that the tarsal claw has three teeth. 
The male, biology, and hosts are unknown. 
The neotype, designated by Elgueta and 
Lanfranco (1994), is in the Museo Nacional 
de Historia Natural, Santiago. 


ACKNOWLEDGMENTS 


The cooperation of the following curators 
is very much appreciated: L. Stange and J. 
Wiley (FSCA); W. Pulawski (CAS); M. E. 
Smethurst and J. M. Carpenter (AMNH); D. 
Wahl (AEI); J. T. Huber (CNC), and G. 
Pagliano, Torino, Italy. Cathy Apgar, Sys- 
tematic Entomology Laboratory, USDA 
(SEL) took the photos and arranged the 
plates. I thank N. M. Schiff, U.S. Forest 
Service, Stoneville, MS, and M. A. Solis 
and T. J. Henry, SEL, Washington, DC, for 
review of the manuscript. 


LITERATURE CITED 


Barriga, R. J. E. 1990. Parasitos depredadores de lar- 
vas de Cerambycidae y Buprestidae (Coleoptera) 
de Chile. Revista Chilena de Entomologia 18: 57— 
39. 

Elgueta, M. and D. Lanfranco. 1994. La familia Au- 
lacidae en Chile (Hymenoptera: Evanioidea). Re- 
vista Chilena de Entomologia 21: 85—97. 

Huber, J. T. and M. J. Sharkey. 1993. Chapter 3. Struc- 
ture, pp. 13-59. /n Goulet, H. and J. T. Huber, eds. 
Hymenoptera of the World: An_ Identification 
Guide to Families. Research Branch, Agriculture 
Canada Publication 1894/E, 668 pp. 

Kieffer, J.-J. 1902. Hymenoptera, Fam. Evaniidae. /n 
Wytsman, P., Genera Insectorum, Fascicule 2, 13 
pp, | plate. Bruxelles. 

. 1903. Les Evaniides, pp. 347—482. Jn André, 


834 
E. Species des Hyménopteres d’Europe & 

d’Algérie. Volume 7, part 2, Paris. 

1911. Etude sur les Evaniides exotiques 
(Hym.) du British Museum de Londres. Annals 
Société Entomologique de France 80: 151—230. 

Philippi, R. A. 1873. Chilenische Insekten. Stettiner 
Entomologische Zeitung 34: 296-316. 

Schletterer, A. 1889. Die Hymenopteren-Gruppe der 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Evaniiden. Annalen des K. K. Naturhistorischen 
Hofmuseums 4: 373-546, plates 19-22. 

Smith, D. R. 2001. World catalog of the family Au- 
lacidae (Hymenoptera). Contributions on Ento- 
mology, International 4(3): 263-319. 

Townes, H. K. 1950. The Nearctic species of Gaster- 
uptiidae (Hymenoptera). Proceedings of the Unit- 
ed States National Museum 100: 85-145. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 835-845 


NEW APHIDOIDEA (HEMIPTERA: STERNORRHYNCHA) IN 
BURMESE AMBER 


GEORGE POINAR, JR. AND ALEX E. BROWN 


(GP) Department of Zoology, Oregon State University, Corvallis, OR 97331, U.S.A. 
(e-mail: poinarg@science.oregonstate.edu); (AB) 629 Euclid Avenue, Berkeley, CA 
94708, U.S.A. (e-mail: alex.brown3 @comcast.net) 


Abstract.—Three new aphid taxa (Hemiptera: Sternorrhyncha) are described from Low- 
er Cretaceous amber from Myanmar (Burma). A new family, the Verrucosidae Poinar 
and Brown, is described for the new genus and species Verrucosa annulata Poinar and 
Brown, which is characterized by 3-segmented antennae, with the third segment com- 
posed of 20 annuli, forewing containing only 3 veins radiating from the main vein (Rs, 
M and distal C), and the forewing membrane covered with scalelike warts. Another new 
family, the Burmitaphidae Poinar and Brown, is described for the new genus and 
species Burmitaphis prolatum Poinar and Brown, and the new genus and species Cau- 
linus burmitis Poinar and Brown. This family is characterized by greatly reduced (stub- 
like) hind wings, 7- segmented antennae, and a greatly reduced rostrum and frons with a 
protruding median tubercle. In B. prolata, the forewing has only 3 veins radiating from 
the main vein and the aedaegus is long and highly sclerotized. In C. burmitis, the forewing 
has 4 veins departing from the main vein and an elongate cauda is present. These new 
taxa, together with previously described aphids from Mesozoic deposits, show a high 
degree of morphological diversity in Cretaceous aphids. 


Key Words: Verrucosidae n. fam., Verrucosa n. gen., Verrucosa annulata n. sp., Bur- 
mitaphidae n. fam., Burmitaphis n. gen., Burmitaphis prolatum n. sp., Cau- 


linus n. gen., Caulinus burmitis n. sp., Burmese amber, Cretaceous 


During an investigation of insects in Bur- 
mese amber, several unique members of the 
Aphidoidea were discovered. Since they 
could not be placed in any extant or extinct 
genera, they are described below in two 
new families. Amber from Burma (Myan- 
mar) occurs in lignitic seams in sandstone- 
limestone deposits in the Hukawng Valley. 
Palynomorphs obtained from the amber 
beds where the fossil piece originated have 
been assigned to the Upper Albian (~ 105— 
100 mya) (Cruickshank and Ko 2003). Nu- 
clear magnetic resonance (NMR) spectra of 
amber samples taken from the same locality 
as the fossils indicated an araucarian (pos- 


sibly Agathis) plant source (Lambert and 
Wu, personal communication). 


MATERIALS AND METHODS 


The amber pieces were repolished to bet- 
ter view the specimens. The amber piece 
containing the holotype of Verrucosa an- 
nulata (B-He-13A) is roughly rectangular, 
measures 14 mm long, 8 mm wide and | 
mm deep. The amber piece containing the 
paratype of Verrucosa (B-He- 
13B) is almost cuboid in outline, measuring 
3 mm long, 2 mm wide, and 2 mm deep. 
The amber piece containing the holotype 
and paratype of Burmitaphis prolatum (B- 


annulata 


836 


He-14A, B) is 6 mm long, 4 mm wide and 
2 mm deep. The amber piece containing an- 
other paratype of Burmitaphis prolatum (B- 
He-14C) is 7 mm long, 3 mm wide, and 1.5 
mm deep. The amber piece containing the 
holotype of Caulinus burmitis (B-He-15) is 
7 mm long, 4 mm wide, and 4 mm deep. 
Observations, drawings, and photographs 
were made with a Nikon SMZ-10 R stereo- 
scopic microscope and Nikon Optiphot TM 
compound microscope (with magnifications 
up to 650X). All measurements are in mi- 
crons unless otherwise noted. 


Verrucosidae Poinar and Brown, 
new family 


Description.—With 3-segmented, ex- 
tremely thick antennae composed of deep 
ringlike structures; rostrum very long with 
extremely long apical segment; proximal 
branch of cubitus in fore wing greatly re- 
duced (possibly absent); media with one 
branch; eyes without ocular tubercles 
(triommatidia); forewing membrane cov- 
ered with scalelike warts; siphunculi well 
developed. 

Etymology.—‘“‘Verrucosus”’ is Latin for 
‘full of warts”’ in reference to the wing ar- 
mature. 

Diagnosis.—There is no aphid family, 
extinct or extant, with the above combina- 
tion of forewing characters. While members 
of the Adelgidae and Phylloxeridae also 
have only 3 veins behind the pterostigma 
reaching or almost reaching the wing mar- 
gin, these represent the M, distal Cu, and 
proximal Cu veins (Annand 1928). In the 
Verrucosidae, the three veins are the Rs, M, 
and distal Cu, with the proximal Cu vein 
absent or greatly reduced, possibly repre- 
sented by a faint, indistinct zone at the base 
of the forewings. 

Type genus.—Verrucosa, n. gen. 


Verrucosa Poinar and Brown, 
new genus 


This new genus is based on two alate fe- 
males (B-He-13A; B-He-13B) in separate 
pieces of Burmese amber. One specimen 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(B-He-13B) is poorly positioned in cloudy 
amber and cannot be adequately described, 
other than to verify the structure of the ros- 
trum, antennae and wing venation. The de- 
scription is based on the holotype of the 
new species (B-He-13A), which is well pre- 
served with only the tip of the abdomen 
somewhat distorted and the right hind leg 
and middle leg missing. 

Description.—With characters listed un- 
der family description; antenna 3-segment- 
ed, with third segment composed of 20 an- 
nuli. 

Etymology.—The generic name 1s taken 
from the Latin “‘verrucosus” for “full of 
warts”’ in reference to the wing armature. 
The gender is feminine. 

Diagnosis.—There is no extinct or extant 
aphid genus with the above characters. 

Type species.—Verrucosa annulata, Nn. 


sp. 


Verrucosa annulata Poinar and Brown, 
new species 
(Figs. 1—5) 


Description.—Characters as listed under 
family and generic descriptions. Alate vi- 
viparous female; brown; dorsum densely 
covered with short setae; body length 737. 

Head: Compound eyes ventrally orien- 
tated, composed of 13—15 ommatidia; ocelli 
not observed; rostrum 4-segmented (broken 
between second and third segments), nar- 
row, length 887, all segments bearing setae, 
length stylet bundle 950; antenna 3 seg- 
mented, first and second segments sube- 
qual, second segment appears subdivided 
into two or three overlapping portions; third 
segment composed of 20 closely placed an- 
nuli; rhinaria not observed; length antennal 
segments; 1, 50; 2, 49; 3, 141. 

Thorax: Forewing length 867; width 
411; membrane transparent but covered 
with small crescent-shaped warts; warts 
denser in pterostigma portion of wing; pter- 
ostigma 277 long, 111 wide; radial sector 
departing from subapical part of pterostig- 
ma, media departing from proximal edge of 
pterostigma, with one fork, stem 418, an- 


Fig. 1. 


VOLUME 107, NUMBER 4 


- clan ae 
7 NS 
a“ = 
/ ~ 
\ aaa 
ve : 7 
. \ S 
‘ \ \ % 
f \ \ i ‘ 
( \ \ 1 \ 
i 4 ns \ 
( \ 1 \ 
{ \ f \ 
\ ay, 
/ 
\ 
\ 
a \ 
Ware \ 
nA of . 
ryt) \ 
Sala aA \ 
oa 44R0% r 


a 
a ? 2 - 
a . ia 4 
ry Ss 3 
e -) 5 ry 
t\p tae wr 2A * 
ACr\ anes rs % a4 a Cy Ae HG 
e t 2 Ln ~ o Mn. 
EA: + 1 . 
Lary . 4 x 9 eS 
Op, B * ie 
Ge) ene “ A 
~Ant a XG 7. 5 QD 4) 4) 
Sa es “ a Ay 
a | a \ n 
VG, io po 6 a OWL - * 
7 rN is Cy oO n 
14 ~ OD Os ome 
Sn SAS aS ra vA 
BD 
re wo a 9 as pea 


Lateral view of alate female holotype of Verrucosa annulata in Burmese amber. Note dark oval 


objects. Tip of right forewing is reconstructed based on venation observed in left forewing and in paratype. Bar 
= 126 pm. 


837 


838 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


qe“: 


ne | 


‘ 


a 


+s 


Figs. 2-5. 


Alate female holotype of Verrucosa annulata in Burmese amber. 2, Head region showing third 


antennal segment composed of 20 annuli. Bar = 85 pm. 3, Detail of pterostigma of forewing showing dense 
concentration of scalelike warts on wing surface. Bar = 


46 pm. 4, Detail of basal portion of forewing showing 
claval fold (arrow). Bar = 40 pm. 5, Spherical object adjacent to aphid. Bar = 33 pm. 


VOLUME 107, NUMBER 4 


terior branch 191, posterior branch 111, dis- 
tal cubitus slightly curved in middle, de- 
parting from main vein 161 from root of 
media; claval fold near wing base, length 
101, width, 10; hind wings not observed; 
legs long, femora and tibia with setae; tarsi 
2-jointed, basal segment short; setae on 
fore- and mid-tarsi; tarsal claws 2, equal, 
simple, slightly curved. 

Abdomen: With at least 6 segments; one 
definite siphunculus observed on left side 
of body, other siphunculus partly obscured, 
no sculpturing evident, length 73, diameter 
of opening 24; oval objects (N = 2) adja- 
cent to specimen, length 63—68; width 40— 
45; surface covered with raised granules. 

Material examined.—Holotype female in 
Burmese amber from the Hukawng Valley, 
deposited in the Poinar amber collection 
maintained at Oregon State University (ac- 
cession number B-He-13A). One paratype, 
same data and depository (number B-He- 
13B). 

Etymology.—The specific epithet “‘an- 
nulata”’ pertains to the annulated condition 
of the third antennal segment. 

Comment.—tThe oval objects adjacent to 
the fossil are similar in size, shape and ar- 
mature and occur nowhere else in the am- 
ber. Could they represent eggs of V. annu- 
lata? The objects are smaller than most 
aphid eggs (Miyazaki, 1987), yet Essig 
(1917) described the egg of Myzocallis 
arundicolens (Clark) as 70 wm in length, 
which is in the size range of the oval ob- 
jects (63-68 ppm). However, aphid eggs are 
normally smooth and these objects have def- 
inite surface irregularities. They could also 
be pollen grains introduced by the aphid 
and may indicate the plant host. Vesiculate 
wingless pollen grains in this size range are 
produced by some conifers, as the North 
American Tsuga canadensis (L.) Carr. 
(Kapp et al. 2000). 


Burmitaphidae Poinar and Brown, 
new family 


Description.—Hind wing reduced to 
stubs; antenna 7-segmented, very short; 


839 


frons with protruding median tubercle; me- 
dia with one branch; rostrum absent: si- 
phunculi absent. 
Etymology.—Burmitaphididae is based 
on the country of origin of the fossils. 
Diagnosis.—The greatly reduced, stub- 
like hind wings separate this family from 
all other extant and extinct aphid genera. 
Type genus.—Burmitaphis, n. gen. 


Burmitaphis Poinar and Brown, 
new genus 


The description is based on two males 
(B-He-14A; B-He-14B) in a single piece of 
amber and one male (B-He-14C) in a sec- 
ond piece of amber. Two specimens (B-He- 
14B and B-He-14C) are poorly preserved- 
however,they have all the diagnostic char- 
acters of the family and genus. The generic 
and specific descriptions are based mainly 
on the holotype (B- He-14A). 

Description.—With characters listed un- 
der family description. Mouthparts atro- 
phied, rostrum absent; with extended cop- 
ulatory organ; forewing with 3 veins de- 
parting from main vein. 

Etymology.—‘‘Burmit-” represents the 
country of origin of the specimen. The gen- 
der is neuter. 

Type species.—Burmitaphis prolatum, n. 
sp. 

Diagnosis.—There is no aphid family, 
extinct or extant with the above combina- 
tion of characters. Especially significant are 
the reduced hind wings. 


Burmitaphis prolatum Poinar and 
Brown, new species 
(Figs. 6, 8—10) 


Description.—Alate male, with charac- 
ters as listed under family and generic de- 
scriptions; general coloration brown; body 
length 832. 

Head: Width across eyes 189; longitu- 
dinal diameter of eye 76; ocelli not ob- 
served; antenna 7- segmented; length seg- 
mentse le SOs 2 Gs) oy) Os 25 2 See 
23: 7, 43; rostrum not observed, greatly re- 
duced or absent. 


840 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Dorsal view of holotype male of Burmitaphis prolatum in Burmese amber. Bar = 252 wm. Upper 
left shows detail of copulatory organ. Bar = 45 wm. 


VOLUME 107, NUMBER 4 


Thorax: Width prothorax 246; length 
forewing 1090; greatest width forewing 409 
(378); membrane transparent but covered 
with small crescent-shaped warts; warts 
denser in pterostigma portion of wing; with 
three well-developed veins; radial sector 
departing from subapical edge of pterostig- 
ma (root not visible), length 88; media de- 
parting from proximal edge of pterostigma 
(root not visible), stem 277 long, anterior 
branch 271 long, posterior branch 164 long; 
distal cubitus (root not visible) departing 
from main vein, 170 from base of media; 
claval fold (158 long by 19 wide) in basal 
part of wing; hind wing represented by nar- 
row stub, 100 (138) long and 19 wide; legs 
short, with retrose hairs running length of 
all tibiae and on some portions of femora; 
tarsi 2 segmented; claws well developed, 
without basal thickening. 

Abdomen: Short, nearly as wide as long, 
334 long (without copulatory organ), 315 
wide; siphunculi not visible; copulatory or- 
gan elongate, directed ventrally at about 
45°; 193 long, flanked by a pair of shorter 
lobes (claspers, parameres?) 34 long; tip of 
copulatory organ slightly swollen. 

Material examined.—Holotype male in 
Burmese amber from the Hukawng Valley, 
deposited in the Poinar amber collection 
maintained at Oregon State University (ac- 
cession number B-He-14A). Two paratypes 
(B-He-14B, B-He-14C), same data and de- 
pository. 

Etymology.—The specific epithet “‘pro- 
latus’”’ is from the Latin “‘prolatus” for ex- 
tended, elongated in reference to the long 
copulatory organ. 

Comment.—Males, even today, are con- 
sidered rare and finding three of the same 
type in one amber deposit is interesting and 
may indicate that parthenogenetic reproduc- 
tion had not yet occurred in this genetic line 
(Heie 1987), or that at this stage of their 
evolutionary development, sexual repro- 
duction was very important as a means of 
obtaining a high degree of genetic diversity 
to cope with the changing landscape (evo- 
lution of Angiosperms). 


841 


Caulinus Poinar and Brown, new genus 


The holotype of the type species (B-He- 
15) is essentially complete with only the 
left hind leg and tip of the cauda missing. 
While the right forewing is present, it is po- 
sitioned in a fracture and could not be clear- 
ly portrayed. 

Description.—Antenna 7-segmented, 
third segment longest; rostrum absent; fore- 
wing with 4 veins departing from main 
vein; media with one branch; small scale- 
like warts on pterostigma and along outer 
margin of forewing; hind wing reduced to 
stub; cauda elongate (tip missing); siphun- 
culi absent. 

Etymology.—‘“‘caulinus”’’ is from the 
Latin ‘“‘caulis’’ for stem or stalk referring to 
the reduced hind wings. The gender is mas- 
culine. 

Type species.—Caulinus bursitis, n. sp. 

Diagnosis.—The wing venation and an- 
tennae are similar to some members of the 
subfamily Pemphiginae. However, the re- 
duced hind wings separate this genus from 
all extinct and extant aphids. 


Caulinus burmitis Poinar and Brown, 
new species 
(Figs. 7, 11) 


Description.—Holotype alate female; 
with characters listed under generic descrip- 
tion. Generally brown, body length 1900. 

Head: With 3 ocelli; compound eyes 
prominent, longitudinal diameter of com- 
pound eye 98; width across compound eyes 
357; rostrum absent; antenna 7-segmented, 
first and second and third and fourth seg- 
ments closely appressed; length of seg- 
ments, 55.92, 42> 5200; 45 98-c5. 1.6; 
84: 7, 119; all visible segments with mul- 
tiple rings except first and second one each 
with only a few rings. 

Thorax: With anteriolateral angles pro- 
jecting outward; length forewing 2000; 
greatest width forewing 805; membrane 
transparent but outer margin covered with 
small scalelike warts; warts denser in pter- 
ostigma portion of wing; pterostigma 511 


842 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


long, 189 wide; radial sector (root not vis- 
ible) departing from subapical part of pter- 
ostigma, 679 long; media (root not visible) 
departing from proximal edge of pterostig- 


\ 


Fig. 7. Ventral view of holotype of Caulinus burmitis in Burmese amber. Bar = 280 ym. 


ma with one fork, stem 448 long, anterior 
branch 531 long, posterior branch 357 long; 
distal cubitus curved at apex, departing 
from main vein 280 from root of proximal 


843 


VOLUME 107, NUMBER 4 


10 


Figs. 8-11. Aphidoidea in Burmese amber. 8. Ventral view of holotype male of Burmitaphis prolatum. Bar 
= 354 um. 9, Detail of basal portion of wing of B. prolatum showing elongate claval fold (arrow) in forewing 
and narrow stublike hind wing (arrowhead). Bar = 45 wm. 10, Copulatory organ of B. prolatum. Arrows show 
basal lobes (parameres, claspers?). Bar = 43 pm. 11. Ventral view of holotype of Caulinus burmitis. Bar = 388 


wm. 


844 


branch, 880 long; proximal cubitus well de- 
veloped, as thick as other veins, slightly 
curved, 350 long, forming nearly 90° angle 
with main vein; faint vein or fold at base 
of main vein, 91 long; no claval fold at base 
of wing; hind wing represented by narrow 
pointed stub, 304 long, no associated mem- 
brane observed; legs relatively short, all tib- 
iae bearing long setae; tarsi 2-segmented; 
claws well-developed, with enlarged base. 

Abdomen: Length 1120, width 476; si- 
phunculi absent; posterior end with elon- 
gate cauda (tip missing). 

Material examined.—Holotype female in 
Burmese amber from the Hukawng Valley, 
deposited in the Poinar amber collection 
maintained at Oregon State University (ac- 
cession number B-He-15). 

Etymology.—The specific epithet “‘bur- 
mitis’”’ is from the country of origin of the 
fossil. 

Diagnosis.—This species has some char- 
acteristics of the subfamily Pemphiginae; 
i.e. wing venation, antennal structure (ex- 
cept for an extra segment) and the absence 
of both siphunculi and a rostrum. The scale- 
like warts along the margin of the forewing 
closely resemble those of the extant Ervio- 
soma lanigerum (Hausmann) (Baker 1915). 


DISCUSSION 


The antennae of the fossil forms reported 
here are unusual. The maximum number of 
antennal segments in extant aphids is 6 (Mi- 
yazaki 1987), which is one less than is 
found in members of the Burmitaphidae. 
Certainly, there is no extant or extinct aphid 
which has the annulated antennal condition 
of V. annulata, and this clade probably be- 
came extinct in the Cretaceous. 

The claval fold, which actually appears 
as a closed cell along the posterior margin 
of the forewing, is posteriorly positioned in 
V. annulata and B. prolatum in comparison 
with those of extant aphids. In fact, in B. 
prolatum, it is positioned at the very base 
of the wing, possibly reflecting the reduced 
condition of the hind wings. On the anterior 
border of the shortened hind wings of B. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


prolatum are some roughened areas that 
could represent vestigial hamuli (Miyazaki 
1987). The location and size of the claval 
fold differs in B. prolatum and V. annulata, 
suggesting that these features could be used 
as diagnostic characters in aphids. 

The presence of reduced stublike hind 
wings in B. prolatum and C. burmitis is in- 
teresting, since this is the first record of this 
degree of hind wing reduction in aphids 
(Miyazaki 1987). At first, it was thought 
that the stubs represented the remains of 
damaged wings. However, on both species, 
the opposing appendages are similar in size 
and shape, and there is no evidence of torn 
wing membranes associated with them. As 
mentioned previously, the basal position of 
the claval fold in B. prolatum and the ab- 
sence of a claval fold in C. burmitis tend to 
support the observations that the hind 
wings are greatly reduced. Such reduced 
wings could be considered hamulohalters, 
which are reduced metathoracic wings in 
male coccids. The Burmitaphidae would 
then represent an ancient group, apparently 
disappearing in the Cretaceous, in which 
the hind wings were reduced to hamulohal- 
ters. 

The plant hosts of these aphids are un- 
known, especially since thus far, only two 
plant groups have been identified from Bur- 
mese amber, a member of the Araucari- 
aceae, on the basis of NMR analysis of the 
amber and a grasslike monocot with char- 
acteristics of a bamboo (Poinar 2004). If the 
oval objects adjacent to V. annuata are pol- 
len grains and can be identified further, they 
might indicate another plant host group in 
the Burmese amber forest. 

On the basis of morphology, the Burmi- 
taphidae would appear to be most closely 
related to extant members of the subfamily 
Pemphiginae. It is interesting that the only 
extant aphids that lack a rostrum are the 
sexuales of the Pemphiginae and the males 
of Stomaphis Walker (Lachnidae) (Miya- 
zaki 1987). Some extant species of pem- 
phigines, such as Smynthurodes betae West- 


VOLUME 107, NUMBER 4 


wood, include bamboos in their range of 
plant hosts (Blackman and Eastop 1994). 

The present fossils, together with others 
from a variety of Mesozoic fossil sites 
(Heie 1996; Heie and Wegierek 1998; Heie 
and Pike 1992; Richards 1966; Essig 1938; 
Heie and Azar 2000; Kononova 1975, 
1976, 1977; Wegierek 2000; Zhang 2000) 
show that aphids were quite diverse in the 
Cretaceous. 


ACKNOWLEDGMENTS 


The authors thank Roberta Poinar for ed- 
itorial comments on earlier drafts of this 
manuscript. We also thank two anonymous 
reviewers for their helpful comments. 


LITERATURE CITED 


Annand, P. N. 1928. A contribution toward a mono- 
graph of the Adeline (Phylloxeridae) of North 
America. Stanford University Publications, Uni- 
versity Series, Biological Sciences 6: 1-146. 

Baker, A. C. 1915. Identity of Eriosoma pyri. Journal 
of Agricultural Research 5: 1115-1120. 

Blackman, R. L. and V. EF Eastop. 1994. Aphids on 
the World’s trees. CAB International, Wallingford, 
UK, 986 pp. 

Cruickshank, R. D. and K. Ko. 2003. Geology of an 
amber locality in the Hukawng Valley, northern 
Myanmar. Journal of Asian Earth Sciences 21: 
441-455. 

Essig, E. O. 1917. Aphididae of California. University 
of California Technical Bulletin 1: 301-346. 

. 1938. Family Aphididae, pp. 17—21. In Car- 
penter, E M. et al., eds. Insects and Arachnids 
from Canadian amber. Toronto, University of To- 
ronto Series 40. 

Heie, O. E. 1987. Palaeontology and Phylogeny, pp. 
367-391. In Minks, A. K. and P. Harrewijn, eds. 
Aphids, Their Biology, Natural Enemies and Con- 
trol, Vol. A. Elsevier, Amsterdam. 

. 1996. Palaeoaphididae and Tajmyraphididae 

in Cretaceous amber from Alberta, Canada (He- 

miptera: Aphidinea). Annals of the Upper Silesian 

Museum, Entomology 6—7: 97—103. 


845 


Heie, O. E. and D. Azar. 2000. Two new species of 
aphids found in Lebanese amber and a revision of 
the family Tajmyraphididae Kononova, 1975 (He- 
miptera: Sternorrhyncha). Annals of the Entomo- 
logical Society of America 93: 1222-1225. 

Heie, O. E. and T. Pike. 1992. New aphids in Creta- 
ceous amber from Alberta, Canada (Insecta: Ho- 
moptera). The Canadian Entomologist 124: 1027— 
1053. 

Heie, O. E. and P. Wegeirek. 1998. A list of fossil 
aphids (Homoptera: Aphidinea). Annals of the 
Upper Silesian Museum, Entomology 8—9: 159— 
192. 

Kapp, R.O., O.K. Davis, and J. E. King. 2000. Pollen 
and spores. 2nd Edition. American Association of 
Stratigraphic Palynologists: College Station, Tex- 
as, 279 pp. 

Kononova, E. L. 1975. A new family of aphids (Ho- 
moptera, Aphidinea) from the Upper Cretaceous 
of the Taymyr. Entomologicheskoe Obozrenie 54: 
60-68 (English translation, Entomological Re- 
view 54: 60—68). 

1976. Extinct aphid families (Homoptera, 

Aphidinea) from the Late Cretaceous. Paleonto- 

logical Zhurnal 3: 117—126 (in Russian). 

. 1977. New species of aphids (Homoptera, 
Aphidinea) from the Upper Cretaceous deposits of 
Taimyr. Entomologicheskoe Obozrenie 56: 72—80 
(English translation, Entomological Review 56: 
72-80). 

Miyazaki, M. 1987. Morphology of aphids, pp.1—25. 
In Minks, A. K. and P. Harrewijn, eds. Aphids, 
Their Biology, Natural Enemies and Control, Vol. 
A. Elsevier, Amsterdam. 

Poinar, Jr, G. O. 2004. Programinis burmitis gen. et 
sp. nov., and P. laminatus sp. noy., Early Creta- 
ceous grass-like monocots in Burmese amber. 
Australian Systematic Botany 17: 497-504. 

Richards, W. R. 1966. Systematics of fossil aphids 
from Canadian amber (Homoptera: Aphididae). 
The Canadian Entomologist 98: 746-760. 

Wegierek, P. 2000. A new genus and species of aphid 
(Hemiptera: Aphidinea) from Cretaceous amber, 
pp 141-145. In Grimaldi, D., ed. Studies on Fos- 
sils in Amber with Particular Reference to the 
Cretaceous of New Jersey, Backhuys. Leiden. 

Zhang, J.-E 2000. On problems of classification and 
descriptions of genera and species of Chinese 
aphids from the Late Mesozoic. Acta Palaeonto- 
logica Sinica 39: 558-572. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 846-850 


A NEW SPECIES OF THE GENUS CONASPIDIA KONOW 
(HYMENOPTERA: TENTHREDINIDAE) FROM JAPAN, 
WITH A KEY TO THE JAPANESE SPECIES 


ICH TOGASHI 


1-chome, Tsurugihonmachi, Hakusan-shi, Ishikawa Prefecture 920-2121, Japan 


Abstract.—Conaspidia ishikawai, n. sp., from Honshu, Japan, is described and illus- 
trated. A key is provided for the five Japanese species of the genus. 


Key Words: 


Conaspidia Konow is a genus of the sub- 
family Tenthredininae and includes about 
20 species in eastern and southeastern Asia. 
Nineteen species were recorded by Wei and 
Nie (1997). In Japan, four species were 
treated by Togashi (1984): C. flavipes 
(Smith, 1874), C. guttata (Matsumura, 
1912), C. hyalina Sato, 1933, and C. mu- 
rotai Togashi, 1984. 

Wei and Nie (1997) divided the 19 spe- 
cies of Conaspidia into five groups based 
on the presence or absence of the macula- 
tion of the forewing, shape of the front mar- 
gin of the clypeus, or the form of the post- 
ocellar area. Accordingly, the Japanese spe- 
cies fall into two of the groups, the sikki- 
mensis group which lacks dark maculae on 
the forewing and the scutellaris group 
which has dark brown maculae on the fore- 
wing. 

In 2004, I captured two females belong- 
ing to the scutellaris group at Mt. Hakusan 
in Honshu. These specimens are very close 
to C. guttata, but they are distinguished 
from C. guttata by the straight radial cross- 
vein (2r) of the forewing, the short petiole 
of the anal cell of the hind wing, and by 
the shape of the sawsheath and serrulae of 
the lancet. Thus, I concluded that these 
specimens represent a new species. I de- 
scribe and illustrate this species below and 


Hymenoptera, Symphyta, Tenthredininae, Conaspidia, new species, Japan 


give a revised key to the Japanese species 
of Conaspidia. 


KEY TO THE JAPANESE SPECIES OF 
CONASPIDIA (FEMALES) 


1. Forewing with dark brown maculae (Fig. 1) 


(GeutellarissstOup) meee ee DD 
— Forewing entirely hyaline, without dark brown 
maculae (sikkimensis group) ............ 4 


i) 


. Mesonotum almost entirely black; mesopleu- 
roniblack -2 5 35 eas. weet aoe ea tee epee ee 3) 
— Mesonotum yellowish brown with five black 
maculae; upper half of mesopleuron yellowish 
DrOWM 2 2c. ein Ges bc ae pais ea eae murotai Togashi 
3. Posterior margin of lateral lobes of mesoscu- 
tum reddish brown; front margin of mesoscu- 
tellum angular and acute (Fig. 9); radial cross- 
vein (2r) of forewing curved (Fig. 10); saw- 
sheath as in Fig. 16 ...... guttata (Matsumura) 
— Lateral lobes of mesoscutum black; front mar- 
gin of mesoscutellum rounded and obtuse (Fig. 
3); radial crossvein (2r) of forewing nearly 
straight (Fig. 5); sawsheath as in Fig. 13 
do fiscveatdesptan Eco top aps hay ngs aes c ray ee ishikawai, n. sp. 
4. Mesonotum mostly yellowish brown; basal 
half of stigma of forewing yellowish brown, 
apical half black .............. hyalina Sato 
— Mesonotum black with a small yellowish spot 
on scutellum; stigma of forewing black 
BN eae Oar os CO athe SPO Ul ETA flavipes (Smith) 


Conaspidia ishikawai Togashi, 
new species 
(Figs. 1-8, 13-15) 


Female.—Length, 10 mm. Head and an- 
tenna yellowish brown with following dark 


VOLUME 107, NUMBER 4 


847 


I 


Fig. 1. 


Conaspidia ishikawai, holotype. 


brown to black: ocellar basin, lateral side of 
frontal area, malar space, and apical half of 
mandible. Thorax dark brown to black with 
following yellowish brown: pronotum, te- 
gula, postspiracular sclerite, central portion 
of lateral lobe of mesoscutum, and posterior 
margin of lateral lobe of mesoscutum. Ab- 
domen yellowish brown with following 
dark brown to black: basal half of Ist ter- 
gite, front margin of 2nd tergite, 4th to 6th 
tergites, and sawsheath. Wings hyaline with 
three dark brown maculae (Fig. 1), veins 
yellow, stigma almost entirely black. Legs 
yellow. 

Head: Dilated behind eyes (Fig. 2); post- 
ocellar area nearly subquadrate; OOL:POL: 
OCL = 1.5:1.0:1.7; interocellar furrow dis- 
tinct and deep; postocellar and lateral fur- 


rows distinct; frontal area concave; median 
fovea distinct, deep, and circular; lateral fo- 
vea distinct, with a conical projection in 
middle; antenno-ocular distance nearly as 
long as distance between antennal sockets; 
clypeus slightly convex, front margin emar- 
ginate; postorbital groove distinct; postgen- 
al carina distinct; malar space narrow. An- 
tenna shorter than costa of forewing (ratio 
1.0:1.2); relative lengths of segments about 
B03 U2e5 2S sile7AeSr ike 1k2: Wpedicel 
nearly as long as wide. 

Thorax: Mesoscutellum rather flattened, 
front margin rounded (Fig. 3); cenchrus 
small, distance between them about 4X as 
long as breadth of one. Venation of fore- 
wing as in Fig. 5; radial crossvein (2r) near- 
ly straight; petiole of anal cell of hind wing 


Q 9 3 
= 


Figs. 2-12. 


6 we 


2-8, Conaspidia ishikawai, holotype. 2, Head, dorsal view. 3, Mesoscutellum, dorsal view. 4, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Mesepisternum, lateral view. 5, Forewing. 6, Hind tarsus, lateral view. 7, Tarsal claw, lateral view. 8, Inner 
foretibial spur, lateral view. 9-12, C. guttata. 9, Mesoscutellum, dorsal view. 10, Forewing. 11, Tarsal claw, 


lateral view. 12, Inner foretibial spur, lateral view. 


shorter than nervulus (cu-a) (ratio 1.0:1.7). 
Legs with hind tibia slightly longer than 
hind tarsus (ratio 1.0:0.9); hind basitarsus 
nearly as long as following 4 segments 
combined; inner foretibial spur as in Fig. 8; 
tarsal claws as in Fig. 7. 

Abdomen: Sawsheath as in Fig. 13; lan- 
cet with 22 serrulae (Figs. 14, 15). 

Punctation: Head covered with fine se- 
tigerous punctures; frontal area and lower 
half of inner orbits covered with medium- 
sized punctures and closely punctured; 
clypeus and labrum moderately, sparsely, 
and shallowly punctured; mesoscutum cov- 
ered with fine and sparse punctures, inter- 
spaces between punctures impunctate, shin- 
ing, but posterior portion of lateral lobes of 
mesoscutum moderately, rather closely, and 
distinctly punctured, interspaces of punc- 
tures subequal to puncture diameters; me- 


soscutellum nearly impunctate, shining, but 
posterior portion sparsely and distinctly 
punctured, interspaces between punctures 
impunctate, shining; mesoscutellar append- 
age and metascutellum nearly impunctate, 
shining; upper half of mesepisternum dis- 
tinctly, closely, and reticulately punctured, 
lower third of mesepisternum moderately 
and sparsely punctured (Fig. 4). Abdominal 
tergites nearly impunctate, shining. 

Male.—Unknown. 

Food plant—Unknown. 

Distribution.—Japan (Honshu). 

Type material.—Holotype: female, Mt. 
Hakusan (altitude: 1,300—1,500 m), Ishi- 
kawa Prefecture, Honshu, Japan, 
17.V1I.2004, I. Togashi leg. Deposited in the 
collection of the National Science Museum 
(Nat. Hist.), Tokyo. Paratype: Same data as 
holotype except 15.VI.2004, T. Ishikawa 


VOLUME 107, NUMBER 4 849 


16 


14 


ee ee 


10th idea 12 icla 


17 


ee ee 


10th ier deth 


Figs. 13-18. 13-15, Conaspidia ishikawai, paratype. 13, Sawsheath, lateral view. 14, Apical portion of 
lancet. 15, 10th—13th serrulae of lancet. 16-18, C. guttata. 16, Sawsheath, lateral view. 17, Apical portion of 
lancet. 18, 10th—12th serrulae of lancet. 


850 


leg (1 2). Deposited in the National Mu- 
seum of Natural History, Smithsonian In- 
stitution, Washington, D.C. 

Remarks.—This species may be separat- 
ed from other Japanese species of Conas- 
pidia in the preceding key. It is very closely 
allied to C. guttata, but it is distinguished 
from the latter by the black posterior mar- 
gin of the lateral lobes of the mesoscutum 
(reddish brown in C. guttata), by the round 
and obtuse anterior margin of the meso- 
scutellum (angular and acute in C. guttata, 
see Figs. 3, 9), by the shape of the inner 
foretibial spur (see Figs. 8, 12), by the al- 
most straight radial crossvein of the fore- 
wing (curved in C. guttata, see Figs. 5, 10), 
by the shape of the sawsheath (see Figs. 13, 
16), and by the structure of the serrulae of 
the lancet (see Figs. 14, 15, 17, 18). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ACKNOWLEDGMENTS 


I express my thanks to David R. Smith, 
Systematic Entomology Laboratory, USDA, 
Washington, D.C., for reviewing the man- 
uscript. I also thank Mr. Takuya Ishikawa, 
Ishikawa Insect Museum, Yahata machi, 
Hakusan-shi, Ishikawa Prefecture, for lend- 
ing me the valuable specimens. 


LITERATURE CITED 


Togashi, I. 1984. The sawfly genus Conaspidia Konow 
in Japan (Hymenoptera: Tenthredinidae). Konta 
52: 580-584. 

Wei, M. and H. Nie. 1997. Studies on the genus Con- 
aspidia Konow (Hymenoptera: Tenthredinidae) 
from China with a key to known species of the 
world. Entomotaxonomia 19 (Supplement): 95— 
117. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 851-858 


NOTES ON THE FRUIT FLIES (DIPTERA: TEPHRITIDAE) OF CALIFORNIA 
JUDITA KINKOROVA 


Department of Zoology, Charles University, Praha, CZ-128 44 Czech Republic (e-mail: 
kinkor @natur.cuni.cz) 


Abstract.—The tephritid fauna in California was studied in May and June 2002 on a 
two-week field trip and in September 2003 during a 16-day round trip through most of 
California. Records are given for 22 species of Tephritidae from 26 collection sites. Es- 
tablishment of three introduced species, Urophora jaculata Rondani, Terellia fuscicornis 
(Loew), and Chaetorellia succinea (Costa), is verified. Rhynencina longirostris Johnson 


is reported from the western United States for the first time. 


Key Words: 


The current knowledge of tephritid fruit 
fly species in California is based primarily 
on the long-term studies of Goeden and co- 
workers, who published a considerable 
number of papers on the biology and tax- 
onomy of the native species. These works, 
along with Foote et al. (1993), were useful 
in developing and implementing field stud- 
ies to determine the presence of purpose- 
fully and accidentally introduced tephritid 
species in California. 

Nine tephritid species have been intro- 
duced into Canada and the USA since 1969. 
Urophora jaculata Rondani (misidentified 
as U. sirunaseva) was the first bioagent re- 
leased to reduce yellow starthistle, Centau- 
rea solstitialis LL. (Maddox 1981), but it 
failed to establish (Julien and Griffiths 
1999). Urophora affinis (Frauenfeld), U. 
guadrifasciata (Meigen), U. sirunaseva 
(Hering), Chaetorellia australis Hering, C. 
acrolophi White and Marquardt, and Ter- 
ellia virens (Loew) were introduced from 
Europe as biological control agents against 
invasive thistle species. Terellia fuscicornis 
(Loew) was an accidental introduction into 
North America (White and Elson-Harris 
1994) and Chaetorellia succinea (Costa) 


Tephritidae, native, introduced species 


was introduced, presumably as a contami- 
nant of C. australis, into California for con- 
trol of yellow starthistle, (Julien and Grif- 
fiths 1999). The main objective of the cur- 
rent study was to determine post-release es- 
tablishment and geographic extent of the 
above-mentioned species by collecting on 
known host plants throughout California. 


MATERIALS AND METHODS 


Field collections of adult tephritids and 
their host plants were conducted throughout 
California, focusing on the southern and 
eastern portions in 2002, and the western, 
central, and southern portions in 2003. Col- 
lections were made in most major ecosys- 
tems in California and from sea level to 
over 1,500 m. Adult flies were collected by 
sweeping stands of knapweeds and/or this- 
tles. Sites where tephritids were collected 
are given in Table 1. The numbers in pa- 
rentheses in the specimen data sections re- 
fer to the collection numbers for the sites 
in the first column of Table 1. These sites 
are located along a 5,600 km route from 
Los Angeles to San Francisco along Hwy 
1, then inland to Napa Valley. The north- 
ernmost site was Paskenta near Mendocino 


852 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. List of localities with dates and descriptions of collections. 
eee eee 
Collection No. Date Locality 
2002 
i May 27 Riverside Co., Aguanga Rd near Hwy 79, Cleveland National Forest 
ii June 6 Mariposa Co., near Yosemite National Park, Hwy 120 
ili June 8 San Luis Obispo Co., ca. 1.6 km west from intersection of Hwy 101 
and 166 
iv June 8 San Luis Obispo Co., ca. 8 km west from Santa Maria, Rd 166 
Vv June 8 Santa Barbara Co., New Cuyama, Rd 166 Ventucopa Rd 33 
vi June 8 San Bernardino Co., Pine Mountain, Summit, Rd 138 
2003 
1 Sep 6 Ventura Co., Monte Pinos east from Lake of the Woods, east of 
Gorman (unpaved road) 
2 Sep 7 San Luis Obispo Co., near Cambria on Hwy 46 about 16 km before 
Hwy 1 
3 Sep 7 Monterey Co., Santa Lucia Mountains south of Los Padres Rd con- 
necting Jolon to Hwy 1 
4 Sep 7 Monterey Co., Santa Lucia, near the sea shore 
5 Sep 8 Monterey Co., military area in marina close to Salinas 
6 Sep 9 Sonoma Co., Sonoma Hwy 12 
7 Sep 10 Lake Co., south from Clear Lake on Hwy 175 ca. 3 km from Hwy 
101 
8 Sep 10 Mendocino Co., exit Hwy 162 from Hwy 101 
9 Sep 11 Butte Co., near Paradise on Hwy 70 
10 Sep 13 San Joaquin Co., north of Clements on Hwy 88 
11 Sep 13 Amador Co., Silver Lake, Eldorado National Forest 
12 Sep 14 Madera Co., on Hwy 145 ca. 8 km from Madera 
13 Sep 15 Kern Co., Caliente (about 50 km from Bakersfield) on Hwy 58 
14 Sep 15 Kern Co., Tehachapi on Hwy 58 
15 Sep 15 Kern Co., Theachapi city 
16 Sep 15 Kern Co., hills on Hwy 202 ca. 16 km from Tehachapi 
17 Sep 17 Riverside Co., on Hwy 86 ca. 3km beyond Indio 
18 Sep 17 Imperial Co., Salten Sea near desert shores 
19 Sep 17 San Diego Co., close to Hwy 78 and Rd S2 at the entrance to Anza 
Borrego National Park 
20 Sep 20 Orange Co., San Juan Capistrano close to Hwy 74 
National Forest. Sites were visited south RESULTS AND DISCUSSION 


through the San Joaquin Valley with short 
side trips into the Sierra Nevada Mountains. 
After crossing the Mohave Desert, the route 
continued across San Bernardino National 
Forest, down to the Salton Sea, over to 
Anza Borrego, and finally returning to Los 
Angeles (Fig. 1). NATIVE SPECIES 
Collected fly specimens were pinned and Subfamily Tephritinae 
identified using Foote et al. (1993). Voucher pre 
specimens are deposited in the Charles Uni- Tribe Terelliini Hendel 
versity Collection, Prague. Plant names fol- Neaspilota signifera (Coquillett).—Near 
low Hickman (1993). Paradise at Hwy 70 (9), Sep 11, 2003, 5 °; 


A total of 53 specimens were collected 
in 2002, and 259 specimens were collected 
in 2003. Altogether 22 species were col- 
lected and identified, three of which were 
species introduced from Europe. 


VOLUME 107, NUMBER 4 853 


Capon 


ig & 


aes 
aie 


San Grtses 6 FS MC Barolo: 
Francisco) eaidakla Yi 


RS 
Ban 
Wana 
SUPA =: Ye i 


=) 


Sen Mguyt 


| rapes Ss a 


Los Angeles: j 


Sra Barer mrance AES a ‘ 
Long Beach \'S. 


- sia caairss) 
San neolas pera Laon Deacey 
wlan Qa Yrter 03 
ae Coens Sen 
Guiles Onn 


Ont Mar 
' S J 

San Diego ¢ 
Coronado” 


PACIFIC OCEAN 


* = collections in 2002, o = collections in 2003. 


Fig. 1. Map of California showing collection localities. 


854 


north of Clements at Hwy 88 (10), Sep 13, 
2003, 1 2; at Hwy 145 ca. 8 km from Ma- 
dera (12), Sep 14, 2003, 3 6, 2 2; hills at 
Hwy 202 ca. 16 km from Tehachapi (16), 
Sep) 153 20035152" 

This is the only species in the genus lack- 
ing supra-alar bristles. Two host plants were 
recorded by Freidberg and Mathis (1986), 
Baeria Fremontii Hoover and Hemizonia 
pungens (Hook & Arn.) Torr. & A. Gray. 
Adult flies were swept from dry plants 
close to Paradise and Clements bordering 
Camanche National Forest. Five specimens 
were collected north of Fresno, near Ma- 
dera, and one specimen on the hills at Hwy 
202 near Tehachapi. Neaspilota signifera is 
a commonly collected species in California. 

Neaspilota wilsoni Blanc and Foote.— 
Santa Maria Hwy close to Hwy 166 and 
Hwy 101, (iii), Jun 8, 2002, 1 3d; Santa Lu- 
cia Mountains south of Los Padres Rd con- 
necting Joion to Hwy | (3), Sep 7, 2003, 3 
Creo ae 

This species was collected only at two 
localities. The only known host plant is 
Haploppapus squarosus (Hook & Arn.) = 
Hazardia squarrosa (Hook & Arn.) E. 
Greene. Neaspilota wilsoni is not as com- 
mon as N. signifera, and was collected at 
the only previously known locality (Foote 
et al. 1993). The species is considered uni- 
voltine; adults occur from June to August, 
but sometimes with a partial second gen- 
eration (Goeden and Headrick 1999). Spec- 
imens were collected in August 2002 and 
September 2003, so apparently a second 
generation developed in 2003. 


Tribe Myopitini Bezzi 


Rhynencina longirostris Johnson.—Santa 
Lucia Mountains south of Los Padres Rd 
connecting Jolon to Hwy 1 (3), Sep 7, 
MTOB, ll. S. 

Rhynencina is easily distinguished from 
other North American genera by the face in 
lateral view is slanted forward at an angle 
to the vertical axis of the head, and the an- 
terior oral margin produced anteriorly well 
beyond the anterior margin of the parafacial 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(Foote et al. 1993). The only known host 
plant, Smallanthus uvedalius (L.) Macken- 
zie ex. Small (Asteraceae), was only re- 
cently discovered by Steck et al. (2003). 

This collection represents a new species 
and genus record for California. The pre- 
viously known distribution included only 
the eastern states of Georgia, North Caro- 
lina, Pennsylvania, and Maryland. 


Tribe Dythricini Hendel 
Subtribe Cecidocharina Hering 


Procecidochares spp.—P. minuta (?) Te- 
hachapi at Hwy 58 (14), Sep 15, 2003, 1 
2; P. (2?) sp., a small species with yellow 
femora, Salton Sea Lake near desert shore 
lS), Seo 17, ZOOS, 1S, 1-2. 

This is a species-rich genus in the United 
States, but the specimens collected in the 
current study were not determined to spe- 
cies level because of their poor condition 
and because of the many sympatric or cryp- 
tic species. 


Tribe Tephritini 


Dioxyna picciola (Bigot).—Santa Lucia, 
near the seashore (4), Sep 7, 2003, 1 c. 

This is one of two American species in 
this genus, It is widespread with a cosmo- 
politan distribution and recorded from 21 
host plant species in five genera. 

Euaresta stigmatica Coquillett—Calien- 
te (about 50 km from Bakersfield) at Hwy 
58 (13), Sep 15, 2003; 10) G; 6 2: Salton 
Sea Lake near desert shore (18), Sep 17, 
2003, 1 3; close to Hwy 78 and Road S2 
at entrance to Anza Borrego National Park 
(19), Sep 17, 2003, 6 d. 

The specimens were collected on Ambro- 
sia spp. in September 2003. Both the fly 
and host plants are common in California. 
The fly is bivoltine and adults occur from 
mid-June to late September (Headrick et al. 
I). 

Neotephritis finalis (Loew).—Yosemite 
IRGl om Jehan, 20 Gn), Un 1, 2002, | So, 2 &. 

Neotephritis finalis is the most common- 
ly encountered tephritid in the United States 
and Canada (Foote et al. 1993). Two spe- 


VOLUME 107, NUMBER 4 


cies of Neotephritis occur in the United 
States, N. finalis and N. rava Foote, and 
they easily can be recognized by the color 
of scutum, and distinct or indistinct wing 
pattern, respectively (Foote et al. 1993). 

Campiglossa genalis (Thomson).—Miili- 
tary area in marina close to Salinas (5), Sep 
8, 2003, 31 dg, 12 2; Sonoma Hwy 12 (6), 
Seo 9, 2003, Zo. Il 2. 

Campiglossa genalis is a widespread spe- 
cies recorded from Canada, e.g., from Brit- 
ish Columbia, Alberta, and Saskatchewan 
(Foote et al. 1993), on a variety of host 
plants in the genera Haplopappus, Senecio, 
Eriophyllum, Layia, Madi, and Venegasia. 
It was collected in great numbers on Hap- 
lopappus spp. at both localites. The species 
is multivoltine and very common (Goeden 
et al. 1994). 

Campiglossa_ variabilis (Doane).—Fish 
Camp, close to Yosemite National Park (11), 
Jun 6, 2002, 1 6. 

This specimen was collected at a high el- 
evation near Yosemite National Park, as 
also reported in Foote et al. (1993). 

Tephritis araneosa (Coquillett)—Santa 
Lucia Mountaints, near the seashore (4), 
Sep 7, 2003, 4 6, 4 2; Silver Lake, Eldor- 
ado National Forest (11), Sep 13, 2003, 4 d. 

This is the nominate species of the ara- 
neosa complex (araneosa, signatipennis, 
candidipennis, leavittensis, and ovatipen- 
nis) and a very common species in Califor- 
nia. 

Tephritis californica Doane.—Near 
Cambria at Hwy 46 about 16 km before 
lskwny il (@), Seo i, ZAUOS, tl 

Only one specimen was swept in the San- 
ta Lucia Mountains on its known host plant, 
Baccharis sp. 

Tephritis ovatipennis Foote.—Monte Pi- 
nos (1), Sep 6, 2003, 1 3; Silver Lake (11), 
Sep 13; 2003) 2.46, 4 2. 

This is closely related to but not as com- 
mon as 7. araneosa. The only known host 
plants belong to the genera Corethrogyne 
and Erigeron. 

Tephritis signatipennis Foote.—Monte 
Pinos (1), Sep 6, 2003, 1 d; Silver Lake, 


855 


Eldorado National Forest (11), Sep 13, 
ZDOOSR ING Zeer 

Tephritis signatipennis is closely related 
to T. araneosa, also collected at the same 
localities. Specimens were collected only at 
sites within its currently known distribu- 
tion. The only known host plant is Ma- 
chaeranthera canescens (Pursh) Gray 
(Wasbauer 1972), that usually only grows 
at high elevations. All three Tephritis ara- 
neosa complex species were collected at the 
same locality at Silver Lake ca. 2,000 m. 

Tephritis stigmatica (Coquillett).— 
Aguanga Rd near Hwy 79, Cleveland Na- 
tional Forest (i), May 29, 2002, 3 6, 2 @. 

Five specimens were collected as pupae 
in flower heads, but the host plant was not 
determined; reared on June 3, 2002. 

Trupanea bisetosa (Coquillett)—Calien- 
te (about 50 km from Bakersfield) at Hwy 
Sis (18), Seo 15; 2008, i Se 

This species was collected on Helianthus 
sp. It is a common species in this part of 
California. A closely related species, T. ni- 
gricornis, is sympatric in southern Califor- 
nia, but they differ in ecological character- 
istics, such as oviposition behavior, where- 
by females oviposit different number of 
eggs at different sites in different develop- 
mental stages of flower heads (Knio et al. 
1996). 

Trupanea californica Malloch.—Santa 
Lucia Mountaints south of Los Padres Rd 
connecting Jolon to Hwy 1 (3), Sep 7, 
2003, 1 3; military area in marina, close to 
Salinas (5), Sep 8, 2003, 1 6; exit Hwy 162 
from Hwy 101 (8), Sep 10, 2003, 1 6, 1 &. 

Trupanea californica was collected at 
three localities, two with host data: Santa 
Lucia Mountains on plants in the genus 
Gnaphalium and close to Salinas on Gna- 
phalium sp. The species is widely distrib- 
uted through California. It is a multivoltine, 
nondiapausing, and oligophagous species in 
California (Headrick and Goeden 1991). 

Trupanea_ femoralis (Thomson).—Near 
Yosemite National Park, Rd 120 (11), Jun 1, 
2002, 1 6, 3 &; Santa Lucia, near the sea- 
shore (4), Sep 7, 2003, 5 2; Paradise at 


856 


Hwy 70 (9), Sep 11, 2003, 4 ¢; north of 
Clements at Hwy 88 (10), Sep 13, 2003, 5 
2; Caliente, about 50 km from Bakersfield 
atiLiwys5s)(L3)s;.Sep 15.) 200382 -sSan 
Juan Capistrano close to Hwy 74 (20), Sep 
PX, POO Wn Ie 

Specimens were collected near the coast 
and at higher altitudes near Paradise, Cle- 
ments, and Tehachapi. The fly is a common 
species in this part of California with many 
host plants. 

Trupanea jonesi Curran.—Silver Lake, 
GUD, Seo 13, 2008, 3 6, i &. 

This is a polyphagous species with host 
plants from 29 genera and is considered one 
of the more common tephritid species in 
California. 

Trupanea nigricornis (Coquillett).—Te- 
hachapi city (15), Sep 15, 2003, 1 @; hills 
at Hwy 202 ca. 16 km from Tehachapi (16), 
1 2; at Hwy 86 about 3 km beyond Indio 
(17), Sep 17, 2003, 1 3; close to Hwy 78 
and Rd S2 at the entrance to Anza Borrego 
National Park (19), Sep 17, 2003, 1 gd, 1 @. 

This species was collected at three local- 
ities in close proximity. 

Trupanea wheeleri Curran.—Monte Pinos 
(1), Sep 6, 2003, 1 6; Tehachapi at Hwy 58 
(14), Sep 15, 2003, 2 3d, 3 2; hills at Hwy 
202 ca. 16 km from Tehachapi (16), Sep 15, 
2003, 10 6, 8 2; San Juan Capistrano close 
to Hwy 74 (20), Sep 20, 2003, 1 @. 

At one locality, Monte Pinos, specimens 
were taken from Chrysothamnus viscidiflo- 
rus (Hook.) Nutt., one of its known host 
plants, which was abundant and blooming. 


INTRODUCED SPECIES 


Chaetorellia succinea (Costa).—Twenty- 
SiX Specimens were collected in 2002, and 
176 specimens in 2003. This species was 
also found at several more localities (not 
included in Table 1) with high abundance 
of Centaurea solstitialis: San Luis Obispo 
Co., San Luis Obispo, May 4, 2002, 2 °; 
Morro Bay Hwy 41, Jun 7, 2002, 3 6, 3 
2; Atascadero Hwy 41, Jun 7, 2002, 1 6, 
3 2; Santa Maria Rd 166 (iii), Jun 8, 2002, 
5 6, 2 2; Santa Maria Hwy (iv), Jun 8, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


2002, 1 6, 2 2; New Cuyama Rd 166 (vy), 
Jun 8, 2002, 1 6, 3 2; near Cambria at 
Hwy 46 about 16 km before Hwy | (2), Sep 
7, 2003, 1 6; Santa Lucia Mountains south 
of Los Padres Rd connecting Jolon to Hwy 
1 (3), Sep 7, 2003, 2 3; Santa Lucia, near 
the seashore (4), Sep 7, 2003, 6 6, 4 2; 
Sonoma Hwy 12 (6), Sep 9, 2003, 4 6, 2 
2; south from Clear Lake at Hwy 175 ca. 
3 km from Hwy 101 (7), Sep 10, 2003, 24 
3,14 2; north of Clements at Hwy 88, sev- 
eral stops on the way (10), Sep 13, 2003, 
36 3, 14 2; Silver Lake, Eldorado National 
lores (iD), Seo 13, ZO0R, 22 o, 3 23 ali 
Hwy 145 ca. 8 km from Madera (12), Sep 
14, 2003, 3 6; Caliente (about 50 km from 
Bakersfield) at Hwy 58 (13), Sep 15, 2003, 
1 6, 2 2; Tehachapi at Hwy 58 (14), Sep 
le 2008e S26 -al2 2 tehachapiicityagis)) 
Sep) 153 2003,.5' 65.3925 hills} ar ElwyeZ 02 
ca. 16 km from Tehachapi (16), Sep 15, 
2002, 2 ¢; Salton Sea Lake near desert 
shore (18), Sep 17, 2003, 1 d. 

In the early seventies, Chaetorellia aus- 
tralis and Urophora sirunaseva were pur- 
portedly released as biological control 
agents against yellow starthistle, Centaurea 
solstitialis, and were established in several 
states (Turner et al. 1995). Balciunas and 
Villegas (2001) studied this species and 
learned that it established on yellow star- 
thistle only at sites where the alternative 
host plant, bachelor button (Centaurea cy- 
anus), was also present. However, they did 
recover Chaetorellia flies at many sites 
where bachelor button was absent. Later, 
those adult specimens were identified as 
Chaetorellia succinea. In this study, the 
widespread distribution of C. succinea 
through California is confirmed on its host 
plant Centaurea solstitialis. 

Urophora jaculata Rondani.—Santa Ma- 
ria Rd 166 (iv), Jun 8, 2002, 4 d, 1 29; New 
Cuyama Rd 166 Ventucopa Rd 33 (v), Jun 
8, 2002, 3 3d, 1 2; Pine Mountain Summit 
(vi), Jun 8, 2002, 1 6, 2 2; south from 
Clear Lake at Hwy 175 ca. 3 km from Hwy 
OMG) eSepalOS2 0035s oe 

Urophora jaculata was introduced into 


VOLUME 107, NUMBER 4 


California in 1969 for the control of yellow 
starthistle Centaurea solstitialis, but it 
failed to establish (Maddox 1981). At that 
time it was mistakenly identified as U. si- 
runaseva (Julien and Griffiths 1999). 
Twelve specimens were collected randomly 
in June 2002 and one female was collected 
near Kelseyville on highway 175, 3 km 
from Hwy 101, in 2003. 

Terellia fuscicornis (Loew ).—Santa Ma- 
ria w of intersection at Hwy 166 and Hwy 
IO, Gd), ditin 8 ATOZ, i es, i Se 

This species was accidentally introduced 
and has been spreading naturally through- 
out the range of its host, Cynara cardun- 
culus L., causing significant damage (Julien 
and Griffiths 1999). Larvae also are report- 
ed to develop in the flower heads of Sily- 
bum marianum (Woods et al. 1996). This 
fly has not been purposefully released for 
biological control. 


ACKNOWLEDGMENTS 


I am grateful to Dr. D. H. Headrick, Cal 
Poly, San Luis Obispo, for help in determi- 
nation of specimens collected in 2002 and 
for access to his research collection of native 
Californian tephritids. I am indebted to Dr. 
M. Rejmanek, University of California, Da- 
vis, for help in host plant identification. This 
study was made possible by the financial 
support provided by Dr. T. A. Miller of the 
University of California, Riverside. 


LITERATURE CITED 


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released Chaetorellia succinea (Diptera: Tephriti- 
dae): Is this natural enemy of yellow starthistle a 
treat to safflower growers? Environmental Ento- 
mology 30(5): 953-963. 

Foote, R. H., E L. Blanc, and A. L. Norrbom. 1993. 
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of America North of Mexico. Cornell University 
Press, Ithaca, New York. xii + 571 pp. 

Freidberg, A. and W. N. Mathis. 1986. Studies of Ter- 
elliinae (Diptera: Tephritidae): A revision of the 
genus Neaspilota Osten Sacken. Smithsonian 
Contribuitions to Zoology No. 439, 75 pp. 

Goeden, R. D. and D. H. Headrick. 1999. Life history 
and description of immature stages of Neaspilota 
wilsoni Blanc and Foote (Diptera: Tephritidae) on 


857 


Hazardia squarrosa (Hooker and Arnott) E. 
Greene (Asteraceae) in Southern California. Pro- 
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ington 101(4): 897-909. 

Goeden, R. D., D. H. Headrick, and J. A. Teerink. 
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ifornia. Proceedings of the Entomological Society 
of Washington 96(4): 612—629. 

Headrick, D. H. and R. D. Goeden. 1991: Life history 
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fornia. Proceedings of the Entomological Society 
of Washington 93(3): 559-570. 

Headrick, D. H., R. D. Goeden, and J. A.Teerink. 
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stages of Euaresta stigmatica (Diptera: Tephriti- 
dae) on Ambrosia spp. (Asteraceae) in Southern 
California. Annals of the Entomological Society 
of America 88(1): 58-71. 

Hickman, J. C. ed. 1993. The Jepson manual. Higher 
plants of California. University of California 
Press. Berkeley and Los Angeles, xv + 1400 pp. 

Julien, M. H. and M. W. Griffiths. 1998. Biological 
control of weeds. A world catalogue of agents and 
their target weeds. 4" edition. CSIRO Entomolo- 
gy, Long Pocket Laboratories, Indooroopilly, 
Brisbane, Queensland 4068. CAB International 
Publishing, Australia, 223 pp. 

Knio, K. M., R. D. Goeden, and D. H. Headrick. 1996. 
Description of immature stages of 7rupanea ni- 
gricornis and T. bisetosa (Diptera: Tephritidae) 
from Southern California. Annals of the Ento- 
mological Society of America 89(1): I-11. 

Maddox, D. M. 1981. Introduction, phenology, and 
density of yellowstarthistle in coastal, intercoastal, 
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cultural Research Results, ARR-W-200. USDA- 
ARS, Oakland, California. 

Steck, G. J., B. D. Sutton, and D. Defoe. 2003. Biology 
of Rhynencina longirostris Johnson (Diptera: Te- 
phritidae). Proceedings of the Entomological So- 
ciety of Washington 105: 542-547. 

Turner, C. E., J. B. Johnson, and J. P. McCaffrey. 1995. 
Yellow starthistle. /n Nechols J. R., L. A. Andres, 
J. W. Beardsley, R. D. Goeden, and C. G. Jackson, 
eds. Biological control in Western United States: 
Accomplishments and Benefits of Regional Re- 
search Project W-84, 1964-1989. Department of 
Natural Resources Publication 3361. University of 
California, Berkeley, California. 

Wasbauer, M. S. 1972. An annotated host catalog of 
the fruit flies of America north of Mexico (Dip- 
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White, I. M. and M. M. Elson-Harris. 1994. Fruit flies 


858 


of Economic Significance: Their Identification and 
Bionomics. CAB International, Wallingford, xii + 
601 pp. 

Woods, D. M., M. J. Pitcairn, D. B. Joley, and C. E. 
Turner. 1996. Terellia fuscicornis, a new biocon- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


trol agent of artichoke thistle in California, p. 28. 
In Bezark, L. G., ed. Biological Control Program 
Annual Summary 1995. Sacramento, California 
Department of Food and Agriculture, Division of 
Plant Industry. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 859-863 


TWO NEW SPECIES OF HAPLOPERLA NAVAS (PLECOPTERA: 
CHLOROPERLIDAE) FROM NORTH CAROLINA, U.S.A. 


B. C. KONDRATIEFF, R. EK KIRCHNER, AND DAVID LENAT 


(BCK) Department of Bioagricultural Sciences and Pest Management, Colorado State 
University, Fort Collins, CO 80523, U.S.A. (e-mail: bkondrat @ceres.agsci.colostate.edu); 
(RFK) 5960 East Pea Ridge, Ridgeview Apartment |, Huntington, WV 25705, U.S.A.; 
(DL) 3607 Corbin Street, Raleigh, NC 27612, U.S.A. 


Abstract.—Two new species of stoneflies, Haploperla parkeri Kirchner and Kondra- 
tieff and H. fleeki Kondratieff, Kirchner, and Lenat, are described from Haywood Co., 
Macon Co., and Hoke/Moore Co., North Carolina, USA, respectively. Diagnostic char- 
acters are given to separate these two species from the three other known eastern North 


American species of Haploperla. 


Key Words: 


Currently, three species of the Nearctic 
and eastern Palearctic genus Haploperla 
Navas are known from eastern North Amer- 
ica. The common Haploperla brevis 
(Banks) is widespread from Alberta to east- 
ern Canada south to Alabama, and north to 
Minnesota (Surdick 1985, Surdick 2004); 
H. chukcho (Surdick and Stark) appears re- 
stricted to an area of southwestern Missis- 
sippi (Hardy et al. 1994): and H. orpha 
(Frison) has been reported from North Da- 
kota to northeastern Canada and United 
States (Surdick 1985, 2004). 

Two additional eastern North American 
species collected by the authors are de- 
scribed below. The descriptions follow the 
style of Surdick (2004). One of these is an 
unpatterned species from the Great Smoky 
Mountains of North Carolina, whereas the 
second one is a patterned species from the 
Coastal Plain of North Carolina. 


Haploperla parkeri Kirchner and 
Kondratieff, new species 
(Figs. 1—2) 
Adult.—Male: Forewing length 5.5—6 
mm. General body color pale yellow in life, 


Plecoptera, Chloroperlidae, Haploperla, new species, USA 


white in alcohol. Antenna pale; head wider 
than pronotum, unpatterned except for three 
dark ocellar rings; abdomen lacking dark 
markings. Pronotum oval, wider than long; 
mesonotum and metanotum unpatterned. 
Wings macropterous, hyaline, representa- 
tive of genus Abdomen without markings. 
Epiproct tablike, oval in dorsal view (Fig. 
1), narrow, slightly recurved in lateral view 
(Fig. 2). 

Female: Unknown. 

Nymph.—Unknown. 

Types.—Holotype 3d: North Carolina, 
Haywood, Co., small stream to Right Fork 
of Cove Creek, Co. Rd 1395, 0.6 mi SW 
Cove Creek Gap, 16 May 1983, R. F Kirch- 
ner and B. C. Kondratieff. Paratypes: Same 
data as holotype, 4 ¢; Macon Co., Shot 
Pouch Creek, 26 May 1994, B. P. Stark, S. 
W. Szczytko, and J. Sandberg, 1 d. 

Holotype deposited in the National Mu- 
seum of Natural History, Smithsonian In- 
stitution, paratypes in the Colorado State 
University (CSUC), R. FE Kirchner Collec- 
tion and B. P. Stark Collection, Clinton, 
Mississippi. 

Etymology.—We honor our friend, Dr. 


860 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-2. 


Charles R. Parker, trichopterist and aquatic 
biologist, Great Smoky Mountains National 
Park. 

Diagnosis.—-Haploperla parkeri can be 
distinguished readily from both patterned 
species, H. chukcho and H. orpha by the 
lack of any pigmentation. It is most similar 
to the usually unpatterned H. brevis, but the 
male terminalia in lateral view, can be dis- 
tinguished easily by the narrow epiproct, 
which is distinctly wedged-shaped in H. 
brevis, as illustrated by Surdick (1985: figs. 
137-139, Surdick 2004: figs. 6.275, 6.277). 
Many male specimens examined from the 
vicinity of the Canadian and Pennsylvania 
type localities of H. brevis, and its recog- 
nized synonyms support the concept of H. 
brevis as clearly presented by Surdick 
(1985, 2004). . 

Remarks.—The type locality (35°37'27"N 
83°3'4"W) is approximately 1,158 m ele- 
vation, and at this location the stream is 
high gradient with many spring seeps. Oth- 
er stoneflies collected with H. parkeri in- 
cluded Sweltsa lateralis (Banks), S. urticae 
Ricker, Malirekus hastatus (Banks), Ocon- 
operla innubila (Needham and Claassen), 


Haploperla parkeri. 1, Male terminalia, dorsal. 2, epiproct, lateral view. 


Isoperla spp., Tallaperla anna (Needham 
and Claassen), and 7. maria (Needham and 
Claassen). Shot Pouch Creek draining Way- 
ah Bald in the Nantahala National Forest is 
a well-known collecting site for rare stone- 
flies (Stark and Stewart 1982a, b). 


Haploperla fleeki Kondratieff, Kirchner, 
and Lenat, new species 
(Figs. 3-7) 


Adult.—General body color straw yellow 
in life, pale yellow in alcohol. Wing vena- 
tion representative of genus. Head with 
dusky marking as Fig. 3, pronotum with 
dusky lateral margins, rugulae dusky (Fig. 
3), meso- and metathoracic scutellum with 
dusky U-shaped and bisecting markings, 
abdomen with dusky longitudinal stripe. 

Male: Body length 6.0—6.5 mm; fore- 
wing length 7.0 mm. Epiproct tip in dorsal 
view elongate, parallel-sided, apex tapered 
(Fig. 4), in lateral view, tip broad, minute 
spicules at base (Fig. 5). Aedeagus in lateral 
view with skeletal rod (Fig. 6). 

Female: Body length 8.0—8.5 mm; fore- 
wing length 7.0—7.5 mm, venation dark. 


VOLUME 107, NUMBER 4 861 


Figs. 3-7. Haploperla fleeki. 3, Head and pronotum. 4, Male terminalia, dorsal. 5, Epiproct, lateral view. 6, 
Aedeagus, lateral view. 7, Female terminalia, ventral view. 


862 


Subgenital plate narrowly rounded at apex 
(icky): 

Pre-emergent nymph.—Typical of genus 
(Surdick 1985), adult color pattern visible. 

Types.—Holotype d: North Carolina, 
Hoke/Moore Co., Little River, Morrison 
Bridge Road, East of Southern Pines, 18 
May 2004, B. Kondratieff, R. Kirchner, R. 
Zuellig, and D. Lenat. Paratypes: Same data 
as holotype, 7 6, 41 2. Holotype deposited 
in the National Museum of Natural History, 
Smithsonian Institution, paratypes in the 
Colorado State University (CSUC), R. E 
Kirchner Collection, and Monte L. Bean 
Life Sciences Museum, Brigham Young 
University (BYUC). 

Etymology.—The patronym honors Eric 
Fleek, Department of Environment and 
Natural Resources, Division of Water Qual- 
ity (DWQ), Raleigh, North Carolina. Eric 
first noted the uniqueness of the nymph in 
benthic samples being processed by his 
agency. 

Diagnosis.—Using the excellent treat- 
ment of the eastern Chloroperlidae by Sur- 
dick (2004), H. fleeki is similar to the two 
other patterned species, H. chukcho and H. 
orpha. The epiproct of H. fleeki is most 
similar to H. chukcho, but in dorsal view, 
much narrower in width, and narrowing to- 
ward the apex (Fig. 4). The epiproct of H. 
chukcho is broad, and broadly rounded at 
the apex (see Surdick 2004: figs. 6.282-— 
6.284). The female appears distinctive with 
the combination of the dusky markings and 
a more acute apex of the subgenital plate 
(Fig. 7). 

Remarks.—The lower Little River is in- 
cluded in the Cape Fear River Basin, in- 
cluding parts of Moore, Hoke, Lee, Harnett 
and Cumberland counties. Land use is 
mostly forest (80%), with small amounts of 
cropland (8%) and pasture (8%). The lower 
Little River is largely in the Sandhills 
ecoregion, but the headwater area lies with- 
in the Piedmont. Sandhills streams usually 
have abundant groundwater storage (due to 
the coarse sandy soils), and maintain good 
flow even during drought periods. The ex- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


istence of piedmont soil in the headwaters 
of the Upper Little River, however, produc- 
es atypical flow characteristics for a San- 
dhills stream. For example, during the re- 
cord drought of 2002, the lower Little River 
had a prolonged period with little flowing 
water, and the macroinvertebrate rating 
crashed from an “Excellent” bioclassifica- 
tion (1988-1998) to only “Good-Fair” in 
October 2002 and January 2003 (North 
Carolina DWQ, Kathy Herring, unpub- 
lished data). Full recovery, however, had 
occurred by the time of the next DWQ sam- 
ple in March 2004. 

According to the North Carolina DWQ, 
the lower Little River has very good water 
quality, characterized by low specific con- 
ductance (median = 36 umhos/cm), low nu- 
trient levels (nitrogen and phosphorus), and 
high dissolved oxygen (median = 8.0 mg/ 
1, minimum = 5.5 mg/l). This site is slightly 
acidic (median pH = 5.8) and humic acids 
usually stain the water. 

More than 84 EPT (Ephemeroptera, Ple- 
coptera, Trichoptera) taxa are known from 
the lower Little River (http://www.esb.enr. 
state.nc.us/bar.html). Many of these taxa are 
sandhills endemics, including the caddis- 
flies, Hydropsyche decalda Ross, Brachy- 
centrus chelatus Ross, Setodes arenatus 
Holzenthal, and the stoneflies, Alloperla 
lenati Kondratieff and Kirchner, /soperla n. 
sp., and Helopicus bogaloosa Stark and 
Ray. 


ACKNOWLEDGMENTS 


We thank Bill P. Stark, Mississippi Col- 
lege, for reviewing the manuscript and pro- 
viding material. Robert E. Zuellig is 
thanked for his help in the collection of ma- 
terial used in this study. Dave Carlson, 
Windsor, Colorado, provided the illustra- 
tions. 


LITERATURE CITED 


Hardy, C. L., B. P. Stark, C. E Boll, N. C. Phifer, Jr., 
and W. T. Tharpe. 1994. A survey for stoneflies 
in the Homochitto National Forest, Southwest 
Mississippi. Proceedings of the Annual Confer- 


VOLUME 107, NUMBER 4 


ence of the Southeastern Association of Fish and 
Wildlife Agencies 48: 368-373. 

Department of Environment and Natural Resources, 
Division of Water Quality, Raleigh, North Caro- 
lina Basinwide Assessment Reports. http:// 
www.esb.enr.state.nc.us/bar.html. Accessed No- 
vember 2004. 

Stark, B. P. and K. W. Stewart. 1982a. Oconoperla, a 
new genus of North American Perlodinae (Ple- 
coptera: Perlodidae). Proceedings of the Entomo- 
logical Society of Washington 84: 746-752. 98. 


863 


. 1982b. The nymph of Viehoperla ada (Ple- 
coptera: Peltoperlidae). Journal of the Kansas En- 
tomological Society 55: 494—498. 

Surdick, R. E 1985. Nearctic genera of Chloroperlinae 
(Plecoptera: Chloroperlidae). Illinois Biological 
Monographs 54:1—102. 

. 2004. Chloroperlidae (The Sallflies), pp. 1— 

60. Jn Stark, B. P. and B. J. Armitage, eds. Stone- 

flies (Plecoptera) of Eastern North America. Vol. 

II. Perlidae, Chloroperlidae, Perlodinae. Bulletin 

of the Ohio Biological Survey, New Series 14(4): 

1-192. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 864-868 


A NEW WESTERN NEARCTIC SPECIES OF CALAMEUTA KONOW 
(HYMENOPTERA: CEPHIDAE) 


DaAvip R. SMITH AND NATHAN M. SCHIFF 


(DRS) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. 
Department of Agriculture, % National Museum of Natural History, Smithsonian Insti- 
tution, PO. Box 37012, MRC 168, Washington, DC 20013-7012 U.S.A. (e-mail: 
dsmith @sel.barc.usda.gov); (NMS) U.S. Forest Service, U.S. Department of Agriculture, 
Southern Research Station, Center for Bottomland Hardwood Research, P.O. Box 227, 
Stoneville, MS 38776 U.S.A. (e-mail: nschiff@fs.fed.us) 


Abstract.—Calameuta middlekauffi, n. sp., is described from southern Oregon and 
California. It is the second species of Calameuta in North America and is differentiated 
from C. clavata (Norton) by head shape and coloration. Illustrations, descriptions, and a 
key are given to separate the two species. The food plant is unknown, but Palearctic 


species of Calameuta are known to feed in grass stems. 


Key Words: 


The genus Calameuta Konow is best rep- 
resented in the Palearctic Region where 
about 20 species are known. Only one spe- 
cies, C. clavata (Norton 1869), is known 
from North America, and it occurs from 
California to Idaho, Oregon, and Washing- 
ton (Ries 1937, Middlekauff 1969, Smith 
1979). For a number of years, we noted a 
few peculiar specimens masquerading un- 
der C. clavata. These were mostly single 
specimens from scattered localities, that ap- 
pear like and key to C. clavata and were 
considered variants of that species. Not un- 
til NMS collected a series from the same 
place could we substantiate that these un- 
usual specimens represent a distinct species 
rather than an occasional aberration of C. 
clavata. The head shape and relatively con- 
stant color pattern are the most distinctive 
characteristics, and we here describe these 
specimens as a new species. From the few 
specimens available, it does not appear to 
be as common as C. clavata, and it is ap- 
parently restricted in distribution to the cen- 


Cephinae, grasses, California, Oregon 


tral valley and coastal ranges of southern 
Oregon to southern California. 

Food plants are not known for North 
American Calameuta. Larvae of Palearctic 
species feed in grass stems, and for C. cla- 
vata, Middlekauff (1969) stated ““unknown 
grasses”” as the food plant but speculated 
that the grass hosts may turn out to be Bro- 
mus laevipes Shear and/or Deschampsia 
danthomioides Trin., which have sympatric 
ranges with C. clavata. 

Calameuta clavata has been placed in 
Cephus Latreille by some North America 
authors (Ries 1937, Middlekauff 1969). 
However, we follow Benson’s (1946) defi- 
nition of the two genera and place it in Cal- 
ameuta, as was followed by Smith (1979). 
Calameuta is distinguished from Cephus by 
the interantennal distance shorter than the 
antennal-tentorial distance; lower interocu- 
lar distance usually less than the eye height; 
female cerci more than half as long as the 
sheath; and the eighth sternite of the male 
without a fringe of long, modified setae. In 


VOLUME 107, NUMBER 4 


North America, C. clavata and the new spe- 
cies below can be distinguished from Ce- 
phus by the bidentate left mandible, with a 
rounded shoulder on the inner side of the 
second tooth. In contrast, the left mandible 
of Cephus is tridentate. 


Calameuta middlekauffi Smith and 
Schiff, new species 


(Figs. 1—5) 

Female.—Length, 6.7—8.5 mm. Antenna 
and head black, sometimes faint yellow 
spot at center of supraclypeal area; mandi- 
ble yellow with extreme base black and 
apex reddish brown. Thorax black with up- 
per corner of mesepisternum and spiracular 
sclerite yeliow. Legs with coxae and tro- 
chanters black, a small yellow spot on low- 
er posterior margin of hind coxa; femora 
yellow, fore- and mid-femora with black at 
extreme base and hind femur with extreme 
apex and base black; fore- and mid-tibiae 
and tarsi yellowish, apical 2 or 3 segments 
of fore- and midtarsi infuscate; hind tibia 
and tarsus black, sometimes hind tibia 
slightly yellowish. Abdomen (Fig. 2) black 
with continuous lateral longitudinal yellow 
stripe of about equal width, yellow on 8th 
segment broader and extending onto pos- 
terior margin of 8th sternite, 9th segment 
with small yellow stripe posteriorly on lat- 
eral margin; segments 4—7 with medial dor- 
sal yellow spots, varying in size and with 
faint to small spots on segments 5 and 8; 
apex of segments 8 and 9 with small yellow 
spots. Wings hyaline; veins and stigma 
brown; outer surface of costa of fore- and 
hind wing yellow. 

Antenna 23—25 segmented. Left mandi- 
ble bidentate, inner tooth with blunt, round- 
ed shoulder on inner margin. Head (Figs. 
3—5) round in lateral view, in lateral view 
and dorsal view with frons protuberant; in 
dorsal view head deeply emarginate behind, 
depth of emargination half distance from 
posterior corner to eye. Eye oval, about 
1.3X higher than wide. Distance from an- 
tennal insertion to tentorial pits 1.3% dis- 
tance between antennal insertions. Lower 


interocular distance slightly shorter than up- 
per interocular distance and subequal to eye 
height. Distances between eye and lateral 
ocellus, between lateral ocelli, and from lat- 
eral ocellus to hind margin of head as 1.0: 
1.0:2.2. Hind tibia with one preapical spine. 
Cercus about as long as sheath. Sheath in 
dorsal view of equal width and bluntly 
rounded at apex, width slightly more than 
width of hind basitarsus; oblong plate (val- 
vifer 2) 2.0 longer than sheath (valvula 3). 

Male.—Length, 6.5—7.0 mm. Color as 
for female except large yellow spot on su- 
praclypeal area; inner surface of midcoxa 
yellow; most of hind coxa yellow with 
black spot only at extreme base; outer sur- 
face of hind tibia may be yellowish. Ab- 
domen (Fig. |) black with medial yellow 
marks on posterior halves of terga 3, 4, 6, 
and usually 7; small if present on 2 and 5. 
Lateral yellow marks on 1—8 separated by 
black or connected below by narrow yellow 
stripes; hypandrium mostly yellow. Ab- 
dominal sternite 8 apically notched at cen- 
ter, with fringe of very short setae on mar- 
gin. 

Type material.—Holotype °, labeled 
“USA: California: Solano Co., Stebbins 
Cold Creek Canyon, 38°27’N 122°42’W, 
15-IV-1-V-96, Nathan M. Schiff, Malaise 
trap.”’ Deposited in the National Museum 
of Natural History, Smithsonian Institution, 
Washington, DC (USNM). 

Paratypes: CALIFORNIA: Same data as 
holotype (3 2, 1 6), same data as holotype 
except 1-14-IV-96 (2 2); Solano Co., Cold 
Cyn. Res., 11 km W Winters, 12-II-1-IV- 
92, S. L. Heydon, MT (2 2); Siskiyou Co., 
Hill Hungry Road, Forest Service Rd. 1, 
May 12, 1996, Nathan M., Schiff (1 @): El 
Dorado Co., Blodgett, 38°54’N 122°42’W, 
12-V-1-VI-96, Nathan M. Schiff, Malaise 
trap (1 2); Mts. near Claremont, Baker (1 
2, 1 3); Upland, 4/17/20 (1 2); Corte Ma- 
dera Cr., [1V-25—1960 (1 2, 1 d); Santa Cla- 
ra Co., Mount Hamilton, 2 May 1976, Lar- 
ry Bezark (1 2); 1.5 mi W on Mix Canyon 
Road, Solano Co., ele. 1,000, HI-30-76, N. 
J. Smith, colr. (1 2); Green Valley, Solano 


$66 


iN 


3 \ SN 


Figs. 1-5. 


otf 


Calameuta middlekauffi. 1, Male abdomen, lateral view. 2, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


4 


Female abdomen, lateral view. 3, 


Head, front view. 4, Head, lateral view. 5, Head, dorsal view. 


Co., IH-24-1954, E. I. Schlinger, collector 
(1 2); Orchard Hills, Alameda Co., [V-14- 
74, coll. R. D. Moon (1 2); Round Mt., 
Shasta Col., V-23-1949, R. M. Bohart (1 
2); Mt. San Jacinto, 4,000’, VI-6-42, R. 
Bohart (1 2). OREGON: 4 mi W of Selma, 
Josephine Co., May 23, 1964, Kenneth 
Goeden (1 @). Deposited in the USNM, 
University of California at Davis, and the 
collection of N. Schiff. 


Etymology.—The specific epithet is in 
honor Dr. Woodrow W. Middlekauff, Uni- 
versity of California, Berkeley, for his ex- 
cellent contributions to the knowledge of 
North American sawflies. 

Remarks.—Calameuta middlekauffi is 
distinguished from C. clavata by a solid lat- 
eral yellow stripe on the female abdomen, 
small yellow markings laterally and on the 
dorsum of segments 3, 4, 6, and usually 7 


VOLUME 107, NUMBER 4 


Figs. 6—10. 
front view. 9, Head, lateral view. 10, Head, dorsal view. 


on the male abdomen, the round head in 
side view, more rounded eyes, protuberant 
frons, and deep posterior emargination of 
the head as seen in dorsal view. In general, 
the size of C. middlekauffi is smaller than 
C. clavata, with most specimens of the lat- 
ter being 10 mm or more in length. 

The female abdomen of C. clavata (Fig. 
7) is banded yellow on segments 3, 4, and 
6, with medial and lateral spots on segment 
7, segment 8 mostly black, and segment 9 
black with the apical half yellow. It lacks 
the solid lateral yellow stripe. The male ab- 
domen has similar yellow markings as the 
female (Fig. 6). The eyes are more elongate, 
with the height 1.6 or more their width; 
the head is more oval in lateral view; the 


Calameuta clavata. 6, Male abdomen, lateral view. 7, Female abdomen, lateral view. 8, Head, 


frons is more evenly rounded and less pro- 
tuberant in lateral and dorsal views; the 
posterior margin of the head in dorsal view 
is less emarginate, the depth of the emar- 
gination less than half the distance from the 
posterior corner of the head to the eye 
(Figs. 8-10); and the oblong plate is 1.6 
the length of the sheath. The male usually 
has more yellow on the supraclypeal area, 
the coxae are yellow except at their bases, 
and the hypandrium is mostly yellow with 
a large round black spot at its base. Though 
the coloration of C. clavata is variable, as 
stated by Middlekauff (1969), the variation 
does not reach the extreme solid lateral 
stripe as in C. middlekauffi. The amount of 
yellow on segments 2, 3, and 5 and 6 of C. 


868 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


clavata varies somewhat, but segment 4 is 
normally entirely black. 

Calameuta middlekauffi has been found 
from southern Oregon south to southern 
California. Most specimens are from the 
coastal range in central California. Cala- 
meuta clavata has a much wider distribu- 
tion, which encompasses that of C. middle- 
kauffi and occurs from Washington and Ida- 
ho south to southern California (as far as 
the Mexican border) in the coastal ranges 
and at high elevations in the Sierras. 

We have examined a number of Palearc- 
tic species of Calameuta and checked the 
literature and are not aware of a Palearctic 
species of Calameuta similar to C. middle- 


kauffi. 


KEY TO SPECIES OF NORTH AMERICAN 
CALAMEUTA 


1. Female abdomen with solid lateral yellow 
stripe (Fig. 2); male abdomen with dorsal yel- 
low spots on segments 3, 5, 6, and usually 7, 
usually not extending laterally, and lateral 
spots on segments 1—8 (Fig. 1); head round in 
lateral view (Fig. 4), frons protuberant (Figs. 
4—5), and in dorsal view with deep posterior 
emargination (Fig. 5) C. middlekauffi 

— Female and male abdomen without solid lateral 
stripe, tergites 3, 4, and 6 usually solidly yel- 
low (Figs. 6—7); head more oval in lateral view 
(Fig. 9) frons not protuberant (Figs. 9-10), and 


in dorsal view with shallower posterior emar- 
enunerutorn (Eng, 10) coe 60000e00000¢ C. clavata 


ACKNOWLEDGMENTS 


We thank Steve Heydon, University of 
California, Davis, for the loan of speci- 
mens. Cathy Apgar, Systematic Entomolo- 
gy Laboratory, USDA, took the Auto-Mon- 
tage photos and arranged the plates. We ap- 
preciate the reviews of the following: S. G. 
Codella, Kean University, Union, NJ; N. J. 
Vandenberg and M. A. Solis, Systematic 
Entomology Laboratory, USDA, Washing- 
ton, DC. 


LITERATURE CITED 


Benson, R. B. 1946. Classification of the Cephidae 
(Hymenoptera Symphyta). Transactions of the 
Royal Entomological Society of London 96: 89— 
108. 

Middlekauff, W. W. 1969. The cephid stem borers of 
California (Hymenoptera: Cephidae). Bulletin of 
the California Insect Survey 11, 19 pp. 

Norton, E. 1869. Catalogue of the described Tenthre- 
dinidae and Uroceridae of North America. Trans- 
actions of the American Entomological Society 2: 
321-368. 

Ries, D. T. 1937. A revision of the Nearctic Cephidae 
(Hymenoptera). Transactions of the American En- 
tomological Society 63: 259-324. 

Smith, D. R. 1979. Symphyta, pp. 1-137. In Krom- 
bein, K. V., P D. Hurd, Jr, D. R. Smith, and B. 
D. Burks, eds. Catalog of Hymenoptera in Amer- 
ica North of Mexico. Vol. 1. Symphyta and Apo- 
crita (Parasitica). Smithsonian Institution Press, 
Washington, DC, xvi + 1198 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 869-882 


TWO NEW SPECIES AND A NEW RECORD OF EUCOENOGENES MEYRICK 
(LEPIDOPTERA: TORTRICIDAE) FROM THAILAND WITH A DISCUSSION 
OF CHARACTERS DEFINING THE GENUS 


NANTASAK PINKAEW, ANGSUMARN CHANDRAPATYA, AND RICHARD L. BROWN 


(NP) Department of Entomology, Faculty of Agriculture Kamphaeng Saen, Kasetsart 
University, Kamphaeng Saen Campus, Nakhon Pathom 73140 Thailand (e-mail: 
agrnsp@ku.ac.th); (AC) Department of Entomology, Faculty of Agriculture, Kasetsart 
University, Bangkok 10900, Thailand (e-mail: agramc @ku.ac.th); (RLB) Mississippi En- 
tomological Museum, Box 9775, Mississippi State, MS 39762, U.S.A. (e-mail: 
moth @ra.msstate.edu) 


Abstract.—Two new species of Eucoenogenes Meyrick (Lepidoptera: Tortricidae: Eu- 
cosmini), FE. bicucullus Pinkaew and E. vaneeae Pinkaew, are described and illustrated 
from montane evergreen forest in Thong Pha Phum National Park, Kanchanaburi Prov- 
ince, Thailand. Epinotia munda Diakonoff, described from a female collected in Sumatra, 
is transferred to Eucoenogenes (n. comb.) based on male and female specimens collected 
in Thailand; the imago and genitalia of both sexes are described and illustrated. Characters 


for defining the genus are discussed and illustrated. 


Key Words: 
notia, Eucosmini 


Eucoenogenes Meyrick, 1939, was pro- 
posed as a replacement name for Caeno- 
genes Meyrick, 1937, which was preoccu- 
pied by Caenogenes Walsingham, 1887. 
Meyrick (1937) described the type species, 
Caenogenes melanancalis, based on two 
specimens reared from larvae mining leaves 
of Eugenia jambolana Lamark (Myrtaceae) 
in India. Meyrick defined the genus by the 
thickened palpus with densely appressed 
scales concealing the third segment, the ab- 
sence of the R, vein in the forewing, and 
absence of the M, vein in the hindwing. 
Clarke (1958) designated a female as lec- 
totype, noting that “‘the male is missing,” 
even though Meyrick indicated that both 
specimens were female in his original de- 
scription. However, Clarke (1955) previ- 
ously noted that Meyrick often misidenti- 
fied sexes because of his reliance on a hand 


new combination, distribution records, generic characters, Acroclita, Epi- 


lens and emphasis on non-genitalic char- 
acters; thus, the sex and present location of 
the second specimen is uncertain. In con- 
trast to Meyrick’s description, Clarke’s fig- 
ure of venation showed that both R, in the 
forewing and M, in the hindwing were pre- 
sent and that all veins were separate except 
the fused anal veins in the forewing and the 
stalked M,-CuA, in the hindwing; this ve- 
nation has been confirmed by a re-exami- 
nation of the venation of the lectotype. 
Diakonoff (1967) considered the female 
genitalia of E. melanancalis to lack char- 
acters of generic significance and consid- 
ered the palpi with appressed scales and 
concealment of the third segment to be the 
only character defining the genus. Based on 
similar palpi, Diakonoff described EF. del- 
tostoma based on a single male specimen 
from the Philippine Islands. This descrip- 


870 


tion of the male genitalia gave an identity 
to the genus that was lacking in Meyrick’s 
description, specifically the possession of 
enlarged spiniform setae originating from 
the sacculus. Subsequently, additional spe- 
cies of Eucoenogenes have been described 
or transferred to the genus based largely on 
the possession of the saccular spiniform se- 
tae. These species vary, some intraspecifi- 
cally, in the degree that the scales of the 
second segment of the labial palpus are ap- 
pressed and cover the third segment; thus, 
this character may be an artifact of individ- 
ual specimens. Other characters for defining 
the genus have not been provided previously. 

Eucoenogenes currently includes 11 spe- 
cies occurring in the Oriental and eastern 
Palearctic regions. Three species have been 
reported previously from Thailand (Kawabe 
1989): ancyrota Meyrick (1907), euphlebia 
Kawabe (1989), and japonica Kawabe 
(1978). In addition to these three species 
and E. melanancalis and E. deltosoma, oth- 
er species of Eucoenogenes include aestuo- 
sa and cyanopsis, described by Meyrick 
(1912) from India, levatana Kuznetsov 
(1997a) and segregana Kuznetsov (1997b) 
from South Vietnam, pythonias (Meyrick 
1910) from Java, and teliferana (Christoph 
1882) from Vladivostok, Russia, the sole 
species occurring outside the Oriental Re- 
gion. 

About 200 species of Olethreutinae were 
collected during a recent (2001—2003) sur- 
vey of Tortricidae in the Thong Pha Phum 
National Park, Thailand (Kanchanaburi 
Province). Of these, two new species of Eu- 
coenogenes are described here and an ad- 
ditional species is transferred from Epinotia 
to Eucoenogenes and reported as a new re- 
cord for Thailand. 


MATERIALS AND METHODS 


Tortricidae were sampled in Thong Pha 
Phum National Park at elevations from 200 
m to 1,000 m in habitats that included 
swamp forest, riparian forest, deciduous 
forest, evergreen forest, dry evergreen for- 
est, and montane evergreen forest. All spec- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


imens were collected with a 20-watt black- 
light suspended in front of a sheet and op- 
erated with a car battery. Collections were 
made on 145 nights in all forest types at 
various elevations in the park. Some collec- 
tion sites were accessible by vehicle, and 
others required 2—3 days of hiking to sam- 
ple. Latitude and longitude were recorded 
with a Magellen GPS 315. Specimens were 
transported in relaxing boxes lined with 
cotton to prevent sliding of specimens and 
scale loss. All specimens were spread in the 
laboratory within a week of collection. 

A Leica 12.5 stereomicroscope with an 
ocular micrometer and fiber optic illumi- 
nation was used to examine, measure, and 
illustrate specimens. Forewing length was 
measured from the outer edge of the tegula 
at wing base to the outermost edge of the 
fringe scales at apex and is given as an av- 
erage of lengths for all measured specimens 
of each sex. Compound microscopes were 
used at magnifications of 125—400% for ex- 
amining microtrichia, cornuti, and other 
characters of genitalia. Scanning electron 
micrographs were made with a JEOL JSM- 
6500 FE-SEM. The “‘Methuen Handbook 
of Colour’ (Kornerup and Wanscher 1983) 
was used as a standard for describing color 
patterns of adults. Genitalia preparation fol- 
lowed the methodology given by J. Brown 
and Powell (1991). Genitalia were illustrat- 
ed on acetate drafting film with the use of 
a drawing tube. Terminology for wing ve- 
nation and genitalic structures follows Hor- 
ak (1984) and Horak and Brown (1991). 
Terminology for forewing patterns and stri- 
gulae follows R. Brown and Powell (1991), 
as modified and discussed by Baixeras 
(2002). Specimens of Eucoenogenes col- 
lected in Thailand were compared with 
identified specimens and genitalia prepara- 
tions of E. aestuosa (3, 2), E. ancyrota (6, 
2), E. cyanopsis (3), E. euphlebia (3d, 2) 
and E. japonica (d, °) in the National Mu- 
seum of Natural History, Smithsonian In- 
stitution, Washington DC (USNM) and 
University of Osaka Prefecture and with 
type specimens of E. deltosoma (d), E. me- 


VOLUME 107, NUMBER 4 


Figs. 1-4. Adults. 1, Eucoenogenes munda, male 
paratype. 4, Labial palpus of Eucoenogenes vaneeae. 


lanancalis (2 genitalia and venation), and 
E. pythonias (3 genitalia). Identities of E. 
cyanopsis (2), E. teliferana (3), E. leva- 
tana (4d), and E. segregana (d) were de- 
termined by examining published illustra- 
tions of imagos and genitalia (Kuznetsov 
I997anbs ZOU): 


Eucoenogenes munda (Diakonoff), 
new combination 
(Figs. 1, 6-8, 12) 


Epinotia (Asthenia) 
1983: 40. 


munda Diakonoff 


Description.—Head: Upper frons orange 
white to pale orange mixed with brownish 
orange, light brown laterally, lower frons 
light brown to brown; labial palpus with 


871 


2, E. bicucullus, male holotype. 3, E. vaneeae, female 


first segment light brown mixed with 
brown, second segment brownish orange, 
with dark brown spots basally, dorsomedi- 
ally, and ventrally at % length, and brown- 
ish-orange spot apically, apical segment or- 
ange white; vertex yellowish white, eye 
bordered posteriorly by row of brown and 
dark brown scales. 

Thorax: Pronotal collar and tegula 
brown mixed with dark brown; mesonotum 
brown mixed with yellowish white and dark 
brown. Forewing (Fig. 1): Length: 7.3 mm 
in males (n = 3), 6.9 mm in females (n = 
1); male costal fold absent; venation (Fig. 
6) with R,—R, stalked % length of Rs, ter- 
men concave between apex and M,, ground 
color brownish orange mixed with brown, 


872 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


inner margin with patch extending from 
basal fascia to pretornal spot and medially 
to 1A+2A, yellowish white mixed with 
brownish grey; costal strigulae 1—9 paired, 
orange white to pale orange, separated by 
dark brown spots or short lines; basal fascia 
with scattered dark brown scales, distal 
margin extending as dark brown line from 
costa near strigula 1 to inner margin; sub- 
basal fascia present as dark brown spot be- 
tween strigulae 2 and 3 and extending from 
costa to R, dislocated apically to form 
brownish orange spot with dark brown mar- 
gins between middle of discal cell and 
1A+2A; median fascia indistinct, present as 
dark brown spot on costa between strigulae 
4 and 5; postmedian and preterminal fasciae 
present as dark brown spots on costa be- 
tween strigulae 6 and 7 and 8 and 9, re- 
spectively; apex with dark brown spot ex- 
tending into adjacent fringe scales; pretor- 
nal subtriangular spot extending from inner 
margin to CuA,, brownish orange with dark 
brown margins; ocellar region with a dark 
brown line extending from R; to CuA,, 
scalloped between veins, outer margin with 
distinct dark brown line, broken by orange 
white, unpaired strigulae between R; and 
M,, M, and M,, and CuA, and CuA,. Un- 
derside light brown, strigulae on costa and 
termen yellowish grey to yellowish white. 
Hindwing (Fig. 7): Brown dorsally, light 
brown ventrally. 

Male genitalia (Fig. 8): Tegumen arms 
narrow ventrally, widened in dorsal %, with 
moderately dense setae dorsomedially; un- 
cus short, apices pointed; soci pendent 
from base of uncus, apically rounded, mod- 
erately setose; gnathos arising from mid- 
length of tegumen, membranous; anellus 
closely surrounding basal 4% of aedeagus; 
aedeagus moderately long, apically tapered, 
dorsoapical % unsclerotized, cornuti mod- 
erately dense; juxta triangular; valva with 
small group of setae basally and large group 
dorsally at midlength, ventromedial area at 
midlength with small, rounded lobe, long 
setae arising from between lobe and ventral 
margin, microtrichia present on medial sur- 


face from near base to rounded lobe, ven- 
trolateral surface of valva with group of 
long, spiniform setae surrounded by micro- 
trichia, some setae with deeply bifid apices 
at high magnification (400), base of neck 
with a long, ventrally projecting, digitate 
process lacking setae or with a few setae 
near middle, cucullus small, rounded, 
densely setose (n = 2). 

Female genitalia (Fig. 12): Sternum VI 
densely microtrichiate on posterior half. 
Sternum VII densely microtrichiate except 
on posterolateral corners, densely setose on 
posterolateral corners and posteromedial 
area, asetose on remainder of sclerite; ter- 
gum VIII with moderately dense scales and 
microtrichia on lateral triangular projec- 
tions; papillae anales with dense setae, 
some lateral setae with papillose bases; la- 
mella antevaginalis reduced, lamella post- 
vaginalis moderately sclerotized, with 
dense microtrichia and moderately dense 
scales; ostium bursae opening into wide, 
cuplike antrum; colliculum small, ringlike; 
ductus bursae with large sclerotized plate 
on anterior half, narrowly encircling ductus 
near middle, incompletely sclerotized on 
ventral side, with two, anterior extensions 
into corpus bursae, apices acute; ductus 
seminalis arising from anterior 4 of ductus 
bursae; corpus bursae with two signa, 
bladelike, apices acute (n = 1). 

Specimens examined.—Thailand: Kan- 
chanaburi Prov.: Thong Pha Phum N.P, 
14°27'41"N 98°37'15"E, 24 Dec. 2001 (1 6, 
genitalia slide NP 61), 14°41'34"N 
98°24'06’E, 12 Jan. 2002 (1 ¢, genitalia 
slide NP 104), 14°41'39"N 98°24'10"E, 13 
Jan. 2002 (1 6, genitalia slide NP 172, 1 
2, genitalia slide NP 173), all collected by 
N. Pinkaew. Deposited in Department of 
Entomology Collection, Kasetsart Univer- 
sity and Mississippi Entomological Muse- 
um. 

Remarks.—This species was described 
based on a single female from Mt. Banda- 
hara at 810 m in Atjeh, Sumatra. The type 
specimen was not examined, but the illus- 
trated imago and genitalia (Diakonoff 1983) 


VOLUME 107, NUMBER 4 


Fig. 5. 
cucullus with expanded femur covered by sex scales 
(arrow). 


Metathoracic leg of male Eucoenogenes bi- 


are identical to the female specimen from 
Thailand. These females share a similar 
seventh sternite and sclerotization of the 
ductus bursae, unlike other known Eucoen- 
ogenes. Specimens from Thailand were col- 
lected during the dry winter season at 980— 
1,000 m in montane evergreen forest. 


Eucoenogenes bicucullus Pinkaew, 
new species 
(ties, Zs S95 13) 


Diagnosis.—The male of this species dif- 
fers from other species in the genus in hav- 
ing a wide, flat hindtibia with dense, seti- 
form scales closely appressed to the medial 
and lateral surfaces. The anal margin of the 
hindwing bears a dark brown hair pencil ba- 
sally and a row of long, yellowish-white 
hairs beyond the wing base. The male gen- 
italia differ from those of other species by 
the cucullus being divided into dorsal and 
ventral lobes. The female is similar to that 
of E. munda, but the seventh sternum is 
quadrate, not rounded, and has microtrichia 
that are restricted to its posterolateral cor- 
ners. 

Description.—Head: Upper frons with 
light brown mixed with brownish orange, 
lower frons with brown to dark brown 
mixed with light brown; labial palpus with 
first segment dark brown at base, remainder 
yellowish white, second segment yellowish 
white to pale orange with dark brown spots 


873 


basally, dorsomedially, ventroapically at % 
length, and apically, apical segment brown 
dorsally, yellowish white ventrally; vertex 
brownish orange to light brown mixed with 
brown laterally. 

Thorax: Pronotal collar with anterior 
short scales brown, posterior long scales 
light brown with orange-white apices; te- 
gula brown mixed with dark brown basally, 
light brown mixed with orange white api- 
cally; mesonotum pale orange mixed with 
light brown to dark brown, with transverse, 
dark brown band medially. Forewing (Fig. 
2): Length: 6.50 mm in males (n = 4), 7.4 
mm in females (n = 2); male costal fold 
absent; R,—R; stalked 7% length of Rs, ter- 
men slightly concave, ground color grayish 
yellow mixed with brownish orange, with 
diffuse dark brown transverse lines, costal 
strigulae 1—9 paired, pale yellow, and sep- 
arated by dark brown, termen with strigulae 
forming pale yellow spots between R, and 
R,, R; and M,, and M, and M.,; basal fascia 
indistinct, with scattered, dark brown scales 
between wing base and strigula | from cos- 
ta to inner margin; subbasal fascia distinct 
between strigulae 2 and 3 from costa to R, 
dislocated apically from middle of discal 
cell to inner margin, dark brown on mar- 
gins, mixed with brownish orange medially; 
median fascia dark brown on costa between 
strigulae 4 and 5, indistinct from near costa 
to inner margin, distal margin extending 
obliquely outward as narrow dark brown 
line bordering brownish-orange scales from 
costa to R,, angled at R;, and extending ba- 
sally to middle of discal cell; silvery striae 
originating from strigulae 5 and 6 and ex- 
tending obliquely to R,, striae from strigu- 
lae 7-9 extending obliquely to R;, confluent 
from R, to tornal area along outer margin, 
striae separating brownish-orange lines 
originating from dark brown spots between 
strigulae, orange line between strigulae 5 
and 6 extending to R,, orange line between 
strigulae 6 and 7 extending to R;, orange 
line between strigulae 7 and 8 confluent 
with orange line from between strigulae 8 
and 9 and extending to R;; ocellar region 


874 


PORT RZAERS R4 


stem of R4+5 (chorda) Sc _ 


CuP 


1A+2A 


Figs. 6-7. Venation of Eucoenogenes munda. 6, 
Forewing. 7, Hindwing. Scale bar: 1 mm. 


with diffuse dark brown lines from R, to 
inner margin near tornus; wing apex dark 
brown between R, and R;; outer margin 
with distinct dark brown line from apex to 
CuA,; fringe orange white mixed with 
brown and dark brown. Underside light 
brown, with yellowish-grey strigulae on 
costa and between veins on outer margin. 
Hindwing: Brown dorsally; male with 
scales narrow between anal margin and 
CuA,, anal margin rolled dorsally in spread 
specimens and bearing row of hair pencils, 
basal % of row dark brown basally, yellow- 
ish white apically; ventrally light brown, 
with dense brown scales on basal % of cos- 
tal area, with yellowish-white, narrow 
scales on rolled anal area; male hindtibia 
(Fig. 5) wide, flattened and densely covered 
dorsally and laterally with specialized long, 
setiform scales adhering closely to surface. 

Male genitalia (Fig. 9): Tegumen arms 
moderately wide, dorsally rounded, with 
moderately long setae medially; uncus 
short, bilobed dorsally, with rounded ven- 
tral projection extending over socii bases; 
socii arising from base of uncus, moderate- 
ly long, apically rounded, densely setose 
dorsoapically; gnathos arising from dorsal 
*3 of tegumen, membranous; anellus closely 
surrounding base of aedeagus; aedeagus 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


moderately long, curved medially, with 
eight cornuti extending along apical 4; jux- 
ta triangular; sacculus sparsely setose ba- 
sally, densely setose medially between bas- 
al opening and neck, ventral margin with 
dense short setae between basal opening 
and neck, microtrichia relatively dense on 
medial surface from near base of sacculus 
to apical margin of basal opening, ventro- 
lateral area of valva with row of 4—7, long, 
spiniform setae, setae with attenuate apices; 
valva deeply incised to form narrow neck; 
cucullus moderately incised medioapically 
forming two, long rounded lobes, dorsal 
lobe densely setose except apical %, ventral 
lobe larger than dorsal lobe, densely setose 
except ventrobasal margin (n = 3). 

Female genitalia (Fig. 13): Sternum VI 
with sparse microtrichia on posteromedial 
margin. Sternum VII with large sclerotized 
plate, scales moderately dense on posterior 
half and lateral areas, becoming more dense 
towards posterior margin, anteromedial area 
scaleless, sparsely setose on posterior half, 
microtrichia restricted to lateral margins; 
tegumen VIII without setae or scales dor- 
somedially, with sparse setae and moderate- 
ly dense scales and microtrichia on lateral 
triangular extensions: papillae anales with 
dense setae, lateral setae with papillose ba- 
ses; sterigma reduced; ostium bursae behind 
sternum VII, antrum forming a wide, scler- 
otized cup, colliculum small, weakly scler- 
otized, ductus bursae encircled by large 
sclerotized plate medially, divided anteri- 
orly to form two triangular projections ex- 
tending into corpus bursae; ductus semin- 
alis arising from near middle of ductus bur- 
sae; corpus bursae with two signa, blade- 
like, rounded at apex, right signum larger 
than left signum (n = 1). 

Holotype.—Male. Thailand: Kanchana- 
burl %Prov-- thong )Phay PhummNEes 
14°41'38’N 98°24'17"E, 30 Jul. 2002, N. 
Pinkaew; [specimen no.] NP10; male [gen- 
italia slide] NP 391. Data given as on three 
labels except for bracketed information. 
Deposited in USNM. 

Paratypes.—Thailand: Kanchanaburi Prov.: 


VOLUME 107, NUMBER 4 


875 


| 
il 4 Cre eeceS 


ve, / an if 
: f ; 


yi 


Figs. 8-10. Male genitalia. 8, Eucoenogenes munda. 9, E. bicucullus. 10, E. vaneeae. Scale bar: | mm. 


Thong Pha Phum N.P., 14°41'35’N 
98°24'19"E, 23 Dec. 2001, (1 6, genitalia 
slide NP 41), 14°41'17"N 9824’02’E, 9 Jan. 
2002 (1 2, genitalia slide NP 141), 
14°41'40"N 98°24'15’E, 5 Feb. 2002 (1 ¢, 


genitalia slide NP 231), 14°41'41’N 
98°24'12"E, 6 Feb. 2002 (1 &, genitalia 
slide NP 180), 14°41'34”N 98°24'06"E, 12 
Jan. 2002 (1 d, genitalia slide NP 105), all 
collected by N. Pinkaew. Deposited in Ka- 


876 


Fig. 11. 


Eucoenogenes vaneeae, ventral view of 
male abdominal segment 8 with hair pencils intact on 
left side. Scale bar: 1 mm. 


setsart University Entomology Collection 
and Mississippi Entomological Museum. 

Etymology.—The name of this species 
refers to the division of the male cucullus 
into two parts. 

Remarks.—The expanded hindtibia with 
specialized sex scales in the male is unique 
among species of Eucoenogenes. These tib- 
ial sex scales may be involved in the pro- 
duction of a pheromone that is disseminated 
by hair pencils on the hindwing, similar to 
pheromone systems in males of other spe- 
cies of Lepidoptera (Grant 1978). This spe- 
cies was collected during dry winter and 
early rainy seasons in montane evergreen 
forest at 980—1,000 m. 


Eucoenogenes vaneeae Pinkaew, 
new species 
(Figs. 3-4, 10-11, 14) 


Diagnosis.—The male of this species dif- 
fers from related species in having long hair 
pencils arising from anterolateral pockets of 
sternum VIII (Fig. 11), a sacullus bearing 
spiniform setae with distal scales having bi- 
fid apices and basal scales having trifid api- 
ces, and a valva that is rotated at the neck 
to produce a ventrally facing cucullus. This 
species is most similar to E. levatana Kuz- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


netsov (1997a) in the form of the spiniform 
scales on the sacculus and the presence of 
a narrow cucullus, but differs in having the 
uncus narrowed medially and the rotated 
valva described above. A forewing costal 
fold is present in E. levatana, but is absent 
in this new species. 

Description.—Head: Upper _ frons 
brownish orange mixed with brown to dark 
brown laterally, lower frons yellowish 
white, dark brown laterally; labial palpus 
(Fig. 4) with first segment orange white to 
pale orange, with dark brown spot apically, 
second segment orange white with small, 
dark brown spot basally, confluent with 
spot on first segment, dorsomedial area with 
small, dark brown spot, narrowly separated 
from large, dark brown, ventroapical spot 
at % length, latter spot narrowly separated 
from large, dark brown apical spot, apical 
segment orange white ventrally, dark brown 
dorsally; vertex dark brown anteriorly, or- 
ange white posteriorly, eye posteriorly bor- 
dered by row of dark brown and light 
brown scales. 

Thorax: Pronotal collar brown to dark 
brown mixed with light brown; tegula dark 
brown mixed with light brown on basal 
half, light brown mixed with brown on api- 
cal half; mesonotum dark brown, with 
transverse light brown band. Forewing (Fig. 
3): Length 5.77 mm in males (n = 4), 6.34 
mm in females (n = 5); male costal fold 
absent; R, and R; stalked % length of Rs, 
termen concave between R; and M,; ground 
color yellowish white mixed with pale or- 
ange and brownish grey; costal strigulae or- 
ange white to orange gray basally, yellow- 
ish white apically, strigulae 1—9 paired and 
separated by dark brown, except some spec- 
imens with strigula 6 single, termen with 
yellowish-white striguia between R, and 
M,;; basal fascia indistinct, with dark brown 
spot divided by orange-white scales be- 
tween costa and R and irregular rows of 
dark brown scales between R and inner 
margin; subbasal fascia with dark brown 
proximal and distal margins enclosing 
brownish orange mixed with orange gray, 


VOLUME 107, NUMBER 4 877 


7 S 
song eet 


1 
' 
oer 
, 
—y 


Figs. 12-14. Female genitalia. 12, Eucoenogenes munda. 13, E. bicucullus. 14, E. vaneeae. Scale bar: 1 mm. 


878 


distinct on costa between strigulae 2 and 3 
and extending to near middle of discal cell, 
dislocated apically between middle of dis- 
cal cell and inner margin; median fascia 
dark brown mixed with orange, extending 
obliquely from costa between strigulae 4 
and 5 to R;, widened at R; and extending 
transversely to 1[A+2A, with narrow, api- 
cally directed spur on R;, broken by ground 
color at M stem in discal cell; pretornal spot 
subtriangular, dark brown mixed with or- 
ange surrounded by dark brown on proxi- 
mal and distal margins, postmedian fascia 
extending obliquely from costa between 
strigulae 6 and 7 to R; near outer margin, 
dark brown on costa, orange with some 
brown on distal margin from strigulae to R;, 
strigulae 5 and 6 with silvery striae becom- 
ing confluent and bordering postmedian 
fascia to R;, strigula 7 with silvery stria ex- 
tending obliquely to Rs, strigulae 8 and 9 
with striae reduced to small silvery spots, 
outer margin with silvery stria (possibly 
confluent striae) extending from M, to 
CuA,, broken near CuA, by ground color; 
ocellar region with dark brown patch be- 
tween R; and CuA,, outer margin with dark 
brown line from apex to CuA,, broken by 
strigula between R; and M,. Underside light 
brown with yellowish-grey strigulae on cos- 
ta and between R; and M, on outer margin. 
Hindwing: Brown dorsally, light brown 
ventrally. 

Abdomen (Fig. 11): Male sternum VIII 
with two, brown hair pencils arising from 
pockets on anterolateral corners. ; 

Male genitalia (Fig. 10): Tegumen arms 
narrow ventrally, widened at midlength, 
with dense long setae along outer margin of 
ventral two-thirds; uncus short, narrowed 
medially, with bifid apex; socii pendant 
from base of uncus to near half of tegumen 
length, densely setose, apically rounded; 
gnathos arising from midlength of tegumen, 
sclerotized basally, membranous apically; 
anellus closely surrounding basal one- 
fourth of aedeagus, extending dorsally to 
one-third length; aedeagus moderately long, 
not greatly tapered apically, with dense 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Female genitalia of Eucoenogenes melan- 
ancalis lectotype. 


Fig. 15. 


bundle of 15—20 cornuti; juxta triangular, 
truncated ventrally; valva with group of 
sparse setae along ventrobasal margin of sa- 
cullus and group of dense setae medially 
between basal opening and neck, ventroap- 
ical margin of sacullus with short and spi- 
niform setae, ventrolateral surface of valva 
with large group of long, spiniform, flat- 
tened setae with multidentate apices basally 
and rounded, spiniform setae with bidentate 
apices towards neck, setiform setae bor- 
dered basally by microtrichia, neck narrow, 
asetose, rotated near middle with cucullus 


VOLUME 107, NUMBER 4 


Figs. 16-17. 
E. cyanopsis, lateral view of sacculus, cleft in bifid 
spiniform setae indicated by arrow. Abbreviations: vs 


Male genitalia of Eucoenogenes. 16, 


= valval spiniform setae, ss = setae of sternum 9. 
Scale bar = 40 um. 17, E. cyanopsis, magnification of 
valval spiniform setae. Scale bar = 4 um. 


facing ventrally, cucullus narrow, with 
dense, short setae basally, acute and sparse- 
ly setose apically (n = 4). 

Female genitalia (Fig. 14): Sternum VI 
densely microtrichiate on posterior 7%. Ster- 
num VII densely scaled and microtrichiate, 
sparsely setose; tergum VIII moderately 
scaled on triangular lateral extensions; pa- 
pillae anales densely setose, lateral setae 
with papillose bases; lamella postvaginalis 
short, sparsely setose and densely microtri- 
chiate; antrum forming long, lightly scler- 
otized cup; colliculum small, not encircling 
ductus bursae; ductus bursae with long 
sclerotized band encircling *4 of ductus cir- 
cumference, completely encircled by nar- 


879 


row band near middle and posterior to in- 
ception of ductus seminalis, apically round- 
ed without projections into corpus bursae; 
corpus bursae with two, bladelike signa, 
smaller signum on posterior margin of cor- 
pus bursae near connection with ductus bur- 
sae (n = 1). 

Holotype.—Male. Thailand: Kanchana- 
bun Proves hones) Ehaysehume Nees 
14°41'31"N 98°24'27"E, 13 May 2002, N. 
Pinkaew; [specimen no.] N96; male [geni- 
talia slide] NP 464. Data given as on three 
labels except for bracketed information. 
Deposited in USNM. 

Paratypes.—Thailand: same data as ho- 
lotype (1 6, genitalia slide NP 452, 2 @), 
same data except 14°41'43"N 9824'34’E, 
15 May 2002, (1 6, genitalia slide NP 519, 
3 2, genitalia slide NP 463), 14°41'36”"N 
98°24'21"E, 6 Nov 2002 (1 6, genitalia 
slide NP 393). Deposited in Department of 
Entomology collection, Kasetsart Universi- 
ty, Mississippi Entomological Museum, and 
USNM. 

Etymology.—tThis species is named after 
the late Vanee Pinkaew, mother of the se- 
nior author. 

Remarks.—This species was collected 
during the dry winter and dry summer sea- 
sons in montane evergreen forest at 980— 
1,000 m in the same habitat as E. munda 
and E. bicucullus. 


DISCUSSION 


New species of Eucoenogenes have been 
described in recent years based largely on 
their similarity with £. deltosoma in having 
male genitalia with enlarged spiniform se- 
tae on the sacculus. As the male of the type 
species, E. melanancalis, is unknown, the 
concept of the genus rests solely with the 
female lectotype. The genitalia of E. me- 
lanancalis are refigured here (Fig. 15) to 
provide better resolution of detail than the 
photograph in Clarke (1958). A re-exami- 
nation of the genitalia reveals that it is 
unique in having two sclerotized s-shaped 
projections on the inner medial surface of 
the eighth tergite, unlike females of other 


880 


species of Eucoenogenes and other Oleth- 
reutinae. The sterigma is laterally fused 
with the seventh sternite, similar to some 
species of Spilonota Stephens that were il- 
lustrated by Clarke (1958), and unlike other 
species of Eucoenogenes. The ductus bur- 
sae is encircled by a long sclerotized band, 
similar to other species of Eucoenogenes. 
The corpus bursae is densely spiculate on 
the lateral margins, giving a halo effect, 
whereas the corpus bursae in other Eucoen- 
ogenes has spicules evenly distributed over 
most of its surface. Characters of the female 
genitalia that associate the lectotype with 
any other genus are lacking, and only the 
long sclerotized ductus bursae is shared 
with females of other species assigned to 
Eucoenogenes. Thus, two options are avail- 
able for describing new species that cannot 
be assigned to another olethreutine genus 
and that are most similar to species cur- 
rently assigned to Eucoenogenes: 1) restrict 
Eucoenogenes to the type species, repre- 
sented by a single female specimen, and de- 
scribe a new genus to contain species cur- 
rently assigned to Eucoenogenes based on 
characters of the male genitalia, or 2) con- 
tinue to describe new species within Eu- 
coenogenes based on characters of the male 
genitalia and refine the definition of this 
group of species until such time that the 
male of E. melanancalis is discovered. 
Strong evidence is lacking to associate E. 
melanancalis with another genus or to dis- 
associate it from other species currently as- 
signed to Eucoenogenes. In addition, spe- 
cies currently assigned to Eucoenogenes 
display a wide range of variation in some 
characters, including the spiniform setae of 
the valvae. Thus, the latter option is adopt- 
ed in order to further define this group of 
species by describing new species as they 
become known. 

A wide diversity of sex scales is present 
in males of some species of Eucoenogenes, 
but none of these can be used to character- 
ize the genus. A forewing costal fold was 
reported for E. levatana and E. segregana 
(Kuznetsov 1997a, b), a pair of hair pencils 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


on the eighth abdominal segment is present 
in E. vaneeae, hindwing hair pencils and 
tibial sex scales are present in E. bicucullus, 
and sex scales are present on abdominal 
segments 3—5 in E. cyanopsis. 

Several superficial characters of Eucoen- 
ogenes munda, E. bicucullus, and E. va- 
neeae are shared with various other species 
assigned to Eucoenogenes. The second seg- 
ment of the labial palpus has dark spots ba- 
sally, dorsomedially, ventroapically and 
apically in E. bicucullus, E. vaneeae, E. 
aestuosa, E. cyanopsis, and E. japonica 
(Fig. 4). The dorsomedial, ventroapical and 
apical spots are confluent in E. munda and 
E. deltosoma, and all spots are absent in E. 
melanancalis, E. euphlebia, and E. ancy- 
rota. Descriptions of E. levatana, E. segre- 
gana, and E. telifera indicate that the labial 
palpi are uniformly colored. The forewing 
venation has R, and R; stalked for varying 
lengths with the stalk approximate and par- 
allel to R3 in EF. munda, E. bicucullus, E. 
vaneeae, E. euphlebia, E. aestuosa, E. cy- 
anopsis, and E. japonica (Fig. 6); veins Ry 
and R, are separate in E. ancyrota, E. del- 
tostoma, and E. melanancalis. The fore- 
wing lacks a well-defined basal fascia and 
has a subbasal fascia that is dislocated api- 
cally between the middle of the discal cell 
and the dorsal margin in E. bicucullus, E. 
vaneeae, E. aestuosa, E. japonica, and E. 
euphlebia (Fig. 3). The forewing dorsum is 
suffused with the ground color that ob- 
scures expression of fasciae in E. munda 
(Fig. 1), E. ancyrota, and E. pythonias. The 
subbasal and basal fasciae are confluent 
without any dislocation in E. melanancalis. 

All known males of species assigned to 
Eucoenogenes, except E. ancyrota, have 
male genitalia with a bifid uncus and soci 
that are elongate, apically rounded, and 
pendant from the base of the uncus. All 
known males of Eucoenogenes have long, 
spiniform setae on the sacculus, except E. 
ancyrota, in which the spiniform setae orig- 
inate from the cucullus, and E. teliferana, 
in which spiniform setae are absent. These 
spiniform setae have longitudinal ridges 


VOLUME 107, NUMBER 4 


and windows between ridges and some 
have deeply bifid apices, similar to lamellar 
scales (Figs. 16, 17). The distinction be- 
tween lamellar, multidentate scales on the 
abdominal sternites and rounded, non-den- 
tate, spiniform setae on the valva is transi- 
tional among various species. Similar sac- 
cular setae are present in some species as- 
signed to Acroclita Lederer, although these 
are absent in the type species, A. subse- 
quana (Herrich-Schaffer), which appears to 
be a very derived species based on both 
male and female genitalia. The Nearctic ge- 
nus Proteoteras Riley also has spiniform 
setae on the valval sacculus, and its rela- 
tionship with southern Asiatic groups re- 
quires investigation. 

All examined females of Eucoenogenes, 
except E. ancyrota and the type species, E. 
melanancalis, have microtrichia on female 
sternum VI and VII. Based on a survey of 
almost 200 species of Olethreutinae col- 
lected in Thailand and more than 300 spe- 
cies and most genera of Olethreutinae in the 
Nearctic and Palearctic regions, microtri- 
chia are present in other genera only on 
sternum VII of Acroclita subsequana and 
on sterna VI and VII of Lathronympha stri- 
gana (FE) (Grapholitin1). 

Females have a ductus bursae that is en- 
circled by a long sclerotized band in all ex- 
amined species except E. euphlebia, which 
has a short ductus bursae and short band. 
The sclerotized band has two anterior pro- 
jections extending into the corpus bursae 
(Fig. 12) in E. munda, E. bicucullus, E. aes- 
tuosa, E. cyanopsis, E. japonica, and E. eu- 
phlebia, especially long and narrow in the 
last species; projections are reduced to 
broadly rounded lobes in E. vaneeae and 
are absent in E. ancyrota and E. melanan- 
calis. Similar projections of the sclerotized 
band of the ductus bursae have not been 
seen in other Olethreutinae. The presence 
of these anterior projections of the sclero- 
tized band appears to be unique to some 
species of Eucoenogenes. 

The presence of sternal microtrichia in 
the female in combination with spiniform 


881 


setae on the male sacculus appear to be the 
best characters for defining the group of 
species currently assigned to Eucoenoge- 
nes, even though neither are known to be 
present in the type species, and both are 
individually present in species of other gen- 
era. The anterior projections of the sclero- 
tized band of the ductus bursae also may 
prove to be a character defining this group 
of species. Eucoenogenes ancyrota and E. 
teliferana do not appear to be congeneric 
with other species assigned to the genus. 
The spiniform spines originating from the 
cucullus in E. ancyrota are not considered 
homologous with the spiniform spines on 
the sacculus in other Eucoenogenes, and 
other characters shared by E. ancyrota with 
various Eucoenogenes species are lacking. 
Nonetheless, this species is maintained in 
Eucoenogenes because no other generic as- 
signment can be made at present. The ge- 
neric placement of E. teliferana will be in- 
definite until specimens of this species can 
be examined. 


ACKNOWLEDGMENTS 


We appreciate the logistical assistance 
provided by personnel with the Thong Pha 
Phum National Park. This research was 
supported by funds provided by the Thai- 
land Research Fund through the Royal 
Golden Jubilee Ph.D. Program (Grant No. 
PHD/0140/2544), the TRF/BIOTEC Spe- 
cial Program for Biodiversity Research and 
Training grant BRT T_145027, and the Mis- 
sissippi Agriculture and Forestry Experi- 
ment Station. The assistance of Kevin Tuck, 
The Natural History Museum, London, in 
loaning types of E. melanancalis and E. py- 
thonias is greatly appreciated. William 
Monroe, Mississippi State University, as- 
sisted with scanning electron microscopy, 
and Joe MacGown and SangMi Lee, Mis- 
sissippi State University, provided assis- 
tance with photography and formatting of 
images. We appreciate the assistance given 
to the senior author by Furumi Komai, Osa- 
ka University of Arts, in assisting with the 
visit to Osaka and examination of speci- 


882 


mens collected in Thailand, and Toshiya 
Hirowatari for his assistance in examining 
types and loan of specimens in the Ento- 
mological Laboratory, Osaka Prefecture 
University. 


LITERATURE CITED 


Baixeras, J. 2002. An overview of genus-level taxo- 
nomic problems surrounding Argyroploce Hiibner 
(Lepidoptera: Tortricidae), with description of a 
new species. Annals of the Entomological Society 
of America 95: 422—431. 

Brown, J. W. and J. A. Powell. 1991. Systematics of 
the Chrysoxena group of genera (Tortricidae: Tor- 
tricinae: Euliini). University of California Publi- 
cations in Entomology 111, 87 pp. + figs. 1-143. 

Brown, R. L. and J. A. Powell. 1991. Description of a 
new species of Epiblema (Lepidoptera: Tortrici- 
dae: Olethreutinae) from coastal redwood forests 
in California with an analysis of the forewing pat- 
tern. Pan-Pacific Entomologist 67: 107—114. 

Christoph, H. 1882. Neue Lepidopteren des Amurge- 
bietes. Bulletin de la Société Imperiale des Natur- 
alistes de Moscou 4: 405-436. 

Clarke, J. K G. 1955. Catalogue of the Type Specimens 
of Microlepidoptera in the British Museum (Nat- 
ural History) Described by Edward Meyrick. Vol. 
I. Trustees of the British Museum, London, 332 


Pp. 


. 1958. Catalogue of the Type Specimens of 
Microlepidoptera in the British Museum (Natural 
History) Described by Edward Meyrick. Vol. III. 
Trustees of the British Museum, London, 600 pp. 

Diakonoff, A. 1967. Microlepidoptera of Philippine Is- 
lands. United States National Museum Bulletin 
257: 1-484. 

. 1983. Tortricidae from Atjeh, Northern Su- 
matra (Lepidoptera). Zoologische Verhandelingen, 
Leiden 204: 1-132. 

Grant, G. G. 1978. Morphology of the presumed male 
pheromone glands on the forewings of tortricid 
and phycitid moths. Annals of the Entomological 
Society of America 71: 423-431. 

Horak, M. 1984. Assessment of taxonomically signif- 
icant structures in Tortricinae (Lep., Tortricidae). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Mitteilung der Schweizerischen Entomologischen 
Gesellschaft 57: 3—64. 

Horak, M. and R. L. Brown. 1991. Morphology, phy- 
logeny and systematics, pp. 1—22. Jn Van der 
Geest, L. P. S. and H. H. Evenhuis, eds. Tortricid 
Pests, Their Biology, Natural Enemies and Con- 
trol. Elsevier, Amsterdam, 808 pp. 

Kawabe, A. 1978. Descriptions of three new genera 
and fourteen new species of the subfamily Oleth- 
reutinae from Japan. Tinea 10: 173-191. 

. 1989. Records and descriptions of the subfam- 
ily Olethreutinae (Lepidoptera: Tortricidae) from 
Thailand. Microlepidoptera of Thailand 2: 23-82. 

Kornerup, A. and J. H. Wanscher. 1983. Methuen 
Handbook of Colour (3rd ed.). Methuen and Co., 
London, 252 pp. 

Kuznetsov, V. I. 1997a. Little known and new species 
of tortricid moths (Lepidoptera, Tortricidae) of the 
fauna of Vietnam. Entomologischeskoe Obozrenie 
76: 186-202. 

. 1997b. New species of tortricid moths of the 

subfamily Olethreutinae (Lepidoptera, Tortricidae) 

from the south of Vietnam. Entomologischeskoe 

Obozrenie 76: 797-812. 

. 2001. Tortricoidea, pp. 11—472. In Ler, P. A., 
ed. Key to the Insects of Russian Far East. Vol. 
V., Pt. 3. Trichoptera and Lepidoptera. Dal’ nauka, 
Vladivostok, 621 pp. 

Meyrick, E. 1907. Descriptions of Indian micro-lepi- 
doptera. III. Journal of the Bombay Natural His- 
tory Society 17: 730-754. 

. 1910. Descriptions of Malayan micro-lepidop- 

tera. Transactions of the Royal Entomological So- 

ciety of London 1910: 430-478. 

. 1912. Description of Indian micro-lepidep- 

tera. XV. Journal of the Bombay Natural History 

Society 21: 852-877. 

. 1937. Exotic Microlepidoptera 5: 159-160. 

Taylor and Francis, London. Reprinted by E.W. 

Classey Ltd., 1969. 

. 1939. New microlepidoptera, with notes on 
others. Transactions of the Royal Entomological 
Society of London 89: 47-62. 

Walsingham, Lord T. de Grey. 1887. A revision of the 
genera Acrolophus Poey and Anaphora Clem. 
Transactions of the Entomological Society of Lon- 
don 1887: 137-173. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 883-886 


A NEW SPECIES OF DALADER AMYOT AND SERVILLE, 
WITH A KEY TO THE MALAYSIAN SPECIES (HEMIPTERA: 
HETEROPTERA: COREIDAE: COREINAE: DALADERINI) 


H. BRAILOVSKY 


Instituto de Biologia, Departamento de Zoologia, UNAM, Apdo Postal 70-153, México 
D. EF 04510, México (e-mail: coreidae @servidor.unam.mx) 


Abstract.—Dalader pulchrus, n. sp., from Brunei, Sabah, and Sarawak is described. 
The antennae, pronotum, and male genital capsule are illustrated. A key to the known 
species of Dalader from Malaysia is presented. 


Key Words: Heteroptera, Coreidae, Coreinae, Daladerini, Dalader, new species, Malaysia 


The daladerine genus Dalader Amyot 
and Serville (1843) comprises 11 species 
and one variety restricted to southeastern 
Asia (Burma, Indonesia, Malaysia and Phil- 
ippines), southern Asia (India and Sri Lan- 
ka), and eastern Asia (China). In Malaysia 
four species are recorded: D. acuticosta 
Amyot and Serville, 1843, D. anthracinus 
Bergroth, 1912, D. planiventris (Westwood, 
1842), and D. shelfordi Distant, 1900b. 

Westwood (1842) described D. planiven- 
tris from Java, later Distant (1902) cited it 
from Burma, India, Sri Lanka, Sumatra, and 
Malaysia, and Hsiao (1977) recorded it 
from China. Amyot and Serville (1843) de- 
scribed D. acuticosta from Borneo and Dal- 
las (1852) and Distant (1902) recorded it 
from India, while Blote (1938) reported it 
from Indonesia (Sumatra and Timor). Dis- 
tant (1900b) described D. shelfordi from 
Kuching, Borneo (now in Sarawak, Malay- 
sia). Bergroth (1912) described D. anthra- 
cinus from Sarawak, Borneo (presently Ma- 
laysia) and Blote (1938) recorded this spe- 
cies from Indonesia (Mahakkam). 

In this article, I describe one new species 
from Malaysia and give a key to the known 
Malaysian species. 

The following acronyms are for collec- 


tions where specimens are deposited: The 
Natural History Museum, London (BMNH); 
Naturhistoriska Riksmuseet, Stockholm, 
Sweden (NHRM); Instituto de Biologia, 
Universidad Nacional Autonoma de México, 
México (UNAM). 

All measurements are in millimeters. 


Dalader pulchrus Brailovsky, 
new species 
(Figs. 1, 5—6) 


Description.—Measurements: First 
male, second female. Head length 1.70, 
1.75; width across eyes 2.20, 2.30; intero- 
cular space 1.80, 1.90; length antennal seg- 
ments: I, 4.50, 4.20; II, 3.80, 3.80; II, 3.20, 
3.00; IV, 2.75, 2.80. Pronotal length 4.20, 
4.30; maximal width across anterior lobe 
3.80, 3.80; maximal width across posterior 
lobe 8.00, 8.60. Scutellar length 2.40, 2.60: 
width 2.40, 2.60. Maximal width of abdo- 
men 9.50, 10.20. Total body length 20.20, 
DATO: 

Male (holotype).—Head: Quadrate; an- 
tenniferous tubercles protruding forward; 
dilated portion of antennal segment III nar- 
rowly obovate, occupying about two-thirds 
length of segment, with upper lobe wider 
than lower (Fig. 1); antenna covered with 


884 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


oo 


Figs. 1-6. Dalader spp. 1-2, Antennae. 1, D. pulchrus. 2, D. horsfieldi. 3-5, Pronotum. 3, D. horsfieldi. 4, 
D. acuticosta. 5, D. pulchrus. 6, Male genital capsule in caudal view of D. pulchrus. 


large erect setae; rostrum reaching anterior 
third of mesosternum. Pronotum: Humeral 
lobes triangular, not broadly expanded, and 
humeral angles subacuminate; anterolateral 


and posterolateral borders finely denticulate 
to spinate (Fig. 5). Legs: Femora and tibiae 
armed with spines and granules; hind femur 
moderately incrassate. Abdomen: Dilated, 


VOLUME 107, NUMBER 4 


rhomboidal; sterna obsoletely granulate. 
Genital capsule: Posteroventral edge with 
short and stout median tubercle (Fig. 6). 
Dorsal coloration: Head, pronotum, scu- 
tellum, clavus, corium, and connexivum 
pale chestnut orange; antennal segments I 
to III dark reddish, tinged with orange, and 
IV dark reddish orange; median longitudi- 
nal stripe on head and pronotum dark yel- 
low to orange yellow; anterolateral and pos- 
terolateral borders of pronotum with spines 
dark yellow to dark brown; hemelytral 
membrane dark ambarine, with basal angle 
black, and general surface mottled with dis- 
coidal or irregular black spots; upper mar- 
gin of connexivum with spines dark brown; 
dorsal abdominal segments shiny orange. 
Ventral coloration: Pale chestnut orange; 
apex of rostral segment IV black; anterior 
lobe of metathoracic peritreme dark yellow, 
and posterior lobe black; legs dark chestnut 
orange with granules and spines dark brown 
to black. 

Female.—Structure and color similar to 
male. Hind femur slightly incrassate, less 
than male; dorsal coloration pale chestnut 
yellow, with punctures pale reddish orange; 
apex of scutellum yellow; genital plates 
pale chestnut yellow; upper margin of con- 
nexival segments VIII and IX with black 
spines; anterior and posterior lobes of meta- 
thoracic peritreme creamy yellow. 

Variation.—1. Antennal segment IV red- 
dish brown. 2, Head dorsally with the me- 
dian dark yellow to orange-yellow longi- 
tudinal stripe (hard to see). 3. Legs dark 
orange with granules, and spines black. 4, 
Hind tibiae with or without irregular black 
rings. 

Type material—Holotype d, Malaysia, 
Sabah, Sipitang, Mendolong (without data), 
S. Aderbratt (NHRM). Paratypes: Malaysia: 
1 2, Brunei (without any other data) 
(NHRM); 1 36, 1 &, Sarawak, Bidi, 1907— 
1909, C. J. Brooks (BMNH, UNAM); | &, 
Sarawak (without data), collected by pool 
of entomologists, Serial No. 2550, per. C. 
R. Wallace (BMNH). 


Discussion.—Dalader pulchrus, de- 


885 


scribed from Malaysia, is related to D. 
horsfieldi Distant, 1900a, described from 
Java. The former is recognized by having 
the dilated portion of antennal segment III 
with the upper lobe clearly wider than the 
lower (Fig. 1), the humeral lobes of the 
pronotum triangular and not broadly ex- 
panded (Fig. 5), the width across the hu- 
meral angles less than 8.40 mm in the male, 
and the upper margin of connexival seg- 
ments III to VI entirely chestnut orange 
with the spines dark brown. In D. horsfieldi, 
the dilated portion of antennal segment III 
has the upper lobe slightly wider than the 
lower (Fig. 2), the humeral lobes broadly 
rounded and expanded (Fig. 3), the width 
across humeral angles wider than 8.80 mm 
in the male, and the upper margin of con- 
nexival segments III to VI dark orange with 
the spines dark brown, and the anterior 
third and posterior angle yellow with or- 
ange reflections. 

Distribution.—Known only from Brunei, 
Sabah, and Sarawak, Malaysia. 

Etymology.—From pulchrus (Latin) = 
beautiful; refering to the attractive appear- 
ance of the species. 


KEY TO THE MALAYSIAN SPECIES 
OF DALADER 


1. Humeral lobes of pronotum lunately produced; 
humeral angles obliquely truncated (Fig. 4) . . 
Me OER D. acuticosta Amyot and Serville 
— Humeral lobes not lunately produced; humeral 
angles subacuminate (Figs, 3,5) ......... 2 
2. Body dorsally and ventrally almost black ... 
Sgethea, co olan: ripiece eoraas D. anthracinus Bergroth 
— Body dorsally and ventrally chestnut orange to 
MOS mU WMO? oossaoedcosccoceo does 3 
3. Antennal segment III narrowly obovate (Fig. 1) 
SEN e CAE CEA Crewe D. pulchrus, n. sp. 
— Antennal segment III broadly dilated (Fig. 2) 


4. Antennal segment II yellow; tibiae yellow 
Hees ne oe Sea aG ibe D. shelfordi Distant 
— Antennal segment II dark reddish tinged with 
orange; tibiae dark chestnut orange 
D. planiventris (Westwood) 


ACKNOWLEDGMENTS 


I thank the following colleagues and in- 
stitutions for the loan of specimens and oth- 


886 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


er assistance relevant to this study: Mick 
Webb (BMNH), Thomas Pape (NRHM) 
and Bert Viklund (NHRM). I give special 
thanks to Ernesto Barrera (UNAM) for the 
illustrations. 


LITERATURE CITED 


Amyot, C. J. B. et A. Serville. 1843. Histoire Naturelle 
des Insectes. Hemipteres. Fain et Thunot, Paris, 
625 pp. 

Bergroth, E. 1912. Notes on Coreidae and Neididae. 
Annales de la Société Entomologique de Belgique 
56: 76-93. 

Blote, H. C. 1938. Catalogue of the Coreidae in the 
Rijksmuseum van Natuurlijke Historie. Part IV. 
Coreinae, Third part. Zoologisches Mededeelin- 
gen 20: 281-283. 

Dallas, W. L. 1852. List of the Specimens of Hemip- 


terous Insects in the Collection of the British Mu- 
seum. Part 2. Taylor & Francis, London, pp. 369— 
SOD 

Distant, W. L. 1900a. XLV. Rhynchotal Notes. VII. 
Heteroptera: Fam. Coreidae. Annals and Maga- 
zine of Natural History, London (6)7: 366-378. 

. 1900b. XIII. Contributions to a knowledge of 

the Rhynchota. Transactions of the Entomological 

Society of London, 1900: 665—697. 

. 1902. The Fauna of British India, including 
Ceylon & Burma. Rhynchota. Vol. 1 (Heterop- 
tera). Taylor & Francis, London, 438 pp. 

Hsiao, T. Y. 1977. Coreidae, pp.198—258. In Hsiao, T. 
Y. et al. A Handbook for the Determination of the 
Chinese Hemiptera- Heteroptera. Vol. I. Science 
Press, Beijing (in Chinese, English Summary). 

Westwood, J. O. 1842. Catalogue of Hemiptera in the 
Collection of the Rev. E W. Hope, M.A., with 
Short Latin Descriptions of the New Species. Part 
Il. J. C. Bridgewater, Oxford Street, London, 26 


Pp: 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 887-891 


A NEW SPECIES OF PHANOLINUS SHARP (COLEOPTERA: 
STAPHYLINIDAE) WITH A KEY AND COMMENTS FOR 
MEXICAN SPECIES 


JOSE Luis NAVARRETE-HEREDIA 


Entomologia, Centro de Estudios en Zoologia, CUCBA, Universidad de Guadalajara, 
Apdo. Postal 234, 45100 Zapopan, Jalisco, México (e-mail: glenusmx @ yahoo.com.mx) 


Abstract.—Phanolinus elisae, n. sp., from México is described. A key and distribution 
data for all four Mexican species of Phanolinus are provided. A lectotype and paralec- 
totype are designated for P. speciosus Sharp, 1884. 


Key Words: 


Phanolinus Sharp, 1884, is a Neotropical 
genus of Xanthopygina Staphylinidae con- 
sisting of 34 described species, most of 
which are large and beautifully colored, 
usually with metallic reflections. The name 
of one species, P. pretiosus Erichson, de- 
scribes this character very well. The species 
are distributed from México to Bolivia and 
northern Brazil; however, the highest diver- 
sity is in South America, especially in Co- 
lombia where nine species occur (Herman 
2001). 

Three species have been recorded in 
México (Navarrete-Heredia et al. 2002), all 
of which were collected from two localities 
in Veracruz State (see comments on those 
species). Recently, two additional speci- 
mens (one male and one female) were col- 
lected in Jalisco State. These specimens 
were compared with type material held at 
The Natural History Museum, and I con- 
cluded that they belong to a new species 
that is described here. A key and comments 
for the Mexican species are provided. 


MATERIALS AND METHODS 


Throughout this paper the abdominal 
segments are referred by their morphologi- 
cally comparable names; roman numerals 


México, Staphylinidae, Phanolinus 


are used to refer to these. The first visible 
abdominal segment is segment III. Total 
length was measured from the anterior mar- 
gin of head to apex of abdominal segment 
IX. Specimens are deposited in the Ento- 
mological Collection of the Centro de Es- 
tudios en Zoologia, Universidad de Gua- 
dalajara (CZUG); types of the species de- 
scribed by Sharp (1884) are in The Natural 
History Museum, London (BMNH). Addi- 
tional specimens examined are deposited at 
CNIN (Coleccion Nacional de Insectos, In- 
stituto de Biologia, Universidad Nacional 
Autonoma de México). A key for non-Mex- 
ican species of this genus was given by 
Scheerpeltz (1968). 


Phanolinus elisae Navarrete-Heredia, 
new species 
(Figs. 1—4) 


Description.—Length, 17.6 mm. Blue vi- 
olet with elytra lilac, in one abdomen ap- 
pears cobalt blue. Surface covered with mi- 
crosculpture consisting of waves, mixed 
with scattered micropunctures; a small 
well-defined area with isodiametric meshes 
at basolateral portions of abdominal seg- 
ments II-VI. 

Head: Subquadrate, slightly wider than 


888 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Phanolinus elisae, holotype, dorsal view. 


VOLUME 107, NUMBER 4 


889 


Figs. 2-4. Phanolinus elisae, male genitalia. 2, Aedoeagus, lateral view. 3, Parameres, ventral view, setae 


removed. 4, Parameres, apex, with setae, ventral view. 


long (1.09—1.12; n = 2); setiferous punc- 
tures denser at posterior angles and along 
inner borders of eyes; middle area of frons 
without setiferous punctures (Fig. 2). Su- 
bocular ridge well developed. First anten- 
nomere long, as long as next two anten- 
nomeres combined, second shorter than first 
and third. First three antennomeres with 
large black setae, surface shining, 4—11 
with large black setae and small dense and 
paler setae; antennomeres 4—11 large, never 
transverse, decreasing in length to 10, 11 
larger than 10. Last palpomeres (maxillary 
and labial) larger than the preceding ones, 
slightly paler than rest and truncate at apex 
that is wider in midline. Right mandible 
with a tooth opposite emargination of left 
mandible. Gular sutures separate but con- 
fluent at basal 4%. Nuchogenal ridge origi- 
nates in upper lateral face of neck and con- 
tinues well developed to base of gena, and 
then remains only as a depression in genae 
in direction of mandible base. 

Thorax: Pronotum slightly larger than 
head; narrower toward base; in lateral view 
anterior angle slightly projected below; disc 
with several setiferous punctures but scat- 


tered, dorsal punctures 12—14 on each side, 
asymmetrical and not in a well-defined line; 
without postcoxal translucid process on hy- 
pomeron. Elytra covered densely with short 
black setae, setiferous punctures denser 
than on prothorax, with two humeral black 
macrosetae and one near scutellum. Two 
large macrosetae on anterior middle of 
prosternum. Tibiae with sparse spines, few- 
er on protibia. First four protarsomeres 
strongly dilated in both sexes, with modi- 
fied pale setae ventrally; last tarsomere as 
long as previous three tarsomeres com- 
bined. Middle and hind tarsomeres 2—4 
moderately dilated and ventrally lobed (but 
less than protarsomeres). 

Abdomen: Segments shining; tergites 
III—V with impressed line on basal portion, 
less evident on tergite V. Pubescence fine, 
scattered, setiferous punctures not as deep 
as elytral punctures. Sternite VIII of male 
emarginated with three black macrosetae on 
each side. Sternite [IX emarginate, with two 
black macrosetae. Aedoeagus as in Figs 2— 
4. Parameres with apex almost reaching 
apex of median lobe (Fig. 2), peg setae 


890 


more abundant near apex (Figs. 3—4), apex 
of parameres with four large setae (Fig. 4). 

Material examined.—Holotype male: 
México: Jalisco, Autlan, Puerto los Mazos, 
BMM, 1,800 m, 13.1X.1999, 19°41.3'N, 
104°23.7'W, J.L. Navarrete-Heredia col., 
Phanolinus? gravidus Shp., J.L. Navarrete- 
Heredia det., and my holotype label: HO- 
LOTYPE, Phanolinus elisae Navarrete-He- 
redia, des. Navarrete-Heredia 2002 
(CZUG). Paratype: 1 2, same data as ho- 
lotype, except: yellow label: PARATYPE, 
Phanolinus elisae Navarrete-Heredia, des. 
Navarrete-Heredia 2002 (CZUG). 

Etymology.—This species is dedicated to 
my daughter, Elisa Margarita, as the ““beau- 
tiful”’ lady that she is to me and to my fam- 
ily. 

Remarks.—Phanolinus elisae is close to 
P. speciosus Sharp and P. gravidus Sharp 
all of which have a similar color pattern; 
however, P. elisae is easily recongnized by 
the basal transverse impressions on tergites 
II-V. Phanolinus speciosus lacks these im- 
pressions, and in P. gravidus they are pre- 
sent as restricted discrete impressions only 
on the lateral portions of tergites III-—IV. 
Also, there are more than 14 pronotal punc- 
tures in P. speciosus and only nine in P. 
gravidus, whereas in P. elisae are 12-14. 
The abdomen in P. speciosus is mostly dull 
due to the meshed micro sculpture, and the 
iridiscent portions are restricted to the cen- 
tral area of the tergites. In P. elisae, the 
abdomen is mostly shining. 


COMMENTS ON THE MEXICAN SPECIES 
OF PHANOLINUS 


Phanolinus obsoletus Sharp, 1884: 364 


This species was described from one 
specimen collected in México without spe- 
cific locality. No biological information 1s 
available. The type specimen is labeled (on 
card, handwritten): Phanolinus obsoletus, 
Type D.S., Mexico, Coll. Guerin-Men., 
aeodeagous dissected and mounted on card/ 
circle with red border, printed: Type/B.C.A. 
Col. 1.2, Phanolinus obsoletus, Sharp/ 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Sharp Coll. 1905-313. As this species was 
described from a single specimen, I added 
a red holotype label: HOLOTYPE, Phano- 
linus flohri Sharp, teste. J.L. Navarrete-He- 
redia 2002. Later, Sharp (1887: 784) cited 
another specimen from Xalapa, Veracruz. 
The label data for that specimen is: Jalapa, 
Mexico, Hoege/B.C.A. Col. 1.2, Phanolinus 
obsoletus, Sharp/ handwritten: Phanolinus 
obsoletus Sharp, °. 


Phanolinus flohri Sharp, 1884: 364 


This species was described from one 
specimen collected in Xalapa, Veracruz. No 
biological information is available. The 
type specimen is labeled (on card, hand- 
written): 2, Phanolinus flohri, Type D.S., 
Jalapa ex Almolonga, Mex., Flohr/ circle 
with red border, printed: Type/B.C.A. Col. 
1.2, Phanolinus flohri, Sharp/ Sharp Coll. 
1905-313. As this species was described 
from a single specimen, I added a red ho- 
lotype label: HOLOTYPE, Phanolinus floh- 
ri Sharp, teste. J.L. Navarrete-Heredia 
2002. 

Additional specimens examined.—Vera- 
cruz, Naolinco, 1.XI.1975, G. Figueroa, 
Phanolinus flohri Shp., J.L. Navarrete-He- 
redia det. 2003 (1 ¢@: CNIN); Xalapa, 
7.XI1.1980, J. Llorente, leg.A. Luis, Phan- 
olinus flohri, J.L. Navarrete-Heredia det. (1 
Ge CZUG): 


Phanolinus speciosus Sharp, 1884: 367 


This species was described from two 
specimens (syntypes), a male and a female, 
both collected in Cordoba, Veracruz. No bi- 
ological information is available. One spec- 
imen is labeled (on card, handwritten): d, 
Phanolinus speciosus, Type D.S., Cordova, 
Mexico, Sallé/circle with red border, print- 
ed: Type/ B.C.A. Col. 1.2, Phanolinus spe- 
ciosus, Sharp/ Sharp Coll. 1905-313. I se- 
lect this specimen as lectotype to ensure an 
accurate application of this name. A red la- 
bel with my lectotype designation is at- 
tached: LECTOTYPE, Phanolinus specio- 
sus Sharp, des. J.L. Navarrete-Heredia 
2002. The female is designated a paralec- 


VOLUME 107, NUMBER 4 


totype and has the following label data (on 
card, handwritten): 2°, Phanolinus specio- 
sus, Type D.S., Cordova, Mexico/ circle 
with red border, printed: Type/ B.C.A. Col. 
1.2, Phanolinus speciosus, Sharp, and my 
yellow paralectotype label: PARALECTO- 
TYPE: Phanolinus speciosus Sharp, des. 
J.L. Navarrete-Heredia 2002. 


DISCUSSION 


The known distribution of the Mexican 
Phanolinus species is scattered and restrict- 
ed primarily to Veracruz State, due primar- 
ily to the fieldwork efforts on that area; 
however, their presence in Jalisco consid- 
erably increases our knowledge of their dis- 
tribution and agrees with patterns observed 
in other Xanthopygina, such as species of 
Xenopygus and Xanthopygus (Navarrete- 
Heredia et al. 2002; Navarrete-Heredia 
2004) known to occur in Sinaloa, Nayarit, 
and or Jalisco in western México. It is pos- 
sible that more specimens will be found in 
other tropical localities of the Pacific coast 
and Veracruz and possibly in San Luis Po- 
tosi due to similar climatic conditions and 
vegetation types. 


KEY TO MEXICAN PHANOLINUS 


1. Tergites I—V without transverse basal impres- 
SOUS werticvee2nteletareunsGuaree memetaseres 
— Tergites HI—V or only tergite III with trans- 


P. speciosus 


verse basal depressions; in P. gravidus restrict- 
ed as discrete impressions on lateral portions 
Ofster ote MD ares ces otis! chee ORNS eRe: 2 
. With 12-14 pronotal dorsal setiferous punc- 
tures; head, pronotum and abdomen blue violet 
P. elisae 


NO 


— With less than 10 pronotal dorsal setiferous 
punctures; head, pronotum and abdomen gold- 


891 


en coppery, bright colored, never blue violet 


3. With 4 widely separated pronotal dorsal seti- 
ferous punctures, distributed on no more than 
*4 of pronotal length P. flohri 

— With 6-8 pronotal dorsal setiferous punctures, 
some widely separated, others close together, 
distributed close to basal border of pronotum 

P. obsoletus 


ACKNOWLEDGMENTS 


I thank M. Brendell for his kind help and 
friendship during my visit to The Natural 
History Museum (London) to study type 
material, H. E. Fierros-L6pez for the draw- 
ings, and two anonymous reviewers for the 
critical revision of the manuscript. 


LITERATURE CITED 


Herman, L.H. Jr. 2001. Catalog of the Staphylinidae 
(Insecta: Coleoptera). 1758 to the end of the sec- 
ond millennium. Bulletin of the American Muse- 
um of Natural History 265: 1—4218. 

Navarrete-Heredia, J. L. 2004. Sinopsis del género 
Xanthopygus Kraatz, 1857 (Coleoptera: Staphylin- 
idae) de México. Acta Zoologica Mexicana (n.s.) 
20(3): 1-13. 

Navarrete-Heredia, J. L., A. E Newton, M. K. Thayer, 
J. S. Ashe, and D. S. Chandler.2002. Guia ilustra- 
da para los géneros de Staphylinidae (Coleoptera) 
de México. Illustrated guide to the genera of Sta- 
phylinidae (Coleoptera) of Mexico. Universidad 
de Guadalajara y Conabio, México. 401 pp. 

Scheerpeltz, O. 1968. Eine neue Art der Gattung Phan- 
olinus Erichson (Col.: Staphylin.), nebst einer 
Ubersicht iiber die bisher bekannt gewordenen Ar- 
ten dieser Gattung. Revista de la Facultad de 
Agronomia (Maracay) 4: 51—68. 

Sharp, D. 1884. Fam. Staphylinidae, pp. 313-392, pls. 
8-9. In Biologia Centrali-Americana. Insecta, Co- 
leoptera. Vol. 1(2). Taylor & Francis, London. 

. 1887. Fam. Staphylinidae, pp. 673-824, pls. 

18-19. Jn Biologia Centrali-Americana. Insecta, 

Coleoptera. Vol. 1(2). Taylor & Francis, London. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 892-902 


KEY TO THE PUPAE OF THE MOSQUITOES 
(DIPTERA: CULICIDAE) OF FLORIDA 


RICHARD E DARSIE, JR. 


Florida Medical Entomology Laboratory, 200 9'" Street SE, Vero Beach, FL 32962, 


U.S.A. (e-mail: rfd@mail.ifas.ufl.edu) 


Abstract.—A key is presented for the known pupae of the mosquito species in Florida. 
This will complement the recent keys to adult females and fourth-instar larvae by Darsie 
and Morris. The pupal stage for all 78 species in Florida are known, except Ochlerotatus 
condolescens (Dyar and Knab). The sources for pupal descriptions are included. 


Key Words: 


Of all the stages in the life cycle of mos- 
quitoes, the pupa and possibly the male and 
its genitalia, are of least interest to practic- 
ing mosquito control agencies. Yet, the 
more stages that are known and can be 
identified the more complete will be the 
knowledge of the species encountered in a 
district. One problem that confronts those 
interested in pupal identification is the prep- 
aration of specimens for study. Whole pu- 
pae are not ordinarily used; instead the pu- 
pal exuviae are employed. They are usually 
prepared as part of an individual rearing 
when larval and pupal exuviae are associ- 
ated with the emerged adults. The rearing 
technique is described by Darsie (1951). 

Gradually a body of knowledge has ac- 
cumulated in many taxonomic works de- 
scribing the pupa along with the other stag- 
es from the fourth-instar larva to the adult. 
Sufficient descriptions have been published, 
in addition to my own unpublished work, 
to devise a key to the 78 species now 
known from Florida, except Ochlerotatus 
condolescens (Dyar and Knab), recently re- 
ported from Florida by Darsie (2003), 
whose larva and pupa are unknown. Major 
nomenclatural changes occurred when Rei- 
nert (2000) elevated Ochlerotatus Lynch 


pupal key, mosquitoes, Florida 


Arribalzaga to generic rank and Reinert et 
al. (2004), studying the tribe Aedini, further 
raised Howardina Theobald and Stegomyia 
Theobold to genus as well as other former 
subgenera and one new genus in the Nearc- 
tic Region not found in Florida. 

Here is a list of the publications which 
contributed to formulating the keys: Arnell 
(1976), Barr (1963), Barr and Barr (1969), 
Belkin et al. (1970), Berlin (1969), Darsie 
(1949, 1951, 2001, 2003, 2005), Darsie and 
Day (2003), Floore et al. (1975), Lacey and 
Lake (1972), Reinert @970amby crdiver te 
g, 1971), Reinert et al. (1997), Zavortink 
(1968, 1972), Zavortink and O’Meara 
(1999). 


MORPHOLOGY OF THE PUPAL EXUVIAE 


It is customary to mount the pupal exu- 
viae, for this, not the whole pupa, is used 
to study the pupal stage. The head capsule, 
prothorax, and mesothorax are closely unit- 
ed into the cephalothorax (CT), whereas the 
metathorax is reduced to a dorsal plate 
known as the metanotum. The scutum of 
the mesothorax is split longitudinally dur- 
ing eclosion, therefore the cephalothorax is 
mounted ventrally so that the external 
halves are uppermost after detaching the 


VOLUME 107, NUMBER 4 


893 


Figs 1. 
lateralia, D—clypeus, E—labrum, F—mandible, G—maxilla, H—maxillary palpus, J 
L—midleg, M—hindleg, N—mesothoracic wing, O—trumpet, P—tracheal trunk, R—scutum. 


metanotum and abdomen from the remain- 
der of the cephalothorax. Hence, the mouth- 
parts are located centrally, the halves of the 
scutum with the trumpets and the mesotho- 
racic wings are lateral in position. 

Head (Fig. 1): The dorsal apotome (A) 
is located anteromedially, attached to the 
clypeus (D) and the bases of the antennae 
(J). The labrum (E), fused basally with the 
clypeus, forms the central, long, narrow 
mouthpart. The mandibles (F), which bor- 
der the labrum on each side, are also long 
and narrow, followed laterally by the max- 
illae (G), which are joined at the bases by 


Cephalothorax of pupal exuvium in slide mounted position. Abbreviations: A—dorsal apotome, C— 


antenna, K—foreleg, 


the maxillary palpi (H). The bases of the 
antennae lie in a fold which develops as a 
result of the cephalothorax being mounted 
ventrally and the long antennae are found 
lateral to the mouthparts. The lateralia (C) 
lie anteriorly, mesad to the antennal bases 
and are mainly located in the fold. The la- 
teralia bear three rather prominent setae, 
1,2,3-CT. 

Thorax (Fig. 1): The pronotal sclerites 
are small and usually misshapen by the 
folding. Setae 4—7-CT are attached to this 
area. The largest sclerite of the cephalotho- 
rax is the scutum (R), split in half by eclo- 


894 


sion. The margin along the split is known 
as the median keel and a creaselike line par- 
allel to the keel is the lateral line. The 
small, dorsomedial sclerite of the scutum 
bears an anterior angle. The scutum has se- 
tae 8,9-CT and a more or less cylindrical 
trumpet. The tubular portion is the meatus 
and the open, distal portion is the pinna. 
The base of the trumpet is connected to the 
tracheal trunks (P) of the respiratory sys- 
tem. The trumpet index is calculated by di- 
viding the length of the organ by the width 
at midlength. Lateral to the mandibles and 
maxillae are the sclerites of the fore- and 
midlegs (K,L). Finally the mesothoracic 
wings (N) are posterior and sclerites of the 
hindlegs (M) are curled beneath them. 

The metanotum consists of two rather 
quadrangular sclerites connected by a 
bridge. They cover the developing halteres 
and bear setae 10—12-CT. 

Abdomen (Figs. 2—4): Abdominal seg- 
ments I-VIII are more or less well devel- 
oped. Segment I is without a sclerotized 
sternum and is modified by the presence of 
float hairs, seta 1-I. The terga and the sterna 
of segments II—-VIII are well developed and 
consist of quadrangular sclerites adjoined 
by short intersegmental membranes. Poste- 
rior to segment VIII is a flap, segment IX, 
a genital pouch in which genitalia of the 
adult develop, and the paddle (Pa). The 
paddles are variously shaped, usually oval. 
Each is supported by an external buttress, a 
thickening along the basal 0.75 of the ex- 
ternal margin, and a midrib, a similar thick- 
ening located medially. The external margin 
of the paddle sometimes bears short or long 
spicules or coarse denticles. The paddle in- 
dex is calculated by dividing the paddle 
length by the greatest width. 

Chaetotaxy of the abdomen: The present 
nomenclature for the abdominal setae fol- 
lows Belkin (1962) and is shown in Figs. 
2—4. Those studying older pupal literature 
will find it helpful to consult a table in Har- 
bach and Knight (1980) comparing various 
past nomenclature with the one in Belkin 
(1962). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


The keys to the pupae of the mosquitoes 
of Florida follow. The reader is referred to 
the Systematic Index of the Mosquitoes of 
Florida by Darsie and Morris (2003) for 
complete species detail. The keys were test- 
ed with pupae from the author’s collection 
in which 60 of 78 species were available. 
For the other 18 species, descriptions and 
illustrations from the literature, except for 
Oc. condolescens, were employed (see lit- 
erature citations above). I have three pupae 
of Oc. canadensis mathesoni Middlekauff 
from Camp Blanding, Clay County, Flori- 
da, collected on II-19—46. They are quite 
similar to the pupa of the typical subspe- 
cles. 

Four illustrations are included to assist 
users of the key, namely, cephalothorax, 
Fig. 1; Anopheles crucians Wiedemann, 
Fig. 2; Ochlerotatus sollicitans (Walker), 
Fig. 3; and Culex nigripalpus Theobald, 
Fig. 4. Figures 2—4 represent the three ma- 
jor genera of mosquitoes in Florida and 
have the setae numbered. These three fig- 
ures will help in dealing with the identifi- 
cation of minor genera. 


KEYS TO THE PUPAE OF THE MOSQUITOES OF 
FLORIDA KEY TO GENERA 


1. Seta 9-III-VI at or very near caudolateral 
angle of tergum, usually distinctly spini- 
form; meatus of trumpet deeply slit to near 
base Anopheles 

~ Seta 9-III-VI distinctly removed from cau- 

do-lateral angle of tergum; meatus of trum- 

pet usually split a short distance from pin- 

ma, iat tall os ae icycwcy vetoes eae Ree eae 2 

Meatus of trumpet with distinct tracheoid 

extending considerable distance from base 


2(1). 


beat ett iarina sh Oem racine ee A eerie aor 3 
- Meatus of trumpet without tracheoid or a 
faint tracheoid near base ............ 7 
3(2). Trumpet without pinna, apical process 
pointed, adapted for piercing plant tissue 
eer AO eta erty AR mht cowie Jere 4 
~ Trumpet with distinct open pinna ...... 5 
4(3). Seta 6-I-VI absent; paddle emarginate api- 
cally, with 2 equal lobes .......... 
4 Td cope of sh i ae Coquillettidia perturbans 
= Seta 6-I-VI present; paddle emarginate 
apically on inner margin only, outer lobe 
deévelopedeeaeare en creer aeaene- Mansonia 
5(3). Seta 8-CT closer to base of trumpet than 


VOLUME 107, NUMBER 4 a05 


Vee ee ee" a 
SS a ee 


Fig. 2. Pupa of Anopheles crucians. A, Cephalothorax. B, Metanotum and abdomen, dorsal left, ventral 


right. Abbreviations: CT—cephalothorax, Pa 


paddle. 


896 


9(8). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


to seta 9-CT, at or near level of trumpet 
Bre atondr Chaeoncl ar oRor eo Gicro amOncaNO tc Uranotaenia 
Seta 8-CT closer to seta 9-CT than to base 
of trumpet, removed far caudad of trumpet 


Paddle with seta 1-P subequal to paddle 


ISIN 6 6 ce ago0d0000 Deinocerites cancer 
Paddle with seta 1-P much shorter than 
BENGKGIE: Gicig og Glave oa oe os 6 Ho S)G.0'6 ol om Culex 


Anal segment with conspicuous cercal seta 
DENG esha eat Toxorhynchites r. rutilus, 
Toxorhynchites r. septentrionalis 
Cercal seta 1-X absent 
Setae 9-VII,VHI subequal, both large with 
numerous branches, paddle small, without 
setae Wyeomyia 
Seta 9-VII rarely subequal to 9-VIII, both 
with fewer branches; paddle normally with 
at least one seta 
Setae 8,9-CT in line perpendicular to me- 
dian keel; paddle without marked infusca- 
tion near external buttress and apex ... 10 
Seta 8-CT usually distinctly more anterior 
than 9-CT, if rarely the two setae in line 
perpendicular to median keel, then paddle 
with marked infuscation near external but- 


tresspandkapexa marys re ener mc urene 11 
. Abdominal seta 1-VI lateral to setae 2,3- 

WAR ests ine aah Biya Sh ee ta ha a cee ee Culiseta 

Abdominal seta 1-VI mesad of setae 2,3- 

VAD oye speek sa a, SB easel Orthopodomyia 
. Abdominal seta 5-II mesad of seta 4-II or 

paddle deeply infuscated near external but- 

MESS ANG! AEX soacbacsnadee Psorophora 


Abdominal seta 5-II lateral of seta 4-II and 
paddle not infuscate ... Aedes, Ochlerotatus, 


Howardina, Stegomyia 


KEY TO THE GENERA AEDES (AE), 


OCHLEROTATUS (OC), HOWARDINA (Hw) AND 


AX). 


STEGOMYIA (ST) 


Seta 6-CT long, stout, longer than seta 7- 


CLM EA Paes hen a crete alaenas apne D 
Seta 6-CT short to moderately long, usu- 
ally much shorter than seta 7-CT ..... 3 


Seta 9-VIII single; paddle with long, mar- 
ginal spicules St. albopicta 
Seta 9-VIII with 3-8 branches; paddle 
margin with short, coarse spicules 


St. aegypti 
Seta 5-VII short, 0.3 or less length of fol- 
lowing tergum 
Seta 5-VII moderately long to long, 0.5 
or more length of following tergum .. 12 
Anterior border of abdominal segment 
VIII almost as wide as posterior border of 
VII; paddle with margin smooth; seta 6- 


5(4). 


6(5). 


WO) 


8(7). 


9(8). 


12(3). 


13(12). 


VII ventral and removed cephalad from 
posterior border ...... Oc. fulvus pallens 
Anterior border of segment VIII definitely 
narrower than posterior border of VII; 
posterior border paddle with marginal and 
submarginal spicules; seta 6-VII dorsal 
and near posterior margin 
Seta 1-II with 14 or more branches; seta 
6-VI single; paddle slightly emarginate 
posteriorly 
Seta 1-II with 12 or fewer branches; seta 
6-VI usually double or triple; paddle 
evenly rounded posteriorly 
Seta 1-III usually with 6 or more branch- 
es; paddle as wide as long or nearly so, 
with prominent marginal and sub margin- 
al spicules, longer than diameter of 1-P 
alveolus Oc. taeniorhynchus 
Seta 1-III with fewer than 6 branches; 
paddle longer than wide, with marginal 
and sub-marginal spicules usually smaller 
than seta 1-P alveolus 
Seta 5-IV—VI single, extremely long, lon- 
ger than following tergum .... Oc. dupreei 
Seta 5-IV—VI usually double, never all 
single, often no longer than following ter- 
UTM, dese eS lls. Aa vol Ma chee fecal oe em 8 
Seta 3-I-III and 11-CT usually all single, 
sum of all branches of the 8 setae no more 


Oc. scapularis 


thie Seca! WN Dre et ete Am auc aie 9 
Setae 3-I,I]J, 11-CT and usually 3-III with 
2 or more branches, sum of all branches 
of the 8 setae not less than 17 ...... 11 


Seta 6-I subequal to seta 7-I or somewhat 
shorter, about equal to median length of 
tergum I Oc. thelcter 
Seta 6-I much longer than seta 7-I and the 
median length of tergum I 
Seta 6-III double; seta 12-CT usually 
double Oc. infirmatus 
Seta 6-III usually single; seta 12-CT with 
3 or more branches Oc. tortilis 
Seta 6-CT with 3 or more branches; api- 
cal margin of paddle spiculate 


WGie B Gace elo Er eae ae eae Oc. tormentor 
Seta 6-CT usually single; apical margin 
of paddle without spicules .. Oc. atlanticus 
Seta 1-VI and usually seta 1-V short, less 


than 0.5 length of following tergum; seta 


B=\Vi usualllyssin'ele sia crc ne 13} 
Seta 1-V,VI moderately long to long, 
more than 0.5 length of following tergum; 
seta 3-V usually double or multibranched 

BE See ee ee EN Gls, inga rors ot core 15 


Seta 1-IV usually 4- or 5-branched; seta 
5-IV longer than tergum V = Oc. hendersoni 
Seta 1-IV usually double or triple (2—4); 


VOLUME 107, NUMBER 4 897 


Fig. 3. Pupa of Ochlerotatus sollicitans. A, Cephalothorax. B, Metanotum and abdomen, dorsal left, ventral 
right. Abbreviations: CT—cephalothorax, Pa 


paddle. 


898 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ausencn 


Us 


Fig. 4. Pupa of Culex nigripalpus. A, Cephalothorax. B, Metanotum and abdomen, dorsal left, ventral right. 
Abbreviations: CT—cephalothorax, Pa—paddle. 


seta 5-IV usually shorter than tergum V single; seta 3-VII with 4 or more branches 

MOMS oan aOR kot ond Ue eeai te. RSM NO eeS 14 Jussi ey, Suetbenes eo aushenuied Gachalea ie cutee eC MV EXCILS| 
14(13). Seta 6-I single; seta 7-I usually double; — Sternum II without apical spicules; seta 

seta 9-VIIIT usually with fewer than 7 3-I usually with 2 or more branches; seta 

branches” See a ee Oc. triseriatus 3-VII usually double or single ...... 16 
= Seta 6-I usually double; seta 7-I usually 16(15). Seta 1-V and usually seta 1-VI much lon- 

triple; seta 9-VIII usually with more than ger and stouter than any other setae on 

Stbranches? +3. eae ee Hw. bahamensis tergum, except seta 5 and sometimes seta 


15(12). Sternum I with apical spicules; seta 3-I LOVEE SS oe Ss ae are ee 17 


VOLUME 107, NUMBER 4 


17(16). 


18(17). 


19(16). 


20(19). 


2(1). 


4(3). 


5(4). 


6(5). 


Seta 1-V,VI only slightly stouter and usu- 
ally only slightly longer than any other 
seta on tergum, except seta 5 and some- 


times seta 10-V,VI, if atall ........ 19 
Seta 5-[V—VI shorter than following ter- 
gum; seta I-VI usually triple ...... 

ng alo. Bea EMOTO MORRO eC ORS Oc. mitchellae 
Seta 5-IV—VI as long as or longer than 
following tergum; seta 1-VI usually sin- 
leper y hese cnet tele tick ct «.d 18 


Seta 9-VII with 5 or more branches 

Oc. sollicitans 
Seta 9-VII with 3 or fewer branches ... 

Oc. c. canadensis, Oc. c. mathesoni 
Seta 1-IV,V longer than following tergum 


Oc. sticticus 
Seta 1-IV,V not as long as following ter- 
fulllineles. a cued eo mcr crete te cre picees otha tren 

Seta 3-IL,[JJ and 5-IV—VI almost invari- 
ably single; seta 5-IV occasionally double 
or triple; seta 9-VIII single, rarely double 


Ae. cinereus 
Seta 3-II, II] and 5-IV-VI never all single; 
seta 9-VIII usually with 3 or more 
branches Oc. thibaulti 


KEY TO THE GENUS ANOPHELES 


Abdominal setae all very short, 1-[V—VI 
0.2 length of following tergum .... barberi 
Abdominal setae long and short, 1-[V—VI 
at least 0.5 length of following tergum 2 
Seta 9-VIIf usually single, rarely forked 
apically; seta 1-[V—VII all single and 
about as long as following tergum 

PES set ESET oS Senos Shee albimanus 
Seta 9-VIII with thickened shaft and 
many lateral branches; seta 1-[V-VII usu- 
ally all branched and shorter than follow- 
ing tergum 
Paddle with coarse, blunt teeth on exter- 


MaAlemMar Cuneewee a ce oy cl cmeets enous walkeri 
Paddle without coarse teeth on external 
IMAGO UMM event ees mile. Sustiencs lie ksrg Muistee 4 


Seta 2-V usually with 4—6 branches; seta 
0-V with 2-11 branches, rarely single; 
seta 11-CT usually with 6 branches ... 5 
Seta 2-V usually single to triple; seta O- 
V single seldom double; seta 11-CT usu- 
ally with 3—5 branches 
Seta O-IV large, usually with 2—5 branch- 
es; seta O-V large, with 3—11 branches 
FN ds ree as eee SRI = HAIG EVA crucians 
triple 
Seta 1,5-I['V with 5—10 branches, usually 
with 5,6 branches; seta 1-V and 5-VI with 
3-8 branches; seta 5-V with 3—5 branches 
bradleyi 


9(8). 


10(9). 


11(10). 


12(10). 


13(7). 


899 


Seta 1-IV with 9—14 branches: seta 1-V 
with 6—10 branches; seta 5-ITV with 12— 
17 branches; seta 5-V with 8—16 branch- 
es; seta 5-VI with 9—13 branches 


EIS StS APE Aber ip, d0+0 Ese ORB selon eo georgianus 
Seta 6-V double or triple; seta 8-CT usu- 
ah? GlotinS Or tiny 2 osencadacasce 8 
Seta 6-V and seta 8-CT single, seldom 
GOUD egies Sygate: Paes Re as Seach 13 
Seta 9-I single, seldom double; length of 
seta 9-VIT usually 7.0 or greater than bas- 
EEA A(t ol cane aNRS Ronotoeo: cro ra Naaey ator te atropos 
Seta 9-I usually with 2 or more branches; 
length of seta 9-VII usually 6.8 or less 
than basal width (quadrimaculatus com- 
DICK) i cas cer th tier octet he a ca eee ee 9 


Dorsal apotome with well developed me- 
dian apical projection; scutum with ante- 
rior angle broad, approximately 90° 

Ser Pl he waarmee er ea orn eien = see maverlius 
Dorsal apotome without apical projection; 
scutum with anterior angle acute .... 10 
Cephalothorax with lateral line on median 
keel long, extending posteriorly to seta 8- 
CT; cephalothorax with postscutal area 
split by dorsal ecdysial opening; seta |- 
VII usually slightly longer than length of 
tereum VILL eee ed tee) eet ee ons 11 
Cephalothorax with lateral line of median 
keel short, extending posteriorly only to 
trumpet base; cephalothorax with posts> 
cutal area intact; seta 1-VII usually 0.75 
or less length of tergum VIII 
Sum of branches for both setae 1-P usu- 
ally 4-12; seta 1-P usually with 2-6 
branches; sum of both setae 9-VIII usu- 
ally 24—48 
Sum of branches for both seta 1-P usually 
2,3; seta 1-P usually single; sum of 
branches for both setae 9-VIII usually 


quadrimaculatus ss 


ET oer oe Oba ore carte smaragdinus 
Sum of branches for both setae 10-CT 
usually 2—5; distribution limited to part of 
Ploniday: eter. och ental Wea neeote< ter abe diluvialis 
Sum of branches for both setae 10-CT 
usually 6-11; distribution in at least 3 
StatestomU SiAuers ses .u-ueist et eine inundatus 
Seta 6-I single to triple; seta 9-IV 0.67 or 
more length of seta 9-V 
Seta 6-I with 5 or more branches; seta 9- 
IV 0.63 or less length of 9-V ... grabhamii 


punctipennis 


KEY TO THE GENUS CULEX 


Seta 5-CT very long, about 5.0 length of 
seta 4-CT; abdominal tergum VIII with 
posterior lobe overlying lateral part of ter- 
gum IX; seta 1-IX absent (subgenus Mi- 
craedes) biscaynensis 


900 


2(1). 


4(3). 


5(4). 


6(4). 


7(6). 


8(7). 


9(2). 


10(9). 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Seta 5-CT no more than 2.0 length of seta 
4-CT; abdominal tergum VIII with pos- 
terior lobe not overlying lateral part of 
tergum IX; seta 1-IX present ........ 2 
Meatus of trumpet with narrow slit ex- 
tending from proximal part of pinna; seta 
2-VI usually lateral of seta 1-VI (subge- 
nus Melanoconion) 
Meatus of trumpet without narrow slit ex- 
tending from proximal part of pinna; seta 
2-VI mesad of seta 1-VI 
Seta 9-VIII very near posterolateral cor- 
ner; posterolateral corner of segment VIII 
gently rounded cedecei 
Seta 9-VIII distinctly removed from pos- 
tero-lateral corner; posterolateral corner 
of segment VIII with distinct point .... 4 


Seta 5-V double or triple and nearly as 
long as following tergum ........... 5 
Seta 5-V usually with at least 4 branches, 
if fewer. then distinctly shorter than fol- 
lowing tergum 
Seta 1-II with 14 or fewer branches; pinna 
of trumpet, including slit, less than 0.3 of 
total trumpet length ............ pilosus 
Seta 1-II with 25 or more branches; pinna 
of trumpet including slit 0.4—0.5 of total 
trumpet length erraticus 
Seta 6-IV,V at least 4-branched; seta 5-V 
usually 5-branched; trumpet index usually 
SASK? WORKIN Cho) oo o00080000500c 
Seta 6-IV,V usually triple; seta 5-V usu- 
ally 4-branched; trumpet index about 8.0 


atratus 


Pinna including slit about 0.35 length of 
trumpet; seta 8-CT single..... 
Pinna including slit about 0.4 length of 
trumpet; seta 8-CT with 3 or more 
branches 
Trumpet lighter between apex of trach- 
eoid and base of pinna, flared apically 


mulrennani 


iolambdis 
Trumpet uniformly dark distal to apex of 
tracheoid and base of pinna, if lighter, 
then remainder of trumpet also lighter, not 
flaredyvapicalllyaeruse inner ne 


peccator 
Trumpet index 7.7; seta 9-VII usually 
double; seta 9-VIII usually 4-branched 


(subgenus Neoculex) territans 


Trumpet index about 7.0; seta 9-VII usu- 
ally with 4 or more branches; seta 9-VIII 
usually 6-branched or more (subgenus 
Gules agués, Ses eo ease 10 
Abdominal tergum I with distinct short 
spicules posterior bahamensis 
Abdominal tergum I without short spic- 
WES DONIEHIO? ccoccccccacccéunsoa 11 


11(10). Seta 2-P absent on paddle; seta 1,6-VI 


12(11). 


usually double restuans 
Seta 2-P present, rarely absent; seta 1,6- 
VI usually with 3 or more branches .. 12 
Trumpet index about 5.0; seta 1-VII usu- 
ally with 4 or more branches 


quinquefasciatus 
Trumpet index 5.5 or more; seta 1-VII 


usually double or triple ........... 13 
13(12). Seta 1-II with 16 or more branches .. . 14 
- Seta 1-IJ with 15 or fewer branches .. 15 
14(13). Posterior border of abdominal sternum II 
with short, sharp spicules; seta 5-[V—VI 
shorter than following tergum nigripalpus 
- Posterior border of abdominal sternum II 
without spicules; seta 5-[V—VI longer 
than following tergum ........ declarator 
15(13). Trumpet index about 7.0; pinna short, 
about 0.13 of total trumpet length ..... 
ERY: Veneta Oe Nth Ba an At ah Sra roe salinarius 
_ Trumpet length about 6.0; pinna longer, 
about 0.16 of total trumpet length . . tarsalis 


2(1). 


KEY TO THE GENUS CULISETA 


. Seta 2-P present; meatus of trumpet with nar- 


row slit extending from proximal part of pinna 
almost to base; seta 2-II longer than seta 3-II 
melanura 
Seta 2-P absent: pinna of trumpet without slit; 
seta 2-II usually much shorter than seta 3-II 
inornata 


KEY TO THE GENUS MANSONIA 


. Trumpet broad, index about 6.0; paddle mod- 


erately broad, index about 2.5 titillans 
Trumpet slender, index about 10.0; paddle nar- 
row, index more than 3.0 dyari 


KEY TO THE GENUS ORTHOPODOMYIA 


. Seta 2-II-VI 0.4—0.6 length of seta 1-II—VI; 


seta 5-III longer and stronger than seta 3-III; 
setae 6,7-I weakly developed ........... 
Seta 2-II-VI less than 0.3 length of seta 1-II— 
VI; seta 5-III not as long nor as strong as seta 
3-III; setae 6,7-I long, strong signifera 


alba 


KEY TO THE GENUS PSOROPHORA 


Posterolateral corner of abdominal segment 
IV with large spines; seta 2-P usually absent 
(subgenus Janthinosoma) 
Posterolateral corner of IV without large 
spines; seta 2-P usually present, sometimes 
indistinct 
Seta 10-CT with 3 or fewer branches; seta 
5-II usually with 4 or fewer branches; seta 
1-III usually with 6 or fewer branches 
mathesoni 


VOLUME 107, NUMBER 4 


3(2). 


6(5). 


7(5). 


8(7). 


9(8). 


Seta 10-CT and seta 5-II usually with 5 or 
more branches; seta 1-III usually with 7 or 
more branches 
Heavily pigmented, appears brown, not yel- 
low; trumpet index less than 3.0; seta 12- 
CT usually with 5 or more branches..... 
AEE ARN Wo Mieerseke, 2 cairedcekt Packets Johnstonii 
Lightly pigmented, appears yellow; trumpet 

index 3.0—4.0; seta 12-CT usually with 4 or 

fewer branches (except horrida)........ 4 


. Seta 12-CT single or double; seta 4-I,[ with 


4 or fewer branches 
Seta 12-CT and 4-I,II usually with 4 or more 
branches horrida 


. Seta 4 closer to seta 5 than to seta | on 


abdominal segment III, usually anterior to 
and subequal to seta 5; seta 5-II,III usually 
with 3 or fewer branches; seta 9-VIII usu- 
ally with 5 or fewer principal branches; 
pupa large (subgenus Psorophora) 
Seta 4 closer to seta 1 than to seta 5 on III, 
seta 4 usually posterior to and much longer 
than seta 5; seta 5-II,III usually with 4 or 
more branches; seta 9-VIII usually with 6 or 
more principal branches; pupa medium or 
Small ies techs pars a eit ego wae hers ok ayn oe q 
Abdominal segment VIII with dark spots 
anteriorly and posteriorly on either side of 
midline, segment usually spotted; without 
dark median stripe on abdomen; with diag- 
onal stripes enclosing conspicuous light ar- 
eas laterally on segment VII howardii 
Abdominal segment VIII without dark spots 
anteriorly and posteriorly, but usually with 
darker median stripe, segment usually rather 
evenly dark laterally and on midine, with 
light pigmentation elsewhere; with dark me- 
dian stripe on abdomen; without pattern as 
above but with diagonal stripes laterally on 
segments V—VII ciliata 
Seta 2-P usually absent; seta 11-CT usually 
with 3 or more branches; seta 4-I usually 
with 6 or more branches 
Seta 2-P usually present; seta 11-CT usually 
single, occasionally double; seta 4-I usually 
with 4 or fewer branches (subgenus Grab- 


cyanescens 


hamia) 
Seta 5-I with 4 or fewer branches; seta 7-I 
with 5 or more branches; seta 6-III,1V and 
1-VI with 3 or more branches 
Seta 5-I with 5 or more branches; seta 7-I 
with 4 or fewer branches; setae 6-ILI, I-VI, 
and 6-IV usually single or double 
Seta 1-P very long, 0.25 length of paddle; 
seta 1-VII usually placed about midway be- 
tween lateral border and middorsal line of 
tergum 
Seta 1-P usually about 0.16 length of pad- 


pygmaea 


discolor 


901 
dle; seta 1-VII usually much closer to mid- 
dorsal line than to lateral border of tergum 
J.storatemee date eee: Me eee ees 2 columbiae 


KEY TO THE GENUS URANOTAENIA 


1. Trumpet index about 11.0 or more; seta 11-CT 
usually doublevor tiiplesee eee eee sapphirina 
— Trumpet index less than 7.0: seta 11-CT usu- 


ally/sinsle SS ea a ee ee: lowii 
KEY TO THE GENUS WYEOMYIA 
Ile Paddle fringed with long, filamentous spic- 


ules on outer and apical 0.2 of inner margin; 

seta 4-VIII less than 0.2 length of paddle 

VS ROP. roel Mn apite, tes e Ee tale cath mitchellii 
= Paddle spicules short; seta 4-VIII 0.5 length 
OF paddle vim: xe! seoeauch ya eac ue (oH aeRO 2 
Seta 2 contiguous with seta 1 on abdominal 
segment VII; paddle index 1.5 
_ Seta 2 far removed from seta | on VII: pad- 

dle index usually greater than 1.5. . 


2(1). 
Sp a heel wae smithii 


vanduzeei 


ACKNOWLEDGMENTS 


I am indebted to T. G. Floore for per- 
mission to use Fig. 2 in this work. The man- 
uscript was reviewed by C. R. Rutledge- 
Connelly and C. N. Mores. This is Florida 
Agricultural Experiment Station Journal Se- 
ries No. R-10547. 


LITERATURE CITED 


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PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 903-913 


REVIEW OF OREGOCERATA RAZOWSKI (LEPIDOPTERA: TORTRICIDAE: 
EULIINI) WITH DESCRIPTIONS OF FOUR NEW SPECIES 


JOZEF RAZOWSKI AND JOHN W. BROWN 


(JR) Polish Academy of Sciences, Institute of Systematic Zoology, Slawkowska 17, 
Krakow, Poland; (JWB) Systematic Entomology Laboratory, PSI, Agricultural Research 
Service, U.S. Department of Agriculture, % National Museum of Natural History, Smith- 
sonian Institution, PO. Box 37012, MRC 168, Washington, DC 20013-7012, U.S.A. (e- 
mail: jbrown @sel.barc.usda.gov) 


Abstract.—Oregocerata is a poorly known genus restricted to the Andes and its asso- 
ciated ranges in northwestern South America (i.e., Colombia, Venezuela, Ecuador, Bolivia) 
usually above 2000 m. Specimens are exceedingly rare in collections; most species are 
known only from the holotype male. We describe and illustrate four new species: O. 
triangulana (TL: Colombia), O. caucana (TL: Colombia), O. quadrifurcata (TL: Colom- 
bia), and O. submontana (TL: Venezuela); and we propose a new combination, Ore- 
gocerata chrysodetis (Meyrick). As currently defined the genus includes 9 species. It is 
assigned to Euliini on the basis of the possession of a characteristic hairpencil on the 


prothoracic leg. 


Key Words: 


Oregocerata Razowski, 1988, is one of 
several poorly known genera of Neotropical 
Euliini apparently restricted to the higher 
elevations of the Andes and its associated 
ranges in northwestern South America. 
Species of Oregocerata have been recorded 
from Colombia, Venezuela, Ecuador, and 
Bolivia, primarily at elevations above about 
2000 m. Specimens are rare in collections, 
with about half the species represented only 
by the holotype. Nothing is known of the 
biology. 

Originally described as monotypic, the 
genus includes four described species: O. 
orcula Razowski, O. cladognathos Razows- 
ki, O. rhyparograpta Razowski and Becker, 
and QO. zonalis Razowski and Becker. Four 
new species and one new combination, O. 
chrysodetis (Meyrick), are proposed herein. 
Superficially, all species share elongate la- 
bial palpi (length 2.0—3.0 times the hori- 


Tortricinae, systematics, Andes, Colombia, Ecuador, Bolivia, Venezuela 


zontal diameter of the compound eye), long 
antennal cilia in the male, and a patch of 
long, flattened orange scales extending 
from the base of the proboscis to the area 
between the bases of the foreleg coxae. The 
male genitalia usually are characterized by 
a short, somewhat digitate process at the in- 
ner base of the sacculus; a narrow, fringed 
flange or group of spiniform setae from the 
dorsum of the phallus (at the junction of the 
phallobase and aedeagus) representing a 
modification of the anellus; and a highly 
modified distal portion of the gnathos. 
However, because the genitalic characters 
have a mosaic distribution among the in- 
cluded species, none is a convincing syna- 
pomorphy for the genus. Females are 
known only for O. orcula and O. zonalis. 
The genus is assigned to Euliini based on 
the possession of a characteristic hairpencil 
of the prothoracic leg in the male (Brown 
1990). 


904 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Male genitalia characters distinguishing species of Oregocerata. See text for description of char- 


acters and character states. 


nnn UU TEESE EIS EIS SSDS an 


Species Sacculus Lobe Anellus 


Gnathos 


Uncus Socii 


a 


orcula absent unmodified moderate unmodified broad distally 
cladognathos weak unmodified long unmodified digitate throughout 
zonalis weak unmodified long reflexed distally broad throughout 
triangulana present with spines very long unmodified digitate throughout 
rhyparograpta present with thorns long unmodified digitate throughout 
submontana present with spines moderate unmodified digitate throughout 
quadrifurcata present unmodified short, blunt expanded distally broad throughout 
caucana present with spines short, curved unmodified digitate throughout 


 ————————— 


The purpose of this paper is to review the 
four described species, describe and illus- 
trate four new species, and propose one 
new combination in the genus. The follow- 
ing institutional abbreviations are used in 
the text: BMNH = The Natural History 
Museum, London, England; UCB = Essig 
Museum of Entomology, University of Cal- 
ifornia, Berkeley, USA; USNM = National 
Museum of Natural History, Washington, 
D.C., USA; and VBC = Vitor O. Becker 
personal collection, Planaltina, Brazil. Oth- 
er abbreviations are as follows: ca. = circa 
(approximately); GS = genitalia slide; n = 
number examined. 


SYSTEMATICS 
Oregocerata Razowski, 1988 


Oregocerata Razowski 1988: 392, 1997: 
92, 1999: 328: Powell et al. 1995: 145; 
Razowski and Becker 2002. Type spe- 
cies: Oregocerata orcula Razowski 1988, 
by original designation. 


The species included in Oregocerata (ex- 
cluding O. chrysodetis that lacks an abdo- 
men) can be separated by structures of the 
male genitalia. A summary of the most con- 
spicuous characters is presented in Table 1. 
In Table 1, ““Sacculus lobe’’ refers to the 
digitate process at the inner base of the sac- 
culus. It is absent in O. orcula, represented 
by a weakly swollen, setose mound in O. 
zonalis and O. cladognathos, and is a prom- 
inent digitate process in the remaining spe- 
cies. Anellus “‘with spines” refers to the 


finely spined membrane attached to the dor- 
sum of the phallus at the junction of the 
phallobase and the aedeagus. It is conspic- 
uous and well defined in O. caucana, O. 
triangulana, and O. rhyparograpta (the 
small projections are more thornlike than 
narrow spinelike in the last), and weak or 
absent in the remaining species. ““Gnathos”’ 
refers to the shape and/or length of the dis- 
tal portion of the gnathos arms. The gnathos 
has an unmodified basal portion and a var- 
iably modified distal part, which is extreme- 
ly short and blunt in O. quadrifurcata, short 
(less that 0.25 the length of the basal por- 
tion of the gnathos) and curved in O. cau- 
cana, and slender and long (0.5—0.7 times 
the length of the basal portion) to very long 
(equal to or longer than the basal portion) 
in the remaining species. ““Uncus”’ refers to 
the shape of the uncus, which is slightly 
modified in some species, but highly diver- 
gent in O. quadrifurcata, with an expanded, 
four-pointed apical process, and somewhat 
reflexed dorsally in O. zonalis. ““Socii” re- 
fers to the shape and relative width of the 
SOc. 


Oregocerata orcula Razowski 
(Figs. 1, 9) 


Oregocerata orcula Razowski 1988: 393 
(description, illustration of male and fe- 
male genitalia); Powell et al. 1995: 145 
(checklist). 


Diagnosis.—Superficially, O. orcula is 
similar to its congeners, with a grayish 


VOLUME 107, NUMBER 4 905 


ce t nm \y 
' \ 
Ss HIN 


Figs. 1-3. Male genitalia of Oregocerata; valvae spread, aedeagus removed. 1, O. orcula, 2, O. cladogna- 
thos, 3. O. zonalis. 


906 


brown forewing lacking distinct pattern el- 
ements. The male genitalia (Fig. 1) are 
characterized by the absence of the digitate 
process from the base of the sacculus, un- 
modified anellus, moderately long distal 
processes of the gnathos, simple uncus, and 
extremely broad distal portion of the socii. 
The last is the most conspicuous autapo- 
morphy for the species. In the female gen- 
italia (Fig. 9) the ventral lobes of the eighth 
tergite are large; and the anterior portion of 
the sterigma is rounded, strongly sclero- 
tized, and slightly asymmetrical. 
Holotype.—6, Bolivia, Cochabamba, In- 
cachaca, tropical cloud area, 2,100 m, 27 
Aug—5 Sep 1956, L. Pena (USNM). 
Paratype.—1l 2, same data as holotype. 


Oregocerata cladognathos Razowski 
(Fig. 2) 


Oregocerata cladognathos Razowski 1999: 
328 (description, illustration of male gen- 
italia). 


Diagnosis.—Oregocerata cladognathos 
is similar to its congeners in forewing size 
and pattern. The male genitalia (Fig. 2) can 
be distinguished by the slightly swollen 
base of the sacculus, unmodified anellus, 
moderately long distal processes of the gna- 
thos, simple uncus, and digitate soci. 

Holotype.—4 , Ecuador, Pichincha Prov- 
ince, km 40 via Quito-Pto. Quito, 25 Mar 
1986, S. McKamey (UCB). 


Oregocerata zonalis Razowski and Becker 
(Figs. 3, 10) 


Oregocerata zonalis Razowski and Becker 
2002: 316 (description, illustration of 
male and female genitalia). 


Diagnosis.—The male genitalia of O. 
zonalis (Fig. 3) are most similar to those of 
O. orcula and O. cladognathos, with the 
basal lobe of the sacculus poorly developed 
and the distal portion of the gnathos rela- 
tively long and slender. O. zonalis can be 
distinguished by the simple transtilla, which 
is bilobed in O. orcula and has a single me- 
dian lobe in O. cladognathos, and the api- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


cally reflexed distal portion of the uncus, 
which is unique within the genus. The socii 
of the three are also distinct: narrow and 
digitate in O. cladognathos, conspicuously 
broadened in the distal portion in O. orcula, 
and moderately broad throughout in O. zon- 
alis. The female genitalia (Fig. 10) have the 
sterigma tapering proximally, broader and 
less strongly sclerotized than in O. orcula. 

Holotype.—d, Ecuador, Loja Province, 
ojay27/ 50521 Dec 19927 Na Becker @is ©): 

Paratypes.—1l d, same data as holotype; 
2 36, Ecuador, Morona, Indanza, 2,800 m, 
24 Dec 1992, V. Becker (VBC); 1¢, Ec- 
uador, Tungurahua, Patata, 300 m, 7 Dec 
1992, V. Becker (VBC). 


Oregocerata triangulana Razowski and 
Brown, new species 
(Figs. 4, 11) 


Diagnosis.—The forewing of O. trian- 
gulana (Fig. 11) is conspicuously longer 
than that of its congeners, and has an ill- 
defined brown trapezoidal patch in the mid- 
dle. The male genitalia (Fig. 4) are similar 
to those of O. cladognathos and O. rhypar- 
ograpta, with elongate distal processes of 
the gnathos and a somewhat straight distal 
portion of the aedeagus. The genitalia can 
be distinguished from those of O. cladog- 
nathos by the longer distal processes of the 
gnathos, conspicuous basal lobe of the sac- 
culus, shorter aedeagus, and spiny lobes of 
the anellus. They differ from O. rhyparo- 
grapta by the longer distal processes of the 
gnathos, the slightly more slender aedeagus, 
and the spiny anellus, which is thorny in O. 
rhyparograpta (i.e., the projections are 
broader and/or shorter). 

Description.—Head: Vertex pale grayish 
brown, rust ventrolaterally; length of labial 
palpus ca. 3 times horizontal diameter of 
compound eye, pale grayish brown on inner 
surface, pale orange brown on outer sur- 
face; fan-shaped patch of flattened orange 
scales extending from base of proboscis to 
between foreleg coxae, with a few scales at 
middle of patch extending ca. 2 times 
length of other scales. Thorax: Dorsum pale 


VOLUME 107, NUMBER 4 907 


EN .. NW 
: cull, Hee 
INN 


aa 


Figs. 4-6. Male genitalia of Oregocerata;, valvae spread, aedeagus removed. 4, O. triangulana, 5, O. rhy- 
parograpta, 6. O. submontana. 


908 


grayish brown. Prothoracic leg with hair- 
pencil. Forewing length 12.0 mm (n = 3), 
broad, expanding terminally, with costa 
uniformly convex, apex short, termen rather 
straight; ground color whitish, weakly suf- 
fused with scattered tiny brown specks and 
faint brown striae, especially in distal half; 
costa with irregular band of brown scales; 
a pale trapezoidal brown blotch near middle 
of wing. Fringe cream, whiter towards tor- 
nus. Hindwing white, tinged with cream on 
periphery, weakly strigulated with pale gray 
brown in anteroterminal portion. Abdomen: 
Male genitalia (Fig. 4; drawn from GS 
USNM 68602) with tegumen rather narrow; 
uncus comparatively short, curved ventrally 
from near base, rounded apically; socii 
moderately large, densely setose, ca. 0.8 
length of basal portion of gnathos; gnathos 
arms relatively slender, distal processes lon- 
ger than basal portion; transtilla broad, shal- 
low W-shaped, weakly sclerotized at mid- 
dle; valva rounded distally, with costa and 
ventral margin weakly curved; a conspicu- 
ous, dorsally projecting, digitate process at 
base of sacculus; phallus short, phallobase 
slightly longer than aedeagus; a short, spiny 
skirt at dorsal junction of phallobase and 
aedeagus (= lobes of anellus); aedeagus 
slender, with a sclerotized distal process; 
vesica with one large and two small, fine 
cornuti. Female: Unknown. 

Holotype.—d, Colombia, Cauca, Para- 
mo de Parace, Lake San Rafael, 29 Jan 
1959, J. E G. Clarke (USNM). 

Paratypes.—2 6d, same data as holotype 
(USNM). 

Etymology.—The specific epithet refers 
to the subtriangular patch of the forewing. 


Oregocerata rhyparograpta 
Razowski and Becker 
(Fig. 5) 


Oregocerata rhyparograpta Razowski and 
Becker 2002: 316 (description, illustra- 
tion of male genitalia). 


Diagnosis.—The male genitalia of O. 
rhyparograpta (Fig. 5) are most similar to 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


those of O. triangulana, with a well-devel- 
oped basal lobe of the sacculus, digitate so- 
cii, and long, slender distal processes of the 
gnathos. O. rhyparograpta can be distin- 
guished by the slightly longer phallus with 
a thorny anellus and the conspicuously 
shorter distal processes of the gnathos. 

Holotype.—d, Ecuador, Pastasa Prov- 
ince, Mera, 1,300 m, Dec 1992, V. Becker 
(WEO@) 


Oregocerata submontana Razowski and 
Brown, new species 
(Figs. 6, 12) 


Diagnosis.—The poorly developed fore- 
wing pattern of O. submontana (Fig. 12) is 
similar to most other congeners. The male 
genitalia (Fig. 6) of O. submontana can be 
distinguished by the shorter uncus, shorter 
socii, shape of the distal portion of the ae- 
deagus, and the absence of cornuti. 

Description.—Head: Vertex pale grayish 
brown, rust ventrolaterally; length of labial 
palpus ca. 3 times horizontal diameter of 
compound eye, pale grayish brown on inner 
surface, pale orange brown on outer sur- 
face; fan-shaped patch of flattened orange 
scales between foreleg coxae and base of 
proboscis, with a few scales at middle of 
patch extending posterad ca. 2 times length 
of other scales. Thorax: Dorsum pale gray- 
ish brown. Prothoracic leg with hairpencil. 
Forewing length 10.5 mm (n = 1), broad, 
expanding terminally, with costa uniformly 
convex, termen rather straight; [specimen 
worn] ground color whitish, weakly suf- 
fused with scattered tiny brown specks; a 
pale brown median fascia from costa ca. 0.6 
distance from base; basal 0.3 with darker 
scaling. [Fringe lacking]. Hindwing white, 
weakly strigulated with pale gray-brown in 
anteroterminal portion. Abdomen: Male 
genitalia (Fig. 6; drawn from GS USNM 
95215) with tegumen rather narrow; uncus 
shorter than in other species, curved ven- 
trally, pointed apically; socii broad, densely 
setose, ca. 0.6X length of basal portion of 
gnathos; gnathos arms relatively slender, 
distal processes less than twice as long as 


VOLUME 107, NUMBER 4 


909 


Figs. 7-8. 
furcata. 


basal portion; transtilla broad, shallow W- 
shaped, weakly sclerotized at middle; valva 
somewhat parallel-sided, with costa nearly 
straight and ventral margin weakly upcur- 
ved in distal 0.3; a conspicuous, dorsally 
projecting, digitate process at base of sac- 
culus; phallus short, phallobase about the 
same length as aedeagus; a short, thorny 
skirt at dorsal junction of phallobase and 
aedeagus; aedeagus slender, with a sclero- 
tized distal process; vesica without cornuti. 
Female: Unknown. 

Holotype.—<d, Venezuela, Lara, Yacum- 


Male genitalia of Oregocerata; valvae spread, aedeagus removed. 7, O. caucana 8, O. quadri- 


bu National Park, 13 km SE Sanare, 4,800’, 
cloud forest, 4—7 Mar 1978, J. B. Heppner 
(USNM). 

Etymology.—tThe specific epithet refers 
to the submontane distribution of this spe- 
cies. 


Oregocerata caucana Razowski and 
Brown, new species 
(Figs. 7, 13) 
Diagnosis.—Oregocerata caucana (Fig. 
13) is most similar to O. chrysodetis (see 
Clarke 1958: 235) in forewing length and 


910 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 9-10. 


maculation: both are pale orange with an 
ill-defined fascia across the middle of the 
wing, angled outward near the lower edge 
of the discal cell. The forewing pattern is 
slightly darker orange in O. caucana, with 
the apical region scaled with orange-brown. 
Because the holotype of O. chrysodetis 
lacks the abdomen, the genitalia cannot be 
compared. On the basis of the aedeagus, O. 
caucana appears to be most closely related 
to O. rhyparograpta and O. triangulana; 
the spiny lobes of the anellus are most sim- 
ilar to those of the latter. Oregocerata cau- 
cana can be distinguished from its conge- 
ners by the short, curved distal processes of 
the gnathos, which are considerably shorter 
than the basal portion of the gnathos (Fig. 
7). The latter is the most conspicuous au- 
tapomorphy for this species. 
Description.—Head: Vertex whitish, 
tinged with pale brownish gray; labial pal- 


Female genitalia of Oregocerata. 9, O. orcula 10. O. zonalis. 


pus pale grayish brown on inner surface, 
pale orange brown on outer surface; fan- 
shaped patch of flattened orange scales be- 
tween foreleg coxae and proboscis, with a 
few longer scales at middle of patch. Tho- 
rax: Dorsum concolorous with head, slight- 
ly browner anteriorly. Prothoracic leg with 
hairpencil. Forewing length 8.7 mm (n = 
1); ground color whitish, densely strigulat- 
ed and suffused with pale orange with 
sparse, small, inconspicuous dark specks; 
an ill-defined, pale fascia across middle of 
wing, angled outward at lower edge of dis- 
cal cell; a row of 5—6 small brown dots 
along termen. Fringe concolorous with 
ground color, with dark brown median line. 
Hindwing whitish, tinged with cream in 
apical third. Abdomen: Male genitalia (Fig. 
7; drawn from GS USNM 68623) with teg- 
umen rather narrow; uncus simple, slender, 
rather short, curved, pointed apically; socii 


VOLUME 107, NUMBER 4 


comparatively slender, densely setose, ca. 
0.8 length of basal portion of the gnathos; 
gnathos arms relatively slender, with distal 
portion extremely short, weakly curved; 
transtilla narrow, inverted U-shaped, weak- 
ly sclerotized at middle; valva broadest at 
base, gradually attenuate distally, with costa 
nearly straight and ventral margin weakly 
curved throughout; sacculus with a com- 
paratively short, dorsally projecting, digi- 
tate process at base; phallus short, fairly 
broad, curved dorsad terminally; phallobase 
about as long as aedeagus; a short, spiny 
skirt at junction of phallobase and aedea- 
gus; vesica with one large and two small 
cornuti, and a sclerotized distal plate. Fe- 
male: Unknown. 

Holotype.—¢d, Colombia, Cauca, Para- 
mo de Parace, Lake San Rafael, 27 Jan 
1959, J. E G. Clarke (USNM). 

Etymology.—The specific epithet refers 
to the Cauca region, Colombia. 


Oregocerata chrysodetis (Meyrick, 1926), 
new combination 


Tortrix chysodetis Meyrick 1926: 248 (de- 
scription); Clarke 1958: 235 (illustration 
of adult). 

‘“Eulia’ chrysodetis: Powell et al. 1995: 
146 (checklist). 


Discussion.—Clarke (1958: 235) illus- 
trated the holotype of Tortrix chrysodetis, 
which lacks the abdomen. Because the gen- 
italia are lost, this species has defied con- 
temporary generic assignment; it was treat- 
ed as “‘Eulia” chrysodetis by Powell et al. 
(1995), indicating uncertain placement. The 
discovery of Oregocerata caucana (de- 
scribed above), with a similar forewing 
shape, size, and pattern, suggests that Eulia 
chrysodetis likely belongs in Oregocerata. 
Although it is possible that O. chrysodetis 
and O. caucana are conspecific, subtle dif- 
ferences in forewing markings suggest oth- 
erwise. 

Holotype.—¢, Colombia, Central Cor- 
dilleras (BMNH). 


911 


Oregocerata quadrifurcata Razowski and 
Brown, new species 
(Figs. 8, 14) 


Diagnosis.—Oregocerata quadrifurcata 
is externally similar to O. caucana; it can 
be distinguished from its congeners by its 
nearly uniform pale brown forewing (Fig. 
14). The male genitalia (Fig. 8) are highly 
divergent from other Oregocerata, with a 
greatly expanded, quadrifurcate distal por- 
tion of the uncus, a pair of short, blunt pro- 
cesses distally from the gnathos, and a 
highly modified base of the sacculus, all of 
which are interpreted as autapomorphies. 
The dorsum of the aedeagus lacks the spiny 
lobes of the anellus present in many spe- 
cies. The species is provisionally assigned 
to the genus on the basis of the digitate pro- 
cess at the base of the sacculus and the 
patch of long, flattened orange scales ex- 
tending from the base of the proboscis to 
between the foreleg coxae, both characters 
of which are characteristic of Oregocerata. 

Description.—Head: Vertex cream, 
weakly tinged with brownish; labial palpus 
elongate, pale grayish brown on inner sur- 
face, pale orange brown on outer surface, 
rust colored near junction of segments II 
and III; fan-shaped patch of flattened or- 
ange scales extending from base of probos- 
cis to between prothoracic coxae. Thorax: 
Dorsum concolorous with head. Prothoracic 
leg with hairpencil. Forewing length 9.0 
mm (n = 1), apex short, sharp; termen 
somewhat oblique; ground color pale gray- 
ish brown in basal area, tinged pale yellow- 
ish brown in remaining portion of wing: 
dorsum and costa with pale orange-brown 
to rust-colored scaling; tiny black dots in 
median and subcostal parts of postbasal 
field, along termen, and scattered in poste- 
rior half of wing; dorsum dotted with white; 
markings darker than suffusion, diffuse, 
represented by medial fascia and subapical 
spots. Hindwing cream colored, lighter to- 
wards base, with weak strigulation in ter- 
minal area. Abdomen: Male genitalia (Fig. 
8: drawn from GS USNM 68623) with teg- 


912 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Figs. 11-14. Adults of Oregocerata. 11, O. triangulana, 12, O. submontana, 13, O. caucana, 14, O. quad- 
rifurcata. 


umen broad; uncus large, curved, greatly 
swollen distally, with large, flattened, 
crown-shaped apex; socii large, compara- 
tively broad, densely setose, extending 
nearly to distal end of gnathos arms; gna- 
thos arms relatively broad, with a pair of 
short, blunt subterminal processes; transtilla 
a slender bridge; valva nearly parallel-sided 
in basal half, weakly attenuate in distal half, 
truncate distally; dorsally projecting pro- 
cess at base of sacculus large, thorny, with 
a smaller subtriangular prominence imme- 
diately distad becoming contiguous with 
sacculus; phallus short, aedeagus slightly 
longer than phallobase, with slender termi- 
nation, curved in basal 0.4; vesica with one 
large curved and one smaller, straight cor- 
nutus. Female: Unknown. 

Holotype-—d, Colombia, Cauca, Para- 


mo de Parace, Lake San Rafael, 27 Jan 

1959, J. E G. Clarke (USNM). 
Etymology.—tThe specific epithet refers 

to the somewhat quadrifurcate tip of the un- 


Cus. 


ACKNOWLEDGMENTS 


We thank the following for allowing us 
to examine specimens in their care: Jerry 
Powell (UCB), Kevin Tuck (BMNH), and 
Vitor Becker (Serra Bonita, Brazil). We 
thank the following for helpful reviews of 
the manuscript: Stuart McKamey, USDA 
Systematic Entomology Laboratory, Na- 
tional Museum of Natural History, Wash- 
ington, D.C. and Daniel Rubinoff, Univer- 
sity of Hawaii, Honolulu. The figures were 
drawn by Jé6zef Razowski and David 
Adamski; the latter prepared the plates. 


VOLUME 107, NUMBER 4 


LITERATURE CITED 


Brown, J. W. 1990. Taxonomic distribution and phy- 
logenetic significance of the male foreleg hairpen- 
cil in the Tortricinae (Lepidoptera: Tortricidae). 
Entomological News 101: 109-116. 

Clarke, J. E G. 1958. Catalogue of the type specimens 
of microlepidoptera in the British Museum (Nat- 
ural History) described by Edward Meyrick, Vol- 
ume 3. Trustees of the British Museum, London. 
600 pp. 

Meyrick, E. 1926. Exotic Microlepidoptera 3(8): 225— 
256. 

Powell, J. A., J. Razowski, and J. W. Brown. 1995. 
Tortricidae: Tortricinae, Chlidanotinae, pp. 138— 
151. In Heppner, J. B., ed. Atlas of Neotropical 
Lepidoptera, Checklist Part Il: Hyblaeoidea—Pyr- 


913 


aloidea—Tortricoidea. Association for Tropical 
Lepidoptera, Scientific Publishers, 
Florida. 
Razowski, J. 


Gainesville, 


1988. New genera and species of the 

Neotropical Archipini (Lepidoptera: Tortricidae). 

Acta Zoologica Cracoviensia 31: 387—422. 

1997. Euliini (Lepidoptera: Tortricidae) of 

Peru with descriptions of new taxa and list of the 

New World genera. Acta Zoologica Cracoviensia 

40: 79-105. 

. 1999. Tortricidae (Lepidoptera) from Ecuador. 
Acta Zoologica Cracoviensia 42: 321—342. 

Razowski, J. and V. O. Becker. 2002. Description of 
new species of some known or new Neotropical 
Euliini genera (Lepidoptera: Tortricidae). SHILAP 
Revista de Lepidopterologia 30: 315-323. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 914-916 


AN ADDITIONAL SPECIES OF THE GENUS MASAAKIA TAKEUCHI 
(HYMENOPTERA: TENTHREDINIDAE) FROM JAPAN 


IcHWI TOGASHI 


1-chome, Tsurugihonmachi, Hakusan-shi, Ishikawa Prefecture 920-2121, Japan 


Abstract.—Masaakia hakusana, n. sp., from Mt. Hakusan, Honshu, Japan, is described 
and illustrated. A key is provided for the four Japanese species of Masaakia. 


Key Words: 
Japan 


Masaakia Takeuchi is a small genus of 
the subfamily Blennocampinae and is en- 
demic to Japan. It contains three described 
species. In 2004, I captured one female of 
this genus from Mt. Hakusan, Honshu, Ja- 
pan. This specimen is similar to M. longi- 
vaginata Takeuchi, but it is easily distin- 
guished from the latter by the shapes of the 
sawsheath in lateral view, the serrulae of 
the lancet, the claw, and the inner foretibial 
spur. I concluded that this specimen repre- 
sents a new species, and I describe and il- 
lustrate this species and give a revised key 
to the species of Masaakia. 


KEY TO THE JAPANESE SPECIES OF MASAAKIA 
(FEMALES) 


1. Tegula black; pedicel longer than wide; apex 
of sawsheath narrowly rounded in lateral view 
(EE Sy ED) ee eacaps pred enc Peni nucaee. Pee eres 2 
— Tegula yellow; pedicel shorter than wide; apex 
of sawsheath truncate in lateral view (Togashi 
ANOS {ti 3) soc csc 06 katayamai Togashi, 2002 
. Antenna entirely black 
Antenna black with undersides of 4th to 9th 
segments and apical half of underside of 3rd 
antennal segment reddish brown ....... 
By shot Met Aca acne aG OES shinoharai Togashi, 1998 
3. Apex of sawsheath pointed in lateral view (Fig. 
12); apex of inner foretibial spur slightly emar- 
ginated (Fig. 7); preapical tooth of tarsal claws 
broad (Fig. 8) longivaginata Takeuchi, 1950 
— Apex of sawsheath narrowly rounded in lateral 
view (Fig. 9); apex of inner foretibial spur 


i) 


Hymenoptera, Tenthredinidae, Blennocampinae, Masaakia, new species, 


deeply emarginated (Fig. 5); preapical tooth of 
tarsal claws slender (Fig. 6) . . . . hakusana, n. sp. 


Masaakia hakusana Togashi, 
new species 
(Figs. 1-6, 9-11) 


Female.—Length, 7 mm. Body black 
with cenchrus milky white and cercus 
brownish black. Antenna entirely black. 
Wings hyaline, stigma and veins dark 
brown to black. Legs black with knees, 
femora, except for black apical halves, and 
tarsi, except for milky white basal half of 
basitarsi, dark brown. 

Head: Transverse, postocellar area trans- 
verse, convex; circumocellar furrow distinct 
but anterior half absent; interocellar furrow 
distinct and deep; postocellar furrow dis- 
tinct; lateral furrows distinct and deep; 
OOL:POL:OCL = 1.4:1.0:1.6; frontal area 
nearly flattened, anterior margin with a 
transverse wall; median fovea deep, nearly 
quadrate in outline; lateral fovea deep, cir- 
cular in outline; supra-antennal tubercles 
distinct; antenno-ocular distance longer 
than distance between antennal sockets (ra- 
tio 2.5:1.0); front margin of clypeus emar- 
ginated; malar space very narrow (Fig. 2); 
postorbital groove distinct (Fig. 2); post- 
genal carina distinct near mandible only 
(Fig. 2). Antenna longer than costa of fore- 
wing (ratio 1.0:0.8), relative lengths of seg- 


VOLUME 107, NUMBER 4 


Figs. 1-8. 


5 


1—6, Masaakia hakusana. 1, Head, dorsal view. 2, Head, lateral view. 3, Antenna, lateral view. 


4, Forewing. 5, Inner foretibial spur, lateral view. 6, Tarsal claw, lateral view. 7-8, M. longivaginata. 7, Inner 


foretibial spur, lateral view. 8, Tarsal claw, lateral view. 


Mmentseabout Wo s1eO: Selle Sal lees eO: 
1.3; pedicel longer than wide (ratio 1.0:0.6) 
(Fig. 3). 

Thorax: Mesoscutellum slightly convex; 
cenchrus large, distance between cenchri 
slightly longer than breadth of one (ratio 
1.2:1.0). Forewing venation as in Fig. 4; ra- 
dial crossvein (2r) slightly curved; hind- 
wing with petiole of anal cell nearly as long 
as nervulus (cu-a). Legs: Apex of inner 
foretibial spur deeply emarginated (Fig. 5); 
hind basitarsus slightly shorter than follow- 
ing 4 segments combined; preapical tooth 
of tarsal claws slender (Fig. 6). 

Abdomen: Sawsheath rather long, apex 
of sawsheath narrowly rounded (Fig. 9); 
lancet with 19 serrulae; apical portion of 


lancet as in Fig. 10; 11th to 13th serrulae 
as in Fig. 11. 

Punctation: Vertex, inner orbits, frontal 
area, clypeus, and labrum with distinct but 
sparse punctures; hind orbits covered with 
rather small, distinct, and dense punctures; 
thorax covered with fine setigerous punc- 
tures; abdominal tergites nearly impunctate, 
shining. 

Male.—Unknown. 

Food plant.—Unknown. 

Distribution.—Japan (Honshu). 

Holotype.—Female, 15.VI.2004, Mt. 
Hakusan (1,300—1,500 m), Ishikawa Pre- 
fecture, Honshu, Japan, I. Togashi leg. De- 
posited in the National Science Museum 
(Nat. Hist.), Tokyo. 


916 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


“Cowal 
2 leth llth 10th 


Figs. 9-13. 9-11, Masaakia hakusana. 9, Sawsheath, lateral view. 10, Apical portion of lancet. 11, 11th to 
13th serrulae of lancet. 12-13, M. longivaginata. 12, Sawsheath, lateral view. 13, 10th to 12th serrulae of lancet. 


Remarks.—This new species is very ACKNOWLEDGMENT 
closely allied to M. longivaginata, but it is 
easily distinguished from the latter by the 
more rounded sawsheath (apex pointed in 
M. longivaginata, see Figs. 9, 12), by the 
deeply emarginated apex of the inner fore- 
tibial spur (apex shallowly emarginated in 
M. longivaginata, see Figs. 5, 7), by the LITERATURE CITED 


slender subapical tooth of the tarsal claws 
P Togashi, I. 2002. Description of a new species of the 


(broad in M. longivaginata, see Figs. 6, 8), genus Masaakia Takeuchi (Hymenoptera: Ten- 


and by the shape of the serrulae (see Figs. thredinidae) from Japan. Proceedings of the En- 
ils i113). tomological Society of Washington 104: 373-375. 


I thank David R. Smith, Systematic En- 
tomology Laboratory, U.S. Department of 
Agriculture, Washington, D.C., for review- 
ing the manuscript. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 917-940 


A REVIEW OF GROUND BEETLE SPECIES (COLEOPTERA: CARABIDAE) 
OF MINNESOTA, UNITED STATES: NEW RECORDS AND 
RANGE EXTENSIONS 


KAMAL J. K. GANDHI, DANIEL W. GILMORE, GEORGE E. BALL, RALPH W. HOLZENTHAL, 
STEVEN A. KATOVICH, JESSICA J. KOEHLE, KIRK J. LARSEN, WILLIAM J. MATTSON, AND 
STEVEN J. SEYBOLD 


(KJKG, RWH, SJS) Department of Entomology, 219 Hodson Hall, 1980 Folwell Av- 
enue, University of Minnesota, St. Paul, MN 55108, U.S.A.; (KJKG, DWG, SJS) De- 
partment of Forest Resources, 115 Green Hall, 1530 Cleveland Avenue, University of 
Minnesota, St. Paul, MN 55108, U.S.A.; (GEB) E. H. Strickland Museum, Department 
of Biological Sciences, CW 405 Biological Sciences Center, University of Alberta, Ed- 
monton, AB T6G 2E9, Canada; (SAK) USDA—Forest Service, Forest Health Protection, 
1992 Folwell Avenue, St. Paul, MN 55108, U.S.A.; (JJK) Department of Fisheries and 
Wildlife, 200 Hodson Hall, 1980 Folwell Avenue, University of Minnesota, St. Paul, MN 
55108, U.S.A.; (KJL) Department of Biology, Luther College, 700 College Drive, De- 
corah, IA 52101, U.S.A.; (WJM) USDA—Forest Service, North Central Research Station, 
Forestry Sciences Laboratory, 5985 Highway K, Rhinelander, WI 54501, U.S.A.; (KJKG, 
SJS) Current mailing address: USDA—Forest Service, Pacific Southwest Research Station, 
Chemical Ecology of Forest Insects, 720 Olive Drive, Suite D, Davis, CA 95616, U.S.A. 
(e-mail: SJS: sseybold @fs.fed.us) 


Abstract.—We report new Minnesota records for 13 genera and 100 species of ground 
beetles (Coleoptera: Carabidae), reflecting 21% and 31% increases, respectively, over 
records in the literature. There are now 76 genera and 433 carabid species recorded from 
Minnesota. New material was collected from sub-boreal forest study sites in northeastern 
and central Minnesota. We also surveyed 16 museum collections and private collections 
in the United States with an emphasis on collections in the north central region and those 
likely to contain Minnesota records. New generic records for Minnesota from the museum 
collections include Paratachys Casey, Nomius Laporte, Platypatrobus Darlington, Gas- 
trellarius Casey, Lophoglossus LeConte, Pseudamara Lindroth, Panagaeus Latreille, 
Dicheirotrichus Jacquelin du Val, Discoderus LeConte, Leptotrachelus Latreille, Tetra- 
gonoderus Dejean, Apenes LeConte, and Axinopalpus LeConte. New species records from 
the field surveys in northeastern and central Minnesota include Notiophilus aquaticus 
(Linnaeus), Sphaeroderus nitidicollis brevoorti LeConte, Trechus crassiscapus Lindroth, 
Bembidion mutatum Gemminger & Harold, Bembidion wingatei Bland, Patrobus foveo- 
collis (Eschscholtz), Patrobus septentrionis Dejean, Pterostichus melanarius (llliger), 
Amara coelebs Hayward, Pseudamara arenaria (LeConte), Bradycellus semipubescens 
Lindroth, Harpalus ventralis LeConte, Agonum affine Kirby, and Agonum trigeminum 
Lindroth. In addition, we highlight the special role of P. melanarius as an invasive beetle. 
Since many of the new records were based on specimens stored in research entomological 
collections, we recognize the significance of institutional holdings as a source of infor- 
mation for studies of native biodiversity. 


Key Words: biodiversity, Carabidae, ground beetles, Minnesota, museum collections, 


918 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


species distributions, sub-boreal forest, invasive species, Pterostichus me- 
lanarius, threatened species, local extinctions, Calosoma scrutator 


Ground beetles (Coleoptera: Carabidae) 
are relatively abundant and diverse in the 
forests and grasslands of Minnesota. Min- 
nesota is unique among neighboring US 
states and Canadian provinces because it 
occurs at a transition point where three bi- 
omes meet: sub-boreal coniferous/decidu- 
ous forests, central deciduous forests, and 
tall grass prairies (Tester 1995, Minnesota 
Department of Natural Resources 2004) 
(Fig. 1). This habitat diversity leads to cor- 
responding biodiversity of the Carabidae in 
the state. In a literature and museum (ex- 
cluding the University of Minnesota Insect 
Collection) survey of America north of 
Mexico, Bousquet and Larochelle (1993) 
reported 63 genera and 323 species of ca- 
rabids inhabiting Minnesota. In a survey of 
northeastern North America, Downie and 
Arnett (1996) documented 28 genera and 
94 species of ground beetles in Minnesota. 
Three species records in the latter work 
were unique, bringing the total in the state 
to 63 genera and 326 species. 

Since the late 1800s, ground beetles have 
been collected by numerous professional 
and amateur entomologists in Minnesota, 
with the earliest collections dating from 
1888 when the Division of Entomology and 
Botany was established at the University of 
Minnesota (Department of Entomology, 
University of Minnesota 2004). Otto Lug- 
ger, a faculty entomologist, developed the 
first insect collection, which is housed at 
the University of Minnesota Insect Collec- 
tion in St. Paul (UMIC). O.W. Oestlund, an 
entomologist with the Minnesota Geologi- 
cal and Natural History Survey and a pro- 
fessor in the Department of Animal Biology 
at the University of Minnesota, also col- 
lected a large number of carabids in the 
state. The carabid collection was further en- 
hanced in the late 1800s and early 1900s by 
C.N. Ainslie, a professor in the Department 


of Zoology, who collected almost entirely 
in Olmsted County. In the early to mid- 
1900’s, C.E. Mickel, a professor in the De- 
partment of Entomology, D.G. Denning, a 
student at the UMIC, W.E. Stehr, a curator 
at the UMIC, M.H. Hatch, an instructor at 
the University of Minnesota (1926-1927), 
and A.A. Granovsky, a professor in the De- 
partment of Entomology (1930-1956), all 
were avid Minnesota insect collectors who 
contributed carabid specimens to the 
UMIC. Between the mid-1960’s and the 
early 1990’s, J.R. Powers, a biology profes- 
sor at Concordia College in Moorhead, 
Minnesota, and his students, made exten- 
sive collections of carabids from the west- 
ern counties. Many of these specimens were 
deposited in the Essig Museum, University 
of California, Berkeley. 

During the last twenty years, carabid sur- 
veys have focused on specific geographic 
areas and habitat types in the state (Fig. 2). 


Coniferous/Deciduous 
Forests 


oT 


oe ee 


< Central Deciduous 
Forests 


mB yes 


Tall Grass 


| Prairies 
| Re 
| ; 


Fig. 1. Map of Minnesota showing the three major 
ecological biomes present in the state. 


919 


VOLUME 107, NUMBER 4 


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For example, Epstein (1982) and Epstein 
and Kulman (1984, 1990) studied carabids 
in oak and cottonwood forests, and grassy 
fields in east-central Minnesota. These sites 
are being re-sampled in 2005 by the first 
author and M.E. Epstein to document long- 
term changes in ground beetle assemblages 
in Minnesota. More recently, Tinerella 
(2000) and Tinerella and Rider (2001) stud- 
ied carabids in the tallgrass prairie region 
of western Minnesota, and reported six new 
state records. MacLean (2002) studied sub- 
boreal forest stands and wetlands in the 
Grand Portage National Monument in ex- 
treme northeastern Minnesota, and reported 
five new state records. Gilmore et al. (2002) 
outlined a research project in the sub-boreal 
forests of northeastern Minnesota and first 
reported the presence of Prerostichus me- 
lanarius (Mlliger), a European invasive spe- 
cies, in Minnesota. Subsequently, Petrice et 
al. (2002) confirmed the presence of P. me- 
lanarius in Minnesota and established eight 
additional records of carabid species in 
hardwood vernal pond habitats in north- 
central Minnesota. Hladilek (2003) con- 
ducted an ecological study of the interac- 
tion of ground beetles with detrital food 
webs in a wheat field on the University of 
Minnesota campus in east-central Minne- 
sota (Ramsey County), but reported no new 
state species records. 

During the summer of 2000, we initiated 
a study of ground beetles in the sub-boreal 
forested region of northeastern Minnesota 
including the Boundary Waters Canoe Area 
Wilderness. A catastrophic windstorm event 
occurred on the 4" of July, 1999, and re- 
sulted in windthrow of trees on more than 
193,035 hectares of forestland in the Su- 
perior National Forest (USDA 2000). This 
study focused on the influences of this 
large-scale wind-disturbance event and 
post-wind-disturbance silvicultural treat- 
ments (e.g., salvage-logging and prescribe- 
burning) on the abundance and diversity of 
forest Coleoptera (Gilmore et al. 2002, 
2005). During the summer of 2002, we ini- 
tiated a separate study to assess the imme- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


diate responses of ground beetles to a se- 
vere wildfire in a Pinus banksiana Lamb. 
forest near Brainerd in central Minnesota. 
Results from a disturbance ecology per- 
spective from both of these field studies 
will be reported elsewhere. 

Our field studies on carabids in Minne- 
sota have revealed a number of species oc- 
currences that were previously unrecorded 
in the state. In addition, we surveyed the 
UMIC and 15 other insect collections, 
housed in academic and research institu- 
tions or in private hands, for new state re- 
cords. In this paper, we list and discuss 
these new records and include information 
on sample locations, collection dates, name 
of the collector, and number of adults col- 
lected in each series. We also provide the 
current distributions and known habitat as- 
sociations of the species that represent new 
state records from our studies in northeast- 
ern and central Minnesota. 


MATERIALS AND METHODS 


Study sites.—In northeastern Minnesota, 
ground beetles were sampled from research 
sites along the Gunflint Trail and in the 
Boundary Waters Canoe Area Wilderness 
in the Superior National Forest in Cook 
County (Fig. 2). The forest is composed of 
paper birch, Betula papyrifera Marsh., 
trembling aspen, Populus tremuloides 
Michx., northern white cedar, Thuja occi- 
dentalis L., jack pine, Pinus banksiana, 
eastern white pine, P. strobus L., red pine, 
P. resinosa Ait., black spruce, Picea mari- 
ana (Mill.) B.S.P., white spruce, P. glauca 
(Moench) Voss, balsam fir, Abies balsamea 
(L.) Mill., and tamarack, Larix laricina (Du 
Roi) K. Koch. We sampled carabids from 
P. tremuloides/B. papyrifera/conifer-, and 
P. banksiana-dominated forests in undis- 
turbed, severely wind-disturbed (67—100% 
tree mortality), wind-disturbed-salvage- 
logged, and wind-disturbed-prescribe- 
burned stands (Gilmore et al. 2002, 2005). 
The burned stands were moderately sal- 
vage-logged prior to burning, and thus had 
been subjected to three disturbance factors. 


VOLUME 107, NUMBER 4 


During 2000, we sampled two sites (repli- 
cates) for each of three treatment types (un- 
disturbed, severely wind-disturbed, and 
wind-disturbed-salvage-logged) in each of 
the two (P. tremuloides/B. papyrifera/co- 
nifer and P. banksiana) forest cover types 
for a total of 12 sites. During 2001—2003, 
we sampled four sites (replicates) for each 
of the four treatment types (undisturbed, se- 
verely wind-disturbed, wind-disturbed-sal- 
vage-logged, and wind-disturbed-prescribe- 
burned) in each of the two (P. tremuloides/ 
B. papyrifera/conifer and P. banksiana) for- 
est cover types for a total of 32 sites 

In central Minnesota, ground beetles 
were sampled near the town of Barrows, 
located ten km southwest of Brainerd in 
Crow Wing County (Fig. 2). The forest is 
composed primarily of oak, Quercus spp., 
and jack pine, P. banksiana, growing on 
sandy soils. On 31 May 2002, a lightning 
strike started a wildfire between Brainerd 
and Barrows, and it quickly consumed over 
285 hectares of this forest type. We sampled 
carabids from unburned and severely- 
burned (100% tree mortality) P. banksiana- 
dominated forest stands. During 2002 and 
2003, we sampled four sites in each of the 
two treatment types (unburned and burned) 
for a total of eight sites. 

Field surveys.—Adult ground beetles 
were collected with standard pitfall traps 
consisting of an outer | L and an inner 500 
mL plastic cup filled with 2—3 cm of pro- 
pylene glycol as a preserving agent 
(Spence and Niemelad 1994). Each trap was 
covered with a 100 cm? plywood roof to 
protect the collecting cup from small mam- 
mal disturbances and flooding during rain. 
During 2000 in northeastern Minnesota, 
six unbaited pitfall traps were placed in 
each of the 12 sites for a total of 72 traps. 
During 2001—2003, six unbaited pitfall 
traps were placed in each of the 32 sites 
for a total of 192 traps. In addition, we 
collected carabid beetles from pitfall traps 
that were baited with various combinations 
of a- or B-pinene that were used to attract 
rhizophagous beetles. During 2000, three 


921 


replicates of each of four bait-types were 
placed in each of the 12 sites for a total of 
144 traps. During 2001—2003, only P. 
banksiana sites were sampled with baited 
pitfall traps, and five of these traps (in- 
cluding an unbaited control trap) were 
placed in 16 sites for a total of 80 traps. 
In northeastern Minnesota, all traps were 
spaced by 20 m, operated from late-May 
to early October, and emptied every 15—20 
days. In the central Minnesota study area, 
four unbaited pitfall traps spaced by 50 m 
were placed in each of the eight sites for 
a total of 32 traps. These traps were op- 
erated from mid-June to mid-September 
and emptied every 15—20 days. 

Adult beetles were identified by the au- 
thors K. J. K. Gandhi and G. E. Ball using 
taxonomic keys provided by Lindroth 
(1961—69), Downie and Arnett (1996). 
Liebherr and Will (1996), and Ball and 
Bousquet (2001). Distribution records of 
the species were cross-checked with Bous- 
quet and Larochelle (1993), Downie and 
Arnett (1996), and other specific references 
noted above. Nomenclature and phyloge- 
netic concepts in this paper generally fol- 
low that of Noonan (1991) and Bousquet 
and Larochelle (1993). Voucher specimens 
from our sub-boreal forest studies in north- 
eastern and central Minnesota have been 
deposited at the UMIC. 

Museum survey.—We searched system- 
atically for new records of 
amongst the specimens in major collec- 
tions. At the UMIC, we surveyed the iden- 
tified and unidentified material in the family 
Carabidae. For the identified beetles, we fo- 
cused on adult carabids collected from all 
counties in Minnesota. Because of the large 
number of unidentified material at UMIC, 
we focused primarily on unidentified cara- 
bids collected from Aitkin, Beltrami, Carl- 
ton, Cook, Itasca, Kittson, Koochiching, 
Lake, Lake of the Woods, Marshall, Pen- 
nington, Polk, Red Lake, Roseau, and St. 
Louis counties of Minnesota. We chose 
specimens from these counties reasoning 


carabids 


that they would include all boreal species 


922 


that might overlap with our own biodiver- 
sity surveys. We searched the database of 
the Insect Collection at the California 
Academy of Sciences (San Francisco, CAS) 
for new Minnesota records amongst the 
identified carabid beetles, and searched the 
complete holdings of the California Collec- 
tion of Arthropods at the Plant Pest Diag- 
nostic Center (Sacramento, CDFA), the En- 
vironmental Division of the Grand Portage 
Band (Grand Portage, Minnesota, GPB), 
Iowa State University (Ames, ISU), the 
Milwaukee Public Museum (MPM), North 
Dakota State University (Fargo, NDSIRO©), 
the Smithsonian Institution (Washington, 
D.C., NMNH), South Dakota State Univer- 
sity (Brookings, SDSU), University of Cal- 
ifornia, Berkeley (UCB), University of Cal- 
ifornia, Davis (UCD), Illinois Natural His- 
tory Survey (Champaign, INHSIC), Uni- 
versity of Wisconsin (Madison, UWM), and 
University of Wyoming (Laramie, UW). 
Furthermore, we searched the private 
ground beetle collections of K.J. Larsen, 
Luther College (Decorah, Iowa, KJL) and 
K.W. Will, University of California (Berke- 
ley, KWW) for new records. 

Collection data for each specimen includ- 
ed county and specific locality, any habitat 
information, collector, collection date, and 
number of specimens in the series (indicat- 
ed parenthetically at the end of the record). 
Some information about the museum spec- 
imens may be incomplete because of the 
limited locality and habitat data recorded on 
specimen labels. If the specimen was only 
labeled “‘Minn.”’ or ““MN,”’ then this is list- 
ed as unknown county and unknown col- 
lector in our results. The records are pre- 
sented in chronological order to reflect the 
historical precedence of the collections, and 
when dates were not present on the labels, 
we have placed them in the chronology 
based on our knowledge of when various 
collectors were active. Unless indicated by 
a specific collection acronym, the record is 
from the UMIC. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


RESULTS 
Tribe Notiophilini 


Notiophilus aquaticus (Linnaeus).—Ot- 
ter Tail Co., O. Lugger (1). Lake Co., Two 
Harbors, M.H. Hatch, VI.28.1927 (1). Crow 
Wing Co., C.E. Mickel, V.14.1928 (1): 
Cook Co., Superior National Forest, baited 
pitfall trap, K.J.K. Gandhi, 48°07'31"N 
90°50'30"W, P. banksiana wind-disturbed- 
prescribe-burned forest, VII.30-VIII.11. 
2003 (1). We collected a total of one adult. 

Notiophilus aquaticus is a transcontinen- 
tal species that ranges from Newfoundland 
to Alaska and south to Arizona (Bousquet 
and Larochelle 1993: 50). This species is 
associated with open areas (Lindroth 1961— 
69: 95). 


Tribe Carabini 


Calosoma affine Chaudoir.—Olmsted 
Co., C.N. Ainslie (1, ISU). Martin Co., Tru- 
man, J.H. Patchin, VII.20.1936 (1). 

Calosoma externum (Say).—OlImsted 
Co., C.N. Ainslie (1, ISU). Ramsey Co., St. 
Paul, Como Park lights, W.E. Hoffman, 
V1II.6.1921 (1). Houston Co., unknown col- 
lector, V.26.1940 (1). 

Calosoma obsoletum Say.—Olmsted 
Co., C.N. Ainslie (1, ISU). Cass Co., Cass 
wakes KE Aiccler  ViEZTE9345 (i) eeaRocK 
Co., Luverne, A.E. Pritchard, IX.13- 
14.1935 (1); PB Nicholson, VI.8.1938, 
WAL WAL SNM) WAIT EI@ SKS WALZ M235 (7). 
Big Stone Co., Odessa Township, W. Stehr, 
VIUI.24.1935 (2, CAS), (1, NMNH). Polk 
Co., D.G. Denning, VIII.13.1936 (1). Mille 
acse Con Re Handford= V2 ssl937 eG»: 
Ramsey Co., R.H. Handford, V.28.1937 (1). 
Beltrami Co., D.G. Denning, VI.22.1937 
(1). Kittson Co., Hallock, D.G. Denning, 
VII.8.1937 (1); A.W.Buzicky, VIII.12.1939 
(1). Nobles Co., C.E. Mickel, VI.2.1938 
(3). Pipestone Co., C.E. Mickel, VI.3.1938 
(3). Lincoln Co., C.E. Mickel, VI.4.1938 
(3). 

Calosoma_ scrutator (Fabricius).—Hen- 
nepin Co., Minneapolis, O.W. Oestlund (1); 
Lake Minnetonka, G. Swanson, VI.10.1933 


VOLUME 107, NUMBER 4 


(1). Fillmore Co., Entomology Class, 
V251936 (@) Houston’ i€o..) BiG) Mas 
V.22.1937 (1). Ramsey Co., E. Thomas, 
IX.15.1937 (1). 

Carabus_ sylvosus Say.—Olmsted Co., 
C.N. Ainslie (1, ISU). Itasca Co., Itasca 
Park, L.W. Orr, VII.2.1928, VIII.22.1931 
(2). Anoka/Isanti Co., Cedar Creek Bog, 
H.L. Gunderson, VIII.24.1948 (1). Anoka 
Co., Cedar Creek Natural History Area, 
Kelp earsens ME Davis, *VILEZ6-2000: 
IX.13.2000 (2, KJL). 

Carabus vinctus (Weber).—OlImsted Co., 
C.N. Ainslie (1, ISU). 


Tribe Cychrini 


Scaphinotus fissicollis (LeConte).—Olm- 
sted Co., C.N. Ainslie (10, ISU). Houston 
Co., Jellison, V.25.1940 (1). 

Sphaeroderus nitidicollis brevoorti Le- 
Conte.—Cook Co., Superior National For- 
est, baited and unbaited pitfall traps, K.J.K. 
Gandhi, 48°02'35"N 90°22'30"W, 
48°02'42"N 90°21'58’W, 48°02'26’"N 
90°23'37'"W, P. tremuloides/B. papyrifera/ 
conifer wind-disturbed forests, VIII.10- 
IDK AD) ZOO; VZV IN AOOUS — M12 7/— 
VIVO 20015 VII2ZO=X. 13.2001, V.31- 
Mille AZO O25 WaIRS AXE 2120025 N23 0= 
IX.15.2003 (36); 48°02'11"N 90°23'01"W, 
48°01'46"N 90°23'53"W, P. tremuloides/B. 
papyrifera/conifer wind-disturbed-salvage- 
logged forests, VIII.21—26.2000, VI.7— 


Die) OLENA. 6- ViIE2O 2001 VIE O- 
Xe SeZ0 OIRO NEZ8=ViE 1722002 vill9o- 
LPXeZ IZ OO 22" VAIS -=Dxer 420035" 21): 
48°01'56"N, 90°23'55"W, 48°01'56"N 


90°23'56"W, 48°02'42"N 90°21'58”"W, P. 
tremuloides/B. papyrifera/conifer undis- 
turbed forests, WVI.7—26.2001, VII.9- 
eS ZOO TE VESIEV. 27-2002) “VIlle- 
[X.21.2002, V.30-IX.15.2003 (81); and 
48°00'33"N 90°25'15’"W, P. tremuloides/B. 
papyrifera/conifer wind-disturbed-pre- 
scribed burned forest, VI.11—25.2001, 
VITRZ0=8: 2001, “Vi2Z7-VIL2-.2002: 
VIII.9—23.2002, VIII.18-1X.14.2003 (10). 
We collected a total of 148 adults. 

Our collection of S. n. brevoorti is a new 


923 


regional record for the north central United 
States. This species had been collected in 
the northeastern United States as far west 
as New York, and in Canada from Nova 
Scotia, through Ontario, Manitoba, and as 
far west as Saskatchewan (Bousquet and 
Larochelle 1993: 78). It is a forest species 
associated with moist habitats such as moss 
and litter (Lindroth 1961—69: 30, Pearce et 
al. 2003: 347). In agreement with Pearce et 
al. (2003), we caught S. n. brevoorti exclu- 
sively in the P. tremuloides/B. papyrifera/ 
conifer forest cover-type, which suggests 
that this species may be dependent upon 
micro-habitats typical of deciduous forests. 


Tribe Elaphrini 


Elaphrus lecontei Crotch.—Itasca Co., 
Itasca Park, Bohall Lake, D.G. Denning, 
V3 1935 (1)? Clay Cos Bluestem State 
Natural Area, mesic prairie, D. Rider and 
G. Fauske, VII.27.1995 (1, NDSIRC). 

Tribe Clivinini 

Clivina americana Dejean.—Ramsey 
Co., Gray Cloud Island, H.H. Knight, 
V.23.1920 (1). Wilkin Co., D.G. Denning, 
VII.11.1937 (1). Houston Co., R. Anderson, 
V.20.1938 (1). 

Clivina bipustulata (Fabricius).—Hen- 
nepin Co., Minneapolis, E. Vaughn, 
IV.23.1942 (1). Houston Co., C.E. Mickel, 
IV2371942-> (1). 2Astkin™ Go:, “Bluth (C-E- 
Mickel, VI.1.1942 (1). 

Clivina impressefrons LeConte.—Wright 
Corn Buttalo;> Se) Partin, \Vil2omo4 7: 
VIII.3.1947 (2, NMNH). 

Dyschirius aeneolus LeConte.—Lake 
Co., Two Harbors, at beach, M.H. Hatch 
(1). 

Dyschirius erythrocerus LeConte.—Un- 
known Co., A. Bolter (1, INHSIC). Olm- 
sted Co., C.N. Ainslie (2); Rochester, C.N. 
Ainslie (1). Nicollet Co., St. Peter, Fish 
Hatchery, S. Kepperley, VIII.11.1925 (1). 
Polk Co., Crookston, Red Lake River, at 
light, D.G. Denning, VII.18.1935 (3). 

Dyschirius globulosus (Say).—Ramsey 
Co., O. Lugger (4). Unknown County, O. 


924 


Lugger (1). Olmsted Co., C.N. Ainslie (5). 
Lake Co., Two Harbors, roots of grass, 
beach-drift, beach, M.H. Hatch, 
WDA ODI. VALSOSIS A, WNL WAT (3). 
Goodhue Co., Frontenac, W.C. Stehr, 
V.29.1930 (2). Winona Co., A.W. Buzicky, 
V.20.1938 (1). Mille Lacs Co., Mille Lacs, 
H.E. Milliron, V.18.1940 (1). Clay Co., 
Trust Lands, six miles east of Felton, hayed 
prairie, unmanaged prairie, C. Locken and 
G. Fauske, VI.7.1996, VI.21.1996, 
VII.28.1996 (4, NDSIRC); Bluestem State 
Natural Area, unmanaged prairie, dry prai- 
rie, hayed prairie, L. DeCock, J. Albertson 
and K. Urlacker, VI.27.1996, VII.10.1996, 
VIII.5.1996 (4, NDSIRC); Bicentennial 
Prairie, J. Albertson and L. DeCock, 
VII.3.1996 (1, NDSIRC); and Blazingstar 
Prairie, J. Albertson and L. DeCock, 
VII.3.1996, (2, NDSIRC). Anoka Co., Ce- 
dar Creek Natural History Area, K.J. Lar- 
sen, VII.6.2000, VIII.16.2000, (2, KJL). 

Dyschirius haemorrhoidalis (Dejean).— 
Mille Lacs Co., Mille Lacs, H.G. Rodeck, 
X.21.1928 (1). 

Dyschirius longulus LeConte.—Polk 
Co., Crookston, light trap, D.G. Denning, 
WILD ULB s) (CY). 

Dyschirius pumilus (Dejean).—Anoka 
Co., Cedar Creek Natural History Area, 
K.J. Larsen, VII.6.2000, VIII.29.2000 (2, 
KJL). 

Dyschirius sellatus LeConte.—Ramsey 
Co., Gray Cloud Island, W.E. Hoffman, 
WIN IZM OA Ci). 

Schizogenus lineolatus (Say).—Olmsted 
Co., C.N. Ainslie (2). 


Tribe Trechini 


Trechus crassiscapus Lindroth.—Cook 
Co., Superior National Forest, unbaited pit- 
fall trap, K.J.K. Gandhi, 48°03’30"N 
90°32'40"W, P. tremuloides/B. papyrifera/ 
conifer wind-disturbed forest, V.30- 
V1I.18.2003 (1). We collected a total of one 
adult. 

Trechus crassiscapus is primarily an 
eastern species, and our collection reflects 
a major westward extension of this species 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


in the United States. It ranges from New- 
foundland to Ontario and south to Massa- 
chusetts (Bousquet and Larochelle 1993: 
120), and is a hygrophilous species associ- 
ated with swamps in forested areas (Lin- 
droth 1961—69: 200). 


Tribe Bembidiini 


Bembidion coloradense Hayward.— 
Wilkin Co., light trap, D.G. Denning, 
VII.11.1937 (1). Itasca Co., Itasca Park, 
light trap, C.E. Mickel, VII.8.1939 (1). Polk 
Co., Crookston, unknown collector, VII.28— 
DS) NSIS) (AD). 

Bembidion coxendix Say.—Benton Co., 
O.W. Oestlund (1). Ramsey Co., Gray 
Cloud Island, W.E. Hoffman, VII.12.1921 
(1). Le Sueur Co., Ottawa, W.E. Hoffman, 
Wil Salo 225@)s 

Bembidion dorsale Say.—Ol\msted Co., 
C.N. Ainslie (2). 

Bembidion fortestriatum (Motschul- 
sky).—Itasca Co., Litthe Winnibigoshish 
Lake, Ky) Cooper, VIA0—TEI935 Ge 
NMNH). Wright Co., Buffalo, S.I. Parfin, 
VIII.3.1947 (1, NMNH). Roseau Co., blue- 
grass, A.G. Peterson, V.22.1968 (1). 

Bembidion inaequale Say.—St. Louis 
Co., Duluth, unknown collector (Chtth?) (1, 
NMNH). Hennepin Co., Minneapolis, at 
light, A.T. Hertig, VII.13.1922 (1). Red 
Lake Co., Plummer, from mud near river, 
off sand near river, D.G. Denning, 
V.16.1933, V.23.1933 (2). Houston Co., 
Southeast tip of county, H.R. Dodge, 
V.24.1936 (1). Unknown Co. (probably 
Ramsey), Gray Cloud, C.E. Mickel, 
IV.21.1939 (1). Washington Co., Afton, En- 
tomology Class, V.3.1947 (1). Houston Co., 
Winnebago Creek Valley, 3-4 m NE Ei- 
tzen, A. Raske, V.30.1960 (1, NMNH). 
Clay Co., Buffalo River State Park, S.J. 
Vick, [X.15.1980 (1, UCB). 

Bembidion mutatum Gemminger & Har- 
old.—Clay Co., Clay County Trust Lands, 
hayed mesic prairie, P. Tinerella, A. Abbott, 
and G. Fauske, VI.24.1997, VII.9.1997, 
VII.28.1997, VIII.13.1997, [X.22.1997 (46, 
NDSRIC); unmanaged prairie, P. Tinerella, 


VOLUME 107, NUMBER 4 


and A. Abbott, VII.28.1997 (10, NDSRIC); 
and grazed mesic prairie, P. Tinerella, and 
A. Abbott, VII.9.1997 (1, NDSRIC). Cook 
Co., Superior National Forest, unbaited and 
baited pitfall traps, K.J.K. Gandhi, 
48°07'21"N 90°51'25”"W, P. banksiana 
wind-disturbed-salvage logged forest, 
VITI.26-I1X.30.2000 (1); 48°05'12"N 
90°47'35"W, P. banksiana wind-disturbed- 
prescribe-burned forest, VII.6-VIII.4.2001 
(1); and 48°00'30"N 90°25'15’W, P. tre- 
muloides/B. papyrifera/conifer wind-dis- 
turbed-prescribe-burned forest, I[X.8— 
29.2001 (1). We collected a total of three 
adults. 

Bembidion mutatum is transcontinental 
from Newfoundland to Alaska extending 
south in eastern North America to New 
York and south in the Rocky Mountains to 
New Mexico (Bousquet and Larochelle 
1993: 146, Purrington et al. 2000: 202). 
This species is generally caught in open 
habitats with sparse vegetation (Lindroth 
1961—69: 387). 

Bembidion pedicellatum LeConte.— 
Hennepin Co., O.W. Oestlund (1). Winona 
Co., Dresback, W.S. Lund, VII.12.1920 (1). 

Bembidion postremum Say.—Olmsted 
Co., C.N. Ainslie (1). Itasca Co., Itasca 
Park, light trap, C.E. Mickel, VII.8.1939 
(1). 

Bembidion texanum Chaudoir.—Henne- 
pin Co., O.W. Oestlund (1). Ramsey Co., 
O.W. Oestlund (1). Olmsted Co., C.N. Ain- 
slie (10). Le Sueur Co., Fish Hatcheries, at 
light, W.E. Hoffman, VII.23.1922 (1). 

Bembidion transversale Dejean.—Un- 
known Co., A. Bolter (3, INHSIC). St. Lou- 
is Co., Duluth, A. Bolter (1, INHSIC). 
Cook Co., Grand Portage, Susie Island, 
Lake Superior, unbaited fence trap, gravel 
beach of Lake Superior, C. Garry and D. 
Schwert, VII.1.1991 (10). 

Bembidion wingatei Bland.—Cook Co., 
Superior National Forest, baited and unbaited 
pitfall traps, K.J.K. Gandhi, 48°02'42"N 
90°21'58’"W, P. tremuloides/B. papyriferalco- 
nifer undisturbed forest, VII.7-VIII.3.2000 
(1); 48°07'51"N 90°51'30"W, P. banksiana 


S25 


undisturbed forest, VUI.6-VIII.30.2001 (2); 
and 48°07'21"N 90°51'25"W, P. banksiana 
wind-disturbed-salvage-logged forest, VI.6- 
V1I.21.2001 (1). We collected a total of four 
adults. 

Bembidion wingatei ranges from New- 
foundland and St. Pierre and Miquelon 
south to North Carolina, and west to Wy- 
oming (Bousquet and Larochelle 1993: 
149). It also has been reported by Petrice et 
al. (2002: 9) in Aitkin and Cass counties of 
Minnesota. This species inhabits subterra- 
nean habitats beneath rocks in grasslands 
and deciduous litter in forest stands (Lin- 
droth 1961—69: 406). 

Paratachys scitulus (LeConte).—Ramsey 
Co., St. Paul, Battle Creek, W.E. Hoffman, 
V.20.1922 (1); St. Paul, University Farm, at 
light, A.A. Granovsky, VI.26.1936 (1). 
LeSueur Co., Fish Hatcheries, W.E. Hoff- 
man, VII.20-30-1922 (1). Hennepin Co., G. 
Kohls, V.1.1937 (1). 


Tribe Psydrini 


Nomius pygmaeus (Dejean).—St. Louis 
Co., Duluth, O. Lugger (6). Unknown Co., 
O. Lugger (2). Unknown Co., unknown col- 
lector and A. Bolter (2, INHSIC). Kooch- 
iching Co., International Falls, J.P. Kelly, 
VII.30.1921 (3). Koochiching Co., Little 
Fork, C.N. Cantwell, VII.18.1922 (1); Cook 
Co., Schroeder, unknown _ collector, 
VIII.20.1970 (2). 


Tribe Patrobini 


Patrobus foveocollis (Eschscholtz).— 
Koochiching Co., T160N, R26W, S33, 
H.M. Kulman, VI.30-VII.10.1971 (1). 
Cook Co., Superior National Forest, baited 
and unbaited pitfall traps, K.J.K. Gandhi, 
48°03'30"N 90°32'40’W, P. tremuloides/ 
B. papyrifera/conifer wind-disturbed for- 
est, VIII.3-I[X.29.2000 (1); 48°00'30’N 
90°25'15"W, P. tremuloides/B. papyrifera/ 
conifer wind-disturbed-prescribe burn- 
ed forest, VI.25-VII.9.2001 (1); and 
48°08'12"N 90°51'25"W, PP. banksi- 
ana wind-disturbed-salvage-logged forest, 


926 


VII.12—25.2002 (1). We collected a total of 
three adults. 

Patrobus foveocollis is a holarctic spe- 
cies, ranging in North America from Alaska 
and British Columbia southward in the 
Rocky Mountains to Colorado, and from 
Newfoundland to Vermont in eastern North 
America (Bousquet and Larochelle 1993: 
159). It is generally found under fallen dead 
leaves and shaded places (Lindroth 1961— 
69: 186). 

Patrobus septentrionis Dejean.—Kooch- 
iching Co., T71B, R24W, S10, H.M. Kul- 
man, V.25-VI.6.1971 (1). Cook Co., Supe- 
rior National Forest, unbaited pitfall trap, 
K.J.K. Gandhi, 48°03'30"N 90°32'40’W, P. 
tremuloides/B. papyrifera/conifer wind-dis- 
turbed forest, VII.9-20.2001 (1). We col- 
lected a total of one adult. 

Patrobus septentrionis is also a holarctic 
species, ranging from Alaska and Washing- 
ton southward in the Rocky Mountains to 
Colorado, and from Newfoundland to New 
Hampshire in eastern North America. It has 
also been reported by Petrice et al. (2002: 
9) in Aitkin and Cass Counties of Minne- 
sota. This species is associated with vege- 
tation in riparian areas along lakes, ponds, 
and streams (Lindroth 1961—69: 184-185). 

Platypatrobus lacustris Darlington.— 
Cook Co., Grand Portage Reservation, un- 
known collector, VII.5.2000 (2, GPB). 


Tribe Pterostichini 


Cyclotrachelus sodalis colossus (Le- 
Conte).—Olmsted Co., six miles east Chat- 
field, J.R. Powers, V.17.1967 (1, UCB). 
Wabasha Co., Lake City, J.R. Powers, 
V1.4.1970 (1, UCB). 

Cyclotrachelus torvus torvus LeConte.— 
Rocke Gos Y luvernte e©- ES eMicke lr 
V1.26.1925 (1). 

Gastrellarius honestus (Say ).—Otter Tail 
(COs, (©), Ibmeryerr (il), 

Lophoglossus scrutator (LeConte).— 
Olmsted Co., C.N. Ainslie (1). 

Poecilus corvus (LeConte).—Norman 
Co., A.A. Nichol, V.20.1923 (1). Unknown 
County, swept from sweet clover, B.A. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Haws, VII.23.1956, VIII.11.1956 (6). Polk 
Co., Crookston, A.E. Grable, VII.17.1960 
(1). Clay Co., Buffalo River State Park, J. 
Allen, [X.19.1980 (1, UCB); Moorhead, J. 
Allen, [X.15.1980 (1, UCB). Wilkin Co., 
potato, R. Johnston, VI.19.1982 (3). We 
also found three specimens of P. corvus in 
the NDSIRC from Clay and Norman Cos. 
(Bluestem and Agassiz Dunes State Natural 
Areas) that had been collected between 
V1I.27 and IX.24.1996 (Tinerella and Rider 
2001). 

Poecilus scitulus (LeConte).—Polk Co., 
Crookston, swept from sweet clover, un- 
known collector, VII.21.1952 (1). 

Pterostichus ebeninus (Dejean).—Un- 
known Co., unknown collector (1, NMNH). 

Pterostichus melanarius (iliger).— 
Wright Co., Monticello, unknown collec- 
tor, VIII.26.1990 (1). Aitkin Co., 1.4 
miles west of Willow River, D.E. Hansen, 
46.334°N 03.096°W, IV.22.1994 (1). Clay 
Co., Trust Lands, hayed prairie, D. Rider 
and “G. ‘Rauske,  YVilllnIA aS Sosa Ge 
NDSIRC); Bluestem Prairie, grazed prai- 
rie, wet prairie, reclaimed prairie, mesic 
prairie, P. Tinerella, C. Jordan and C. Da- 
vis, [X.4.1997, VIII.5.1999 (5, NDSIRC); 
and Blazingstar Prairie, C. Jordan and C. 
Davis, VIII.30.1999 (1, NDSIRC). Ram- 
sey Co., St. Paul, pitfall trap, 70% etha- 
nol, wheat, E.E. Hladilek, VI.29.2000, 
V1.30.2000 (5). Anoka Co., Cedar Creek 
Natural History Area, K.J. Larsen, 
VII.6.2000, VII.19.2000 (2, KJL). Cook 
Co., Superior National Forest, baited and 
unbaited pitfall traps, K.J.K. Gandhi, 
48°03'42"N 90°33'55"W, 48°02'42"N 
90°21'58"W, P. tremuloides/B. papyrifera/ 
conifer undisturbed forests, VII.6- 
VIII.27.2000, VIII.3-X.5.2000, VIII.21- 
X.5.2000 (20); 48°03’30"N 90°32'40"W, 
P. tremuloides/B. papyrifera/conifer 
wind-disturbed forest, VIII.3-[X.29.2000 
(3); 48°03'48"N 90°32'40"W, P. tremulo- 
ides/B. papyrifera/conifer wind-dis- 
turbed-salvage-logged forest, VII.6- 
[X.30.2000, VIII.3-1X.30.2000 (86); 
48°07 SIN 9075130 Wa) 48073 1N 


VOLUME 107, NUMBER 4 


927 


Table 1. Habitat, locality, and seasonal data on Pterostichus melanarius caught in baited and unbaited pitfall 
traps in Populus tremuloides/Betula papyrifera/conifer and Pinus banksiana sites in Cook Co., Minnesota during 


2001—2003. 


Forest Cover-type 


P. trem./B. pap./ 
Conifer 


Pinus banksiana 


Total Number 
of Adults 


Silvicultural 
Treatment 


Undisturbed 


Wind-disturbed 


Wind-disturbed- 


salvage logged 


Wind-disturbed- 
prescribe burned 


Undisturbed 


Wind-disturbed 


Wind-disturbed- 
salvage logged 


Wind-disturbed- 
prescribe burned 


Number 
Latitude/Longitude Trapping Period of Adults 
48°01'56"N 90°23'56"W V.21—VI.25.2001 3 
48°03'42”"N 90°33'55"W VII.9—VII.7.2001 | 
48°02'42"N 90°21'58”W VIII.20—X.13.2001 3 
48°01'56"N 90°23'55”W VI.1-1[X.21.2002 442% 
V.29-IX.14.2003 
48°01'50"N 90°24'20"W VI.7—VI.27.2001 D, 
48°03'30"N 90°32'40"W VI.25—VII.9.2001 2 
48°02'26"N 90°23'37"W VIII.20-IX.8.2001 17 
48°01'50"N 90°24'20"W VI.27-VIII.3.2002 BOs 
V .30—VIII.5.2003 
48°01'45"N 90°24'18"W V.30-1X.29.2001 659 
48°03'48"N 90°32'40"W V .28-IX.21.2002 
48°01'46"N 90°23'53"W VI.19-IX.15.2003 
48°02'11”N 90°23'01"W 
48°00'30"N 90°25’ 15”W V.29-IX.29.2001 2,622 
48°05'01"N 90°46'52”W V.28-IX.21.2002 
48°05'05"”N 90°49'17"W V.29-IX.14.2003 
48°03'30"N 90°35'00"W 
48°07'31"N 90°51'50"W VI.6-IX.22.2001 211 
48°07'38"N 90°51'36"W V.29-IX.14.2002 
48°07'51"N 90°51'24"W VI.4-IX.10.2003 
48°07'29"N 90°51'40"W 
48°07'12"N 90°50'50”"W V1I.6—VII.5.2001 10 
VII.13—VII.27.2002 
VII.28—VIII.1 1.2003 
48°07'21”"N 90°51'25"W VI.6-IX.22.2001 794 
48°07'40"N 90°50'00"W VI.13-IX.14.2002 
48°07'27"N 90°51'00"W VIT.9-IX.10.2003 
48°05'00"N 90°48'12”W V.31-IX.22.2001 5,561 
48°05'12”N 90°47'35”W V.29-IX.14.2002 
48°07'51”"N 90°51'30"W VI.4-IX.11.2003 
48°07'31"N 90°50'30"W 
10,362 


* Includes number of adults from both 2002 and 2003. 


90°51'50"W, P. banksiana undisturbed 
forests, VIII.3-VIII.27.2000, VIII.4- 
xXe5e2 000) VIII.20-X.1.2000 (CIB) 
48°07'12”N 90°50'50"W, P. banksiana 
wind-disturbed forest, VIII.28-X.5.2000 
@)euweand "48207, 21N) »9Os5 1425 Ww, 
48°07'27"N 90°51'00"W, P. banksiana 
wind-disturbed salvage-logged forests, 
VIII.3-1X.30.2000, VIII.4-1X.30.2000, 
VIII.20-IX.30.2000 (38). We also collect- 
ed 10,362 specimens of P. melanarius in 
our northeastern Minnesota study site dur- 


ing the summers of 2001—2003 in baited 
and unbaited pitfall traps (Table 1). we 
collected a total of 10,523 adults. Anoka 
Co., Carlos Avery Wildlife Management 
Area, Old Game Farm Road, A.K. Am- 
bourn, 45°19’'N 93°07'W, VII.21.2002 (1). 
Crow Wing Co., Barrows, south of Brai- 
nerd, near Hwy. 371, unbaited pitfall trap, 
K.J.K. Gandhi, 46°24’N 94°08'W, P. 
banksiana forest, VII.25-VII.8.2002 (1). 

Pterostichus melanarius is an introduced 
from Europe, and has 


species western 


928 


achieved a transcontinental distribution in 
North America (Lindroth 1961—69: 491, 
Bousquet and Larochelle 1993: 174, Will et 
al. 1995: 66, Purrington et al. 2000: 201). 
This beetle is associated with disturbed and 
open habitats such as cultivated agricultural 
lands and managed forest landscapes (Lind- 
roth 1961—69: 492). It is a generalist pred- 
ator of economically important pests (Lee 
1998), but it also known to feed on conifer 
seeds (Lindroth 1961—69: 1116). In Min- 
nesota, this species has been reported from 
Aitkin and Cass Counties (Petrice et al. 
2002). Three records in the UMIC (Wright, 
Aitkin and Ramsey Counties), three records 
from the NDSIRC (Clay County), one re- 
cord from KJL (Anoka County), and our 
collections from Cook County pre-date the 
report by Petrice et al. (2002). Prerostichus 
melanarius is known to aggressively colo- 
nize new habitats. In Minnesota, it has rap- 
idly established populations even in the re- 
mote areas of the Superior National Forest, 
although it was absent in similar forest- 
types in Ontario (Pearce et al. 2003). We 
collected 10,523 adults in 2000—2003 in- 
dicating the abundance of this species in 
northeastern Minnesota. It was the most 
abundant carabid in our survey there. We 
trapped only one specimen of P. melanarius 
in our survey in central Minnesota. 

Pterostichus permundus (Say).—Wa- 
bashay Gor dake Citys eRe. Bowens: 
IX.3.1961, VIII.20.1962 (3, UCB); Dum- 
fries, JER] Powers, VIE23: 1991 @) UCB): 
Clay Co., Moorhead, R.A. Woehl, 
IX.11.1972 (1, UCB). Redwood Co., Red- 
wood Falls, A.C. Rustand, [X.18.1993 (1, 
UCB). 


Tribe Zabrini 


Amara basillaris (Say).—Unknown 
County, O. Lugger (1). Olmsted Co., C.N. 
Ainslie (1). 

Amara chalcea Dejean.—Anoka Co., 
Cedar Creek Natural History Area, K.J. 
Larsen, VII.6.2000, VII.19.2000, (2, KJL). 

Amara coelebs Hayward.—Big Stone 
Co., O. Lugger, VII.19.1910 (1). Unknown 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Co., unknown collector (1, INHSIC), (7, 
NMNH). Hennepin Co., O.W. Oestlund (2). 
Olmsted Co., C.N. Ainslie, May (3); C.N. 
Ainslie (9). Unknown Co., A. Fenyes (2, 
CAS). Ramsey Co., W.E. Hoffman, 
IfI.21.1922 (1); St. Anthony Park, O. Lug- 
ger, H.H. Knight, D. Murray, W.D. Buch- 
anany, and) Wee) Hottimany  leAnioZ2 
LEZ61922- EZ Ss 19205 TiEZ Sano 
V.5.1920, III.23.1937 (18); St. Paul Univer- 
sity Farm, W.E. Hoffman and C.E. Mickel, 
II.27.1922, V.19.1922 (3). Clearwater Co., 
Wake itascay S-Ay Grahams ViltvelS23nG): 
Lake Co., Two Harbors, M.H. Hatch, 
V1I.29.1927 (1). Hennepin Co., Entomology 
Class, IV.30.1938 (1). Pope Co., one mile 
south Sedan, J. Hafsted, IV.29.1961 (1, 
UCB). Clay Co., Moorhead, B. Wermager, 
V.3.1961 (2, UCB). Lyon Co., J.T. Bush, 
IV-13—-1963, female specimen, unable to 
verify species status completely (1, ISU). 
Traverse Co., two miles north Browns Val- 
leva wilake p iraverse.. bo a olleison: 
V1I.20.1974 (1, UCB). Cook Co., Superior 
National Forest, baited and unbaited pitfall 
traps, K.J.K. Gandhi, 48°03'30"N 
90°32'40"W, P. tremuloides/B. papyrifera/ 
conifer wind-disturbed forest, VI.25- 
VIL9.2001 (1); 48°03'48"N 90°32'40"W, 
48°01'46"N 90°23'53”"W, 48°01'45’N 
90°24'18"W, P. tremuloides/B. papyrifera/ 
conifer wind-disturbed-salvage-logged for- 
ests, VI.7—25.2001, V.28-VII.12.2002 (5); 
ALSO SOINE GODS MS WE ESOS OLIN 
90°46'52"W,. 48°05’05’N_ 90°49'17"W, PP. 
tremuloides/B. papyrifera/conifer wind-dis- 
turbed-prescribe-burned forests, VI.7- 
VIS 2001 V2 8-N 117520 02 ave 
V1I.18.2003, VII.14-VIII.18.2003 (10); 
ASSO L2IUING 9 ORS Se Wee Se OTe TEN 
90°51'00"W, P. banksiana wind-disturbed- 
salvage-logged forests, VII.5-VII.16.2001, 
VI.27-ViIL.13-2002, .VI:23-ViIL9-2003 G6): 
and 48°05’00"’N 90°48'12”W, 48°07'31"N 
90°50’30”"W, 48°05'12”N 90°47'35"W, 
48°04'59"N 90°47'35"W, P. banksiana 
wind-disturbed-prescribe-burned forests, 
V.31-VI1.6.2001, V.29-VIHI.12.2002, VI.4- 


VOLUME 107, NUMBER 4 


VIII.27.2003 (23). We collected a total of 
44 adults. 

Amara coelebs ranges from Wisconsin to 
British Columbia, and south to Colorado 
(Bousquet and Larochelle 1993: 195). It is 
a prairie species occurring on dry, grassy 
and sandy areas (Lindroth 1961—69: 725). 
In our study, it was caught only in the dis- 
turbed forest stands. 

Amara crassispina LeConte.—Hubbard 
Co., unknown collector, [X.10.1980 (1, 
KWW). 

Amara ellipsis (Casey).—Clay Co., 
Moorhead, J. Allen, [X.15.1980 (1, UCB). 

Amara pallipes Kirby.—Kanabec Co., 
Mora, C.R. Yeager, VI.24—30.1934 (1). 
Anoka Co., Cedar Creek Natural History 
Area, burned, K.J. Larsen, M. Davis, 
VII.27.2000 (1, KJL). 

Amara pennsylvanica Hayward.—Ram- 
sey Co., St. Paul, University Farm lights, S. 
Kepperley, VII.23.1924 (1). Rock Co., Lu- 


verne, C€.E. Mickel, IX.13—-14.1935 (2). 
Houston Co., unknown collector, V.23— 
24.1936 (1). 


Amara torrida (Panzer).—Otter Tail Co., 
© ys Eusger (G)s Martin) Co: ‘S:S> Easter, 
VilI1321926" (1) Pope Cos Sedan, DIG: 
Demming, WIP Cb). Io: (os 
Crookston, light trap, D.G. Denning, 
VII.19.1935 (1). Ramsey Co., St. Paul, Uni- 
versity Golf Course, light trap, A.A. Gra- 
novsky, VII.20.1936 (1). Roseau Co., Win- 
naska, on Timothy grass, unknown collec- 
tOrV MEI SAO S27): 

Pseudamara arenaria (LeConte).— 
Koochiching Co., emergence trap, L.C. 
Thompson, VI.14.1972 (1). Cook Co., Su- 
perior National Forest, unbaited pitfall trap, 
K.J.K. Gandhi, 4803'48"N 90°32'40'W, P. 
tremuloides/B. papyrifera/conifer wind-dis- 
turbed-salvage-logged forest, V.29- 
VI.18.2003 (1). We collected a total of one 
adult. 

Pseudamara arenaria is primarily a 
northeastern species ranging from New 
Brunswick to Ontario in the north and West 
Virginia to Illinois in the south (Bousquet 
and Larochelle 1992:190). It is reported to 


929 


be an open-habitat, riparian, and cavernic- 
olous species (Lindroth 1961—69:650). 


Tribe Pangaeini 
Panagaeus fasciatus Say.—Hennepin 
Co., O. Lugger (1). Ramsey Co., St. An- 
thony Park, O. Lugger (3). Olmsted Co., 
C.N. Ainslie (2). Clay Co., Bicentennial 
Prairie, burned dry prairie, P. Tinerella and 
C. Davis, VII.27.2000 (1, NDSIRC). 


Tribe Chlaeniini 


Chlaenius erythropus Germar.—Un- 
known County, O. Lugger (1). Olmsted 
Co., C.N. Ainslie (4). Ramsey Co., A.T. 
Herty, IX.2.1921 (2). Anoka Co., K.S. Liu, 
V.8.1937 (1). Houston Co., H.E. Milliron, 
PM: ‘Schroeder, HES. Welford) D:G> Den- 
ning, (Ee. Mickel Vik Jens '@. Kohls EF: 
Thomas, M. Gotschall, H.C. Ma, I. Tarshis, 
and “RH. Dagey,  V-13:119373 V2161937- 
V.22-2381937R NA 9ST eive2 Gn S7- 
V.20.1938 (18). Goodhue Co., unknown 
collector, V.21.1937 (1). Wabasha Co., H.E. 
Gustafson, V.21.1937 (1). 

Chlaenius prasinus Dejean.—Unknown 
Coz, Cliffty(akes Cor Chitttake?) ayer 
Hoffman, V.30.1922 (2). 

Chlaenius purpuricollis purpuricollis 
Randall.—Red Lake Co., Plummer, off 
sand near river, D.G. Denning, V.23.1933 
(1). We also found 113 specimens of C. 
purpuricollis in the NDSIRC from Clay and 
Polk Cos. (Bluestem, Blazingstar, and Ag- 
assiz Dunes State Natural Areas, Clay 
County Trust Lands, and Bicentennial Prai- 
rie) that had been collected between 
VII.27.1995 and [X.17.1999 (Tinerella and 
Rider 2001). 

Chlaenius pusillus Say.—Olmsted Co., 
C.N. Ainslie (2). 

Tribe Licinini 

Badister ocularis Casey.—Traverse Co., 
O.W. Oestlund (1). Ramsey Co., St. Paul, 
Minnesota Farm, A.A. Granovsky, 
V.29.1936 (1). 

Dicaelus furvus carinatus Dejean.— 
Ramsey Co., St. Anthony Park, O. Lugger 


930 


(4). Olmsted Co., May, June, C.N. Ainslie 
(3). Fillmore Co., G. Kohls, [V.24.1927 (1). 
Goodhue Co., Frontenac, W.C. Stehr, 
V.29.1930 (1). Fillmore Co., Preston, S.I. 
Parfin, V.8.1948 (1, NMNH). 


Tribe Harpalini 


Acupalpus partiarius (Say).—Lake Co., 
won anb ons sbedchhnes Velie uuhlatehe 
VI1.28.1927 (1). Houston Co., R.H. Daggy, 
V.22.1937 (1). Mille Lacs Co., C.E. Mickel, 
and H.E. Milliron, VI.2.1937, V.10.1970 
Oye Grows Wine ion, (CE Mickel: 
V.14.1938 (1). 

Anisodactylus agricola (Say).—Wabasha 
Co., seven miles southwest Wabasha, J.R. 
Powers, V.6.1972 (1, UCB). 

Anisodactylus carbonarius (Say).—Ram- 
sey Co., St. Anthony Park, O. Lugger (1). 
Olmsted Co., C.N. Ainslie (2). 

Anisodactylus melanopus (Haldeman).— 
Wabasha Co., Lake City, W.O. Powers, 
War So (Cl, WKEls})). 

Bradycellus atrimedeus (Say).—Lac Qui 
Parle Co., Lac Qui Parle Park, E.U. Bals- 
baugh, Jr, [V.16.1967 (2, SDSU). 

Bradycellus badipennis (Haldeman).— 
Koochiching Co., T160, NR26, WS33, pit- 
fall trap, H.M. Kulman, IX.25-X.13.1971 
CD), 

Bradycellus insulsus (Casey).—Polk Co., 
Crookston, Red Lake River, at light and 
malaise trap, unknown collector, VII.20— 
2M MNOSD), IDX ZW IGS) (Z))s 

Bradycellus neglectus (LeConte).—Kitt- 
SOnea COs ElallockaaINee we Nichelsone 
VII.15.1941 (1). Polk Co., Crookston, B.A. 
Haws, VII.20.1956 (1). Roseau Co., Roo- 
sevelt, A.G. Peterson, V.15.1969 (1). 

Bradycellus semipubescens Lindroth.— 
Washington Co., H.E. Milliron, V.7.1938 
@)F Crow, Wines Co™ = Millemibacsy GE: 
Mickel, V.8.1940 (1). Koochiching Co., 
T7ON, R24W, S14, H.M. Kulman, VIII.11— 
22.1971 (1). Cook Co., Superior National 
Forest, baited pitfall trap, K.J.K. Gandhi, 
48°07’ 21°°N 90°51’25“°W, P. banksiana 
wind-disturbed-salvage-logged forest, 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


VI.5—23.2003 (1). We collected a total of 
one adult. 

Bradycellus semipubescens ranges from 
Newfoundland to Alberta and southward to 
New York and Michigan (Bousquet and 
Larochelle 1993:226). The ecology of this 
species was previously unknown (Lindroth 
1961—69:899), but in recent years it is re- 
ported to be an inhabitant of marsh and 
boggy areas (Liebherr and Song 2002:134). 
In our study sites, it was caught in a sea- 
sonally flooded, wind-disturbed-salvage- 
logged conifer forest. 

Dicheirotrichus cognatus (Gyllenhal).— 
Ramsey Co., St. Paul, University Farm, 
C.T. Schmidt, VIII.27.1926 (1). Lake Co., 
roots of grass, M.H. Hatch, VI.23.1927 (1). 
Polk Co., Crookston, light trap, D.G. Den- 
ning, IX.23.1936 (1). Itasca Co., Grand 
Rapids, North Central Experimental Re- 
search Station, malaise trap, unknown col- 
lector, V.8.1973 (1). Cook Co., malaise trap, 
Ese, Hovland sexe Snlo7a aC) @layaeo- 
Moorhead, T.L. Wanless, VI.1.1997 (1, 
UCB). 

Discoderus parallelus (Haldeman).— 
Jackson Co., Wm. S. Marshall, June 1896 
(2, UWM). Crow Wing Co., Garrison, D.G. 
Denning, VI.8.1935 (16); Garrison, on 
beach, B. Armstrong, VI.8.1935 (1); C.E. 
Mickel, VI.3.1938 (2). Rock Co., Luverne, 
A.E. Pritchard, [X.13—-14.1935 (1). Mille 
Lacs Co., C.E. Mickel, VI.2.1937 (1); Mille 
acs) Wake, (ReEe  Dagey-maVile OS 5m): 
Mille Lacs, unknown collector, VI.2.1935 
(1). Traverse Co., Lake Traverse, six miles 
NE Browns Valley, B. Tollefson, 
VI1.10.1974 (1, UCB). Anoka Co., Cedar 
Creek Natural History Area, K.J. Larsen, 
VII.28.2000, VIII.29.2000 (2, KJL). 

Harpalus desertus LeConte.—Traverse 
Co., Lake Traverse, two miles north of 
Browns Valley, J.R. Powers, VI.10.1974 (1, 
UCB). 

Harpalus ellipsis LeConte.—Itasca Co., 
57, WROAS S26 Dies Raul Vile OM 7s 
(1). Carlton Co., Cloquet Forestry Center, 5 
km west of Cloquet, 46°42'25’N, 


VOLUME 107, NUMBER 4 


92°31'35"W, UMN Silviculture Class, 
WOO): 

Harpalus erythropus Dejean.—Hennepin 
Co., O.W. Oestlund (2). Olmsted Co., C.N. 
Ainshe (3). Unknown Co., unknown collec- 
tor (1, NMNH). Washington Co., Marine, 
St. Croix River, unknown collector, 
V.14.1922 (1). Carver Co., W.E. Hoffman, 
VII.16.1922 (3). Ramsey Co., St. Paul, at 
lichiee Sa i<epperley> sVllaIIeIS2 S52) st: 
Paul, University Farm, C.T. Schmidt, 
VIII.16.1926 (1). Goodhue Co., Frontenac, 
W.C. Stehr, V.29.1930 (1). Anoka Co., Ce- 
dar Creek Natural History Area, K.J. Lar- 
Semen VIE 2 0005 V IL29: 20001 2K): 

Harpalus paratus Casey.—Big Stone 
Co., O. Lugger, VII.20.1910 (1). Hennepin 
Co., O.W. Oestlund (1). Washington Co., 
Marine, St. Croix River, unknown collector, 
Wal O29 @)) RamseyaiCo:, St. ePauls vat 
light, S. Kepperley, VII.11.1925 (1). Hen- 
nepin Co., Fort Snelling, Flood Plain For- 
est, C.T. Schmidt, V.10.1928 (1); Goodhue 
Cor Caleschindiy V29NS30V@): 

Harpalus ventralis LeConte.—Crow 
Wing Co., Barrows, south of Brainerd, near 
Hwy. 371, unbaited pitfall trap, K.J.K. Gan- 
dhi, 46°24'N 94°O8'W, P. banksiana natu- 
rally burned forest, VIH.9—18.2002 (1). 

Harpalus ventralis ranges from North 
Dakota to Utah and New Mexico (Bousquet 
and Larochelle 1993: 234). It is generally 
associated with prairie habitats on sandy 
soils (Lindroth 1961—69: 781—782). 

Selenophorus ellipticus Dejean.—Anoka 
Co., Cedar Creek Natural History Area, 
Ke warsenwaleZO 2 000GE Kgs): 

Selenophorus hylacis (Say).—Ramsey 
Co., St. Paul, A.A. Granovsky, VI.25.1934 
(CD): 

Stenolophus infuscatus (Dejean).—Olm- 
sted Co., C.N. Ainslie (1). 

Stenolophus rotundicollis (Haldeman).— 
Ramsey Co., Ammunition plant, old field, 


pital, trap,  M-E. Bpstem, VII-29- 
WAIN ES): I ke{0) (CE 
Tribe Platynini 
Agonum aeruginosum Dejean.—Un- 


known Co., O. Lugger (2). Ramsey Co., 


931 


Golf Ponds, W.E. Hoffman, VII.28.1921 
(1). Houston Co., unknown collector, 
V2 29 Sia): . 

Agonum affine Kirby.—Unknown Co., 
A. Bolter (2, INHSIC). Ramsey Co., O.W. 
Oestlund (1); St. Anthony Park, at light, 
W.E. Hoffman, VI.25.1921 (1); St. Paul, In- 
dian Mounds Park, W.E. Hoffman, 
V.7.1922 (1); University Farm lights, W.E. 
Hoffman, VI.10.1922 (1); St. Paul, Univer- 
sity Farm lights, W.E. Hoffman, VI.12— 
1331922 1) Hennepin Coz) Oak ‘Grove; 
A.A. Nichol, V.13.1922 (1). Fairbault Co., 
A. Hertig, W.E. Hoffman, VI.19—20.1922 
(2). Nicollet Co., St. Peter, H.H. Holland, 
VIII.6.1922 (1). Polk Co., Crookston, light 
trap, D.G. Denning, VI.23.1931 (1). Itasca 
Co., Itasca Park, at light, C.E. Mickel, 
VII.4.1939 (1). Cook Co., Superior Nation- 
al Forest, unbaited pitfall trap, K.J.K. Gan- 
dhi, 48°08'15"N 90°51'36"W, P. banksiana 
wind-disturbed-salvage logged forest, 
VII.6-VIII.3.2000 (1). We collected a total 
of one adult. 

Agonum affine ranges throughout North 
America (Bousquet and Larochelle 1993: 
255). It is a hygrophilous species, associ- 
ated with vegetation at the margins of 
standing water (Lindroth 1961—69: 605). 

Agonum moerens Dejean.—Ramsey Co., 
St. Paul, University Farm lights, W.E. Hoff- 
man, V.25.1922 (1). 

Agonum nigriceps LeConte.—Lake Co., 
Two Harbors, M.H. Hatch, V.30.1927 (1). 

Agonum nutans (Say).—Ramsey Co., St. 
Paul, University Golf Course, A.A. Gra- 
novsky, VI.26.1936 (1). We also found 31 
specimens of A. nutans in the NDSIRC 
from Clay Co. (Bluestem State Natural 
Area) that had been collected between 
V1I.3.1996 and IX.4.1997 (Tinerella and 
Rider 2001). 

Agonum trigeminum Lindroth.—Chisago 
Coz “Center! -'Cityse SEL Vi alan 
VII.21.1972 (1). Chisago Co., Wyoming 
City, “HM. Kulmany VUES A972): 
Koochiching Co., H.M. Kulman, T69N 
R25 W'S17; VILALON972; EX.12- 1972s); 
T7ON R25W S14, VIII.10.1972 (3); T70N 


932 


R25W S32, VIII.10.1972 (1); T7ON R25W 
S33, VUI.10.1972 (1); T7ON R24W S14, 
VI.20-VIL.1.1971, VII.14.1972, VII.28.1972 


(4); T7ON R24W S14, VII.14.1972, 
WAS MODI (ClO)  UOQIN INAS STs 
Via O72 2) aaaONS RZ Weiss. 
Vile A O72) ON, IRZONVa SSS 
VII.14.1972 (1); T7ON R24W S14, 
WOO “AZie TONG, IRAN SB 
WIL SOSMIIZ (Os UGQIN IRAN SIT, 
VI.30.1972 (4); and T69N R23W S26, 


V1.30.1972 (6). Cook Co., Superior Nation- 
al Forest, baited and unbaited pitfall traps, 
K.J.K. Gandhi, 48°01'45"N 90°24'18’W, P. 
tremuloides/B. papyrifera/conifer wind-dis- 
turbed-salvage-logged forest, VI.7- 
VII.26.2001, VIIT.19-IX.15.2003 (6); 
48°06'29"N 90°50'12”W, P. banksiana 
wind-disturbed forest, X.5.2000 (1); and 
48°07’21"N  90°51'25"W, 48°08'12"N 
90°50'25"W, 48°07'21"N 90°51'25’W, P. 
banksiana wind-disturbed-salvage-logged 
forests, V.25-VI1.6.2001, VI.5-VIII.2.2001, 
VITI.25-VIII.12.2002, VIII.27-1X.14.2002, 
VIUI.27-1X.10.2003 (5). We collected a total 
of twelve adults. 

Agonum trigeminum is an eastern species 
that ranges from New Brunswick and North 
Carolina west to Michigan and Wisconsin 
(Bousquet and Larochelle 1993: 258). In 
Minnesota, this species has also been re- 
ported in Aitkin and Cass counties (Petrice 
et al. 2002: 9), and Cook county (MacLean 
2002). Agonum trigeminum 1s generally as- 
sociated with vegetation growing at the 
margins of bodies of water (Lindroth 1961— 
69: 601). 

Platynus cincticollis (Say).—Wabasha 
Co., Lake City, J.R. Powers, VI.4.1970 (1, 
WGB)): 


Tribe Ctenodactylini 
Leptotrachelus dorsalis (Fabricius).— 
Houston Co., C.E. Mickel, V.20.1938 (2). 
Tribe Cyclosomini 


Tetragonoderus fasciatus (Haldeman).— 
Unknown Co., O. Lugger (20). Unknown 
Co., unknown collector (2, SDSU). Kandi- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


yohi Co., O.W. Oestlund (1). Olmstead Co., 
C.N. Ainslie (4). Ramsey Co., University 
Farm, St. Paul, at light, A.A. Granovsky, 
V1.26.1936 (1) and H. Knutsen, [X.2.1937 
(2). Goodhue Co., C.E. Mickel, V.27.1939 
(I). 


Tribe Lebiini 


Apenes lucidulus (Dejean).—OlImsted 
Co., C.N. Ainslie (1). 

Apristus subsulcatus (Dejean).—Jackson 
Co., Wm. S. Marshall (1, UWM). Olmsted 
Conse New Amnshie@)sltasca \Comaltasea 
Park, DeSoto Lake, C.E. Mickel, VI.2.1937 
(1). Mille Lacs Co., C.E. Mickel, VI.2.1937 
(2). 

Axinopalpus biplagiatus (Dejean).— 
Washington Co., Newport, Bailey’s Nurs- 
ery, under apple bark, R.H. Daggy, 
II.2.1935 (1). Ramsey Co., St. Paul, Uni- 
versity Golf Course, light trap, A.A. Gra- 
novsky, VII.20.1936 (1). Mille Lacs Co., 
Mille Lacs Lake, R.H. Daggy, VI.2.1937 
(1). 

Calleida purpurea (Say).—Ramsey Co., 
St. Anthony Park, O. Lugger (2). Unknown 
County, O. Lugger (1). Traverse Co., O.W. 
Oestlund (1). Unknown Co., Stromberg (1, 
INHSIC). We also found 33 specimens of 
C. purpurea in the NDSIRC from Clay Co. 
(Clay Co. Trust Lands and Bicentenial Prai- 
rie) that had been collected between 
VIIL.15.1996 and VII.30.1999 (Tinerella 
2000). 

Cymindis interior Lindroth.—Anoka Co.., 
Cedar Creek Natural History Area, K.J. 
Larsen, VIII.1.2000, VIII.16.2000 (2, KJL). 

Cymindis planipennis LeConte.—St. 
Louis Co., Duluth, unknown collector (1, 
INHSIC). Hennepin Co., O.W. Oestlund 
(1). Traverse Co., O.W. Oestlund (1). Polk 
Co., D.G. Denning, VII.6.1936 (1). Anoka 
Co., C.E. Mickel, V.6.1939 (1). 

Cymindis platicollis (Say).—Olmsted 
Co., C.N. Ainslie, V.30.1905 (2). Nicollet 
Co., St. Peter, H.H. Holland, VIII.26.1922 
(1). We also found two specimens of C. pla- 
ticollis in the NDSIRC from Norman Co. 
(Agassiz Dunes State Natural Area) col- 


VOLUME 107, NUMBER 4 


lected on X.13.1997 (Tinerella and Rider 
2001). 


Tribe Galeritini 


Galerita bicolor (Drury).-Goodhue Co.., 
Cannon Falls, on sawdust, W.C. Stehr, 
IX.28.1929 (1). Houston Co., A.G. Peter- 
son, T. Knigin, I. Tarshis, E. Ivy, O. Elster, 
C. Reif, RM. Schroeder, H.E. Milliron, R.H. 
Daggy, and C.E. Mickel, R. Cottrell, 
i222 atOS Gs W239. NIB. WPA NDT 
V.20.1938, V.25—26.1940, V.26.1940 (24). 
Wabasha Co., H.E. Gustafson, V.21.1937 
(1). Winona Co., I. Tarshis, V.25.1940 (1). 


DISCUSSION 


We present new state records for 100 ca- 
rabid species from Minnesota collected by 
us or revealed in our survey of 12 institu- 
tional or personai collections (Table 2). We 
found no new state records and in some 
cases no specimens from Minnesota at the 
CDFA, MPM, UCD, and UW. The species 
that are new state records represent 20 
tribes and 43 genera, are reported to inhabit 
sub-boreal to prairie landscapes (Lindroth 
1961-69), and reflect the diversity of land- 
scapes and habitats present within Minne- 
sota (Tester 1995). In our field survey we 
collected 14 species that have never been 
reported before from Minnesota in the peer- 
reviewed literature. Nine of these species 
had been collected earlier and deposited in 
museum collections by other workers; four 
of those nine had also been recently re- 
ported in non-peer-reviewed literature 
(MacLean 2002, Petrice et al. 2002). Thir- 
teen of twenty new records reported by Ti- 
nerella (2000), Tinerella and Rider (2001), 
MacLean (2002), and Petrice et al. (2002) 
were present in the museum collections sur- 
veyed by us or were already reported in the 
literature. When our results (100 species) 
are combined with those in monographic 
works by Bousquet and Larochelle (1993) 
and Downie and Arnett (1996) (326 spe- 
cies), and with the unique collection data 
from the recent reports noted above (7 spe- 


933 


cies), there are now 76 genera and 433 ca- 
rabid species recorded from Minnesota. 

About half (45%) of the new state re- 
cords from the museum survey represent 
specimens that have not been collected 
again since 1950 (Table 2). This may reflect 
historical differences in collecting intensity 
in Minnesota vs. other states and provinces. 
For example, in a recent large-scale study 
in Iowa, Larsen et al. (2003) reported the 
collection of small numbers of four species 
(Clivinia impressefrons, Bembidion postre- 
mum, Chlaenius pusillus, and Apenes luci- 
dulus), which since 1950 have not been col- 
lected in Minnesota and deposited in col- 
lections. Chlaenius pusillus was also re- 
cently collected in Wisconsin (Purrington et 
al. 2000). Alternatively, the long duration 
since the last collection of certain species 
may be a consequence of local extinctions 
resulting from habitat alteration by humans. 
For example, the large distinctive ground 
beetle, Calosoma scrutator, has not been 
added to museum collections since 1937, 
1939, and 1956, in Minnesota, Wisconsin 
and South Dakota, respectively, and a live 
specimen has not been seen in recent years 
in these areas (G. Noonan, MPM, personal 
communication). Calosoma scrutator is 
known to be associated with open hard- 
wood forests (Lindroth 1961—69: 46), 
which have become scarce within the his- 
toric range of this beetle in southern Min- 
nesota (Fillmore, Hennepin, Houston, and 
Ramsey Counties). Local extinctions may 
have especially occurred for other ground 
beetle species associated with undisturbed, 
native habitats such as 
forests with large eastern white pines, Pinus 
strobus, or various wetland habitats that are 
threatened throughout Minnesota (Minne- 
sota Department of Natural Resources 
2005). 

A number of our newly reported carabid 
species such as Carabus sylvosus, Elaphrus 
Bembidion dorsale, B. mutatum, 


late-successional 


lecontei, 
B. postremum, B. wingatei, Poecilus scitu- 
lus, Amara torrida, Bradycellus semipubes- 
cens, Harpalus ventralis, Stenolophus ro- 


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VOLUME 107, NUMBER 4 


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VOLUME 107, NUMBER 4 


tundicollis, Agonum trigeminum, and Cy- 
mindis planipennis have been documented 
in the literature from states and provinces 
adjacent to Minnesota such as Iowa, North 
Dakota, Ontario, South Dakota, and Wis- 
consin (Bousquet and Larochelle 1993, Pur- 
rington and Larsen 1997, Purrington and 
Maxwell 1998, Purrington et al. 2000, Wer- 
ner and Raffa 2000, Larsen et al. 2003). A 
number of other newly reported species for 
Minnesota such as Notiophilus aquaticus, 
Bembidion inaequale, B. mutatum, Patro- 
bus foveocollis, P. septentrionis, Dicheiro- 
trichus cognatus, and Agonum affine have 
transcontinental distributions. Hence, it is 
not surprising that species such as these 
were eventually found in Minnesota. This 
likely also reflects an historically uneven 
effort in the collection of ground beetles 
among states. 

Our collecting in the sub-boreal forests 
of northern Minnesota has extended consid- 
erably the ranges of Sphaeroderus nitidi- 
collis brevoorti and Trechus crassiscapus 
within the United States, and also has ex- 
tended the ranges of Bembidion wingatei, 
Patrobus septentrionis, Pterostichus melan- 
arius, and Agonum trigeminum within the 
state. Previously, these latter four species 
were collected by others from southern and 
central Minnesota (records in UMIC, Petri- 
ce et al. 2002). Hence, it appears that these 
species are associated with both the central 
deciduous and northern coniferous/decidu- 
ous forest biomes. 

From our museum survey, Pterostichus 
melanarius has been present in Minnesota 
at least since 1990. It was recorded in 1993 
in Iowa (Winneshiek Co.) (Purrington and 
Larsen, 1997), in 1990 in southern Minne- 
sota (Wright Co., UMIC), and in 1995 in 
western Minnesota (Clay Co., NDSIRC). 
We suspect that it appeared in Minnesota 
even earlier because it had been document- 
ed in adjacent states in the upper Midwest 
and in Ontario and Manitoba (Lindroth 
1961-69, Bousquet and Larochelle 1993, 
Will et al. 1995). For example, P. melan- 
arius was collected as early as 1948 in Fort 


9877 


William in western Ontario (Lindroth 
1961-69), 1956 in Winnipeg in Manitoba 
(Lindroth 1961—69), and 1980 in Milwau- 
kee, Washington, and Waushara Counties in 
Wisconsin (records from MPM). In Michi- 
gan, the species had become evident in ur- 
ban and agricultural habitats to the extent 
that it was noted in the extension literature 
as a generalist predator of economically im- 
portant pests (Lee 1998). In a 1996—1997 
study on the Michigan-Wisconsin border, P. 
melanarius was the fourth most abundant 
of the 59 species of Carabidae collected in 
mixed northern hardwood forest sites (Wer- 
ner and Raffa 2000). Hladilek (2003) re- 
ported that in 2000 it occurred at approx. 
5% of all ground beetles trapped in pitfall 
traps placed in a wheat field in east-central 
Minnesota. Furthermore, we have trapped 
P. melanarius in extremely high numbers 
in the Superior National Forest, suggesting 
that a long enough time interval had tran- 
spired to allow it to colonize a relatively 
remote and northern portion of Minnesota. 

In 2001-2003, Pterostichus melanarius 
was most active in northern Minnesota be- 
tween mid-July and mid-August (Table | 
and Gandhi et al. unpublished data). This 
seasonal activity pattern is similar to that 
reported for populations of P. melanarius 
in a wheat field in east-central Minnesota 
(peaked in late June) (Hladilek 2003), the 
boreal-prairie transition forests in Canada 
(Carcamo et al. 1995, Niemela et al. 1997), 
and hemlock-northern hardwood forests in 
Michigan and Wisconsin (peaked in late 
July to mid-August) (Werner and Raffa 
2003). In northern Minnesota, P. melanar- 
ius Was most prevalent in prescribed-burned 
forest sites that had experienced previous 
wind disturbance and salvage logging (Ta- 
ble 1). This suggests that recently disturbed 
areas in the sub-boreal forests may be col- 
onized aggressively by exotic species. We 
do not know if the response of P. melan- 
arius was dictated simply by the burning 
that occurred at these sites or by the com- 
bination of the three disturbances (wind, 
salvage logging, burning). We also do not 


938 


know whether P. melanarius will displace 
other native species, e.g., the congener P. 
coracinus Newman, within this sub-boreal 
forest habitat (Werner and Raffa 2003). 

Our trapping and museum survey have 
increased the generic and species records of 
carabids in Minnesota by 21% and 31%, 
respectively. This underscores how little 
known the local historical and current dis- 
tributions of Carabidae are in some areas of 
North America. Purrington et al. (2000) 
summarized increases in known state cara- 
bid fauna of 4% (Illinois), 4% (Wisconsin), 
and 7% (lowa). Furthermore, Werner and 
Raffa (2000) reported only one new state 
record for both Michigan and Wisconsin in 
a collection of 47,590 adult carabid beetles. 
In Minnesota, we have documented an in- 
crease in the species list that is almost an 
order of magnitude greater than that of Pur- 
rington et al. (2000) and Werner and Raffa 
(2000), and this emphasizes the need for 
both further field collections and careful ex- 
amination of museum collections on a re- 
gional basis. 

We find it significant that a number of 
these genera and species, although collect- 
ed, identified, and deposited in museums in 
the earlier part of the last century, remained 
undocumented in literature. Patrobus sep- 
tentrionis, Poecilus corvus, Amara crassis- 
pina, Chlaenius purpuricollis purpuricollis, 
Harpalus paratus, Agonum nutans, Platyn- 
us cincticollis, Calleida purpurea, and Cy- 
mindis platicollis (all collected between 
1905 and 1980 and present in our museum 
survey) were recently reported to be new 
state records by Tinerella (2000: 193), Ti- 
nerella and Rider (2001: 319), MacLean 
(2002: 6), and Petrice et al. (2002: 9). Pter- 
ostichus tenuis (Casey), another new record 
reported by Petrice et al. (2002: 9), actually 
had been previously documented in Min- 
nesota by Downie and Arnett (1996: 149), 
and was also collected in our field study in 
northeastern Minnesota. Since these muse- 
um specimens were collected and deposited 
much earlier than when these studies were 
conducted, we stress the importance of ac- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


cessing the identified and unidentified ma- 
terial at local and regional museums prior 
to reporting new state species records. This 
approach will ensure an enhanced and more 
accurate understanding of species distribu- 
tions for future workers. In addition, the 
time and effort expended in collection and 
curation by previous workers will be rec- 
ognized. 


CONCLUSIONS 


We document the occurrence of 13 pre- 
viously unreported genera and 100 previ- 
ously unreported species of Carabidae in 
Minnesota, and report a major range exten- 
sion for one species in the United States 
and for four species within Minnesota. Fur- 
ther, we describe the seasonal activity and 
numerical dominance of P. melanarius in 
remote sub-boreal forest sites in northern 
Minnesota, suggesting that newly disturbed 
areas in these forests may be colonized by 
invasive species. The results of our field 
and museum surveys should stimulate and 
provide the basis for future biodiversity 
studies of carabids in Minnesota. Our study 
underscores the significant benefits of aca- 
demic insect collections that act as reposi- 
tories of distributional, ecological, and tax- 
onomic information about species present 
in native landscapes. In the future, we hope 
that researchers will take greater advantage 
of such insect collections, and that the uni- 
versities and other institutions will receive 
greater funding and resources to improve 
and maintain their collections for studies 
such as ours. 


ACKNOWLEDGMENTS 


We thank P. Anderson, B. Babcock, J. 
Donnay, L. Fruend, A. Graves, D. Kasten- 
dick, H. Krause, N. La Trace, J. McGovern, 
M. Platta, and the Minnesota Department of 
Natural Resources fire-suppression staff for 
field assistance. We are grateful to J. and 
M. Albers and R. Maki (Minnesota De- 
partment of Natural Resources), T. Mc- 
Cann, D. Neitzke, T. Norman, P. Johnson, 
and M. Theimer (USDA-Forest Service, 


VOLUME 107, NUMBER 4 


Gunflint Ranger District, Superior National 
Forest), and J. Zasada (USDA-Forest Ser- 
vice, North Central Experiment Station, re- 
tired) for assistance on the project. Museum 
assistance was received from C. Bellamy 
(California Department of Food and Agri- 
culture), R. Brett and D. Kavanaugh (Cali- 
fornia Academy of Sciences), P. Clausen 
and A. Graves (University of Minnesota), 
G. Courtney (Iowa State University), R. 
Davidson (Carnegie Museum of Natural 
History), T. Erwin and D. Furth (Smithson- 
ian Institution), C. Favret (Illinois Natural 
History Survey), G. Fauske (North Dakota 
State University), D. Dean and S. Turner 
(University of Wyoming), L. Kimsey (Uni- 
versity of California, Davis), S. Krauth 
(University of Wisconsin), G. Noonan (Mil- 
waukee Public Museum), D. Shpeley (Uni- 
versity of Alberta), J. Sakamato (John Hop- 
kins University), M. Towerton (South Da- 
kota State University), and K. Will and C. 
Barr (University of California, Berkeley). 
We also thank the Grand Portage Band and 
D. MacLean for providing a new ground 
beetle record. Funding for this project was 
provided by two grants from the Dayton 
Natural History Fund of the Bell Museum 
of Natural History, the Alexander P./Lydia 
Anderson Graduate Fellowship, a Doctoral 
Dissertation Graduate School Fellowship, 
and Doctoral Dissertation Research Award, 
University of Minnesota, and a Sigma Xi 
Grant-in-Aid of Research, Sigma Xi Re- 
search Society (all to KJKG); the Under- 
graduate Research Opportunity Program, 
University of Minnesota; the Minnesota 
Agricultural Experiment Station (Project 
MN-17-070 to SJS, Project MIN-42-034 to 
DWG); two grants from the USDA-Forest 
Service-Forest Health Protection, Evalua- 
tion Monitoring Program (01-GD- 
11244225-196, 01-DG-11244225-196) to 
DWG and SJS; a Joint Fire Science Pro- 
gram Grant (O00-2-23) to J. Zasada and 
DWG-; and the USDA-Forest Service North 
Central and the Pacific Southwest Research 
Stations. 


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PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 941—946 


LIVILLA VARIEGATA (LOW) (HEMIPTERA: STERNORRHYNCHA: 
PSYLLIDAE) NEW TO NORTH AMERICA, WITH RECORDS OF THREE 
OTHER PALEARCTIC PSYLLIDS NEW TO NEWFOUNDLAND 


A.G. WHEELER, JR. AND E. RICHARD HOEBEKE 


(AGW) Department of Entomology, Soils, and Plant Sciences, Clemson University, 
Clemson, SC 29634-0315, U.S.A. (e-mail: awhlr@clemson.edu); (ERH) Department of 
Entomology, Cornell University, Ithaca, NY 14853-0901, U.S.A. 
erh2 @cornell.edu) 


(e-mail: 


Abstract.—Livilla variegata (LOw) is reported from Newfoundland as the first North 
American record of this European psyllid. Adults were collected from Laburnum spp. 
(Fabaceae) in and near St. John’s in July 2004. The psyllid is assumed to have been 
introduced with the shipment of ornamental laburnums from Europe. A taxonomic diag- 
nosis and description are provided to facilitate its recognition in the Nearctic fauna. We 
also give the first records from Newfoundland of the Palearctic psyllids Psylla buxi (L.) 
(new Canadian record), Cacopsylla mali (Schmidberger), and C. peregrina (Forster). 


Key Words: insect detection, nonindigenous species, Cacopsylla mali, Cacopsylla per- 


egrina, Psylla buxi, new records, Laburnum 


Alien insects continue to enter North 
America and become established in our fau- 
na. Only a small proportion of non-native 
species should be considered invasive—that 
is, capable of dominating ecosystems, caus- 
ing economic or environmental damage, or 
impairing human health. Non-native insects 
(and mites), however, are estimated to cause 
40-50% of all crop losses in the United 
States (Sailer 1983). It is desirable to doc- 
ument the establishment of all exotic insects 
in North America, regardless of their pre- 
sumed economic, environmental, or medi- 
cal importance. Since 1993, we have em- 
phasized the detection of exotic insects in 
the Atlantic Provinces of Canada. 

On a recent trip to Newfoundland, we 
collected the European psyllid Livilla var- 
iegata (L6w), which is reported here as new 
to North America. We also give records 
from Newfoundland of the Palearctic psyl- 
lid Psylla buxi (L.) as the first for Canada 


and record two other Palearctic psyllids, 
Cacopsylla mali (Schmidberger) and C. 
peregrina (Forster), as new to Newfound- 
land. 


Livilla variegata (L6Ow) 


Livilla Curtis, a western Palearctic genus, 
comprises 43 species that are found mainly 
in the Mediterranean basin and feed only 
on genistoid legumes (Fabaceae), such as 
species of Chamaecytisus, Cytisus, and 
Genista (Percy 2002). 

Livilla variegata was described in the ge- 
nus Floria from Bosnia and Herzegovina 
(as Yugoslavia) (LOw 1881); F. alpina Cer- 
utti is considered a synonym (Burckhardt 
1983). The psyllid also is known in Europe 
from Austria, Czech Republic, France, Ger- 
many, Hungary, Italy, Romania, Slovakia, 
Spain, Switzerland, and United Kingdom 
(England, Scotland, Wales) (Hodkinson and 
Hollis 1987, Lauterer and Malenovsky 


942 


2002, Percy 2003, Malenovsky and Kment 
2004). It is a relatively recent addition to 
the British fauna, apparently having been 
introduced unintentionally with nursery 
stock (Hollis 1978, Hodkinson and Hollis 
1980), and was detected recently in Austria, 
Czech Republic, Germany, Hungary, and 
Slovakia (Lauterer and Malenovsky 2002, 
Malenovsky and Kment 2004). An apparent 
northward spread in Switzerland since the 
1970s has been attributed to climatic chang- 


es (global warming) (Burckhardt and Miihl-— 


ethaler 2003). Its apparently recent estab- 
lishment elsewhere in Central Europe also 
might be due to global warming (D. Burck- 
hardt, personal communication). 

Except for an association with golden 
chain, Laburnum species (Hollis 1978; 
Hodkinson and Hollis 1980, 1987; White 
and Hodkinson 1982; Burckhardt 1983; 
Burckhardt and Miihlethaler 2003; Male- 
novsky and Kment 2004), little is known 
about the life history and habits of L. var- 
iegata. In Italy, nymphs are found in April 
and May and adults from April to August; 
populations are thought to be univoltine, 
with either eggs or early instars Overwin- 
tering (Lauterer and Malenovsky 2002, 
Malenovsky and Kment 2004). White and 
Hodkinson (1982) briefly described the fifth 
instar, and Maryanska-Nadachowska et al. 
(1994) reported on the chromosomal length 
and karyotype. 

Diagnosis.—Livilla variegata is a mem- 
ber of the subfamily Arytaininae, which 
also includes the Old World legume-feeding 
genera Arytaina Forster and Arytainilla Lo- 
ginova. Both genera include a species that 
has been accidentally introduced into North 
America [1.e., Arytaina genistae (Latreille) 
and Arytainilla spartiophila (Forster) | 
(Pfeiffer 1986, Wheeler and Hoebeke 
2004a). Livilla variegata can be readily 
separated from Arytainilla spartiophila by 
the large genal cones that are at least as 
long as the vertex along the midline (genal 
cones in A. spartiophila at most 0.25 times 
the length of the vertex). Also, the forewing 
of L. variegata is oblong oval, broadest in 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


the apical third, yellowish, and without a 
distinct pattern but often with the apical 
portion suffused with gray to pale yellowish 
brown (wing of similar shape in A. spartio- 
phila but entirely pale yellow throughout). 
From Arytaina genistae, L. variegata is dif- 
ferentiated by the oblong-oval forewing 
(forewing elliptical and broadest at or be- 
fore middle in A. genistae), which lacks a 
distinct pattern (in A. genistae, the apical 
cells of the forewing with distinct, longi- 
tudinal dark brown to black pattern; Wheel- 
er and Hoebeke, 2004a: 178, fig. 1). Fur- 
thermore, Arytaina genistae and Arytainilla 
spartiophila feed on Cytisus scoparius (L.) 
Link, Scotch broom, whereas L. variegata 
is known only from Laburnum species. 

Description of adult (Fig. 1).—The fol- 
lowing description is taken from Hodkinson 
and Hollis (1987): Coloration: Mature 
specimens with dorsum of head and thorax 
orange yellow with paler longitudinal mark- 
ings; genal cones orange yellow; abdominal 
sclerites dark brown, intersegmental mem- 
branes yellow; genitalia orange yellow; 
forewing membrane clear to pale yellow 
basally, apical suffusion gray to pale yel- 
lowish brown, veins pale yellow to pale 
brown; antenna dirty yellow, apices of fla- 
gellomeres 1—4 and whole of flagellomeres 
5—8 dark brown; legs dirty yellow. Struc- 
ture: Head with genal cones slender, slight- 
ly longer than vertex, with narrowly round- 
ed apices. Forewing length: male (2.76— 
3.06 mm), female (2.82—3.59 mm). Fore- 
wing oblong oval; costal break and 
rudimentary pterostigma present; veins del- 
icate; dense fine spicules throughout all 
cells; vein Rs weakly curved to margin; 
vein M evenly curved; vein M,,, reaching 
wing apex; cell cu,, strongly arched. Meta- 
tibia with 5 thick black apical spurs; basal 
metatarsus with | black spur. Male proctig- 
er, paramere, aedeagus, and female termin- 
alia illustrated by Hodkinson and Hollis 
(1987:78, 80). 

New Nearctic record.—We first collected 
L. variegata on Laburnum species on the 
main campus of Memorial University of 


VOLUME 107, NUMBER 4 


Riga de 


Newfoundland, St. John’s, and later found 
it on ornamental laburnums at the Univer- 
sity’s Botanical Garden at St. John’s and 
other nearby localities on the Avalon Pen- 
insula. Adults were found by beating labur- 
num branches over a shallow insect net. We 
also observed adults, some of them teneral, 
mostly on midribs of lower leaf surfaces. 
Two adults were on laburnum petioles, one 
was on a petal, and another was on the up- 
per surface of a leaf. Six mating pairs and 
ten nymphal exuviae were observed on 
lower leaf surfaces. We did not find nymphs 
during our collecting (9-14 July 2004). 
Material examined.—CANADA: New- 
foundland: St. John’s, Bannerman Park, 10 
July (143 2, 9 3); Bowring Park, 13 July 
(18 2, 11 3); Memorial University of New- 


Livilla variegata adult female (above) and male (below), lateral view. Scale line = 1.0 mm. 


foundland, main campus and Botanical 
Garden (Mount Scio Rd.), 9-10 July (71 &, 
40 3); South Harbour area, Waterford Riv- 
er Walk, 10 July (11 2, 5 6); Torbay, Rt. 
20212 Julyi@seadiSae ): 

Voucher specimens of L. variegata have 
been deposited in the Canadian National 
Collection of Insects, Ottawa, Ont.; Cornell 
University Insect Collection, Ithaca, NY; 
and National Museum of Natural History, 
psyllid collection, Beltsville, MD. 


ADDITIONAL EUROPEAN PSYLLIDAE IN 
NEWFOUNDLAND 


Cacopsylla mali (Schmidberger).—Brit- 
tain’s (1919) from Nova Scotia 
were the first for this Old World psyllid in 
North America. It also has been recorded in 


records 


944 


Canada from New Brunswick (Maw et al. 
2000); an apparently overlooked record is 
Prince Edward Island (USDA 1929b). No 
published U.S. records are available for C. 
mali, but it is established in the Northeast 
(A. T. Eaton, personal communication; 
A.G.W. and E.R.H., personal observations). 
Our collections from apple (Malus pumila 
P. Mill.) and crabapple (Malus spp.) in 2004 
are the first for Newfoundland: Carbonear, 
11 July (24 &, 18 3); St. John’s, Bowring 
Park, 13 July (16 2, 13 3); Memorial Uni- 
versity of Newfoundland, 14 July (7 @, 10 
36); Torbay, Rt. 20, 12 July (24 @, 26 2). 

Cacopsylla peregrina (Forster).—A re- 
cent addition to the North American fauna, 
C. peregrina previously has been recorded 
in Canada from British Columbia (Maw et 
al. 2000) and Nova Scotia (Wheeler and 
Hoebeke 2004b). U.S. records are limited 
to California, Oregon, and Washington 
(Wheeler and Stoops 2001). The first re- 
cords from Newfoundland are based on our 
collections from hawthorn, Crataegus lae- 
vigata (Poir) DC and C. monogyna Jacq., 
in 2004: St. John’s, Bowring Park, 13 July 
(4 2, 3 3d); Harbourside Park, 9 July (14 
2, 11 3); Memorial University of New- 
foundland, 14 July (12 2, 8 3); South Har- 
bour area, Waterford River Walk, 9-10 July 
(2 teneral adults, 5th instars; not collected). 

Psylla (Asphagidella) buxi (.).—First 
reported in North America from New York 
(Riley 1890), P. buxi also is recorded from 
California, Connecticut, Iowa, New Jersey, 
Ohio, and Oregon in the checklist of Ne- 
arctic psylloids (Hodkinson 1988). Addi- 
tional state records are Delaware (Milliron 
1954), Maryland (USDA 1935), North Car- 
olina (David L. Stephan, personal commu- 
nication), Pennsylvania (USDA 1953), 
Rhode Island (USDA 1960), Virginia 
(USDA 1962), and Washington (USDA 
1929a). 

This pest of ornamental boxwood was 
not listed from Canada by Maw et al. 
(2000). Our collections from Buxus sem- 
pervirens L. in 2004 apparently are the first 
for Canada: St. John’s, Bowring Park, 13 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


July (13 2, 28 3); Memorial University of 
Newfoundland, 14 July (4 2, 6 2). 


DISCUSSION 


Livilla variegata is the only species of 
the genus known from laburnums. The 
most speciose genistoid legumes (30 or 
more species) tend to have the greatest 
number of associated arytainine psyllids 
(Percy 2002). Laburnum is a small genus 
of only three or four species found in the 
Mediterranean region and adjacent Asia 
(Everett 1981). 

Like other psyllids that develop on gen- 
istoid legumes (Percy 2002, 2003), L. var- 
iegata tolerates the quinolizidine alkaloids 
of its hosts. It might sequester alkaloids as 
a defense against generalist predators, as is 
known for a laburnum-feeding aphid (Szen- 
tesi and Wink 1991). 

The shipment and planting of ornamental 
laburnums beyond their native range appear 
to be responsible for the addition of L. var- 
iegata to the British fauna (Hollis 1978, 
Hodkinson and Hollis 1980, White and 
Hodkinson 1982). Hodkinson and Hollis 
(1980) commented that in southern England 
the introduced L. variegata was spreading 
rapidly and becoming increasingly com- 
mon, suggesting a considerable period be- 
tween its establishment and date of first col- 
lection (May 1978, in Hayes, Middlesex). 
When Hollis (1978) first reported it from 
Britain, records were available not only 
from additional localities in Middlesex but 
also from London, Kent, Surrey, Oxford- 
shire, and Wales. 

Evidence points to a similar origin and 
mode of entry for L. variegata in North 
America: Europe (either the British Isles or 
the continent) via shipment of laburnum 
nursery stock. The psyllid belongs to an 
Old World genus and develops on non-na- 
tive plants, Laburnum spp. Planted as or- 
namentals in Europe are common golden 
chain, L. anagyroides Medik.; Scotch la- 
burnum, L. alpinum (Miller) Bercht. and J. 
Presl.; and their hybrid, L. x watereri 
(Kirchner) Dippel (as L. vossii) (Scheller 


VOLUME 107, NUMBER 4 


1974). Laburnums were introduced into the 
United States as early as the eighteenth cen- 
tury (Leighton 1976), but they are used less 
frequently as ornamentals in North America 
than in Europe (Everett 1981). Laburnums, 
however, are common in gardens, parks, 
and yards in St. John’s, Newfoundland, 
where L. alpinum, L. anagyroides, and L. 
X watereri are planted (Peter Scott, person- 
al communication). 

Livilla variegata might have become es- 
tablished many years before we detected it 
in 2004. The Canadian Psylloidea are poor- 
ly known, and only three native species pre- 
viously have been recorded from New- 
foundland (2 spp.) and Labrador (1 sp.) 
(Maw et al. 2000). Moreover, we did not 
observe feeding symptoms on foliage, 
which might attract attention from growers, 
pest-control specialists, or entomologists. 

With the addition of four unintentionally 
introduced Palearctic species, the majority 
of Psylloidea known from Newfoundland, 
excluding Labrador, are adventive (67%). 
All four species can be added to the exten- 
sive list of European insects recorded from 
the port city of St. John’s. European insects 
in Newfoundland tend to be concentrated 
on the Avalon Peninsula and often are re- 
stricted to the St. John’s area (Lindroth 
1957). Hamilton and Langor (1987) pointed 
out that Newfoundland has the world’s larg- 
est proportion of imported leafhopper spe- 
cies and that St. John’s has the largest num- 
ber of immigrant leafhoppers. A high pro- 
portion of immigrant species also charac- 
terizes the fauna of certain other insect 
groups in Newfoundland (e.g., Lindroth 
1957, Morris 1983). 


ACKNOWLEDGMENTS 


We thank Murray H. Colbo (Department 
of Biology, Memorial University of New- 
foundland [MUN]) for his help and hospi- 
tality, Peter J. Scott (Department of Biolo- 
gy, MUN) for comments on laburnums 
grown in the St. John’s area, Alan T. Eaton 


(Department of Plant Biology, University of 


New Hampshire, Durham) for allowing us 


945 


to refer to his unpublished records of C. 
mali in northeastern states, David L. Ste- 
phan (Department of Entomology, North 
Carolina State University, Raleigh) for in- 
formation on the occurrence of P. buxi in 
North Carolina, Cecil L. Smith (Department 
of Entomology, University of Georgia, Ath- 
ens) for checking the UGA collection for 
specimens of P. buxi, Gary L. Miller (Sys- 
tematic Entomology Laboratory, ARS, 
USDA, Beltsville, MD) for confirming the 
identification of L. variegata and providing 
a copy of a reference, Peter H. Adler (De- 
partment of Entomology, Soils, and Plant 
Sciences, Clemson University) and Daniel 
Burckhardt (Naturhistorisches Museum, 
Basel, Switzerland) for suggestions that im- 
proved an earlier draft of the manuscript, 
and Kent Loeffler (Department of Plant Pa- 
thology, Cornell University) for photo- 
graphing L. variegata. 

This research was supported by the Cor- 
nell University Agricultural Experiment 
Station federal formula funds, Project No. 
NYC-139413 to ERH, received from Co- 
operative State Research, Education, and 
Extension Service, U.S. Department of Ag- 
riculture. Any opinions, findings, conclu- 
sions, or recommendations expressed in this 
publication are those of the authors and do 
not necessarily reflect the view of the U.S. 
Department of Agriculture. 


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cha: Psyllidae). Proceedings of the Entomological 
Society of Washington 106: 176-180. 

. 2004b. New records of Palearctic Hemiptera 
(Sternorrhyncha, Cicadomorpha, Heteroptera) in 
the Canadian Maritime Provinces. Proceedings of 
the Entomological Society of Washington 106: 
298-304. 

Wheeler, A. G., Jr. and C. A. Stoops. 2001. Cacopsylla 
peregrina (Foerster) (Sternorrhyncha: Psylloidea: 
Psyllidae): first U.S. records of an Old World spe- 
cialist on hawthorns (Crataegus spp.). Proceed- 
ings of the Entomological Society of Washington 
103: 103-109. 

White, I. M. and I. D. Hodkinson. 1982. Psylloidea 
(nymphal stages) Hemiptera, Homoptera. Hand- 
books for the Identification of British Insects. Vol. 
II, 5(b). Royal Entomological Society of London, 
London. 50 pp. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 947-959 


FIRST RECORDS OF ALIEN INSECTS IN CONNECTICUT 
(ORTHOPTERA: TETTIGONIIDAE; COLEOPTERA: BUPRESTIDAE, 
CHRYSOMELIDAE; DIPTERA: RHAGIONIDAE, TEPHRITIDAE; 
HYMENOPTERA: MEGACHILIDAE) 


CHRIS T. MAIER 


Department of Entomology, Connecticut Agricultural Experiment Station, P.O. Box 
1106, New Haven, CT 06504, U.S.A. (e-mail: Chris.Maier@ po.state.ct.us) 


Abstract.—Entomologists, nursery inspectors, and others have repeatedly discovered 
exotic insects new to Connecticut. Here, I report the first captures and distribution of 
eight non-native species in Connecticut: Meconema thalassinum (De Geer) (Orthoptera: 
Tettigoniidae), Agrilus cyanescens (Ratzeburg) (Coleoptera: Buprestidae), Lilioceris lilit 
(Scopoli), Pyrrhalta viburni (Paykull) (Coleoptera: Chrysomelidae), Rhagio tringarius 
(L.) (Diptera: Rhagionidae), Rhagoletis meigenii (Loew) (Diptera: Tephritidae), Anthi- 
dium manicatum (L.), and Megachile sculpturalis Smith (Hymenoptera: Megachilidae). 
Based on the weekly capture of adults in a Gressitt and Gressitt Malaise trap in northern 
Connecticut in 2003, Rhagio tringarius flew from mid-July to mid-September, although 
adults were collected earlier and later elsewhere in the state. In the same trap samples, 
females of Rhagoletis meigenii were captured mainly in July, with a few taken in August. 
In the laboratory, adults of Rhagoletis meigenii were reared from fruits of Berberis 
thunbergii De Condolle (Berberidaceae) sampled in Connecticut and New York ‘in the 
fall of 2003, confirming that this species develops in congeneric hosts in Europe and 
North America. 


distribution, exotic insects, Meconema thalassinum, Agrilus cyanescens, Lil- 
ioceris lilii, Pyrrhalta viburni, Rhagio tringarius, Rhagoletis meigenti, An- 
thidium manicatum, Megachile sculpturalis 


Key Words: 


Connecticut has a long history of coping 
with pestiferous insects accidentally intro- 
duced from foreign countries. Factors that 
have favored the arrival and establishment 
of exotic insects in the state include the 
presence of ports-of-entry, importation of 
varied products from around the world, a 
well-traveled populace, and a climate mod- 
erated by the ocean in coastal areas. During 
the past decade, entomologists, nursery in- 
spectors, and others have detected several 
potentially important orchard or forest pests 
in Connecticut, including the small Japa- 
nese cedar longhorned beetle, Callidiellum 


rufipenne (Motschulsky) (Cerambycidae) 
(Maier and Lemmon 2000, Maier 2001); 
the spruce needleminer, Batrachedra pini- 
colella (Zeller) (Batrachedridae) (Maier 
2005a), the apple tortrix, Archips fuscocu- 
preanus (Walsingham) (Tortricidae) (Maier 
and Mastro 1998, Maier 2003); and the 
green pug, Pasiphila rectangulata (L.) 
(Geometridae) (Maier 2005b). Here, I re- 
port additional alien insects detected during 
other surveys in the state. Many of these 
foreign species should be targeted for future 
surveys to determine their economic status 


in North America. 


948 


MATERIALS AND METHODS 


The following information is given for 
adults of exotic insects collected or reared 
in Connecticut: county, town (municipali- 
ty), precise location (if available), date(s) of 
capture, number of specimens (or number 
of each sex for Tettigoniidae, Rhagionidae, 
Tephritidae, and Megachilidae) in parenthe- 
ses, collecting method if not netted or cap- 
tured by hand, and the location of vouchers 
that are not deposited at the Connecticut 
Agricultural Experiment Station (CAES). 
Generally, the collection data are organized 
alphabetically by county, then by town, and 
lastly by year of capture. Information that 
did not appear on labels, but was added for 
clarity, is given in brackets. All dates listed 
under new records are given in the same 
format, with the month in lower case roman 
numerals (e.g., 17.vi.2004). To save space, 
the following abbreviations are used for 
trapping methods mentioned more than 
once: AMB, unbaited yellow Pherocon® ap- 
ple maggot, Rhagoletis pomonella (Walsh), 
(Trécé Inc., Adair, OK 74330) sticky trap 
on Japanese barberry, Berberis thunbergii 
De Condolle; GGM, Gressitt and Gressitt 
(1962) Malaise trap (John W. Hock Co., 
Gainesville, FL 32606) in a forest; GGMO, 
Gressitt and Gressitt Malaise trap at border 
of an apple, Malus domestica Borkhausen, 
orchard and a deciduous forest; RSA, red 
sticky trap (Great Lakes IPM, Vestaburg, 
MI 48891) on the trunk of live white ash, 
Fraxinus americana L.; SBA, 46-cm wide 
sticky band on girdled trunk of white ash 
in forest; SC, Sante canopy trap (Sante 
Traps, Lexington, KY 40502) at ground 
level in eastern white pine, Pinus strobus 
L.-white ash forest; SCS, Sante canopy trap 
at ground level in swamp with deciduous 
trees and shrubs; and SM, Sante Malaise 
trap (Sante Traps) in an old eastern white 
pine-spruce, Picea sp., plantation. 

Maier (1984) and Maier and Webb 
(1987) discussed the use of Gressitt and 
Gressitt Malaise traps that were used to 
capture insects between 1980 and 1985. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Other Gressitt and Gressitt Malaise traps 
operated for lengthy periods and checked 
once or twice weekly were situated at the 
border of an apple orchard and deciduous 
forest in Guilford, New Haven Co., from 
May to October 1995-1997 and in Meri- 
den, New Haven Co., over the same months 
in 1996. In 2003, a Gressitt and Gressitt 
Malaise trap placed in a calcareous swamp 
in Canaan, Litchfield Co., was checked 
weekly from May to September to deter- 
mine the flight period of the two exotic flies 
mentioned herein. The two kinds of Sante 
traps and the sticky bands on girdled ash 
trees were checked weekly from early April 
to late October 2004 during a survey for the 
exotic emerald ash borer, Agrilus planipen- 
nis Fairmaire. In the same survey, red 
sticky traps were inspected every 7—10 days 
between May and September 2004. All of 
the sticky bands and colored traps were 
coated with Tangle-trap insect trap coating 
(The Tanglefoot Co., Grand Rapids, MI 
49504). 

In addition to sampling with traps, I ex- 
amined insect collections in the Department 
of Entomology, Connecticut Agricultural 
Experiment Station, New Haven; in the 
Yale Peabody Museum, New Haven 
(YPM); and in the University of Connecti- 
cut, Storrs (UCONN). One specimen cited 
here was donated to the Illinois Natural 
History Survey, Champaign, Illinois 
(INHS). Some records are based on speci- 
mens in the private collections of Chris T. 
Maier (CTM), Guilford, and Michael C. 
Thomas (MCT), Cromwell, Connecticut. 

To confirm that larvae of North Ameri- 
can populations of Rhagoletis meigenii 
(Loew) develop in fruits of Berberis, 200 
fruits of B. thunbergii were sampled in Ca- 
naan, Connecticut, on i0 September 2003, 
and 500 fruits of the same barberry were 
collected in Highlands, Orange Co., New 
York, on 11 September 2003. In the labo- 
ratory, barberries from each sample were 
placed on a screen over a container with 
moist sand and vermiculite (1:1) for one 
month to allow larvae to emerge from 


VOLUME 107, NUMBER 4 


fruits and to form puparia in the mixture 
below. Then, the mixture with puparia was 
sealed in a plastic container and refriger- 
ated at 4 + 1°C for five months; afterward, 
the container was returned to the labora- 
tory where adult flies emerged within two 
months. 


RESULTS AND DISCUSSION 
Meconema thalassinum (De Geer) 
(Orthoptera: Tettigoniidae) 


This European tettigoniid, commonly 
known as the oak-bush cricket, has been in 
North America since at least 1957 when it 
was discovered on Long Island, New York 
(Gurney 1960a, 1960b). Since then, John- 
stone (1970), Sismondo (1978), Smith 
(1979), and Hoebeke (1981) have reported 
M. thalassinum in other parts of New York 
or in Rhode Island. This nocturnal tettigo- 
niid eats other small insects (Marshall and 
Haes 1990). 

In a mesic deciduous forest in Hamden, 
New Haven Co., I collected the first two 
specimens (both males) of M. thalassinum 
in a Gressitt and Gressitt Malaise trap in 
1980. Subsequently, I have captured adults 
of M. thalassinum in seven of eight Con- 
necticut counties, indicating that it is wide- 
spread (Fig. 1). Sticky bands or traps on the 
trunks of upright trees captured 21 of 34 
(61.8%) specimens, and flight interception 
traps caught 9 (26.5%). Adults were cap- 
tured between 10 July and 9 September. 

New state records.—Fairfield Co., Fair- 
field, near jct. Hoyden Hill Road and Hoy- 
dens Lane, 10—16.vii.2004 (4 ¢), SBA. 
Hartford Co., Farmington, | km W jet. 
State Route 4 and Hawley Road, 23—29.vii 
CD) 30a evi (lS). and» 6— 
12.viii.2004 (2 2), SBA. Litchfield Co., 
Barkhamsted, 0.8 km NNE jet. State Routes 
219 and 318, 13—-19.viii (2 2) and 27.viii— 
2.ix.2004 (1 2), SBA; Litchfield, 0.8 km 
NNE jet. State Route 118 and E. Litchfield 
Road No. 2, 10—16.viii.2004 (1 6), RSA; 
Torrington, Drakeville, State Route 272 by 
Stillwater Pond, 10—16.viii.2004 (1 3), 


949 


RSA. Middlesex Co., Cromwell, State 
Route 372, near 75 Cromwell Executive 
Ctr, 5—11.viii.2004 (1 3), RSA; Middle- 
town, near Dooley Pond, 27—28.viii.2002 (1 
2), Sante canopy trap among eastern red 
cedars, Juniperus virginiana L., in shrubby 
area on dry hillside, CTM. New Haven Co., 
Guilford, 4 km NNW jet. State Routes 77 
and 80, near Beaver Head Corner, 7.1x.2001 
(1 ¢), GGM, CTM, 3-9.ix.2004 (1 2), 
SBA; Guilford, 4 km NNW jet. State 
Routes 77 and 80, edge of Beaver Head 
Swamp, 29.vii.2002 (1 2), GGM; Guilford, 
4.5 km NW jet. State Routes 77 and 80, 
890 Beaver Head Road, 21.viii.2004 (1 ¢): 
Hamden, | km NW jet. State Routes 10 and 
40, Lockwood Farm, 18—19.vii.1980 (2 ¢), 
GGM, CTM [first state record], 
14.vii.2004 (1 6, 1 2); Hamden, 0.7 km 
WSW jet. State Routes 10 and 22, 25.vii— 
2.viil.2004 (1 3), SBA; Naugatuck, 0.3 km 
S jct. Hunters Mountain and Old Derby 
Roads, 14—20.vii (2 ¢) and 10—16.viii.2004 
(1 3) , SBA; New Haven, exit 8 (Middle- 
town Avenue) of Interstate Highway 91, 
16—22.vii.2004 (1 6), RSA; Wallingford, 
Wharton Brook State Park, 27—29.vii (1 3), 
30.vii (1 6, 1 2), and 9—11.viii.2002 (1 3), 
GGM, CTM. New London Co., Waterford, 
Civic Triangle, 4—10.viii.2004 (1 2), RSA. 
Windham Co., Ashford, Natchaug St. For- 
est, 0.3 km W jet. Perry Hill Road and Lau- 
rel Lane; 20.viii.2004 (1 2), CTM. 


Agrilus cyanescens (Ratzeburg) 
(Coleoptera: Buprestidae) 


Records, mostly under the synonym 
Agrilus coeruleus (Rossi), reported by Frost 
(1922), Fisher (1928), Wellso et al. (1976), 
Bright (1987), Davies (1991), and Sikes 
(2004) indicate that this European buprestid 
occurs in Ontario, Quebec, Wisconsin, 
Michigan, Massachusetts, and Rhode Is- 
land. Frost (1922) mentioned that the ear- 
liest North American specimen was taken 
in Sturgeon Bay, Wisconsin, in July 1920. 

I captured the first Connecticut specimen 
on 12 June 2003, although I had seen adults 


950 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Agrilus cyanescens 


Pyrrhalta viburni 


Rhagio tringarius 


Megachile sculpturalis 


Anthidium manicatum 


Fig. 1. Distribution of Meconema thalassinum, Agrilus cyanescens, Lilioceris lilii, Pyrrhalta viburni, Rhagio 
tringarius, Rhagoletis meigenii, Anthidium manicatum, and Megachile sculpturalis in Connecticut. The distri- 
butional map for L. Jilii also shows the reported occurrence in southwestern Connecticut (Fairfield Co.). 


VOLUME 107, NUMBER 4 


of A. cyanescens at least two years earlier 
during a local bioblitz. More adults were 
collected at other locations during 2003 and 
2004 (Fig. 1). I suspect more specimens, 
perhaps earlier ones than those documented 
here, will be found in out-of-state collec- 
tions that have yet to be examined. In Con- 
necticut, capture dates ranged from 10 May 
to 8 July. In my collection, I also have spec- 
imens that were captured on the foliage of 
honeysuckle, Lonicera sp. (Caprifoliaceae) 
in Mercer Co., Pennsylvania (two adults), 
and in York Co., Maine (three adults), dur- 
ing 2003. 

In Europe, larvae of A. cyanescens de- 
velop in galleries under the bark of L. nigra 
L. and other Lonicera spp. (Bright 1987, 
Rejzek 2001). Literature summarized by 
Bright (1987) and Rejzek (2001) gives 14 
other genera as hosts, but, until these rec- 
ords are confirmed, they must be considered 
dubious. Although I have observed adults 
of A. cyanescens feeding on the foliage of 
Lonicera spp., larval hosts have yet to be 
identified definitively in North America. 

New state records.—Fairfield Co., West- 
port, near jct. State Routes 33 and 136, 
10.v.2004 (1); Wilton, jct. U.S. Highway 7 
and State Route 33, near Norwalk River, 
8.vil.2003 (3), CIM. Hartford Co., Berlin, 
Shuttle Meadow Avenue, 8.vi.2004 (2), 
CTM. Hartford Co., Farmington, | km W 
jct. State Route 4 and Hawley Road, 
10.vi.2004 (1). Litchfield Co., Canaan, 1 
km NNE jet. Belden and Sand Roads, Rob- 
bins Swamp, 12.vi.2003 (3), on foliage of 
Lonicera morrowii Gray [first state rec- 
ord]. New London Co., Preston, 0.5 km 
NW jet. State Routes 2A and 12, 17.v.2004 
(1), CTM. Tolland Co., Willington, 1.2 km 
SW exit 70 (State Route 32) of Interstate 
Highway 84, 25.v.2004 (2), CTM. Wind- 
ham Co., Killingly, exit 91 (U.S. Highway 
6) of Interstate Highway 395, 25.v.2004 
(1), CTM; Windham, Willimantic, | km 
SW jct. U.S. Highway 6 and State Route 
32, 25.v.2004 (1), CTM. 


Lilioceris lilii (Scopoli) 
(Coleoptera: Chrysomelidae) 


In North America, the lily leaf beetle, L. 
lilii, was found first in Montreal, Canada, 
in 1945 (Brown 1946, LeSage 1983). The 
first find in the United States was at Boston, 
Massachusetts (Livingston 1996). In Con- 
necticut, this chrysomelid was first reported 
from Fairfield Co. (Ellis 2001), but this re- 
cord has not been published formally, and 
specimens have been discarded. This Eur- 
asian beetle apparently now occurs in all 
New England States, in northern New York 
(University of Rhode Island Plant Sciences 
Department 2002), and in Ontario and Que- 
bec (LeSage 1991). Livingston (1996) and 
Haye and Kenis (2004) summarized the life 
history, noting that larvae and adults mainly 
feed on Lilium spp. and Fritillaria spp. (Lil- 
jaceae). 

An adult specimen of L. /ilii was taken 
on an unidentified lily in Simsbury, Hart- 
ford Co., on 4 August 2001. This beetle 
represents the first state record based on an 
existing voucher. In addition to the: 2001 
specimen, a few adults have been collected 
at other Connecticut localities (Fig. 1). In 
2004, I observed more than 50 larvae feed- 
ing on unidentified lilies growing at The 
Arnold Arboretum, Jamaica Plain, Suffolk 
Co., Massachusetts, and later collected an 
adult. 

New state records.—Hartford Co., Sims- 
bury, 5 Gretel Lane, 4.vii.2001 (1), on un- 
identified lily [first state record with 
voucher]; Simsbury, Simscroft Road, 
28.vii.2003 (1), on unidentified lily. New 
Haven Co., East Haven, 25.iv.2004 (3), on 
unidentified lily, 1.v.2004 (2), on unidenti- 
fied Asian lily. Windham Co., Thompson, 
Quaddick Road, 21.vii.2004 (15), feeding 
on unidentified Asian lily. 


Pyrrhalta viburni (Paykull) 
(Coleoptera: Chrysomelidae ) 


In North America, Becker (1979) docu- 
mented (with specimens) the first reproduc- 
ing population of the Eurasian viburnum 


952 


leaf beetle, P. viburni, at Ottawa, Ontario, 
and Hull, Quebec, where beetles were as- 
sociated with Viburnum spp. (Caprifoli- 
aceae) in 1978. Later, Wheeler and Hoe- 
beke (1994) collected P. viburni in Nova 
Scotia, and they also reported it from Brit- 
ish Columbia. In the United States, Weston 
and Hoebeke (2003) noted that it was de- 
tected first in Maine in 1994, and subse- 
quently in New York, Pennsylvania, Ohio, 
and Vermont. Becker (1979) and Weston 
and Desurmont (2002) discussed its host 
preferences among Viburnum spp. grown in 
North America. Both larvae and adults feed 
on the foliage of viburnum. 

My 2004 records (Fig. 1) represented 
beetles found on four cultivars of three spe- 
cies of Viburnum. These nursery plants 
were imported from New York, a beetle 
stronghold (Weston and Hoebeke 2003). 
Sampling in subsequent years will deter- 
mine whether the accidental importation of 
P. viburni has produced an established pop- 
ulation. 

New state records.—Middlesex Co., 
Cromwell, Goodrich Heights, 21.vii.2004 
(3), on foliage of container-grown Vibur- 
num opulus L. “Nanum’ imported from 
New York. New Haven Co., Meriden, Yale 
Avenue, 8.vii.2004, on foliage of container- 
grown Viburnum lantana L. ‘Mohegan’ (2), 
V. opulus L. “Roseum’ (2), and V. trilobum 
Marshall ‘Wentworth’ (1) imported from 
New York [first state records]. Many bee- 
tles were observed at this last site, but only 
five were collected. 


Rhagio tringarius (L.) 
(Diptera: Rhagionidae) 


Chillcott (1965) first reported the Euro- 
pean R. tringarius, as well as R. lineola F, 
from eastern North America. At the time of 
his report, R. tringarius was known only 
from nine specimens collected by J. R. 
Vockeroth at Lockeport, Nova Scotia, in 
July 1958. Pechuman and Hoebeke (1983) 
later found a 1980 specimen from New 
York and a 1982 specimen from Massachu- 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


setts. Two other European species, Rhagio 
scolopaceus (L.) (Thompson 1969) and R. 
strigosus Meigen (Pechuman and Hoebeke 
1983), also now inhabit eastern North 
America. Rhagio scolopaceus still has a vi- 
able population at The Arnold Arboretum, 
Jamaica Plain, Suffolk Co., Massachusetts, 
where I collected three males on 28 May 
1999 and one female on 2 July 2004. Chill- 
cott (1965) and others have suggested that 
the soil-inhabiting larvae of exotic rhagion- 
ids reached this country in the soil of im- 
ported plants. 

In the Department of Entomology, 
CAES, there is a specimen of R. tringarius 
taken in Norfolk, Litchfield Co., Connecti- 
cut, on 13 July 1915 by M. P. Zappe. This 
specimen represents the earliest collection 
in North America. This record, predating by 
43 years the previous “‘first record’? (Chill- 
cott 1965), suggests the possibility of other 
overlooked records in small, but very old, 
collections in the Northeast. Maier and 
Webb (1987) previously reported 16 species 
of rhagionids from Connecticut; the addi- 
tion of R. tringarius brings the total to 17. 

Rhagio tringarius is widespread in Con- 
necticut (Fig. 1), where it has been partic- 
ularly abundant since 1990. Between 1980 
and 2004, I collected 270 specimens of R. 
tringarius, and only 7 of R. vertebratus and 
R. hirtus (two similar Rhagio spp. without 
distinct maculations in wings) in Connect- 
icut; thus, among these three flies, R. trin- 
garius accounted for 97.5% of all speci- 
mens. At Lockwood Farm of CAES (Ham- 
den, New Haven Co.), a Gressitt and Gres- 
sitt Malaise trap operated in a mesic 
deciduous forest from April to September 
1980-1983 caught no adults of R. hirtus, R. 
tringarius, or R. vertebratus, and another at 
Beaver Head Swamp (Guilford) checked 
during the same months of 1983-1985 cap- 
tured only one R. hirtus (Maier and Webb 
1987). A series of Malaise traps operated at 
the border of a sprayed apple orchard and 
deciduous forest in Guilford from May to 
September 1995—1997, and one operated at 
a field-orchard border in Meriden during 


VOLUME 107, NUMBER 4 


8 


No. Adults 
aw 


a) 
July 


Bon: 
2003. 


the same months in 1996, caught 63 adults 
of R. tringarius and none of the other large 
rhagionid species with unpatterned wings. 
Furthermore, in 2003, a Gressitt and Gres- 
sitt Malaise trap placed in a calcareous 
swamp (Canaan, Litchfield Co.) from late 
May to September intercepted 46 adults of 
only R. tringarius, with the flight concen- 
trated in July and August for both sexes 
(Fig. 2). Although many adults of R. trin- 
garius were caught on sticky bands on ash 
trees in 2004, their seasonal abundance was 
not plotted because many were eaten by 
slugs. In Connecticut, the earliest capture 
was between 28 May and 3 June (sticky 
band on white ash) and the latest was be- 
tween 23 and 25 September (Gressitt and 
Gressitt Malaise trap). 

In addition to Connecticut records, I col- 
lected one female of R. tringarius in Ken- 
nebunk, York Co., Maine, on 3 July 2003, 
and one female in Manchester, Bennington 
Co., Vermont, on 21 July 2004. 

New state records.—Fairfield Co., 
Greenwich, Orchard St., 16.vi.1979 (1 2), 
uv light, YPM; Redding, Huckleberry 


Is 28) 30 © 


/ \ Females 


\ 
Ca 
a 


iSy920 2:27 a SrON), Ai 4 


August September 


Seasonal abundance of Rhagio tringarius in a calcareous swamp in northern Connecticut during 


Swamp, 7.vii.1990 (1 3d), 2.viii.1991 (1 Q), 
CTM. Hartford Co., Farmington, Rattle- 
snake Mountain, 18.vi.2004 (1 36), CTM: 
Farmington, | km W State Route 4 and 
Hawley Road, 4—10.vi (1 @), 11—17.vi (2 
3), 18-24.vi (4 6, 2 Q), 25.vi-l.vii (4 6, 
UD) savin (Crs 2), Salant Gl tei. 
5 @), 16-22.vii (4 2), 23-29.vii (1 2), and 
30.vii—5.viii.2004 (1 2), SBA; Hartford, 
Keney Park, 12.vi.1999 (1 2), CTM; New- 
ington, Rt. 176, Roaring Brook Nature Cen- 
ter, 19.vii.1961 (1 2), UCONN; Southing- 
ton, 3 km ESE jet. Interstate Hwy. 84 and 
State Route 10, Lewis Farms, 14.vi.2000 (3 
3,2 2), CTM: South Windsor, [Route] 291 
site, 5.vi.2001 (1 2), UCONN; West Hart- 
ford, Albany Ave., 16.vii.1973 (1 ¢); West 
Hartford, Linbrook Road, nr. Trout Brook, 
16.vii.1973 (1 2), UCONN; [West Hart- 
ford], Hartford Reservoir No. 6, 7.vii.1966 
(1 3). Litchfield Co., Barkhamsted, 0.8 km 
NNE jet. State Routes 219 and 318, 28.v— 
3.vi (2 9), 16-22.vii (1 36, 1 2), 30.vii- 
5.vili (1 2), 6-12.viii (1 2), 20-26.vili (1 
2), 27.viii—2.ix.2004 (1 2), SBA; Canaan, 
| km NNE jet. Belden and Sand Roads, 


954 


Robbins Swamp (see Fig. 2 for seasonal 
distribution of specimens caught in Malaise 
trap); Canaan, 0.2 km N jet. State Route 
126 and Page Road, 19.vi.2003 (1 ¢); Nor- 
folk, 13.vii.1915 (1 &) [first North Amer- 
ican record]; Norfolk, 0.6 km W Westside 
and Windrow Roads, 18—20.viu.2003 (1 
2); Salisbury, near Beeslick Pond, 8— 
10.viii.1989 (2 od), Gressitt and Gressitt 
Malaise trap in stand of eastern red cedar 
near shrubby calcareous fen, CTM; Salis- 
bury, W side of Washinee Lake, 6.viii.1998 
(1 2), Gressitt and Gressitt Malaise trap in 
calcareous fen, CTM; [Salisbury], Twin 
Lakes, M. A. White prop., 8/9.vii.1994 (1 
6), UCONN; Sharon, 4 km N Cornwall 
Bridge, 27—31.viii.1984 (1 ¢), GGM, 
CTM. Middlesex Co., Middlefield, 0.3 km 
W South Street and Powder Hill Road, Ly- 
man Farm, 18.vi.2001 (1 6, 2 @), CTM; 
Middletown, near Dooley Pond, 9.vi.1999 
Cd 6). New Haven Co., Branford, 
30.vii.1985 (1 2); Branford, U.S. Highway 
PP Eni opmOrchard ye l2svielO Oia Gacy). 
CTM; Guilford, 3 km S jct. State Routes 77 
and 80, Bishop’s Orchard, 4—6 August (1 
2), 11-13.viii (1 2), 14-15.vii (1 2), and 
16-17.viii.1995 (1 @), 2-4.vii (1 3, 2 Q), 
5-8.vii (1 6, 4 2), 12-15.vii C1 3d), 16- 
18.vii (1 3), 23-25.vii (2 2), 26—29.vii (1 
2), 30.vii-1.viii (4 2), 9-12.viii 1 2), 13- 
15.vili (2 2), and 23-26.viii.1996 (1 Q), 
17-19.vi (1 do), 1-3.vii A @), 4-7.vii 
2), 8-10.vii BY), 22-24.vii (2 2), 29- 
31.vii (1 3), 5-7.viii (1 @), 15-18.vini C1 
2), 29.viii-l.ix (1 @), and 23—25.ix.1997 
(1 5), GGMO; Guilford, 4 km NNW jet. 
State Routes 77 and 80, edge of Beaver 
Head Swamp, 31.vii.2002 (1 2), CTM; 
Guilford, 4.5 km NW jet. State Routes 77 
and 80, 890 Beaver Head Road, 14.v1i.2003 
(1 2), INHS; Hamden, 1 km NW jet. State 
Routes 10 and 40, Lockwood Farm, 14.vi 
QZ) aul Swit IOS © 6, 7 2), BAM MONS 
(1 6, 1 2), CTM; Meriden, 1 km NNE jet. 
Thorpe Ave. and Fleming Rd., High Hill 
Orchard, 2—4.vii G 2), 5—8.vii (1 6, 1 @), 
OF llevan @Ss, 3 2) lo=hsivan Gis 12): 
19-22.vii (1 2), 23-25.vii (1 @), 30.vii— 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Ix (2 2) Vsvmir (@ Y), Isls waitin Cl 
Oe 23-2 Ovi (2) 2) 2 — 29 Svan Ga) 
30.vili—2.1x 2 2), 3—S.ix (1 2), 6—9.1x C1 
©), and) 13=161x. 1996) (1 2) §GGMOo; 
19.vi.1996 (1 2); North Branford, Totoket 
Mtn., 1.3 km NNE jet. State Routes 22N 
(Forest Road) and 80, 11.vi (1 6), 12—14.vi 
(iS) andl 25=26v1 200272 52) GGVE 
CTM; North Branford, 0.9 km NNW jet. 
State Route 80 and Great Hill Road, 18— 
24.vi (6 6,5 @), 25.vi-1.vii (10 2), 2-8.vii 
Q6. 71 2, Salsa GG, 4 Y), ike svi 
Gy 2r23=29 vin 2) sands 30 hai 
5.vili.2004 (2 2); North Haven, 14.v1i.1992 
(1 @), 27.viii.1992 (1 ¢), uv light; Orange, 
1 km NNW jet. State Route 34 and Dog- 
burn Road, 25.vi-1.vii (33 2), 9-15.vii 1 
OQ O=22 wi OO4S (INGA s a SBAR OIE 
land Co., Coventry, 0.5 km SE [U.S.] Rt. 
44, 29.vii.2001 (1 2), UCONN; Tolland, 
0.5 km SW jet. New and Grant Hill Roads, 
Kollas Orchard, 16.vi.1995 (1 ¢°), CTM. 


Rhagoletis meigenii (Loew) 
(Diptera: Tephritidae) 


Foote et al. (1993) noted that the wide- 
spread European species, R. meigenii, has 
been present in North America since at least 
1986 when it was collected in New Hamp- 
shire. Specimens also are known from 
Maine and Nova Scotia (A. Norrbom, per- 
sonal communication). In Connecticut, the 
first adult was hand-collected in Windsor, 
Hartford Co., 1998. Most subsequent ones 
were captured in Malaise or canopy traps, 
usually in mesic or wet forested areas 
where the likely host plants, Berberis spp., 
were common. This tephritid appears to be 
widespread in western and central Con- 
necticut (Fig. 1). 

In spring 2004, adults (18 ¢6, 21 2) 
emerged from the mixture placed beneath 
fruits of B. thunbergii sampled in Connect- 
icut in 2003. Similarly, adults (4 ¢, 7 @) 
were reared from fruits of B. thunbergii col- 
lected in New York in 2003. Therefore, this 
species uses congeneric larval hosts in both 
North America and Europe (Foote et al. 
1993). 


VOLUME 107, NUMBER 4 


70 


60 


50 


40 


30 


No. Females 


20 


10 


ASHI 55 422% 9 
June 


Fig. 3: 
during 2003. 


In a calcareous swamp in northern Con- 
necticut, adults flew mainly in July, with a 
few specimens captured in August (Fig. 3). 
At Barkhamsted, Litchfield Co., the 14 fe- 
males that were captured in a Sante Malaise 
trap flew between 2 and 29 July 2004. 

New state records.—Fairfield Co., Mon- 
roe, Cutlers Farm Road, 10—14.vui.2004 (1 
2), AMB; Newtown, Tory Lane, 10— 
14.vii.2004 (1 2), AMB. Hartford Co., 
Windsor, Valley Laboratory, Conn. Agric. 
Exp. Sta., 25.vi.1998 (1 °) [first state rec- 
ord]. Litchfield Co., Barkhamsted, 0.8 km 
NNE jet. State Routes 219 and 318, 2—8.vil 
Gaon O=lSsavai(G) 2) sll6=22 vane(Gr 2) rand. 
23—29.vii.2004 (1 2), SM; Canaan, Rob- 
bins Swamp, | km NNE jet. Belden and 
Sand Roads, Robbins Swamp (129 2), see 
Fig. 3; Canaan, 1 km S Falls Village, 
10.ix.2004 (berry collection), chilled and 
reared to adults; Litchfield, White Memo- 
rial Conservation Center, 1—7.vii.2004 (1 
36,9 2), AMB. New Haven Co., Guilford, 
3.8 km NNW jet. State Routes 77 and 80, 
near Beaver Head Swamp, 24.vii.2000 (1 
Om ORvi= levails ll <2) and) Qsvine-Z002), (2); 


1G 28 
July 


Seasonal abundance of females of Rhagoletis meigenii in a calcareous swamp in northern Connecticut 


SOVMAGT Walsy 120,12 


August 


GGM; Guilford, 4 km NNW jet. State 
Routes 77 and 80, 9.vii.2000 (1 Q), 
29.vii.2002 (1 2), CTM; North Branford, 
Totoket Mtn., 1.3 km ENE jet. State Routes 
22N and 80, 2-3.vii.2002 (1 2), GGM; 
North Branford, Totoket Mtn., 3 km NNE 
ject. State Routes 22N and 80, 10- 
12.vii.2001 (1 2), GGM, CTM; North 
Branford, 0.9 km NNW jet. State Route 80 
and Great Hill Road, 2—8.vii.2004 (2 @), 
SBA. Tolland Co., Tolland, Nye-Holman 
St. Forest, 30.vi.2004 (3 2), GGM, CTM. 


Anthidium manicatum (L.) 
(Hymenoptera: Megachilidae) 


This Old World bee has the largest dis- 
tribution of any species of Anthidium (Hoe- 
beke and Wheeler 1999), having acciden- 
tally been introduced into several conti- 
nents. In North America, A. manicatum was 
recovered first in central New York in 1963 
(Jaycox 1967). It subsequently has been 
found at new localities in New York, Penn- 
sylvania, Michigan, Ohio, Ontario, and 
Quebec (Pechuman 1967, Severinghaus et 


956 


al. 1981, Smith 1991, Hoebeke and Wheel- 
er 1999, Payette 2001, Miller et al. 2002). 
Although I found this adventive bee at only 
two locations in New London Co. in 2004 
(Fig. 1), probably it is more widespread. In 
North America, adults are associated main- 
ly with species of mints (Lamiaceae), al- 
though they sometimes use floral hosts in 
Asteraceae, Lythraceae, and Scrophulari- 
aceae (Pechuman 1967, Hoebeke and 
Wheeler 1999, Miller et al. 2002). 

New state records.—New London Co., 
East Lyme, near jct. State Route 156 and 
Liberty Way, 18.viii.2004 (1 63), CTM 
[first state record]; Waterford, Harkness 
Memorial State Park, 25.ix (1 2) and 
2.x.2004 (1 6, 1 2), on blossoms of Stach- 
ys byzantina K. Koch, CTM. 


Megachile sculpturalis Smith 
(Hymenoptera: Megachilidae) 


This eastern Asian bee, known as the gi- 
ant resin bee, steadily has increased its 
range across eastern North America. Man- 
gum and Brooks (1997) first discovered this 
cavity-nesting bee in North Carolina in 
1994. Subsequently, Batra (1998), Kondo et 
al. (2000), Ascher (2001), and Mangum and 
Sumner (2003) reported it from another 14 
states, the District of Columbia, and Ontar- 
io, Canada. Mangum and Sumner (2003) 
did capture M. sculpturalis in Connecticut 
in 2001, but here I report one taken at a 
different location nearly a year earlier and 
others at additional sites (Fig. 1). I also 
have observed adults that were hovering 
near large oaks, Quercus sp., at the Roger 
Williams Zoo, Providence, Providence Co., 
Rhode Island. 

Ascher (2001) and Mangum and Sumner 
(2003) summarized the floral hosts used for 
foraging and male-patrolling. In Connecti- 
cut, M. sculpturalis used two additional flo- 
ral hosts—sea-lavender, Limonium caroli- 
nianum (Walter) Britton (Plumbaginaceae), 
and goldenrod, Solidago sp. (Asteraceae). 

New state records.—Fairfield Co., Strat- 
ford, Long Beach, 7.viii.2003 (1 ¢), prey 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


of asilid, Proctacanthus rufus Williston, 
CTM. Hartford Co., Farmington, Hill-Stead 
Museum, 20.vii.2004 (1 6), on blossoms of 
Lythrum salicaria L., 24.vii.2004 (1 ¢), on 
blossoms of Lythrum salicaria, MCT; West 
Hartford, Sunset Farm Road, 3.vi1i.2004 (1 
3), MCT. Litchfield Co., Norfolk, Great 
Mountain Forest, 10.vii.2004 (1 2), on 
blossoms of Asclepias syriaca L., MCT. 
Middlesex Co., Haddam, Higganum Mead- 
ows Wildlife Area, 27.viii.2000 (1 2), on 
blossoms of Solidago sp., MCT [first Con- 
necticut record]. New Haven Co., Conn. 
Agric. Exp. Sta., 123 Huntington Street, 
23.vii.2001 (1 ¢), 19.vii.2002 (2 3). New 
London Co., Groton, Bluff Point Coastal 
Reserve, 24.viii.2004 (1 6), on blossoms of 
Limonium carolinianum, CT; Voluntown, 
Pachaug State Forest, field along Erickson 
Wildlife Marsh, 17.vii.2004 (1 2), on blos- 
soms of Asclepias syriaca, MCT. 


ACKNOWLEDGMENTS 


I thank Timothy Abbey, Richard Cowles, 
Rose Hiskes, Gale Ridge, Michael C. 
Thomas, and Kenneth Welch, CAES, for 
providing distributional records from Con- 
necticut. Julia Daigler, Morgan Lowry, Sha- 
lyn Zappulla, and Tracy Zarrillo assisted 
with trapping. Allen Norrbom (Systematic 
Entomology Laboratory, USDA, Washing- 
ton, DC) confirmed my identification of 
Rhagoletis meigenii and graciously sup- 
plied his unpublished distributional records. 
Donald W. Webb (Center for Biodiversity, 
Illinois Natural History Survey, Cham- 
paign, Illinois) allowed me to use his un- 
published key to Rhagio and confirmed my 
identification of Rhagio tringarius. Donna 
Ellis (Department of Plant Science, Univer- 
sity of Connecticut, Storrs) provided data 
on an early state record of the lily leaf bee- 
tle. Jane O’Donnell (University of Con- 
necticut) and Raymond Pupedis (Yale Pea- 
body Museum, New Haven) kindly allowed 
me to examine collections under their care. 
Kirby Stafford and Michael Thomas 
(CAES) reviewed an earlier version of the 
manuscript. The USDA Forest Service 


VOLUME 107, NUMBER 4 


(Durham, New Hampshire) provided partial 
funding for this project. 


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VOLUME 107, NUMBER 4 


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PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 960-967 


A NEW SPECIES OF PAIWARRIA (LEPIDOPTERA: LYCAENIDAE: 
EUMAEINI) FROM WESTERN ECUADOR 


JASON P. W. HALL AND KEITH R. WILLMOTT 


(JPWH) Department of Entomology, National Museum of Natural History, Smithsonian 
Institution, Washington, DC 20560-0127, USA (e-mail: hallja@si.edu); (KRW) McGuire 
Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University 
of Florida, Gainesville, FL 32611-8525, USA (e-mail: kwillmott @ flmnh.ufl.edu) 


Abstract.—A new eumaeine lycaenid species, Paiwarria chuchuvia, n. sp., is described 
and illustrated from wet premontane forest in the western Ecuadorian Andes. Its sister 
species, Paiwarria episcopalis (Fassl), appears to be distributed allopatrically in the central 
and eastern Andes. The generic placement of both species is discussed. 


Key Words: 


During nearly fifteen years of studying 
lycaenid butterflies in Ecuador, as part of 
our “Butterflies of Ecuador’? project (see 
http://www.butterfliesofecuador.com), we 
have collected nearly 100 undescribed spe- 
cies. Nearly 130 Ecuadorian lycaenid spe- 
cies in total still require description, ap- 
proaching one quarter of the country’s ly- 
caenid fauna. We continue here our lycaen- 
id descriptive work (Torres et al. 1996, Hall 
et al. 2005) by describing from the western 
Ecuadorian Andes a new species in the ge- 
nus Paiwarria Kaye, one of four genera in 
the Eumaeus section (Robbins 2004) of the 
Eumaeini (Eliot 1973). The other genus in 
this section with undescribed Ecuadorian 
species is Mithras Hitibner, but these will be 
treated elsewhere. 


METHODS 


Morphological terms for genitalia largely 
follow Eliot (1973) and Robbins (1991), 
and terminology for wing venation follows 
Comstock and Needham (1918), with cells 
named for the vein above, or anterior. Light 
microscopy examination was done using an 
Olympus SZH. The digital images in Figs. 


Andes, Colombia, Ecuador, Fasslantonius, Paiwarria, P. episcopalis 


1—2 were taken using a Nikon Coolpix 995, 
and those in Figs. 3—4 were taken using a 
Nikon D1X attached to a Microptics Digital 
Imaging System. 


Paiwarria chuchuvia Hall and Willmott, 
new species 
(Figs. 1A, B; 3; 4) 


Description.—Male: Forewing length 19 
mm. Forewing costal margin convex, distal 
margin approximately straight; hindwing 
anal margin convex then slightly indented 
before tornus, apex rounded and distal mar- 
gin slightly undulating towards tornus, a 
long tail at vein Cu, and a short tail at vein 
Cu,; forewing with four radial veins. Dor- 
sal surface: Both wings pale iridescent 
green, with a broad black apex on forewing 
extending from tornus to just beyond mid- 
point of costa, a uniformly broad black bor- 
der around distal margin of hindwing, and 
a gray hindwing anal margin, distal margin 
of green area angled at vein M, on forewing 
and undulating on hindwing, tips of long 
(and probably short) tails white; forewing 
androconial cluster appears to be a scent 
pad versus a scent patch (sensu Clench 


VOLUME 107, NUMBER 4 


961 


Figs. 1-2. 
Rio Chuchuvi, W. Ecuador (USNM). 1B, Paratype 2 P. chuchuvia, Anchayacu, W. Ecuador (JHKW). 2A, 
Paralectotype 6 P. episcopalis, Rio Aguacatal, W. Colombia (BMNH). 2B, 2 P. episcopalis, Rio Abanico, E. 
Ecuador (JHKW). 


1975, Robbins 1991) and consists of a very 
small, circular, dense patch of elongate, 
smooth-tipped brown scales (half width of 
surrounding green scales) in upper distal 
corner of discal cell; visible fringe on both 
wings black. Ventral surface: Ground color 
of forewing bright iridescent green, with 
broad black anal and distal margins, suba- 
pex a mixture of green and black scales, a 
large triangular area of black scales towards 
base of both cells Cu, and M,, area in discal 
cell occupied dorsally by scent pad largely 
bare, revealing a dark reddish brown sub- 
stance between two wing membranes, a 
large rectangular scent pad immediately be- 
low this bare area (containing similar scales 


Paiwarria adults (dorsal surface on left, ventral surface on right). 1A, Holotype d P. chuchuvia, 


to dorsal scent pad) occupies center of dis- 
cal cell end; ground color of hindwing 
bright iridescent green, with two uniformly 
narrow black stripes along anal margin be- 
low vein Cu, extending from wing margin 
to near tornus, a broad black discal band 
that extends vertically from near costal mar- 
gin to near tornus, tapering sharply towards 
tornus, a uniformly broad, slightly curved, 
postdiscal black band that extends from 
costal margin to same point near tornus, 
and a uniformly broad, curved, submarginal 
black band that extends from apex to vein 
Cu,, all three broad black bands scattered 
with dark iridescent green scales that are 
most readily visible at an oblique angle, cell 


962 


3B 


Figs. 3—4. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Paiwarria genitalia. 3, Male genitalia of P. chuchuvia in lateral (A) and ventral (B) views. 4, 


Female genitalia of P. chuchuvia in ventral (A) and lateral (B) views. Scale bar = 1 mm. 


Cu, with a very thin pale bluish line at dis- 
tal margin, a large black rectangle proxi- 
mally, and a large, round, dark red spot 
proximal to that, tails black, long (and prob- 
ably short) tails with a white tip. 

Head: Labial palpus black, with white 
scaling at outer base of segment two, sec- 
ond and third segments elongate, third seg- 
ment pointed slightly downwards; eye 
brown and setose, surrounded by iridescent 
green scaling; frons entirely iridescent 
green; antenna 40% length of forewing, 
segments brown with darker sclerotization 
around tip and white scaling at base, elon- 
gate clubs black. 

Body: Dorsal surface of thorax (includ- 
ing tegula) and abdomen pale iridescent 
green, ventral surface black; all legs black. 

Genitalia (Fig. 3): Uncus rectangular in 
lateral view, with a pointed ventral poste- 
rior corner, additional sclerotization along 
inner posterior margin, and a shallow “V”’- 
shaped medial indentation dorsally; gnathos 
smoothly rounded at elbow, constricted in 
diameter before tip; tegumen flattened dor- 
sally to accommodate a dorsolateral pair of 
large, elongate brush organs, with a broadly 


triangular, slightly inwardly curved, poster- 
oventral projection four to six times width 
of lower portion of vinculum; vinculum ex- 
tends at nearly 45° almost seamlessly into 
a long and narrow saccus; valvae in lateral 
view approximately oval shaped, with 
smoothly convex ventral and dorsal mar- 
gins, bluntly triangular posterior tips and a 
desclerotized medial area, valvae in ventral 
view laterally bulbous medially before 
gradually tapering to pointed tips, valvae 
joined at anterodorsal margin by membra- 
nous tissue; aedeagus long, uniformly very 
narrow and approximately straight, with a 
pointed posterior tip, ductus ejaculatorius 
exits anterior region of aedeagus from an 
elongate dorsal area immediately before 
rounded anterior aedeagal tip; single cor- 
nutus, positioned in posterior mouth of ae- 
deagus when vesica uneverted, posteriorly 
bulbous with a serrate dorsal margin, taper- 
ing to a long and narrow, flattened anterior 
rod; eighth abdominal tergite rectangular. 
Female: Forewing length 17 mm. Wing 
shape similar to male, but hindwing slightly 
more elongate, with a more prominent tor- 
nal lobe. Dorsal surface: Differs from male 


VOLUME 107, NUMBER 4 


by a paler black ground color on both 
wings, considerably broader black margins 
on both wings, particularly in apex, a duller 
and slightly more turquoise green irides- 
cence, a more angular distal margin to fore- 
wing green, and submarginally positioned 
semicircles of white scaling in three tornal 
cell spaces below vein Cu,. Ventral surface: 
Ground color of both wings pale brown, 
with a faint, dull greenish tinge to basal 
third of wings, basal greenish-brown area 
on hindwing well-defined; both wings with 
three narrow, dirty white lines, approxi- 
mately evenly spaced and parallel to distal 
margin, one immediately distal to discal 
cell end, one submarginal and one inbe- 
tween, basalmost line slightly thinner, more 
crisply defined and brighter white, lines on 
forewing less prominent and confined to 
middle of wing between radial veins and 
middle of cell Cu,, basalmost line on 
hindwing forms a ““W” shape in middle of 
cell Cu, before extending proximally to- 
wards anal margin, distal two lines on 
hindwing interrupted by red markings in 
cells Cu, to M,, with middle line extending 
proximally to anal margin and outer line ex- 
tending directly to anal margin, largest sub- 
tornal red marking in cell Cu, and smallest 
in cell M,, three large black spots occupy 
tornus distal to red markings and below 
vein Cu,, very thin white line extends along 
entire distal margin of hindwing. 

Head: Differs from male as follows: Sec- 
ond and third segments of labial palpus 
more elongate, third segment at least twice 
as long; eye surrounded by white scaling; 
frons black with white scaling along lateral 
margins; antenna 30—40% of forewing 
length, anntennal segments with a narrow 
nudum area visible ventrally. 

Body: Differs from male as follows: Dor- 
sal surface of thorax and abdomen duller 
iridescent green, ventral surface grayish; all 
legs brown, with white scaling on ventral 
surface of tibiae and white banding on tar- 
sal segments. 

Genitalia (Fig. 4): Bursa copulatrix oval, 
with a pair of sclerotized invaginated signa 


963 


that have one large and two very small an- 
teriorly directed spines towards anterior tip; 
ductus seminalis exits bursa copulatrix im- 
mediately posteriorly to junction of ductus 
bursae and is lightly sclerotized along ven- 
tral base; anterior ductus bursae a sclero- 
tized, convex-concave tube that forms a 
broad, “‘horseshoe’’-shaped, sclerotized 
bursal pouch at junction with bursae copu- 
latrix, junction of anterior and posterior 
portions of ductus bursae slightly descler- 
otized and flexible, posterior ductus bursae 
a slightly curved sclerotized tube that 
broadens gradually towards posterior open- 
ing and has a faint dorsal desclerotized 
band along its entire length; lamella post- 
vaginalis with a “‘U’’-shaped ventral mar- 
gin, a smoothly uneven posterior margin, 
and a broad band of dorsally folded scler- 
otization along posterior margin. 

Type material.—Holotype ¢, ECUA- 
DOR: Esmeraldas, Rfo Chuchuvr, km. 
12.5 Lita-San Lorenzo rd., 0°53.01’N 
18°30.90'W, 800-900 m, July (R. Aldas) 
(National Museum of Natural History, 
Smithsonian Institution, Washington, DC, 
USA [USNM]). 

Paratypes, ECUADOR: Esmeraldas, | 
2, same locality data as holotype, | No- 
vember (K. R. Willmott) (Collection of Ja- 
son P. W. Hall and Keith R. Willmott, 
Washington, DC, USA [JHKW]); 1 2, An- 
chayacu, km. 15 Lita-San Lorenzo rd., 900 
m, 1 July (K. R. Willmott) (JHKW). Jm- 
babura, | 2, Rio Verde, km. 16 Lita-Ibarra 
rd., 1450 m., 3 August (K. R. Willmott) 
(Museo Ecuatoriano de Ciencias Naturales, 
Quito, Ecuador [MECN]). 

No additional specimens have been lo- 
cated in other major World museums (as 
listed in Hall 1999, 2002). 

Etymology.—The species name is a fem- 
inine noun derived from the name of the 
type locality. 

Diagnosis.—Paiwarria chuchuvia can be 
genuinely confused only with P. episcopal- 
is (Fass!). The fact that both species share 
a unique male ventral wing pattern, con- 
brilliant iridescence 


sisting of a green 


964 


crossed on the hindwing by broad, evenly 
spaced black bands, and a patch of andro- 
conial scales at the end of the forewing dis- 
cal cell on both wing surfaces, suggests that 
they are sister species. Fass] (1912) de- 
scribed P. episcopalis from three male 
specimens, all from Rio Aguacatal in west- 
ern Colombia (2000 m), and stated that they 
were deposited in the collections of Bra- 
bant, Fass] and Courvoisier. As these spec- 
imens reside in multiple European collec- 
tions and exhibit significant wing pattern 
variation (primarily in the prominence of 
green scaling overlying the black bands on 
the ventral hindwing), Faynel and Balint 
(2004) deemed it prudent to designate a sin- 
gle putative male syntype in the Muséum 
National d’ Histoire Naturelle, Paris, France 
(MNHN), to be a lectotype. Although this 
MNHN male seems quite likely to be a syn- 
type, it is not the ex. Fass] coll. specimen, 
as Faynel and Balint (2004) hypothesized, 
because this specimen is currently housed 
in the Senckenberg Museum in Frankfurt, 
Germany (G. Lamas, personal communi- 
cation). As it is also not the ex. Brabant 
coll. specimen, which belongs to The Nat- 
ural History Museum in London, England 
(this paralectotype is illustrated here in Fig. 
2A), we suggest it is the ex. Courvoisier 
coll. specimen. Although Faynel and Balint 
(2004) suggested that the ex. Courvoisier 
coll. specimen might be in the Naturhisto- 
risches Museum in Basle, Switzerland, 
where some Courvoisier specimens appar- 
ently reside (Horn et al. 1990), G. Lamas 
(personal communication) reports finding 
no such specimen in a cursory examination 
of that collection. The fact that the MNHN 
male bears a handwritten Fass] label with 
the locality ““Villa Elvira,” his base of op- 
erations in the Rio Aguacatal (Fassl 1914), 
and the date “4/7/08,” the year the type 
series was collected (Fass] 1912), strongly 
suggests that it might be the third missing 
syntype of P. episcopalis. 

The male of P. chuchuvia differs from 
that of P. episcopalis in its considerably 
larger size (19 mm compared to 13—15 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


mm), and by having a more pointed fore- 
wing apex (at least compared to the BMNH 
paralectotype), a slightly elongate hind- 
wing, a larger forewing scent pad on both 
wing surfaces, a broader black distal margin 
on the dorsal forewing, slightly longer 
hindwing tails, a broad black band along 
the entire distal margin of the dorsal hind- 
wing, two large black triangles at the base 
of cells Cu, and M, on the ventral forewing, 
green scaling (visible at an oblique angle) 
covering all instead of only some black ar- 
eas above vein Cu, on the ventral hindwing, 
a discal black line on the ventral hindwing 
that is twice as broad, and a large red spot 
in the tornus of the ventral hindwing. 

These two species are so strongly sexu- 
ally dimorphic that Draudt (1920) described 
the female of P. episcopalis as the distinct 
species Thecla phacana (Balint and Salazar 
2003; Robbins 2004). However, similarities 
in wing shape, palpal length, the extent of 
dorsal green iridescence, and the banding 
pattern on the ventral hindwing allow us to 
match the sexes of both species with con- 
fidence. The female of P. chuchuvia differs 
from that of P. episcopalis by exhibiting the 
same type of size and wing shape differ- 
ences of the males, and by having consid- 
erably broader black margins on both dorsal 
wings, a slightly more vertically positioned 
postdiscal white line on the ventral fore- 
wing that does not extend to the wing mar- 
gins, a more proximally displaced postdis- 
cal white line on the ventral hindwing, and 
considerably more elongate red spots in the 
tornus of the ventral hindwing, with larger 
black spots distally. Some female speci- 
mens of P. episcopalis, particularly those 
from the central Andes, have more exten- 
sive hindwing tornal red than the figured 
specimen, but the red is never as extensive 
as it is in P. chuchuvia. Central Andean fe- 
males of P. episcopalis also seem to have 
a slightly more proximally positioned white 
postdiscal line on the ventral hindwing than 
east Andean females. 

The genitalia of these two species are 
very similar. The male genitalia of P. chu- 


VOLUME 107, NUMBER 4 


chuvia differ only by having a more broad 
and bluntly triangular tip to the valvae, and 
the female genitalia differ only by having 
additional small anteriorly directed spines 
on the signa and a broader band of dorsally 
folded sclerotization along the posterior 
margin of the lamella postvaginalis. Balint 
and Salazar (2003) reported finding in fe- 
male P. episcopalis a sclerotized line down 
the center of the bursa copulatrix between 
the signa, but we found no such line in the 
one female specimen of P. episcopalis that 
we dissected. Balint and Salazar (2003) also 
described and purported to figure, in their 
Fig. 5, an “‘anteriorly pointed, sclerotized, 
ventral element of the bursal pouch” on the 
bursa copulatrix, but again we could find 
no such structure. The only structure fitting 
this description that we can discern in their 
Fig. 5 is a single signum floating beneath 
the bursal pouch. 

Systematic placement.—Kaye (1904) de- 
scribed the new monotypic genus Paiwar- 
ria, for Papilio venulius Cramer, using such 
a short and vague description, which con- 
centrated on very weakly diagnostic char- 
acters of the head, wing shape, venation and 
tail position, that the generic name was only 
infrequently used subsequently. Bridges 
(1994) additionally included in Paiwarria 
the species telemus (Cramer) and antinous 
(C. & R. Felder) (mistakenly as a synonym 
of telemus), with a question mark, presum- 
ably based on the combination of telemus 
with Paiwarria by Lamas et al. (1991) in 
their Pakitza butterfly checklist. Most re- 
cently, Robbins (2004) additionally includ- 
ed in Paiwarria the species aphaca (Hew- 
itson) (a combination also used by Brown 
and Freitas 2000), episcopalis and umbra- 
tus (Geyer), to form a total of six described 
species for the genus. He placed Paiwarria 
with Eumaeus Hiibner, Theorema Hewitson 
and Mithras in his Eumaeus section of the 
Eumaeini, because the species in these gen- 
era share male genitalia with brush organs 
(when present) that actually or nearly (in 
Paiwarria) surround the genital capsule 
(Robbins 2004). However, an alternative 


965 


generic placement has been proposed for 
episcopalis by Balint and Salazar (2003). 
They recently erected the new monotypic 
genus Fasslantonius for episcopalis, with- 
out mentioning Paiwarria as a potentially 
close relative to Fasslantonius in either the 
diagnosis or subsequent discussion, instead 
concentrating on comparing Fasslantonius 
with Thereus section genera. As the name 
Fasslantonius was published after the text 
for Robbins’ (2004) checklist was in press, 
it could not be included in that publication. 
However, as Robbins (2004) provided a 
comprehensive new generic classification 
for the Eumaeini, which included the de- 
scriptions of new genera for all those spe- 
cies groups that lacked generic names, it 
can reasonably be inferred that Fasslanton- 
ius would have been treated by him as a 
synonym of Paiwarria if the name could 
have been included in the checklist. 

Based on external characters, including 
wing shape, dorsal and ventral wing pat- 
tern, and the shape and position of the fore- 
wing scent pad, the Paiwarria of Robbins 
(2004) can broadly be divided into three 
species clusters: telemus and relatives, ep- 
iscopalis, and umbratus. Having examined 
the male and/or female genitalia of repre- 
sentatives from each of these three Paiwar- 
ria groups, the male and female genitalia of 
Theorema sapho (Staudinger), and both 
genitalia dissections and illustrations of var- 
ious Eumaeus (in Constantino and Johnson 
1997) and Mithras species (in Johnson and 
Constantino 1997, Balint and Moser 2001), 
we can make the following preliminary ob- 
servations concerning the systematic place- 
ment of episcopalis and the new species 
chuchuvia. Many of the genital characters 
cited by Balint and Salazar (2003) to di- 
agnose Fasslantonius are present in Pai- 
““ap- 
pendix angularis” to the tegumen, a dor- 
sally dentate aedeagal cornutus, and a 
notched tip to the papillae anales, and both 


warria species, including a rounded 


episcopalis and chuchuvia do indeed seem 
to be closely related to the Paiwarria tele- 
mus group of species. The expanded Pai- 


966 


warria is a morphologically rather hetero- 
geneous grouping, but Robbins (personal 
communication) informs us that he regards 
the presence of anterior vinculum processes 
that abut the inner side of the brush organs, 
and a groove on the outer surface of the 
vinculum that is almost flush with the an- 
terior edge of those processes, to be uni- 
versal synapomorphies for the genus within 
the context of the Eumaeus section. We ten- 
tatively follow the conservative arrange- 
ment of Robbins (2004) for now, and place 
the new species chuchuvia next to episco- 
palis in Paiwarria. However, their unique 
male ventral wing pattern, strong sexual di- 
morphism in ventral wing pattern, very 
long palpi, unusual male forewing scent 
pads, and “‘horseshoe’’-shaped bursal 
pouch and very prominently undulating 
ductus bursae in the female genitalia clearly 
set episcopalis and chuchuvia apart from 
the remainder of Robbins’ (2004) Paiwar- 
ria species, and Fasslantonius may yet be 
worth retaining if it can be shown that the 
telemus group and/or the telemus group + 
umbratus are monophyletic, a task that is 
beyond the scope of this paper. 

Biology.—Paiwarria chuchuvia appears 
to be confined to premontane forest, and is 
currently known from about 800 to 1,450 
m. We know of P. episcopalis specimens 
from the eastern slope of the Andes and in 
the Cauca valley of Colombia (1,000 to 
2,000 m), suggesting that P. chuchuvia may 
allopatrically replace P. episcopalis on the 
western slope of the western Andean cor- 
dillera. Paiwarria chuchuvia females were 
recorded flying across ridgetop lightgaps 
two meters above the ground from 1300 to 
1430 h. 

Distribution.—Paiwarria chuchuvia is 
currently known from only a few localities 
in a small area of northwestern Ecuador, but 
it is presumably more widespread along the 
western slope of the Andes from western 
Colombia to western Ecuador. 


ACKNOWLEDGMENTS 


We thank the many curators in Europe 
and the Americas who kindly allowed us to 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


examine the lycaenid collections under their 
care (see list in Hall 1999, 2002); Robert 
Robbins for giving us access to the para- 
lectotype of P. episcopalis on loan from the 
BMNH; The National Geographic Society 
(Research and Exploration Grant #575 1-96) 
and The National Science Foundation (Bio- 
diversity Surveys & Inventories Grant 
#0103746) for research support; the Museo 
Ecuatoriano de Ciencias Naturales and the 
Ministerio del Ambiente, Quito, for assist- 
ing with the procurement of collecting and 
export permits in Ecuador; and Gerardo La- 
mas and Robert Robbins for detailed com- 
ments on the manuscript. 


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Santo. Boletim do Museu de Biologia Mello Lei- 
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Clench, H. 1975. Introduction, pp. 1-72. In Howe, W. 
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Comstock, J. H. and J. G. Needham. 1918. The wings 
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Eliot, J. N. 1973. The higher classification of the Ly- 
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. 2002. A phylogenetic revision of Calydna and 
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PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 968-970 


NOTE 


Notes on the Ethology of Bohartia martini Adisiomarto and Wood (Diptera: Asilidae) 
in Wyoming, USA 


Bohartia martini Adiseomarto and 
Wood, 1975 is an extremely small, rarely 
observed species, having only been collect- 
ed previously in Wyoming at the Mesa 
Conservation Area. [Paratype—*‘l male, 
Mesa Conserv. Area, So. of Pinedale;” 
(Adisoemarto and Wood 1975).] [Author’s 
note: Dr. Wood returned 10 additional spec- 
imens of Bohartia martini to the University 
of Wyoming, College of Agriculture Insect 
Collection that were part of the type series 
and were so labelled. | 

Recognition characters for B. martini are 
as follows: Blackish species. Length 6—8 
mm. Mystax mostly white. Second antennal 
segment usually with some reddish or 
brown bristles. Thorax white to slightly yel- 
lowish and silvery-gray pollinose; scutellar 
bristles black. Wing hyaline with dark 
brown veins. Abdomen reddish brown to 
black. Femora black, tibiae reddish at least 
at base dorsally, fore tibia with fringe-like 
hairs on posterior surface. Male hypan- 
drium bulging ventrally near base. 

A small population of Bohartia martini 
was studied briefly in the Red Desert of 
Wyoming, 3.3 mi N of the ghost town of 
Tipton Junction off Tipton Road in Sweet- 
water County. Observations were made in- 
termittently from 27 June to 9 July, 1978. 
These few notes are being offered because 
no biological data on members of the genus 
have been published previously. 

Vegetation in the study area was com- 
posed of typical arid land species: Agro- 
Pyron spicatum (Pursh) Scribn. & Sm., 
Oryzopsis hymenoides (R & S) Ricker ex 


Piper, Artemesia tridentata Nutt., Tetra- 
dymia canescens DC., Arenaria hookeri 
Nutt. var. hookeri, Salsola kali L., Eriogon- 
um brevicaule Nutt. ssp. micranthum (Nutt) 
Reveal, Leptodactylum pungens (Torr.) 
Nutt., Cirsium pulcherrimum (Rydb.) K. 
Schum., Chrysothamnus nauseosus (Pall ex 
Pursh) Britt., Chrysothamnus viscidiflorus 
(Hook.) Nutt., and Haplopappus armerioi- 
des (Nutt.) Gray (Fig. 1). Bohartia martini 
shared the habitat with a population of Di- 
cropaltum mesae (Tucker), a slightly larger 
species, but no interactions were observed. 
Surface temperatures on site ranged from 
91 to 111°F when observations were made. 
Specimens were observed landing on the 
soil surface, on twigs lying on the surface 
(Fig. 2), on plant debris, on grass stems, 
and on dead twigs of Artemisia to a height 
of 3.6 cm (1.5 in). 

Foraging.—Prey were collected from the 
air in short capture darts, not exceeding 15 
cm (6 in). On one occasion an asilid was 
observed to hover briefly while manipulat- 
ing prey, which it subsequently released. 
Prey may be manipulated during feeding: 
the asilid rears backwards using its fore and 
hind tarsi for manipulation, while balancing 
itself on the middle legs [This same type of 
manipulation is exhibited by Lasiopogon 
cinereus (Cole) (Lavigne and Holland 
1969)]. In one sequence at 10:33.30, a fe- 
male captured a small leafhopper (Cicadel- 
lidae) at the end of a 15 cm capture dart. 
She subsequently moved several times, fly- 
ing less than 30 cm each time. Prey was 
manipulated at 10:43 and 10:44. At 10:47 


Figs. 1-2. 


— 


1 (top) Rangeland site near Tipton Junction, Wyoming, where behavioral observations of Bohartia 


martini were made. 2 (bottom), A female of B. martini resting on soil surface with fore tarsi placed on piece 


of debris. 


969 


VOLUME 107, NUMBER 4 


970 


she made an approximate 30 cm (12 in) 
flight during which she discarded the prey. 
A total of 19 prey were collected on four 
separate dates by foraging Bohartia. These 
represented two Insect Orders. However all, 
but the single leafhopper, were Coleoptera 
[Melyridae: Dastinae], suggesting a fixation 
on this beetle. Four of the beetles were tak- 
en by males and the rest by females 

Mating.—There apparently is no court- 
ship exhibited by males of this species. A 
single complete mating was observed at I: 
54 pm on 27 June. The female flew into the 
air, having been disturbed by the jumping 
of a grasshopper. The male caught her in 
mid-air and the pair tumbled to the soil, 
rolled and came up mated in the tail-to-tail 
position. They flew into the shade (94°F) 
and at 1:58, the pair suddenly separated and 
flew off in different directions. A second 
mated pair in the tail-to-tail position was 
observed at 3:07 on 3 July on the hardpan 
surface where the temperature was recorded 
as being 110°F 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


LITERATURE CITED 


Adisoemarto, S. and D. M. Wood. 1975. The Nearctic 
species of Dioctria and six related genera (Dip- 
tera, Asilidae). Quaestiones Entomologicae 11: 
505-576. 

Lavigne, R. J. and F R. Holland. 1969. Comparative 
behavior of eleven species of Wyoming robber 
flies (Diptera: Asilidae). University of Wyoming 
Agricultural Experiment Station Science Mono- 
graph 18, 61 pp. 


Robert J. Lavigne and Stephen Bulling- 
ton, (RJL) Professor Emeritus, Department 
of Renewable Resources, Entomology Pro- 
gram, University of Wyoming; Honorary 
Research Associate, South Australian Mu- 
seum of Natural History; current address: 
P.O. Box 1010, Mt Barker, SA 5252, Aus- 
tralia (e-mail: rjlavigne @netspace.net.au); 
(SB) Pest Survey Specialist, USDA, APHIS, 
PPO, 401 E. Louther St., Suite 102, Car- 
lisle, PA 17013, U.S.A. (e-mail: swb@key- 
net.net) 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 971-972 


NOTE 


Scoloposcelis discalis Van Duzee, 1914, a Synonym of Anthocoris galactinus Fieber, 
1837, and Xylocoris umbrinus Van Duzee, 1921, a Synonym of Piezostethus 
californicus Reuter, 1884 (Hemiptera: Heteroptera: Anthocoridae) 


Scoloposcelis discalis was described by 
Van Duzee (1914) from Lakeside, San Di- 
ego County, California. Van Duzee (1921) 
placed the species in the genus Xylocoris 
Dufour (1831). Besides the California lo- 
cation, it has been reported from Arizona 
(Drake and Harris 1926) and the Hawaiian 
Islands (Van Duzee 1936). Kelton (1976) 
designated a lectotype male from the type 
series. | have examined specimens of Xy- 
locoris discalis from the Hawaiian Islands 
identified by Van Duzee and a series of 
specimens collected later there that are 
identical to those identified by Van Duzee. 
These specimens are in the collections of 
the Bishop Museum. The male clasper of 
the Hawaiian specimens clearly identifies 
the material as Xylocoris (Proxylocoris) 
galactinus (Fieber), a European species. It 
has been introduced into many regions 
around the world and is commonly found 
associated in stored grain (Lattin 2000). 
Van Duzee (1905) first reported X. galac- 
tinus from North America. Henry (1988) 
listed the species from Alberta, British Co- 
lumbia, California, Florida, Georgia, Idaho, 
Illinois, Manitoba, Missouri, New Jersey, 
New York, Ontario, Quebec, and Saskatch- 
ewan and I have identified specimens from 
Oregon. Péricart (1972) provided a detailed 
treatment of X. galactinus in Europe, Kel- 
ton (1978) included this species in his study 
of the Anthocoridae of Canada and Alaska, 
and both authors included an illustration of 
the distinctive male clasper. Adults of this 
species are known only in the macropterous 
state. 

Scoloposcelis discalis Van Duzee (1914) 
is here considered to be a synonym of An- 
thocoris galactinus Fieber (1837) (now Xy- 


locoris (Proxylocoris) galactinus), new 


synonymy. In addition, Van Duzee (1914) 
described Piezostethus flaccidus from a 
macropterous female and a brachypterous 
female from Descanso and Alpine, Califor- 
nia. Kelton (1977) synonymized P. flacci- 
dus Van Duzee with Xylocoris galactinus 
and designated the macropterous female as 
the lectotype. Because X. galactinus is 
known only from macropterous specimens, 
the identity of the brachypterous female 
Van Duzee (1914) included in his type se- 
ries of P. flaccidus remains uncertain. All 
records of X. discalis from the Hawaiian Is- 
lands are here referred to X. (Proxylocoris) 
galactinus. 

Xylocoris umbrinus Van Duzee (1921) 
was described from a male from Bryson, 
Monterey County, and a female from Cay- 
ton, Shasta County, California. Reuter 
(1884) described Piezostethus californicus 
from a single female from Mariposa, Mar- 
iposa County, California. Earlier specimens 
of X. umbrinus in the Oregon State Uni- 
versity Collection have been identified by 
P.D. Ashlock, H.M. Harris, and J.D. Lattin. 
There was a specimen identified by Harris 
as X. californicus from Klamath Falls, 
Oregon. Recently, a review of the species 
of Xylocoris from western North America 
included the examination of original de- 
scriptions, published literature, and a num- 
ber of specimens from this region. Reuter’s 
description of Piezostethus californicus was 
quite close to that of Xylocoris umbrinus 
Van Duzee. Ultimately, it was recognized 
that only one species was involved with 
Piezostethus californicus Reuter, 1884, hay- 
ing priority over Xylocoris umbrinus Van 
Duzee. 1921, new synonomy. Details of 
the similarity follow: size of macropterous 
P. californicus the same as the type of X. 


O72 


umbrinus (3 mm); dorsum shiny; hemylytra 
brown yellow to piceus brown without 
white areas except hyaline white mem- 
brane; rostrum reaching middle coxae, tho- 
racic scent gland with slender basal stem, 
slightly enlarged at bend, apex of gland not 
reaching anterior margin of metapleuron, 
apex acute. Van Duzee (1921) stated that 
the male type came from Bryson and the 
female allotype from Cayton, California, 
and that he had taken X. californicus from 
Bryson and Cayton, California—both spe- 
cies from the same localities! The species 
is now known from Alberta, British Colum- 
bia, California, Idaho, Manitoba, Oregon, 
and Saskatchewan (Anderson 1962, Kelton 
1978, Henry 1988). I have examined spec- 
imens from Washington and Lewis et al. (in 
press) also reported it from Washington. 

Acknowledgments.—My thanks go to A. 
Ramsdale, Bishop Museum, and R. Bau- 
mann, Brigham Young University, for send- 
ing their collections of Anthocoridae for 
study, and to L. Parks for typing of the 
manuscript. 


LITERATURE CITED 


Drake, C. J. and H. M. Harris. 1926. Notes on Amer- 
ican Anthocoridae with descriptions of new forms. 
Proceedings of the Biological Society of Wash- 
ington 39: 33—46. 

Dufour, L. 1831. Description et figure du Xylocoris 
rufipennis, Hémiptére nouveau. Annales des Sci- 
ences Naturelles 22: 423-426. 

Fieber, E X. 1836, 1837. Beitriig zur Kenntnis der 
Schnabelkerfe (Rhyncota). Jn Weitenweber, W. R., 
ed. Beitrage zur gesammten Natur-und Heilwis- 
senschaft 1(1) 1836: 97-111, 1837: 337-355. 
Barth, Prague. 

Henry, T. J. 1988. Family Anthocoridae, pp. 12—28. In 
Henry, T. J. and R. C. Froeschner, eds. Catalog of 
the Heteroptera, or True Bugs, of Canada and the 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Continental United States. E.J. Brill, Leiden, 958 
PP. 

Kelton, L. A. 1976. Three new species of Xylocoris 
from North America, and a note on the status of 
species in the genus Scoloposcelis Fieber (Heter- 
optera: Anthocoridae). The Canadian Entomolo- 
gist 108: 193-198. 

. 1977. A new species of Elatophilus Reuter 

from Ontario and new synonymy for Piezostethus 

flaccidus Van Duzee (Heteroptera: Anthocoridae). 

The Canadian Entomologist 109: 1017-1018. 

. 1978. The insects and arachnids of Canada. 
Part 4. The Anthocoridae of Canada and Alaska. 
Heteroptera. Anthocoridae. Research Branch, 
Canada Department of Agriculture. Publication 
1639. 101 pp. 

Lattin, J. D. 2000. Minute pirate bugs (Anthocoridae), 
pp. 607—637. In Schaefer, C. W. and A. R. Panizzi, 
eds. Heteroptera of Economic Importance. CRC 
Press, Boca Raton, 828 pp. 

Lewis, T. M., D. R. Horton, and D. H. Boers. (In 
press.) New United States records for Anthocori- 
dae (Hemiptera: Heteroptera). Pan-Pacific Ento- 
mologist. 

Péricart, J. 1972. Hémiptéres. Anthocoridae, Cimici- 
dae, et Microphysidae de |’Ouest—Palearctique. 
Masson et C* Editeurs, Paris, 402 pp. 

Reuter, O. M. 1884. Monographia Anthocoridarum Or- 
bis Terrestris. Helsingforsiae. 204 pp. (also, Acta 
Societate Scientiarum Fennicae 1885: 555-758). 

Van Duzee, E. P. 1905. Hemiptera taken in the Adi- 
rondack Mountains. New York State Museum 
Bulletin 97: 546-556. 

. 1914. A preliminary list of the Hemiptera of 

San Diego County, California. Transactions of the 

San Diego Society of Natural History 2: 1—57. 

1921. Characters of eight new species of 

North American Anthocoridae or flower bugs. 

Proceedings of the California Academy of Scienc- 

es, Fourth Series 9: 137-144. 

. 1936. A report of some Heteroptera from the 

Hawaiian Islands with descriptions of new spe- 

cies. Proceedings of the Hawaiian Entomological 

Society 9: 219-229. 


John D. Lattin, Department of Botany 
and Plant Pathology, Oregon State Univer- 
sity, Corvallis, OR 97331-3902, U.S.A. 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 973-976 


NOTE 


A Note on the Densities of Ixodes scapularis (Acari: Ixodidae) and White-tailed Deer 
on the Campus of the National Institute of Standards and Technology, Maryland, USA 


In recent years Montgomery County has 
been among the leading counties in the state 
of Maryland for confirmed cases of Lyme 
disease (data from Maryland Department of 
Health and Mental Hygiene). Most instanc- 
es of human Lyme disease result from the 
bite of a blacklegged tick, Ixodes scapularis 
Say, particularly in the nymphal stage, in- 
fected with the spirochete Borrelia burg- 
dorferi Johnson, Schmid, Hyde, Steigerwalt 
and Brenner (Lane et al. 1991). Larvae and 
nymphs of /. scapularis use a wide variety 
of vertebrate hosts (mammals, birds, liz- 
ards), but adults feed primarily on white- 
tailed deer, Odocoileus virginianus (Zim- 
mermann) (Lane et al. 1991). It is generally 
thought that the deer is the key host species 
in maintaining dense populations of J. sca- 
pularis (Wilson et al. 1985). 

In 2003, we were requested to survey 
tick populations on the National Institute of 
Standards and Technology (NIST) campus 
after employees began noticing the pres- 
ence of host-seeking ticks. By the mid 
1990s an increasingly dense population of 
deer had severely degraded natural habitats 
and ornamental plantings at the 2.38 km? 
NIST fenced campus in Gaithersburg, 
Montgomery County, MD. Deer numbers at 
NIST exceeded 120/km?’, roughly a tenfold 
greater density than levels considered the 
threshold for overabundance (Tilghman 
1989). Since 1995, the Humane Society of 
the U.S., initially in conjunction with Lori 
Thiele, a Masters Degree student at the Uni- 
versity of Maryland, has conducted a con- 
traceptive-based program to reduce to the 
deer population at NIST. Deer numbers 
gradually declined, but were still overabun- 
dant in 2003 (90 deer/km?’). We report the 
results of surveys conducted in 2003 and 
2004 to document the occurrence and dis- 


tribution of /. scapularis at NIST and the 
presence of B. burgdorferi. 

Nine sample sites (all >100 m from one 
another) were selected in wooded habitats 
at NIST. Eight of the sites were woodlots 
of mature hardwoods (predominantly Quer- 
cus spp. and Liriodendron tulipifera L.) and 
one site was mostly planted white pine, Pi- 
nus strobus L. In 2003 and 2004, the un- 
derstory in the woodlots was nearly barren 
except for Nepal microstegium (Japanese 
stiltgrass), Microstegium vimineum (Tri- 
nius) A. Camus, an invasive grass, that has 
become the principal ground cover in the 
woodlots. Furthermore, the expansive mats 
of M. vimineum appear, despite the overly 
dense deer population, to be undamaged by 
deer feeding. There was little indication of 
tree regeneration, aside from a few oak 
seedlings. Nymphs were sampled by flag- 
ging once in June and again in early July 
2003, and larvae were sampled twice in late 
July 2003. Adults were sampled in October 
2003. Nymphs were sampled twice more in 
June 2004. Using a 0.5 by 0.5 m flag of 
flannel crib cloth, each site was flagged by 
a person walking slowly for 30 s and tra- 
versing about 10 m. By flip-flopping the 
flag cloth, the flagger covered about 10 m°. 
This was repeated 10 times on non-overlap- 
ping routes, so that about 100 m?* were 
flagged at each sample site on each sample 
date. Nymphs and adults were collected in 
vials, and in the laboratory they were iden- 
tified and preserved in 70% isopropyl al- 
cohol. Larvae removed from flag 
cloths on pieces of transparent tape that 
were affixed in notebooks. In the laboratory 
the larvae were identified and counted. In 


were 


order to determine the prevalence of B. 
burgdorferi in ticks at NIST, 
nymphs and adults were tested by PCR with 


collected 


974 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Table 1. Despite an abundance of white-tailed deer, the primary host of adult /. scapularis, few nymphs and 
adults were captured by flagging at NIST. Note that larvae were found at all sample sites. 


Sample Sites 
Ne 2 3 4 5 6 7 8 9 
Nymphs? 
2003 1.0 10) 1.0 0) 0) 1.0 0.5 1.0 0) 
2004 0 5) 0.5 O 0) 0 0) 1.5 0) 
Adults 0) 0) l 0) 0) 0) ] 0) 2) 
Larvae 20.5 6.0 30.5 212.0 87.5 109.5 37.0 44.0 6.0 


«Larvae and nymphs averaged for 2 sampling dates, adults sampled once. Larvae and adults sampled only in 


2003. 


the Bbsl primer set, as described by Cyr et 
al. (2005). Negative controls were included. 

In 2003, a total of nine J. scapularis 
nymphs were found at five sample sites, 
representing four separate woodlots, and in 
2004 a total of seven nymphs were captured 
at three sites (Table 1). In contrast, larvae 
were captured at all sample sites in 2003, 
although larval numbers were not high 
(>100 at only two sites). Four adults were 
captured at three of the five sites where 
nymphs were found earlier in 2003. Re- 
markably, all adult ticks were positive for 
B. burgdorferi in PCR tests, but the nymphs 
were negative. 

Under most circumstances, population 
densities of J. scapularis are generally 
thought to be positively correlated with 
population densities of white-tailed deer 
(Wilson et al. 1985). As deer populations 
increase or decrease in density, so do the 
tick populations. However, the situation at 
NIST was far from typical. Few ticks were 
found despite a high density of deer. The 
occurrence of larvae, sometimes in modest 
numbers (>100), at all sample sites, very 
few nymphs at just six of nine sample sites 
and adults at three of nine sites, suggests 
poor survival of immature /. scapularis at 
NIST. Possible explanations for the low 
numbers of /. scapularis nymphs and adults 
are: 1) a shortage of small vertebrate hosts, 
2) excessive drying at the leaf litter/ ground 
level. Larval and nymphal /. scapularis 
feed on a wide range of vertebrates, includ- 
ing birds and lizards (Main et al. 1982, 


Lane et al. 1991), whose populations can 
be impacted by a cascade of negative ef- 
fects stemming from an overabundance of 
deer (C6té et al. 2004). Chipmunks and 
squirrels were observed only infrequently at 
NIST. Population density estimates for 
these rather visible host species at NIST are 
lacking, as are estimates for the nocturnal 
white-footed mouse, Peromyscus leucopus 
(Rafinesque), an important host of larval 
and nymphal /. scapularis and principal res- 
ervoir of B. burgdorferi (Lane et al. 1991). 
Larvae and nymphs of /. scapularis will 
feed on white-tailed deer (Telford et al. 
1988). In view of the low density of J. sca- 
pularis at NIST, the abundance of deer (as 
available hosts) may not offset other per- 
turbations of the woodlot ecosystems that 
negatively affect the ticks. The severely de- 
graded understory and litter layer may in- 
fluence micrometeorological factors in 
ways that are deleterious to tick host ac- 
quisition and survival (Harlan and Foster 
1990). 

In the northeastern states, B. burgdorferi 
iS maintained in nature by reservoir hosts, 
principally P. leucopus, being infected by 
being bitten in the spring and early summer 
by infected /. scapularis nymphs. Later in 
the summer, larvae of the next generation, 
which are largely free of B. burgdorferi, be- 
come infected by feeding on infected mice. 
Thus, infection of unfed nymphs is due to 
the infection status of the host on which 
they fed as larvae and the infection of unfed 
adults a consequence of nymphal feeding. 


VOLUME 107, NUMBER 4 


Of the ticks we captured at NIST, none of 
the nymphs tested positive for B. burgdor- 
feri, whereas all the adults were positive. 
The computer simulations of Mount and 
Haile (1997) indicate that at least an esti- 
mated 87 I. scapularis nymphs/ha are need- 
ed to maintain B. burgdorferi transmission 
in an otherwise suitable ecosystem. Our 
collections averaged ~0.9 nymphs/100 m2, 
which if multiplied by a factor of ~10 be- 
cause flagging and dragging are considered 
to capture only a small fraction of the ticks 
present (Daniels et al. 2000), indicate that 
the J. scapularis population at NIST ex- 
ceeded the transmission threshold. Higher 
infection rates are expected in adult /. sca- 
pularis, because they have had two chances 
to have had fed on an infected host. Taken 
together the nymphal and adult infection 
rates are anomalous. The small sample size, 
reflective of the sparse tick population, 
could, due to chance, give a somewhat dis- 
torted depiction of the actual NIST situa- 
tion. 

According to Andrén (1994), as habitats 
are degraded and fragmented, some com- 
ponent host species tend to disappear. Ost- 
feld and Keesing (2000) and LoGiudice et 
al. (2003) suggest that “‘species-poor com- 
munities tend to have mice, but few other 
hosts, whereas species-rich communities 
have mice, plus many other hosts, which 
should dilute the impact of mice by feeding 
but rarely infecting ticks.”’ Thus, in frag- 
mented habitats larval and nymphal ticks 
that feed successfully are more likely to do 
sO, on a competent reservoir host and be- 
come infected (Ostfeld and Keesing 2000, 
LoGiudice et al. 2003). At NIST, the infec- 
tion rate for B. burgdorferi was low for un- 
fed nymphal /. scapularis and high for un- 
fed adults, contrary in part to what might 
be expected with the aforementioned sce- 
nario. Population density data for P. leu- 
copus and other small vertebrate hosts are 
needed for a clearer understanding of the 
epidemiology of B. burgdorferi at NIST. 
Deer damage to leaf litter and understory 
vegetation may contribute to microclimatic 


S75 


conditions detrimental to free-living ticks, 
adding further complexity to the situation 
at NIST. 

Despite a ~25% drop from its peak, the 
deer population at NIST remains overly 
dense. With continued reduction of the deer 
herds, the habitat at NIST may more fully 
recover in time. A detailed ecological study, 
assessing host species diversity and abun- 
dance, is needed to elucidate tick-host-path- 
ogen relationships in the greatly deer-per- 
turbed ecosystems at NIST. 

Acknowledgments.—This survey would 
not have been possible without Rhonda 
Hurt, formerly Wildlife Manager of the Na- 
tional Institute of Standards and Technolo- 
gy, Gaithersburg, MD. We thank her for her 
generous cooperation and assistance, and 
for her observations on the changes over 
time in the vertebrate fauna at NIST. We 
also express our appreciation to Kenneth 
Young, Art Abrams, Hasani Martin, and 
Michael Theis, USDA, ARS, Animal Par- 
asitic Diseases Laboratory, Beltsville, MD, 
for sampling and counting ticks. 


LITERATURE CITED 


Andrén, H. 1994. Effects of fragmentation on birds 
and mammals in landscapes with different pro- 
portions of suitable habitat: A review. Oikos 71: 
355-366. 

Coté, S. D., T. P. Rooney, J.-P. Tremblay, C. Dussault, 
and D. M. Waller. 2004. Ecological impacts of 
deer overabundance. Annual Review of Ecology, 
Evolution, and Systematics 35: 113-147. 

Cyn ES Me Gs Jenkins ReaD Hall Ew sWMasters: 
and G. A. McDonald. 2005. Improving the spec- 
ificity of 16S-rDNA-based polymerase chain re- 
action for detecting Borrelia burgdorferi sensu 
lato-causative agents of human Lyme disease. 
Journal of Applied Microbiology 98: 962—970. 

Daniels, T. J., R. C. Falco, and D. Fish. 2000. Esti- 
mating population size and drag efficiency for the 
blacklegged tick (Acari: Ixodidae). Journal of 
Medical Entomology 37: 357—363. 

Harlan, H. J. and W. A. Foster. 1990. Micrometeoro- 
logical factors affecting field host-seeking activity 
of adult Dermacentor variabilis (Acari: Ixodidae). 
Journal of Medical Entomology 27: 471—479. 

Lane, R. S., J. Piesman, and W. Burgdorfer. 1991. 
Lyme borreliosis: Relation of causative agent, its 
vectors and hosts in North America and Europe. 
Annual Review of Entomology 36: 587—609. 


976 


LoGiudice, K., R. S. Ostfeld, K. A. Schmidt, and E 
Keesing. 2003. The ecology of infectious disease: 
Effects of host diversity and community compo- 
sition on Lyme disease risk. Proceedings of the 
National Academy of Sciences 100: 567-571. 

Mount, G. A. and D. G. Haile, and E. Daniels. 1997. 
Simulation of blacklegged tick (Acari: Ixodidae) 
population dynamics and transmission of Borrelia 
burgdorferi. Journal of Medical Entomology 34: 
461-484. 

Main, A. J., A. B. Carey, M. G. Carey, and R. H. 
Goodwin. 1982. Immature /xodes dammini (Acari: 
Ixodidae) on small mammals in Connecticut. Jour- 
nal of Medical Entomology 19: 655—664. 

Ostfeld, R. S. and E Keesing. 2000. The function of 
biodiversity in the ecology of vector-borne zoo- 
notic diseases. Canadian Journal of Zoology 78: 
2061-2078. 

Telford, S. R., II, T. N. Mather, S. I. Moore, M. L. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


Wilson, and A. Spielman. 1988. Incompetence of 
deer as reservoirs of the Lyme disease spirochete. 
American Journal of Tropical Medicine and Hy- 
giene 39: 105-109. 

Tilghman, N. G. 1989. Impacts of white-tailed deer on 
forest regeneration in northwestern Pennsylvania. 
Journal of Wildlife Management 53:524—532. 

Wilson, M. L., G. H. Adler, and A. Spielman. 1985. 
Correlation between abundance of deer and that 
of the deer tick, Ixodes dammini (Acari: Ixodidae). 
Annals of the Entomological Society of America 
78: 172-176. 


J. E Carroll and 1 lb. Cyr, USS) Depari- 
ment of Agriculture, Agricultural Research 
Service, Animal Parasitic Diseases Labo- 
ratory, Beltsville, MD 20705, U.S.A. (e- 
mail: jcarroll@ anri.barc.usda.gov) 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 977-978 


NOTE 


New and Additional Records of Mayflies (Ephemeroptera) from the Southwestern 
United States, including a New Country Record 


There are currently 64 species of may- 
flies (Ephemeroptera) known from Arizona 
(Lugo-Ortiz and McCafferty 1995a) and 78 
species documented from New Mexico 
(McCafferty et al. 1997). These numbers 
include various recent synonymies and ad- 
ditions to the two states (Lugo-Ortiz and 
McCafferty 1995b, McCafferty and Sill- 
dorff 1998, Baumgardner and McCafferty 
2000, Jacobus and McCafferty 2002, 
McCafferty et al. 2002, Jacobus and Mc- 
Cafferty 2003, Wiersema and McCafferty 
2004). An additional two species are herein 
reported from Arizona, one species from 
New Mexico (and its first report in the 
United States), and one species newly re- 
ported from Texas. All specimens listed are 
deposited in the Texas A&M University In- 
sect Collection. 


BAETIDAE 


Camelobaetidius mexicanus (Traver and 
Edmunds).—This species is known from 
numerous localities throughout Mexico, and 
scattered localities in Texas and Kansas 
(Lugo-Ortiz and McCafferty 1995b), and 
most recently from Idaho (Lester et al. 
2002). 

New state records: AZ: Yavapai Co., West 
Clear Creek at Clear Creek Campground, 
ca. 1 mi. from Hwy. 260, ca. 4 mi SE Camp 
Verde (N34°30'54.7”; W111°45'45.4", Elev. 
3,270 ft.), 26.v.2004, DE Baumgardner, | 
larva (DB 04-32). AZ: Graham Co., Gila 
Reweat Ft | Thomas i(N33°02'59:0": 
W109°58'01.2”, Elev. 2,660 ft.), 27.v.2004, 
DE Baumgardner, 3 larvae (DB 04-38). 

Camelobaetidius warreni (Traver and 
Edmunds).—This is a wide-ranging baetid 
mayfly, known from as far south as Costa 
Rica, and throughout much of the western 
United States (Lugo-Ortiz and McCafferty 


1995b). Its presence in Texas was not un- 
expected. 

New state record: TX: Jeff Davis Co., 
Limpia Ck. @ Hwy. 118, ca. 1.5 mi. W. Ft. 
Davis (N30°36'05"; W103°54'57”: Elev. 
4,980 ft.), 15.v.2002, DE Baumgardner, 22 
larvae (DB 02-08). 


HEPTAGENIIDAE 


Epeorus albertae (McDunnough).—This 
is acommon and widely distributed western 
Intermountain species (McCafferty et al. 
1993). It was known previously from New 
Mexico (McCafferty et al. 1997) and Col- 
orado (McCafferty et al. 1993). 

New state records: AZ: Navajo Co., North 
Fork White River at SR 55, Whitewater 
(town) (N33°49'47.4"; W109°57'36.5”, Elev. 
5,170 ft.), 23.v.2004, DE Baumgardner, 3 
larvae (DB 04-26). AZ: Gila Co., Christo- 
pher Creek at Hwy. 260, Christopher Ck. 
Campground, ca. 20 mi. E. Payson 
(N34°18'27 17 W119 02 27-3" Blevas:500 
ft.), 26.v.2004, DE Baumgardner, 47 larvae 
(DB 04-35). 


LEPTOPHLEBIIDAE 


Traverella promifrons Lugo-Ortiz and 
McCafferty.—Traverella promifrons was 
described originally by Allen (1973) as 
“Traverella sp. C’’, based upon larvae from 
Mexico and Honduras. It was described for- 
mally by Lugo-Ortiz and McCafferty 
(1996) from larvae collected in Belize and 
Costa Rica. It was documented most re- 
cently in Guatemala (McCafferty et al. 
2004). Its presence in New Mexico repre- 
sents a new record for the USA. 

New country record: NM: Catron Co., 
Willow Creek at Willow Creek Camp- 
ground (Hwy. 159), ca. 35 mi. E. Glen- 
wood, Gila National Forest (N33°24'37"; 


978 


W 108°34'20", Elev. 7,400 ft.), 17.v.1987, 
R. Leschen, 5 larvae. 

Acknowledgments.—I thank Michael 
Meyer (Purdue University) for confirming 
the identification of Traverella promifrons, 
David Bowles (National Park Service) for 
the donation of material, and W.P Mc- 
Cafferty (Purdue University) for reviewing 
an earlier draft of the manuscript. 


LITERATURE CITED 


Allen, R. K. 1973. Generic revisions of mayfly 
nymphs. |. Traverella in North and Central Amer- 
ica (Leptophlebiidae). Annals of the Entomologi- 
cal Society of America 66: 1287-1295. 

Baumgardner, D. E. and W. P. McCafferty. 2000. Lep- 
tohyphes zalope (Ephemeroptera: Leptohyphidae): 
A polytypic North and Central American species. 
Entomological News 111: 49-59. 

Jacobus, L. M. and W. P. McCafferty. 2002. New syn- 
onyms for three North American Ephemeroptera 
species. Journal of the Kansas Entomological So- 
ciety 75: 59-60. 

. 2003. Revisionary contributions to North 
American Ephemerella and Serratella (Ephemer- 
optera: Ephemerellidae). Journal of the New York 
Entomological Society 111: 174-193. 

Lester, G. T., W. P. McCafferty, and M. R. Edmondson. 
2002. New mayfly (Ephemeroptera) records from 
Idaho. Entomological News 113: 131—136. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1995a. An- 
notated inventory of the mayflies of Arizona. En- 
tomological News 106: 131-140. 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


. 1995b. Taxonomy of the North and Central 
American species of Camelobaetidius (Ephemer- 
optera: Baetidae). Entomological News 106: 178— 
192. 


1996. New species of Leptophlebiidae 
(Ephemeroptera) from Mexico and Central Amer- 
ica. Annales de Limnologie 32: 3-18. 

McCafferty, W. P., R. S. Durfee, and B. C. Kondratieff. 
1993. Colorado mayflies (Ephemeroptera): An an- 
notated inventory. Southwestern Naturalist 38: 
252-274. 

McCafferty, W. P., C. R. Lugo-Ortiz, and G. Z. Jacobi. 
1997. Mayfly fauna of New Mexico. Great Basin 
Naturalist 57: 283-314. 

McCafferty, W. P. and E. L. Silldorff. 1998. Reared 
association and equivalency of Baetis adonis and 
B. caelestis (Ephemeroptera: Baetidae). Entomo- 
logical News 109: 261—265. 

McCafferty, W. P., M. D. Meyer, and G. T. Lester. 
2002. Significant range extensions for southwest- 
ern Nearctic mayflies (Ephemeroptera: Baetidae). 
Entomological News 113: 211-214. 

McCafferty, W. P, D. E. Baumgardner, and J. L. 
Guenther. 2004. The Ephemeroptera of Central 
America. Part 1: Guatemala. Transactions of the 
American Entomological Society 130: 201—219. 

Wiersema, N. A. and W. P. McCafferty. 2004. New 
specific synonyms and records of North American 
Centroptilum and Procloeon (Ephemeroptera: 
Baetidae). Entomological News 115: 121-128. 


David E. Baumgardner, Department of 
Entomology, Texas A&M University, Col- 
lege Station, TX 77843-2475, U.S.A. (e- 
mail: dbaumgardner@ tamu.edu) 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, p. 979 


NOTE 


Apobaetis futilis (McDunnough), a New Combination in Nearctic 
Baetidae (Ephemeroptera) 


The species Pseudocloeon futile Mc- 
Dunnough (Ephemeroptera: Baetidae) was 
transferred to the genus Plauditus Lugo-Or- 
tiz and McCafferty along with some other 
Nearctic species formerly assigned to the 
genus Pseudocloeon Klapalek (Lugo-Ortiz 
and McCafferty 1998). Plauditus futilis has 
been reported from southern Alberta 
(McDunnough 1931), central Texas (Baum- 
gardner et al. 1997), and southeastern 
Oklahoma (Baumgardner and Kennedy 
1999). The larva remains unknown. 

Based on our examination of the holo- 
type and additional material, we recognize 
that Plauditus futilis instead belongs to the 
relatively infrequently collected genus Apo- 
baetis Day, based on characters that in- 
clude: anteriorly divergent turbinate eyes; 
the metanotum with a broad, dorsally pro- 
jecting, metascutellar hump and a deeply 
emarginate posterior margin; a prominent, 
blunt projection between the genital for- 
ceps; and divergent genital forceps, each of 
which has an elongate basal segment and a 
second segment with relatively uniform 
thickness (Edmunds et al. 1976, Waltz and 
McCafferty 1986, Meyer and McCafferty 
2003). Therefore, we transfer Plauditus fu- 
tilis to Apobaetis, as Apobaetis futilis 
(McDunnough), new combination. Two 
other Apobaetis species are known from 
North America: A. etowah (Traver) and A. 
lakota McCafferty (McCafferty 2000). 

Material examined.—HOLOTYPE, male 
adult, Alberta, Milk River, August 18, JH 
Pepper [Canadian National Collection of 
Insects, Ottawa, Ontario, Canada]; 11 male 
adults, 5 female adults, Oklahoma, Push- 
mataha Co., Kiamichi R at unnamed road 


in Tuskahoma, 6-VIII-1993, DE Baumgard- 
ner [Purdue University Entomological Re- 
search Collection, West Lafayette, Indiana, 
USA]. 

Acknowledgments.—David Baumgard- 
ner (College Station, Texas) donated some 
material. This study was funded in part by 
CanaColl Grant 178 to LMJ. 


LITERATURE CITED 


Baumgardner, D. E. and J. H. Kennedy. 1999. Mayflies 
(Insecta: Ephemeroptera) of the Kiamichi River 
Watershed, Oklahoma. Journal of the Kansas En- 
tomological Society 72: 297-305. 

Baumgardner, D. E., J. H. Kennedy, and B. C. Henry, 
Jr. 1997. New and additional records of Texas 
mayflies (Ephemeroptera). Transactions of the 
American Entomological Society 123: 55—69. 

Edmunds, G. EF, Jr., S. L. Jensen, and L. Berner. 1976. 
The Mayflies of North and Central America. Uni- 
versity of Minnesota Press, Minneapolis, 330 pp. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1998. A new 
North American genus of Baetidae (Ephemerop- 
tera) and key to Baetis complex genera. Entomo- 
logical News 109: 345-353. 

McCafferty, W. P. 2000. A new Nearctic Apobaetis 
(Ephemeroptera: Baetidae). Entomological News 
111: 265-269. 

McDunnough, J. 1931. New species of North Ameri- 
can Ephemeroptera. The Canadian Entomologist 
63: 82-93. 

Meyer, M. D. and W. P. McCafferty. 2003. New syn- 
onym of Apobaetis etowah (Traver) (Ephemerop- 
tera: Baetidae). The Pan-Pacific Entomologist 79: 
249. 

Waltz, R. D. and W. P. McCafferty. 1986. Apobaetis 
etowah (Traver), a new combination in Nearctic 
Baetidae (Ephemeroptera). Proceedings of the En- 
tomological Society of Washington 88: 191. 


Luke M. Jacobus and W. P. McCafferty, 
Department of Entomology, Purdue Uni- 
versity, West Lafayette, IN 47907, U.S.A. 
(email: konchu@ purdue.edu) 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 980-981 


Book REVIEW 


A Field Guide and Identification Manual 
for Florida and Eastern U.S. Tiger Bee- 
Hleseaul Mes Choates Im 2003.) 197 spp: 
U.S. $34.95. University Press of Florida, 
Gainesville, FL. ISBN 0-8130-2583-4. 


Tiger beetles have long been one of the 
most popular insect groups with amateur 
collectors and professional entomologists. 
In recent years, these beetles have been dis- 
covered by a much broader audience, in- 
cluding general naturalists, conservation- 
ists, and wildlife biologists. This increased 
interest has helped drive the publication of 
state and regional field guides devoted to 
tiger beetles (e.g., Graves and Brzoska 
1991, Knisley and Schultz 1997, Leonard 
and Bell 1999, Acorn 2001). 

The most recent contribution to this 
genre is an attractive new field guide to 
eastern U.S. tiger beetles by Paul Choate of 
the University of Florida. This book treats 
the species of tiger beetles known to occur 
east of the Mississippi River, with a special 
focus on the fauna of Florida. Florida is a 
particularly appropriate choice for a tiger 
beetle guide with its 26 recorded species, 
including 5 endemics or near-endemics (one 
of these species is also found in southeast- 
ern Georgia, while two others also occur in 
Cuba). Among modern tiger beetle workers, 
Choate is uniquely qualified to write this 
book; he has studied the tiger beetles of 
Florida since 1975 and actually discovered 
and named one of the state’s rarest species, 
the endemic Cicindela highlandensis 
Choate. 

For those who are new to tiger beetles, 
this book provides a wealth of helpful in- 
formation. Every species is illustrated by a 
dorsal habitus photograph of a pinned spec- 
imen (most of which are in color) and there 
are brief written accounts which describe 
each species’ habitat associations. For many 
species, the author includes color photo- 
graphs of live beetles in characteristic hab- 


itats. General photographs of beetle habitats 
and collecting areas are also provided. Dis- 
tribution maps are presented for each spe- 
cies: by county for the species which occur 
in Florida, and also by state for those spe- 
cies which are more widely distributed. The 
author also shares helpful hints for collect- 
ing and photographing tiger beetles. 

Professional entomologists and advanced 
amateurs will also find much that is useful 
in this work, including a technical charac- 
terization of the subfamily Cicindelinae, 
keys to Nearctic tiger beetle genera, a syn- 
opsis of the most important taxonomic lit- 
erature for the U.S. fauna, a discussion of 
species and subspecies categories in Cicin- 
delinae, a illustrated review of important 
morphological characters for this subfami- 
ly, and a key to adults of species of tiger 
beetles from the eastern U.S. In addition, 
there is a fascinating discussion of the geo- 
logical history of Florida which provides 
valuable insights into the evolution of the 
peninsula’s unique tiger beetle fauna. 

The author also provides a brief and 
thoughtful discussion of tiger beetle con- 
servation issues, focusing on the globally 
rare and highly localized species C. high- 
landensis, C. marginipennis DeJean, and C. 
puritana Horn. 

There is relatively little that could be im- 
proved in future editions of this guide. Most 
of the illustrations are excellent, but unfor- 
tunately the habitus photograph of Mega- 
cephala virginica (L.) is cropped too close- 
ly. Other habitus photos have problems 
with illumination or with color reproduc- 
tion. Some of the species which are metallic 
green in the field appear dark blue or violet 
in these photographs. A more critical omis- 
sion is the lack of a scale bar or indication 
of body size for any of the species. 

The coverage of subspecific taxa is not 
consistent. For example, the subspecies of 
C. rufiventris DeJean are treated, but not 


VOLUME 107, NUMBER 4 


those of C. patruela DeJean. With other 
species such as C. dorsalis Say and C. scu- 
tellaris Say, some of the eastern subspecies 
are illustrated but not others. In a field 
guide which is intended for a more general 
audience, it would have been helpful to pro- 
vide illustrations of ail of the eastern color 
morphs for the most variable species (e.g., 
C. scutellaris), even if some of those forms 
are not currently recognized as valid sub- 
species. 

Two species which might be included in 
future editions of this guide are C. denikei 
Brown, found in northern Minnesota (Cof- 
fin and Pfannmuller 1988, Pearson et al. 
1997, Freitag 1999), and C. pamphila 
LeConte, which has occasionally been re- 
ported from Louisiana and Mississippi 
(Graves and Pearson 1973). 

Aside from these minor omissions, this is 
a beautiful and well-produced field guide 
which should occupy a prominent place in 
the library of all cicindelophiles. More im- 
portantly, it will serve for years to come as 
a valuable companion on countless field 
trips in search of tiger beetles. 


LITERATURE CITED 


Acorn, J. 2001. Tiger Beetles of Alberta: Killers on 
the Clay, Stalkers on the Sand. University of Al- 
berta Press, Edmonton, xix + 120 pp. 


981 


Coffin, B. and L. Pfannmuller. 1988. Minnesota’s En- 
dangered Flora and Fauna. University of Minne- 
sota Press, Minneapolis, Minnesota, xv + 473 pp. 

Freitag, R. 1999. Catalogue of the Tiger Beetles of 
Canada and the United States. NRC Research 
Press, Ottawa, vii + 195 pp. 

Graves, R. C. and D. W. Brzoska. 1991. The tiger bee- 
tles of Ohio (Coleoptera: Cicindelidae). Bulletin 
of the Ohio Biological Survey (New Series) 8(4): 
i-iv + 1-32. 

Graves, R. C. and D. L. Pearson. 1973. The tiger bee- 
tles of Arkansas, Louisiana, and Mississippi (Co- 
leoptera: Cicindelidae). Transactions of the Amer- 
ican Entomological Society 99(2): 157—203. 

Knisley, C. B. and T. D. Schultz. 1997. The Biology 
of Tiger Beetles and a Guide to the Species of the 
South Atlantic States. Virginia Museum of Natural 
History, Martinsville, Virginia, viii + 210 pp. 

Leonard, J. G. and R. T. Bell. 1999. Northeastern Tiger 
Beetles: A Field Guide to Tiger Beetles of New 
England and Eastern Canada. CRC Press, Boca 
Raton, x + 176 pp. 

Pearson, D. L., T. G. Barraclough, and A. P. Vogler. 
1997. Distributional maps for North American 
species of tiger beetles (Coleoptera: Cicindelidae). 
Cicindela 29: 33-84. 


Jonathan R. Mawdsley, Research Asso- 
ciate, Department of Entomology, National 
Museum of Natural History, Smithsonian 
Institution, P.O. Box 37012, Washington, 
D.C. 20013-7012, U.S.A. (e-mail: jonathan. 
mawdsley @nfwf.org) 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 982—983 


Book REVIEW 


The Insects: Structure, Function and Bio- 
diversity. Dunston P. Ambrose. 2004. 821 
pp. Kalyani Publishers, Ludhiana, India 
(infokalyanipublishers.com). ISBN 81- 
272-1853-7. 


An excellent textbook of general ento- 
mology has just been published, from a part 
of the world where entomology is of enor- 
mous importance yet where education and 
training in basic entomology have not been 
sufficiently emphasized. This book, “The 
Insects: Structure, Function and Biodiver- 
sity,’ is by one of the preeminent students 
of the Reduviidae (Hemiptera). Its title, not 
(I believe) by coincidence, is reminiscent of 
R. E Chapman’s “The Insects: Structure 
and Function’’; and indeed the first is to 
some extent modeled on the second. The 
first 400, and the final 120 (of 820) pages 
of Ambrose’s book, like all of Chapman’s 
book, eschews the usual insect order-based 
organization, and instead is organized by 
system (“‘Excretory,” “‘Muscular,”’ etc.). 
Most of these chapters are devoted to phys- 
iology and, to a lesser extent, to biochem- 
istry (and endocrinology). But there is a 
long chapter (50 pp.) on behavior, two oth- 
ers on phylogeny and systematics (50 pp.), 
and 50 pages more on ecology and conser- 
vation. The book ends with nearly 70 pages 
on collecting insects, and studying them in 
the field and in the laboratory. Many of the 
topics here are not mentioned in other texts, 
and some of the topics mentioned else- 
where are not covered in nearly the detail 
as they are here. These two chapters are ex- 
cellent, should inspire and guide young en- 
tomologists, and are of value everywhere, 
not just in India. 

This universal value is true of the book 
itself. This is by no means a regional or 
parochial text. Many of the examples cho- 
sen are of Indian insects to be sure, but 
most are not. The world literature has been 
scoured and used, as has of course the au- 


thor’s own extensive work on Indian redu- 
vilds (systematics, biology, ecology, phys- 
iology). The many references are mostly 
from the last few decades of the last cen- 
tury, indicating the years spent by the au- 
thor on the book. As further indication of 
the universality of the book, the 100+ pag- 
es devoted to the groups of insects do not 
emphasize the Indian groups, but give brief 
accounts of all major (and the more inter- 
esting minor) families (somewhat on the 
model of the various Borror et al., editions). 

Coverage of topics is remarkably com- 
plete. For example, the chapter ““‘Excretory 
System,’ covers in 8.5 pages these topics 
(subheads): Malpighian Tubules; Other Or- 
gans; Other Functions of Malpighian Tu- 
bules; Diffusion, Absorption and Resorp- 
tion [of salts and water]; Diuresis and Insect 
Urine; Gut Motility; Nitrogenous Excre- 
tion; Storage Excretion; Diet Influence; and 
Detoxification. These 8.5 pages include 3 
figures, and 2 tables, and are followed by 
28 references (including research papers 
from 1965 through 1994). 

Here and there in the book a few passag- 
es mention the literature, or merely refer to 
it (e.g., in ““Cytogenetics’’), rather than dis- 
cussing or explaining it. But this cursory 
treatment is very much the exception. 

This book will be followed by another on 
the economic aspects of entomology. How- 
ever, the present volume stands alone as a 
remarkable text—remarkable in the number 
of subjects covered, in the depth to which 
those subjects are treated, in the clarity of 
the writing, and in the aptness of the figures 
and tables. This book rivals some American 
texts, and surpasses others: It would be a 
fine text for serious American entomology 
students. This achievement is all the more 
remarkable when one considers the diffi- 
culties with time, support (financial and lo- 
cal), and materials (library especially) that 
its author has, so successfully, overcome. 


VOLUME 107, NUMBER 4 


The book is divided into five sections, as 
follows: Structure and Functions (13 chap- 
ters, ~230 pp.), Behaviour (1 chapter, 54 
pp.), Biosystematics (7 chapters, 200 pp.), 
Ecology (2 chapters, 64 pp.), and Experi- 
mental Entomology (2 chapters, 65 pp.). It 
concludes with a 67-page (!) Glossary 
(which alone rivals several dictionaries of 
entomology), a 45-page Subject Index, and 
a 33-page Taxonomic Index. And the book 


983 


opens with a comprehensive Introduction 
that lays out the importance of insects, and 
provides a quick overview of the book it- 
self. 

I recommend this book for browsing, for 
closer reading and, indeed, for teaching. 


Carl W. Schaefer, Department of Ecology 
and Evolutionary Biology, University of 
Connecticut, Storrs, CT 06269-3043, U.S.A. 
(e-mail: schaefer@ uconnvm.uconn.edu) 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 984-987 


SOCIETY MEETINGS 


1,088th Regular Meeting—January 6, 2005 


The 1,088th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby 
room of the National Museum of Natural 
History, Washington, D.C., by Past Presi- 
dent Eric Grissell, at 7:04 p.m., standing in 
for new President Jason Hall. The meeting 
was attended by 18 members and 13 guests. 
The minutes of the 1,087th meeting were 
approved as read. 

There were three new applicants for 
membership: Donald Wright, Penelope Gul- 
lan, and Deborah Mead. No new members 
were present and no visitors were intro- 
duced. 

For exhibits, Edd Barrows passed around 
a mystery parasitized cottony mass for 
identification—probably braconids. Eric 
Grissell passed around Rhus peppercorns 
related to his presentation. 

Dave Furth introduced the evening’s 
speaker, outgoing ESW President Eric Gris- 
sell, reading him a good-bye poem. Dr. 
Grissell then told the story of his life and 
times, wrought with his characteristic dry 
black humor, in a talk entitled, ‘‘From Cal- 
ifornia to South Africa on a Wild Rhus 
Chase, or How to Nearly Solve a Problem 
in 40 Years.”” The study system included 
two species of seed-feeding torymids, two 
plant species, and three continents. The 
mystery began in 1961, when an African 
species of a Megastigmus wasp hatched 
from a Brazilian Schinus plant, despite the 
fact that the genus Megastigmus is absent 
in South America and Schinus is not native 
to Africa. Schinus has been exported world- 
wide and is now a huge problem in the 
Florida everglades. Two putative species of 
Megastigmus occurred on native Rhus and 
imported Schinus in South Africa. Through 
extensive collecting, rearing, and reciprocal 
host tests during a 3-month trip to South 
Africa, Dr. Grissell showed that the two 


wasp species are synonyms. What remains 
unknown is the history of the African Me- 
gastigmus—Brazilian Schinus relationship: 
was it established in Africa and transported 
to the United States or did the wasp and 
plant arrive independently and the same re- 
lationship developed in parallel on the two 
continents? This is a smaller problem than 
he started with, and the one he passes on to 
the next generation of suckers. 

The meeting was adjourned around 8:30 
p-m. Refreshments were provided by the 
Society. 


Respectfully submitted, 
Stuart H. McKamey, 
Recording Secretary 


1,089th Regular Meeting— 
February 3, 2005 


The 1,089th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby 
room of the National Museum of Natural 
History, Washington, D.C., by President Ja- 
son Hall at 7:07 p.m. The meeting was at- 
tended by 16 members and 15 guests. The 
minutes of the 1,088th meeting were ap- 
proved as read. 

There were no new applicants for mem- 
bership or new members present but four 
visitors were introduced. 

In miscellaneous business, President- 
Elect Steve Lingafelter discussed budgetary 
considerations of holding the ESW Annual 
Banquet in the Natural History Museum. 

For exhibits, Daniel Perez displayed 
posters from recent meetings on a project 
related to his, on the arthropods of Hispan- 
iola. Dave Furth had a two-volume set of a 
series of Australian catalogs, including 
“Mites of Australia: A Checklist and Bib- 
liography” by R.B. Halliday, and “*Oriba- 
tid Mites: a Catalogue of Australian Gen- 
era and Species” by M.J. Colloff and R.B. 
Halliday. Warren Steiner had a new popular 


VOLUME 107, NUMBER 4 


book, “Go Wild in New York City” by B. 
Matsen, a summary of natural areas and 
common organisms, and he mentioned 
plans for a Washington, D.C. version. 

Dave Furth introduced the evening’s 
speaker, Smithsonian Research Associate 
Dr. Daniel Perez, who presented his work 
on a “Survey of the Orthopteroid Insects of 
Hispaniola.”” He began by summarizing the 
complex geological history of the island, its 
four major mountain ranges and consequent 
faunistic regions, and its startlingly high 
levels of endemism in plants (30%), noting 
that its insect fauna is the least known of 
the Greater Antilles. Before beginning his 
survey in 1992, only three orthopterists had 
collected there. With the help of an NSF 
grant from 2002—2004, the survey collected 
over 15,000 orthopteroids from a wide, rep- 
resentative swath of the Dominican Repub- 
lic. Dr. Perez’s work more than tripled the 
number of known grasshoppers, and the 
project added eight undescribed genera of 
katydids, many new species of walking 
sticks, and the first Caribbean record of the 
family Pyrgomorphidae. The specimens are 
divided among the National Museum of 
Natural History, the Philadelphia Academy 
of Sciences, and the Dominican Republic 
Nationa! Museum, to which the project con- 
tributed four cabinets with associated draw- 
ers and unit trays. 

The meeting was adjourned at 8:40 p.m. 
Refreshments were provided by the Society. 


Respectfully submitted, 
Stuart H. McKamey, 
Recording Secretary 


1,090th Regular Meeting—March 3, 2005 


The 1,090th regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby 
room of the National Museum of Natural 
History, Washington, D.C., by President Ja- 
son Hall at 7:02 p.m. The meeting was at- 
tended by 17 members and 7 guests. The 
minutes of the 1,089th meeting were ap- 
proved with modification. 


\O 
(oe) 
Nn 


There were no new applicants for mem- 
bership or new members present, and no 
visitors were introduced. 

For exhibits, Edd Barrows had the book 
“Jungle Bugs: Masters of Camouflage and 
Mimicry” by B. Purser. 

John Brown introduced the evening’s 
speakers: Edd Barrows, Christaine Bird, 
Daniel Balogh, Daniel Kjar, and Catherine 
McCall, all staff, students, or associates with 
the Laboratory of Entomology and Biodi- 
versity, Georgetown University. Their pre- 
sentation, “* “Arthropoversity’ of the Capital 
Area, A—Z: Acari to Zoraptera”’ provided re- 
sults of years of study on the fauna of Rock 
Creek Park and, principally, Dyke Marsh. 
Their searchable database of more than 
12,000 entries on over 2,500 species is on- 
line at http://biodiversity.georgetown.edu. 
There were four facets of the evening’s pre- 
sentations: Edd Barrows introduced the 
team, study areas and Malaise traps set 
across three habitats; Ms. McCall discussed 
the richness and abundance patterns of noc- 
tuids (73 species); Ms. Bird and Mr. Balogh 
discussed the same issues, but with respect 
to ichneumonids (about 150 species); and 
Daniel Kjar addressed the ant fauna with re- 
spect to the possible effect of non-native 
plants. The latter project employed 4 years 
of pit-fall and Berlese funnel samples at 60 
random sites. These were combined with flo- 
ral and soil data and visualized using 
Arcview software. He found regularity in 
seasonal abundance across some ant species, 
33 ant species total, and the second U.S. re- 
cord of a Japanese ant. As the percent in- 
vasive plants increased, so did ant diversity, 
but soil moisture and tree richness explained 
most of the trend. He used Discover Life 
software to provide a key to the ants and to 
map their occurrence. 

The meeting was adjourned at 8:10 p.m. 
Refreshments were provided by the Society. 


Respectfully submitted, 
Stuart H. McKamey, 
Recording Secretary 


986 


1,091st Regular Meeting—April 7, 2005 


The 1,091st regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby 
room of the National Museum of Natural 
History, Washington, D.C., by President Ja- 
son Hall at 7:10 p.m. The meeting was at- 
tended by 19 members and 10 guests. The 
minutes of the 1,090th meeting were ap- 
proved with modification. 

There were no new applicants for mem- 
bership or new members present, but five 
visitors were introduced. 

In miscellaneous business, President- 
Elect Steve Lingafelter announced that the 
Annual Banquet will be held June 2 at the 
Natural History Museum. 

For exhibits, Ed Cohen had prints of un- 
identified insects. Two ESW members not- 
ed the recent death of the British, world fa- 
mous entomologist Miriam Rothschild. 
Dave Furth had four new books: “*Hoverfly 
Fauna of the Bakony Region (Diptera: Syr- 
phidae)” by T. Sandor (in Hungarian), 
“Catalogo dei Lepidoptera Tortricidae del- 
la Fauna Italiana” by P. Trematerra, “‘Las 
Polillas de la Fruta en Chile (Lepidoptera: 
Torticidae; Pyralidae)” by R.H. Gonzalez 
Rodriguez, and “World Catalogue of In- 
sects, Vol. 3 Dytiscidae (Coleoptera) by 
A.N. Nilsson. 

Dave Furth introduced the evening’s 
speaker, Dr. Michael Engel of the Ento- 
mology Division, University of Kansas, 
who gave a presentation entitled “600 Mil- 
lion Years on Six Legs.” This consisted of 
an overview of the major diversifications 
and extinctions of all hexapods, including a 
Devonian, extinct marine taxon that appears 
to be the sister-group to insects (that 1s, 
closer than the extant Entognatha), against 
a backdrop of the geological history of the 
Earth. Most of the excellent fossils from the 
Permian are from the Wellington Forma- 
tion, in Kansas. Major finds there include 
the largest insect (the dragonfly-like Me- 
ganeuropsis, with one wing measuring 13 
inches) and the earliest evidence of beetles 


PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


(an elytron of proto-Coleoptera from the 
Paleozoic). The end of the Permian was 
marked by massive extinction of higher 
taxa. Most modern, recognizable higher 
taxa evolved with the angiosperms in the 
Cretaceous period, from which there are 
many amber and mineral deposits. The rich- 
est is Burmese amber, dating to about 100 
my bp, and which provides the earliest fos- 
sil pompilid and zorapteran. Dr. Engel’s 
particular interest is in Apidae, whose evo- 
lution of eusociality is often debated. The 
apid fossils, which even include a few 
swarms, suggest a single evolutionary ori- 
gin. 

The meeting was adjourned at 8:31 p.m. 
Refreshments were provided by the Society. 


Respectfully submitted, 
Stuart H. McKamey, 
Recording Secretary 


1,092nd Regular Meeting—May 5, 2005 


The 1,092nd regular meeting of the En- 
tomological Society of Washington (ESW) 
was called to order in the Cathy Kerby 
room of the National Museum of Natural 
History, Washington, D.C., by President- 
Elect Steven Lingafelter at 7:10 p.m. The 
meeting was attended by 17 members and 
13 guests. The minutes of the 1,091st meet- 
ing were read and approved with modifi- 
cation by Acting Recording Secretary Da- 
vid Furth. 

There were no new applicants for mem- 
bership or new members present; five visi- 
tors were introduced. 

In miscellaneous business, President- 
Elect Lingafelter again reminded members 
that the banquet ticket deadline was soon 
and encouraged members to buy tickets for 
the ESW Banquet to be held on 2 June at 
the Natural History Museum. 

For exhibits Diane Calabrese talked 
about a 1931 book entitled ““Grasshoppers 
Come” by David Garnett. Dave Furth con- 
tinued his tradition by showing 3 new 
books from the Smithsonian’s Entomology 
library: “‘Cetoniidae of the World’—a 3 


VOLUME 107, NUMBER 4 


volume catalogue by M. Krajik; “‘Faune de 
Belgique Syrphides” by L. Verlinden; and 
‘Atlas of the Butterflies of the Iberian Pen- 
insula and Balearic Islands (Lepidoptera: 
Papilionoidea & Hersperoidea”’ by E. 
Garcia-Barros et al. 

Dave Furth introduced the evening’s 
speaker, Dr. Irina Brake, formerly a post- 
doctoral student at the Museum Koenig 
(Bonn) and doctoral student at the Frei 
Universitat Berlin, currently an E. Schlinger 
Postdoctoral Fellow and recent Global Bio- 
diversity Facility Fellow in the Department 
of Entomology at the Smithsonian. Dr. 
Brake delivered a talk entitled *““Thievishly 
and Trashy: the Biology of the Milichi- 
idae.”’ Her talk discussed the fascinating bi- 
ology of these acalypterate Diptera that are 
a monophyletic group of flies with three 
subfamilies and related to Chloropidae, 
Acartophthalmidae, and Carnidae. Differ- 
ences in various morphological characters 
such as the spermathecal duct separate these 
families. The adult flies feed on various 
flowers, especially of Umbelliferae, Aster- 
aceae, Asclepiadaceae, etc., and often act as 
pollinators. The basal clades are probably 


987 


nectar feeders and some feed on the hon- 
eydew of aphids. Some Milichia species are 
associated with ants and feed on regurgi- 
tated ant food [gross]. Some species of 
Pholeomyia are associated with fungus 
ants. In other species the females are clep- 
toparasites, that is, they feed on the cap- 
tured prey of mantids, spiders, pentatomids, 
dragonflies, etc. In some of these cases they 
actually hitch a ride on the predators and 
clean their mouthparts. Some of the larvae, 
especially in the basal clades, are saproph- 
agous and others in more advanced groups 
are coprophagous, mymecophilous and may 
live inside ant nests covering themselves 
with excreta [gross again]. A lively ques- 
tion and answer session followed the talk 
that drew out several mymecophilous mem- 
bers of the audience as well as a group of 
culicidologists from the Walter Reed Bio- 
systematics Unit who knew of milichiids 
from their perspective. 

The meeting was adjourned at 8:00 p.m. 
Refreshments were provided by the Society. 


Respectfully submitted, 
David G. Furth, 
Acting Recording Secretary 


PROC. ENTOMOL. SOC. WASH. 
107(4), 2005, pp. 988-995 


PROCEEDINGS 
of the 
ENTOMOLOGICAL SOCIETY 
of 
WASHINGTON 


Volume 107 


OFFICERS FOR THE YEAR 2005 


President Jason P. W. Hall 
President-Elect Stephen W. Lingafelter 
Recording Secretary Stuart H. McKamey 
Membership Secretary Hollis B. Williams 
Treasurer Michael G. Pogue 
Program Chairs John W. Brown, David G. Furth 
Custodian Jon A. Lewis 
Editor David R. Smith 
Past President E. E. Grissell 


Published by the Society 
WASHINGTON, D.C. 
2005 


TABLE OF CONTENTS, VOLUME 107 


ARTICLES 


ALBUQUERQUE, GILBERTO S.—See TAUBER, CATHERINE A. .....................0200055 
ANUBRIEWVHGs A —See NOMIRO VODA Ree Bs ee aeeiaae sacra osu biras case eee REE 
NSIEINING WRUEVNINE (SS SO NIMUNING, IN LOIS DNISVAN ccuntonsoooscoodsonnaenegddeceden ocgosasabboccsascoosces 
BALE GEORGE E:— See GAN D HIRIKCAN TATE ia KGa reer pes Ceeetee eee ee eee eC eeeEE eee eee 
BARBA-ALVAREZ, R. and J. BUENO-SORIA—New species of the genus Polycentropus 
Curtis @inchopteray Poly centropodidae)siromylVexi cOmeereer a sesteee eee ee eee eee eee eee nee eee 
BARROWS, EDWARD M., ANNE M. MCINTYRE, and OLIVER S. FLINT, JR.—Alderfly 
(Neuroptera: Sialidae) flight periods, sex ratios, and habitat use in a Virginia freshwater tidal 
manrsh wows torestaanditheliae COLONES see Ree ER ee eee eee ee een eee a eee nee eee eee 
BEAULIEU, FREDERIC and TERRY A. WHEELER—Diptera diversity in a homogeneous hab- 
itat: Brachycera associated with sedge meadows (Cyperaceae: Carex) in Quebec, Canada ..... 
BERENBAUMGIMs Re——See  EIARRIS @ Ne di len) 27s ee nee ree en earner Cee ener enn ee ee eee rere 
BRAILOVSKY, HARRY—A revision of the genus Acanthotyla Stal with the description of 
five new species and synonymical note (Heteroptera: Coreidae: Colpurini) .................... 


543 

21 
623 
917 


663 


VOLUME 107, NUMBER 4 989 


BRAILOVSKY, H.—A new species of Dalader Amyot and Serville, with a key to the Malay- 


sian species (Hemiptera: Heteroptera: Coreidae: Coreinae: Daladerini) ......................--- 883 
BROOKS, SCOTT E. and TERRY A. WHEELER—Ethiromyia, a new genus of Holarctic 

Dolichopidinae (Diptera: Dolichopodidae) ...................0 00 cc cece ccc ccc ceececeeeeeeeees 489 
BROWN TAERXeE See) POINARS GEORGE IRS eee ee aeeee eee eee eee ee eee 835 
BROWN: J OFINGW See RAZOWSEKI, JOZER) = ).5- 9 See ee ee ee 903 
BROWN, RICHARD L.—See PINKAEW, NANTASAK ...........000. 00.0 cece cece cece cee ccceeeeees 869 
BUENO-SORIA See BARBARA VAREZ Rete es tetee narrate nee eee ee eee 663 


BURCKHARDT, DANIEL, PAUL HANSON, and LUIS MADRIGAL—Diclidophlebia lucens, 
n. sp. (Hemiptera: Psyllidae) from Costa Rica, a potential control agent of Miconia calvescens 
(Melastomatacede) mmiETa wai te aie ge Fas te ites ee At Uh eS 741 
BURNS, JOHN M. and DANIEL H. JANZEN—What’s in a name? Lepidoptera: Hesperiidae: 
Pyrginae: Telemaides Hiibner 1819 [Pyrdalus Mabille 1903]: New combinations Telemaides 
corbulo (Stoll) and Telemaides oiculus (Mabille)—and more 
CAUMASURNONDER==SeelwAIMIAIN SIMI SUVA EAU ene see een it ee 623 
CAMBRA, ROBERTO A., VICTOR H. GONZALEZ, and WILLIAM T. WCISLO—Descrip- 
tion of the male, host associations, and new distribution records for Lophostigma cincta (du 


Buysson) x (lymenopterasMutilidac)preessaeeeeeeee eee eee eee ceeeen eee eee eee eee eee eee eee eeenee 229 
CARPENTER, TERRY L.—Notes on the life of Dr. Clara Southmayd Ludlow, Ph.D., medical 

EntomolooistliSS2 192A) ese UG se etaee deal cin Tea ae seamen Roe ale RER a eA Baa eee 657 
CHANDRAPATYA, ANGSUMARN—See PINKAEW, NANTASAK ..............0000000eeee eee 869 
GAPS, GUILEERM® L:—See PINTO; © MIGUEL 2222.22.52 2c ceccene- cee cesseeee sn eeeeeenseeens 3) 
COSTANTINO, JESSICA E.—See SCARBROUGH, AUBREY G. ................... 2222. ee eee 789 
COOKAERRY 1 ==See DERR, DAWN PR) ware. nleritaek see note terete coniclelale salstele Sorselom area seca seins 762 
DATES WiW==S Cer ZHIAIN Gs SYCAIEIN ieise5 Sec ncaic 8 Poa ate ac vie ard ereoieicttnnols oleletelaters atutots aiSerete care tateleretaererats 218 
DARSIE, RICHARD E, JR.—Key to the pupae of the mosquitoes (Diptera: Culicidae) of 

EL Ore La ee eas oT ae Te Cee tere eB EDTA Sw eteiale elerolent itdera leet cvayateve ul Bvotavclere etacera ale eta rneteterer toler 892 
DAVIDSON, JOHN A.—See MILLER, DOUGLASS R. ..... 00... cece eee eee eee eee eee eee eee ees 123 
DERR, DAWN P. and JERRY L. COOK—Morphology of the antenna of Caenocholax fenyesi 

Pierce (Strepsiptera: Myrmecolacidae) based on scanning electron microscopy ..............- 762 
DeWALT, R. EDWARD and B. D. HEINOLD—Summer emerging Ephemeroptera, Plecoptera, 

and Trichoptera of Abrams Creek, Great Smoky Mountains National Park .................... 34 
PIE DRICHME SE——SeevNOVINOWV AD Ve oe Re cese ere crciare otaiete caverns aictolorelovavctetatete svatateralctoratersratetateretetaratateiare 21 
DUARTE, MARCELO—See ROBBINS, ROBERT K.. ................ 0.0 c cece cece e cece eee eee eeeee 398 
EBENSASTRID=SeelGAMEZ-VIRUESYSAGRARTON wes sesn eee tee eee ee eee eee eee 642 


EDMISTON, JAMES E and WAYNE N. MATHIS—A review of two Nearctic species of the 
shore-fly genus Philygria Stenhammar: P. debilis Loew and P. nigrescens (Cresson) (Diptera: 


[SypyonalnGk®)) JoceqacoponpconccveqanponEoncodosouBoneBescacceoodonddcdaadtossobponseconsSsodHdoqascoasbedoc 7 
ESPINASA, LUIS—A new genus of the subfamily Cubacubaninae (Zygentoma: Nicoletiidae) 

fieoron, Were, IMIG SSD) SoccgosocoanacoocoLaaudaccadeopdocodooodn ARON SaosoonesogSoeacSconossSeccesasur 510 
FLINT, OLIVER S., JR.—See BARROWS, EDWARD MM. .............. 00. c cece cece cece ence eens 693 
GAIMARIG STEPHEN D=—SeehYAIN Gs DING) Wiese ere cee -elereielsiniete tie eleleielelan lelaleleleieteielalela\slelelel=iaisiel= 49 


GANDHI, KAMAL J. K., DANIEL W. GILMORE, GEORGE E. BALL, RALPH W. HOL- 
ZENTHAL, STEVEN A. KATOVICH, JESSICA J. KOEHLE, KIRK J. LARSEN, WIL- 
LIAM J. MATTSON, and STEVEN J. SEYBOLD—A review of ground beetle species (Co- 


leoptera: Carabidae) of Minnesota, United States: New records and range extensions ........ 917 
GAMEZ-VIRUES, SAGRARIO—See PEREDO, LUIS CERVANTES .............-0.00e0eeeeee 362 
GAMEZ-VIRUES, SAGRARIO and ASTRID EBEN—Predatory behavior of Repipta flavicans Stal 

(Hemiptera: Reduviidae), a natural enemy of Diabroticina (Coleoptera: Chrysomelidae) ......... 642 
GARCIA ALDRETE, ALFONSO N.—A new ptiloneurid genus (Psocoptera: Ptiloneuridae) 

EN OLAMIE) OMIT CAMs erie eierelatayetcletotelaters alates cle atclateteloyetatete efetelstormtataletacslele|efn/alelotatets slala\a\eisieraleieieintald 267 


GATES, M. W., S. N. MYARTSEVA, and M. E. SCHAUFF—A new Baryscapus Forster 
(Hymenoptera: Eulophidae) parasitic on Diorhabda elongata Brullé (Coleoptera: Chryso- 
melidae) and implications for the biological control of saltcedar (Tamaricaceae: Tamarix spp.) 
in the southwestern United States .............00.. cece cccc een e cece ccc es ness ec ccccsereeterecsscstecee 28 

GILMORE, DANIEL W.—See GANDHI, KAMAL J. K. .... 2.0... e eee eects 917 

GODOY, CAROLINA—A new genus of brachypterous leafhoppers (Hemiptera: Cicadellidae: 
Cicadellinae: Proconiini) from Costa Rica ........... 02. cce ccc e cece cence eee tee e nee eenecenscneceaes 259 


990 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


GONZALEZ, VICOR H.—See CAMBRA, ROBERTO A. ....................5555 dodges ROI 229 
GRIMALDI, DAVID—See SCHAWAROCH, VALERIE ..............00 0c s eee eeeeeeeeeeeeeee eee 323 
GRISSELL, E. E. and G. E. HEVEL—First report of Theocolax ingens Xiao and Huang (Hy- 

menoptera: Pteromalidae) in the Western Hemisphere, with a synopsis of the genus ......... 254 
HALL, JASON P. W.—A review of the Metacharis syloes group (Lepidoptera: Riodinidae), 

with the description of two new species from west of the Andes ....................0+seeeee es 200 
HALL, JASON P. W. and KEITH R. WILLMOTT—A new species of Paiwarria (Lepidoptera: 

Lycaenidae: Eumaeini) from western Ecuador ................0.: ee sseeee eee eect e eee ees 960 
HANSON, PAUL—See BURCKHARDT, DANIEL ............... 0.000 ccc ce cece cece eect eee 741 


HARBACH, RALPH E., RAMPA RATTANARITHIKUL, and BRUCE A. HARRISON—Bai- 
maia, a new subgenus for Anopheles kyondawensis Abraham, a unique crabhole-breeding 


anopheline an) southeastermPAtsian rei elec cls- cee ractetrieeie eles eee eee ee eset tle a 750 
HARRISON, BRUCE A.—See HARBACH, RALPH E. ............... 0.2 c ccc eee cnet e neces 750 
HARRISON, T. L.—A new species of Douglasiidae (Lepidoptera) from the eastern Nearctic .. 596 
HARRISON, T. L. and M. R. BERENBAUM—Rutaceae-feeding Agonopterix Hiibner (Lepi- 

Gloyieerms IS ero evistavelere)) rin WMNMVONS So opn6 600655586900000900000000000000 vasacnDREEBAUAnAbOOSGCHD NDE CeDo AOD 162 
HASTRITER, MICHAEL W. and MICHAEL E WHITING—Records of fleas (Siphonaptera) 

Olt CATO RSS TROON IGENG) 5o5008000000500006000000000000000008000000000000000000060000000000000500800000 417 
HEINOLD, B. D.—See DeWALT, R. EDWARD. ....22 2.220000... cece ec c eee ec e ccc ee tnneeeneeecees 34 


HELLENTHAL, RONALD A. and ROGER D. PRICE—Two new species of Myrsidea Water- 
ston (Phthiraptera: Amblycera: Menoponidae) from the jewel-babblers (Passeriformes: Eu- 
petidac)iinona New, Guineas... tees e een. cee ee eee eee eee eee eee cee eer ace terete 485 
HELLENTHAL, RONALD A., ROGER D. PRICE, and JASON D. WECKSTEIN—The genus 
Ramphasticola Carriker (Phthiraptera: Amblycera: Menoponidae) from the toucans (Picifor- 


mes: Ramphastidae), with description of a Mew SPeCieS ............ 2.20. eee eeee eter eee eee 565 
HENR YS LEOMAS See WHEBIEER WAS G2yRe sieeeeceres ee eee reer ila-cceeee rece ireere rere 209 
HESPENHEIDE, HENRY A.—Weevils of the genus Archocopturus Heller and Zygopsella Cham- 

pion: Sibling species and mimetic homoplasy (Coleoptera: Curculionidae: Conoderinae) ...... 671 
ISIBWABIL, (Ge 13S GINISSIBIUL, 186 186 sonaccccsce00sn0s000008050005000 sec 0acqdcb0edpoGDseDSBUSG0CO0RL 254 
HINOJOSA-DIAZ, ISABEL A. and CHARLES D. MICHENER—A new bee of the genus 

Chilicola Spinola (Hymenoptera: Colletidae: Xeromelissinae) from Central Mexico ......... 1 
HIROSE, YOSHIMI—See JOSE, JOSEPHINE .......20.... 002.00. eee e cece cece cee tence eeeeeeeeess 782 
HODGES, GREG S.—See MILLER, DOUGLASS R. ................ 0.000 c esse cece ees 123 


HOEBEKE, E. RICHARD and A. G. WHEELER, JR.—Establishment of three European flea 
beetles in Nova Scotia: Longitarsus ganglbaueri Heikertinger, L. jacobaeae (Waterhouse), 


and L. rubiginosa (Foudras) (Coleoptera: Chrysomelidae: Alticinae) ....................0eee ee 319 
HOEBEKE, E. RICHARD—See WHEELER, A. G., JR. .......... 2... e eee e neces 941 
HOLZENTHAL, RALPH W.—See GANDHI, KAMAL J. K. ............... 0.0... cece eee ee 917 
HONDA, JEFFREY Y.—See JOSE, JOSEPHINE .................... 0000s cece cece cece eee eeeeeeeees 782 
ISIUJAING, IMDIN-—See ZABUNINIG, YOAILIIN  Sescccbongnbcco0d0ccccndcsadencoadoaonnnsobococconosanecoacsc00E 376 
HUANG, MIN and YALIN ZHANG—Two new leafhopper species of Bolanusoides Distant 

(Hemiptera: Cicadellidae: Typhlocybinae: Typhlocybini) from China ...............--+..+-+++:- 428 
HUANG, YIAU-MIN—Cornetius, a new subgenus of Aedes, and a redescription of Aedes 

(Cornetius) cozi Cornet (Diptera: Culicidae) ............ 0... c cece cence cece eee cnet teen nee SyIl7/ 
HUSBAND, DAVID O.—See HUSBAND, ROBERT W. ............... 0.600 e cc ceeeeeeeet eee eee WA 


HUSBAND, ROBERT W. and DAVID O. HUSBAND—A new species of Dorsipes Regenfuss 
(Acari: Podapolipidae), ectoparasite of Amara latior Kirby (Coleoptera: Carabidae) from Ar- 


ADIN ho AER A See See gH CO SE SROO Cen Som ona BnoddmonSSondass cASose Sonnod SR anadgoEdodoCOsRHoWaGasSooREN 71 
JANZEN, DANIEL H.—See BURNS, JOHN M. ...........22.... 00000 c cnn c cence cece eee enenee 770 
JENNINGS, JOHN T. and DAVID R. SMITH—The taxonomic placement of several New 

World and Oriental gasteruptiid wasps (Hymenoptera: Gasteruptiidae) ......................++- 686 
TOISINISOING 1K Uo R= SlSS TLAIBKOINIINS, Yo IR, Gocoaonsconccacccanse esos qos ansoouscasconcausga0D0000R00E 554 
JOSE, JOSEPHINE, YOSHIMI HIROSE, and JEFFREY Y. HONDA—Two new species of 

Trichogramma (Hymenoptera: Trichogrammatidae) from the Ryukyu Islands, Japan ......... 782 
KCANNDE:, ISIMRVAIN| (C—S=See IMMILIDIEIRE, (GUNIRNT IL “s56600000s0000050056000060s 009 sac cnEecncoDDaea200a00008 700 
KATOVICH, STEVEN A.—See GANDHI, KAMAL J. K. ............ 0.60. e cece ence eee 917 


KIMSEY, LYNN S.—Revision of the northern South American tiphiid genus Merithynnus 
Kimsey, 1991 (Hymenoptera: Tiphiidae: Thynninae) .................... 0.0 e cece eee nee eee eee 576 


VOLUME 107, NUMBER 4 99] 


KINKOROVA, JUDITA—Notes on the fruit flies (Diptera: Tephritidae) of California 


BAe 5 851 

KIRCHNERS RE Sees KONDRWAGIE REG BGs ey.c- eee eee ee een eee eee eee 859 

KLAUS, ANGELA V.—See SCHAWAROCH, VALERIE ................0220eeeeeeeececcecceceeves 323 

KOEHEE, JESSICA J:—See GANDHI, KAMAL J. K. .....0....022e0sceecee-oeeaceeeseeeesesesesee 917 
KONDRATIEFE B. C., R. E KIRCHNER, and DAVID LENAT—Two new species of Hap- 

loperla Navas (Plecoptera: Chloroperlidae) from North Carolina, U.S.A. ..................---. 859 


KULA, ROBERT R. and GREGORY ZOLNEROWICH—A new species of Epimicta Forster 
(Hymenoptera: Braconidae) from North America and new distribution records for Epimicta 
FRG Lf ULTUS NN AATLO Ip es She chnssera state ere vase Sete See le kas st eet «Saye ee eee 78 

LaBONTE, J. R., A. D. MUDGE, and K. J. R. JOHNSON—Nonindigenous woodboring Co- 
leoptera (Cerambycidae, Curculionidae: Scolytinae) new to Oregon and Washington, 1999— 

2002: Consequences of the intracontinental movement of raw wood products and solid wood 
packinSematenialls pai cclajerrslossacicios claw srerisire waaw. te sets Neck nse ae Sob eR ast ls Rlctoraroele eee eto se TERE 554 


LARSEN, KIRK J:—See GANDHI, KAMAL J. K. 2.0.0.2... ccc cece ccc c cece cc ceeeecsceeess SN 
PENATSDAVID See KONDRATIEREWB 1@s Basse seecos eee eceee cece nearer ee Cerner ee ee eeeeeee 859 
ERICONG— Sco RUEDA IEEOROIDO UM. Aoceeate eee escent oe ece ce ane cect oie oe eee eke 604 


MAIER, CHRIS T:—First records of alien insects in Connecticut (Orthoptera: Tettigoniidae; 
Coleoptera: Buprestidae, Chrysomelidae; Diptera: Rhagionidae, Tephritidae; Hymenoptera: 


Mies achilid ac) Ae tk airs Soe sept eterd otsteuattig aioe Ses see poet be aateneemd aneeaeha ane Seiicee oreeeaee eae Senin 947 
MARINO, PABLO I.—See SPINELLI, GUSTAVO R. ............. ccc cee eee eee e ee eeeeeeeeeeneneeees 108 
MATHIS, WAYNE N.—See EDMISTON, JAMES E .............. 0s sceeeeeeeeeeeeeeeeeeeeeeeeeeeee 7 
MATHIS, WAYNE N. and MANUEL A. ZUMBADO—Description of Scatella savegre, a new 

species from Costa Rica in the Triseta group (Diptera: Ephydridae) ...................2........ 386 
MATTSON, WILLIAM J.—See GANDHI, KAMAL J. Kyo... 2c. cece cece ec eceeeeeee eee e eee eee 917 
MAWDSLEY, JOHATHAN R.—Extirpation of a population of Cicindela patruela DeJean (Co- 

leoptera: Carabidae: Cicindellini) in suburban Washington, D.C., USA ........................ 64 


MAWDSLEY, JONATHAN R.—Additional historic records of Cicindela dorsalis Say and Cicin- 
dela puritana Horn (Coleoptera: Carabidae: Cicindelini) from the Chesapeake Bay region, USA 


pag bobs botao bod upOoR Do Ud DOnOOn TOG DAt earns Ha acorcr anne der Onn pop roncnoonadnconondooa oo ancentcss canoaacnops 808 
McCAFFERTY, W. P.—See RANDOLPH, Ree Peis eeeeu Wiyohaeuga tn clnwiee its aanchstaeine ede teaeee Caen 190 
McCAFFERTY, W. P. and L. SUN—Mystaxiops, a new genus of small minnow mayflies 

(Ephemeroptera: Baetidae) from Papua New Guinea ............ 0.00.00 cece cece e eee eens 536 
MCINTYRE, ANNE M.—See BARROWS, EDWARD M. ............0. 0000 ee cece cece cece cece eeee 693 
METLEVSKI, JAN—Contribution to the taxonomy and faunistics of the genus Meropleon Dyar 

(Le IGloy year INETNGES) ccoocasen0086006 5000000095050 80500 ano ao ISDA Sac HooR bos coSbacaboaTcoprensocencc 812 
MICHENER, CHARLES D.—See HINOJOSA-DIAZ, ISABEL A. ..........0.00e00e0eereeeeee ees | 
MILLER, DOUGLASS R., GARY L. MILLER, GREG S. HODGES, and JOHN A. DAVID- 

SON—Introduced scale insects (Hemiptera: Coccoidea) of the United States and their impact 

OM, WeSs Ame. gocoq92000000000e0000 doc ododaeqasddocoun soa DbdoocnoO2OaqDSbd0 Jann ddasoDen6ssSuq0 IDNA 123 
MILLER, GARY IL.—See MILLER, DOUGLASS R. .......... 0.00.00. cccc cee e eee n nee eeeeeeneneesee 123 
MILLER, GARY L., MANYA B. STOETZEL, and ETHAN C. KANE—A systematic study 

of the genus Diuraphis Aizanberg (Hemiptera: Aphididae) ................0.-se esses sees sree 700 
IMIUIDYGIS, VA. IDS ILAIBXOINMINE, Us RG” Goscenagoossooasdecnuncopecosoacconebudoaboag=obedcnposopcre +4 554 
INMDYAIRISE Vite SaN=—Seel GAME Sa WS Wa terse clctare clelsie(olne aisle ara ciel olsvelol lm alata evrterelelelee)n\ai« <]eisielele sels efei-l= 28 
NAVARRETE-HEREDIA, JOSE LUIS—A new species of Phanolinus Sharp (Coleoptera: Sta- 

phylinidae) with a key and comments for Mexican species .............0se0seeee eee eee sees esses 887 
NEUNZIG, H. H. and M. A. SOLIS—Tumoriala, a new Neotropical phycitine genus (Lepi- 

doptera: Pyralidae) 225 oc sn scence ese ss oc steiinc tenn Meese pindin sen deiensieinieh <sldalg rele aminacle na dais aaah 84 
NEUNZIG, H. H. and M. A. SOLIS—A review of the Neotropical genus Difundella Dyar 

(Lepidoptera: Pyralidae: Phycitinae) ..............ceseeee eee ence e ence eens seer eeeeeeee eres er eee eeaes 303 
NOVIKOV, D. V., G. A. ANUFRIEV, and C. H. DIETRICH—New genera and species of 

21 


leafhoppers (Hemiptera: Cicadellidae) from Kyrgyzstan .......-. 66. e seer eee e eee e eee e ene e eee ees 2 
PINKAEW, NANTASAK, ANGSUMARN CHANDRAPATYA, and RICHARD L. BROWN— 

Two new species and a new record of Eucoenogenes Meyrick (Lepidoptera: Tortricidae) from 

Thailand with a discussion of characters defining the QeNUS ....... 6.6.6 e eee e ee eee eee eee eee e ees 869 
PEREDO, LUIS CERVANTES and SAGRARIO GAMEZ-VIRUES—Three species of facul- 

tative Myodochini (Lygaeoidea: Rhyparochromidae) associated with figs in Mexicolmer cae. 362 


992 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


PINTO, C. MIGUEL and GUILLERMO L. CLAPS—First record of Cuterebra almeidai (Gui- 
maras and Carrera) from Argentina, new host records for Cuterebra apicalis Guérin-Méne- 
ville, and a list of Cuterebra (Diptera: Oestridae) in the collection of the Instituto-Fundacién 


MicvelicillosmucumantyAreentinale-coccree rr rEerererrctcece rere Eee errr tee err seer errr SWZ 
PINTO, JOHN D.—Descriptions of additional New World Trichogrammatidae (Hymenoptera): 

The genus Nicolavespa and a new species of Haeckeliania ..................0ccc cece ence eee e eee 627 
PLAKIDAS, JOHN D.—A new species of Porricondyla (Diptera: Cecidomyiidae) from south- 

westem Rens iivainia f..55 caches eae se Melee resettle se Neeteie/alaveralebelets Sevonenacsarm nrevabaee etc arcitiarara cro erates 652 
POINAR, GEORGE, JR.—Fossil Trigonalidae and Vespidae (Hymenoptera) in Baltic amber ... 55 
POINAR, GEORGE, JR.—A Cretaceous palm bruchid, Mesopachymerus antiqua, n. gen., n. 

sp. (Coleoptera: Bruchidae: Pachymerini) and biogeographical implications ................... 392 
POINAR, GEORGE, JR.— Culex malariager, n. sp. (Diptera: Culicidae) from Dominican am- 

ber: The first fossil mosquito vector of Plasmodium .................. ccc cece eee eee eee eeeeeaees 548 
POINAR, GEORGE, JR. and ALEX E. BROWN—New Aphidoidea (Hemiptera: Sternorrhyn- 

Chain IB URIMES exam ET Lee sw ee eect ss ee PSE HC ett eT E eabs onan en MO Re 835 
PRICE, ROGER D.—See HELLENTHAL, RONALD A. ...... 2... e eee cece nee enees 485 
PRICE, ROGER D.—See HELLENTHAL, RONALD A. ........ 0.0... c cic ecce nee ee neee 565 
RANDOLPH, R. P. and W. P- McCAFFERTY—The mayflies (Ephemeroptera) of Alaska, in- 

cludingovammewaspeciesiof Heptas enitdacmeess--eeeeeeeee CE erect eee eee Gee eae eee eee eee 190 
RATTANARITHIKUL, RAMPA—See HARBACH, RALPH E. ................. 22. cceceeeeeee eee 750 
RAZOWSKI, JOZEF and JOHN W. BROWN—Review of Oregocerata Razowski (Lepidop- 

tera: Tortricidae: Euliini) with descriptions of four new species ................... esse eee eee eee 903 
RIDER, DAVID A. and LE-YI ZHENG—Checklist and nomenclatural notes on the Chinese 

Pentatomidae (Heteroptera), III. Phyllocephalinae, Podopinae .....................0eceeeeeeeee ee 90 
ROBBINS, RICHARD G.—The ticks (Acari: Ixodida: Argasidae, Ixodidae) of Taiwan: A syn- 

ONYMIC /CHECKISE ces PATS Neo rdls cee Beene aoe netane ate arene Baa eee mentale geal a eels aside teeta ace area 245 
ROBBINS, ROBERT K.—Phylogenetic relationships among the species of Panthiades Hiibner 

CLyCROIGESs INNSSMESS 1B UTES) S6600000000000008000000000000000 0b 0nRPHODODHAGdOOOCODNOAGAdAOODONC 501 


ROBBINS, ROBERT K. and MARCELO DUARTE—Phylogenetic analysis of Cyanophrys 
Clench, a synopsis of its species, and the potentially threatened C. bertha (Jones) (Lycaenidae: 


Mh china’S HE UMA SIM) Ore A ane Saar ey AS eats ate ae Geta ean UGA Mees Ree OSE Se 24 A OY 398 
ROBINSON, HAROLD and NORMAN E. WOODLEY—A new species of Harmstonia (Dip- 

tera Dolichopodidae) siromvB oliviaseres-eeeeeeeee re eee eee ree eee eee eer eee eee eee eee eecer errr 436 
RONDEROS, MARIA M.—See SPINELLI, GUSTAVO R. 1.2... 0.0.0... cece cence ene e eee eees 108 
RUEDA, LEOPOLDO M., RICHARD C. WILKERSON, and CONG LI—Anopheles (Anoph- 

eles) lesteri Biasas and Hu (Diptera: Culicidae): Neotype designation and description ....... 604 
SAHIN, FIKRETTIN—See YAMAN, MUSTAFA ............... 02. ceccen eee e eee cece ee eeeeeeenneees 623 
SAINI, MALKIAT S. and DAVID R. SMITH—Revision of the southeastern Asian sawfly genus 

Busarbia Cameron (Hymenoptera: Tenthredinidae) .............. 00... cece cece eee n eect eeeeeneees 346 
SCARBROUGH, AUBREY G. and JESSICA E. COSTANTINO—The genus Ommatius Wie- 

demann, Dilatipennis species group (Diptera: Asilidae) ............... 0. ee eee eee eee eee eee ee eees 789 
SCHAWEEIM2E See" GATES Mi. Wit eit nacnacerisacresarccinete ster ciociee sete atareye nee eiocietecie seria eecee 28 
SCHAUFE M. E.— Ammonoencyrtus carolinensis, n. comb. (Hymenoptera: Encyrtidae), a 

parasite of lobate lac scale Paratachardina lobata (Chamberline) (Hemiptera: Kerriidae) .... 115 


SCHAWAROCH, VALERIE, DAVID GRIMALDI, and ANGELA V. KLAUS—Focusing on 
morphology: Applications and implications of confocal laser scanning microscopy (Diptera: 


Campichoetidae, Camillidae, Drosophilidae) .............. 0... c cece cece cnn ence ec ne eee eeeneneeeeeen 323 
SCHIFE NATHAN Mi—==See SIMIDEL DAVID Re saaseeeccce- se cccc eee: scene seria semen eceieeitereeriielel 864 
SEYBOLD, STEVEN J.—See GANDHI, KAMAL J. K. ....... 0... c cece ccc c cee e nen e eee e eee eees 917 
SHENG, MAO-LING—The genus /schnoceros Gravenhorst (Hymenoptera: Ichneumonidae) in 

(1000 ae eee ser eerie eee ecco? Per na niorian Hernaer cen a Guar conanbta ga anes anadamubsopoaboan case 432 
SMITH, DAVID R.—Review of the genus Acordulecera Say (Hymenoptera: Pergidae) of the 

West Indies, and the first records of Symphyta from Montserrat and St. Kitts ................. 99 
SMITH, DAVID R.—A new sawfly (Hymenoptera: Pergidae) feeding on guava, Psidium gua- 

java Ten(Myrtaceae) hn Costati Gar <x ase eee eee eee eee 214 
SMITH, DAVID R.—Two new fern-feeding sawflies of the genus Aneugmenus Hartig (Hy- 

menoptera: Tenthredinidae) from South America ..............0. cee cece cece eee ce eee en ee eeeeeeeees 273 


SMITH, DAVID R.—Review of the Aulacidae (Hymenoptera) of Chile and adjacent Argentina .. 820 


VOLUME 107, NUMBER 4 


SMITH, DAVID R. and NATHAN M. SCHIFF—A new western Nearctic species of Calameuta 
Konowa (Etymeno pterasCephidac) iperer eee eee eee eee ee Getter eee EEE EEE EEE EEE EEE eee eee eee eeeeee 
SMITH, DAVID R.—See SAINI, MALKIAT S. 
SIMIGUSE IDAWAID IR —=Syeo USININIINCGIS), WOJSIN| 1B Goacescsscnnoppnacnncsnoasnoeeeenoscnsaanesacccesso. 
SMITH, DAVID R. and IAN C. STOCKS—A new trigonalid wasp (Hymenoptera: Trigonali- 
dae) siromkeastern™ NorthwAmenricameeee er eee rere eee neon ne eee ee eee En eee eee 
SO EIS IME ZV Sree INI S| UINVACE A a I) eB ts erecasreccaraeeadadenooceU Cc ntcnenGAnbSBdse cncbassnGsacssnneo: 
SOETS MwA See NE UINZIG,, Eb EUs” as\ststerarac crass tarefera'a oye/ctale 5 stove elvoislele stots Slalerorelc aroma arelne ns oie 
SPINELLI, GUSTAVO R., PABLO I. MARINO, and MARIA M. RONDEROS—tThe fourth 
instar larva and pupa of the Neotropical biting midge Forcipomyia (Forcipomyia) rioplatensis 
Marino and Spinelli (Diptera: Ceratopogonidae) .................... cece cece cece ceneeeeeeceeennees 
STEINER, WARREN E., JR.—Studies on the darkling beetles (Coleoptera: Tenebrionidae) known 
from Grand Bahama Island, with descriptions of new species of Branchus and Adelina ....... 
STOCKSSIAN (€.—See) SMEEHY DAVID) Re 2. Fate cece asaccaclecn sects seeuibenctcnicucanseseeniecas 
SUOIAA SIL, IMVAINNON sy Syee) WMMDEILI SIRE (GYNIROY IE, Gaooadoodpodoanscoscodoosb6nansboobsssosspneosae 
SUINGIE = ScemMCCABBE RD YG oWir Be arcis doen ek a santttelersiescrnd lates a Sic cicie siete Sector teioeia So elcrsre a cents 
TAUBER, CATHERINE A., GILBERTO S. ALBUQUERQUE, and MAURICE J. TAUBER— 
Characteristics of the Loyola Navas male (Neuroptera: Chrysopidae: Apochrysinae) ......... 
TAUBER, MAURICE J.—See TAUBER, CATHERINE A. .............. 0. cece cece cece cece cece eees 
TOGASHI, ICHIJI—Description of a new species of Eriotremex Benson (Hymenoptera: Siri- 
GES) nO Grek Aaa a chabR ashe a const tase nba nrarr iar. Acer Canna acncaAdnacaakicocanooaaelocan: 
TOGASHI, ICHIJI—A new species of the Eriocampa ovata group (Hymenoptera: Tenthredi- 
nidae) feeding on Alnus matsumurae Call. (Dicotyledoneae: Fagaceae) from Japan ........ 
TOGASHI, ICHIJI—Description of a new species of the genus Apethymus Benson (Hymenop- 
tera: Tenthredinidae) feeding on Quercus acutissima Carruthers (Fagaceae) in Japan ........ 
TOGASHI, ICHIJI—An additional species of the genus Masaakia Takeuchi (Hymenoptera: 
enthnedimidac) srombapanwereeeeeeeeee eee eeeEe ee ee eee Eee eee eee eee eee eee eee eee ce arene ee ee ee ree 
TOGASHI, ICHIJI—A new species of the genus Conaspidia Konow (Hymenoptera: Tenthre- 
dinidae) from Japan, with a key to the Japanese species ........... 0... cece cece ee eee e eee eee e ee es 
WCISLO, WILLIAM T.—See CAMBRA, ROBERTO A. ....................00.000cceeeeeeee eee es 
WECKSTEIN, JASON D.—See HELLENTHAL, RONALD A. ..................000000000eeeee ee 
WHEELER, A. G., JR. and THOMAS J. HENRY—Description of the adult and fifth instar of 
a myrmecomorphic plant bug, Bicuspidatiella conica Maldonado (Hemiptera: Miridae: Der- 
ASOECOMIMES)), WTO THOWES ON IS IBIS Soooconcoccsucososcoueoedbsdo sos uddosocoO ODO dgHoUaoEOsCaScoSECan. 
WHEELER, A. G., JR. and E. RICHARD HOEBEKE—Livilla variegata (L6w) (Hemiptera: 
Sternorrhyncha: Psyllidae) new to North America, with records of three other Palearctic 
psyllidsinewatomNewloundlanditeseereree eee Coen eeeeee eee cer eneee sree eee eee rere ee Ene rere 
WHEELER, A. G., JR.—See HOEBEKE, E. RICHARD ................00 2.2... cece eee e eee e eee 
WHEELER, A. G., JR.— Blissus minutus (Blatchley) and Toonglasa umbrata (Distant): Sel- 
dom-collected native chinch bugs (Hemiptera: Lygaeoidea: Blissidae) as colonists of the 
African bunchgrass Eragrostis curvula, and their association with other grasses in the south- 
Gian, WOhmirerel Ciao oes aen sons duoaes sudan oboscoseanpdeesceanaaonaosrocsn speed aodnopsosdonodasbaudasopenor 
WHEBDERa MER RYoAe “See BEAULIEU, EREDERIC laiisnassueceesue decd e-ceaceseaseee «eee 
WHEELER, TERRY A.—See BROOKS, SCOTT E. ............... 0. cece cece cece ence ee cceneenee 
WHITING, MICHAEL E—See HASTRITER, MICHAEL W. ...............000 0.00 e cece cece cece 
WILKERSON, RICHARD C.—See RUEDA, LEOPOLDO M. ................... 255.002 s esses eee 
VWYHULIMOME BINNS! R——Syee IVE, UWANSTOIN| 12 NWA | 565505 sccedgn cc doceccodboseduensonesacdeosacocs 
WOODLEY, NORMAN E.—See ROBINSON, HAROLD ............... 0.6.0 c cece eee 
YAMAN, MUSTAFA, IRFAN ASLAN, ONDER CALMASUR, and FIKRETTIN SAHIN— 
Two bacterial pathogens of Helicoverpa armigera (Hiibner) (Lepidoptera: Noctuidae) ....... 
YANG, DING and STEPHEN D. GAIMARI—Review of the species of Elaphropeza Macquart 
(Diptera: Empididae: Tachydromiinae) from the Chinese mainland .................000200ss000 
YONINIG, UIN(GAII— See ZABVAIN(G, IOIEINS Cos ncqaodasopeclcoondoccnocbdbaobOosondosoosSaucusmeeoncoso: 
YOUNG, DANIEL K.—Taxonomic notes on South American Pogonoceromorphus Pic (Cole- 
optera), including transfer from Pyrochroidae (Pyrochroinae) to Anthicidae (Eurygeniinae) .... 
ZHANG, LI-JIE and XING-KE YANG—A new species of the genus Agetocera Hope (Cole- 
optera: Chrysomelidae: Galerucinae) from China .............0.0seceeee een ee cere eters eens e ee eneees 
ZHANG, YALIN and MIN HUANG—Two new leafhopper genera, Direnaia and Xaniona 
(Hemiptera: Cicadellidae: Typhlocybinae: Typhlocybini) from China .................ss0e20eee 


209 


94] 
319 


336 
176 
489 
417 
604 
960 
436 


623 


49 
119 


357 


119 


376 


994 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 


ZHANG, YALIN and WU DAI—A taxonomic review of Matsumurella Ishihara (Hemiptera: 


Eicadelllidaes|Deltocephalin'ac)) irom Chinaeeessneesee eae eee eee ee eee reer see er errr 218 
PHANG. YAIIN—See HUANG, MIN: see cuncccc cri cee encase rae ecae state serssicbetniercian etn atoteteraisiets teieteys 428 
ZHENG, E=YI—See RIDER, DAVID) As one ee sie desea ste sete cieere s St oreratereini= Shebelsie o)siatersyalstete este siete vats 90 
ZOLNEROWICH, GREGORY—See KULA, ROBERT R. ............... 0. cece e cece cence eee 78 
ZUMBADO, MANUEL A.—See MATHIS, WAYNE N. ...........-.... 0s eee cece cece ener ener eees 386 

NOTES 
BARBER-JAMES, H. M.—See McCAFFERTY, W. P. ...............02 2c ceee cece ee eee c cee ee eeeeeee 238 
BAUMGARDNER, DAVID E.—New and additional records of mayflies (Ephemeroptera) from 

the southwestern United States, including a new country record ................... eee eee eee 977 
BITO, DARREN and DAVID R. SMITH—Larva and possible food plant of Ancyloneura var- 

ipes (Cameron) (Hymenoptera: Pergidae) in Papua New Guinea ....................0...2222205- 463 
BULLINGTON, STEPHEN—See LAVIGNE, ROBERT J. ............... cece eee cece cee eee eeeeeee 968 


CARROLL, J. E and T. L. CYR—A note on the densities of Ixodes scapularis (Acari: Ixodidae) 
and white-tailed deer on the campus of the National Institute of Standards and Technology, 


Maryland: USA ssszseseisa dis ee aye eee cere ic See ae Sa eto acetal ciel See ee eee ners 973 
CGYRY Tb —=See CARRO Wyo & watcde cetis ee eet eee aCe eee Beeson oe ae mere 973 
DURDEN, EAN@E A:—See NEEDERS MARKUP) fo2sec ceases: seeeeocce esses se seebe es eens 471 
FAVRET, COLIN—A new non-destructive DNA extraction and specimen clearing technique 

forsaphids (Hemupteral mssaiek seems aeckinsintis Aaa see ee eee en ie net Aeon RAR ene beee 469 
FERRERIA, PAULO S. FIUZA—See HENRY, THOMAS J. 02.2... 0... ccc eee eee eens BS 
GANO) QIESSes INIUIEIDYA, JI SHOOIIDKO) IMIS Gocooccopaccogvacv00cqdeaoonsapaccacodendeosnooanndoancandes 235 
GODDARD, JEROME—See VARNADO, WENDY C. ............-. 0s eee ccn eee c eee e cette eee e eee 476 
HARRISON, BRUCE A.—See VARNADO, WENDY C. .............:0 0c eee cece cece este eee e ee eees 476 


HENRY, THOMAS J. and PAULO S. FIUZA FERRERIA—Froeschneropsidea, a replacement 
name for the preoccupied genus Froeschnerisca (Hemiptera: Heteroptera: Miridae: Deraeo- 


cornnae: Hy alioGimt) «onc cisiaas nacneses tease aaa ae as SE eee one ee eae ese eee 735 
JACOBUS, LUKE M. and W. P. McCAFFERTY—Validation of Potamocloeon macafertiorum 

Fuco-Ortizi(= phemeropteray bactidac) ieee eee ee eee ee ee eee eee Eee eee Ca eee eee eee eee eee 474 
JACOBUS, LUKE M. and W. P- McCAFFERTY—Apobaetis futilis (McDunnough), a new 

combination in Nearctic Baetidae (Ephemeroptera) .................. 000 cece eee cee cece eee eee 979 


JACOBUS, LUKE M., W. P. McCAFFERTY, and ROBERT W. SITES—A new synonym and 
new Thailand records of Cincticostella femorata (Tshernova) (Ephemeroptera: Ephemerelli- 


GAG) iP rset le, cent ive: unre «Ath Reet hays, eR 0. Me ReR RAN ST APRA! FAR. kd Shae. Bags ere tala. cena RY eee 734 
KORECKI, JAMES A.—See NELDER, MARK P. ................0.00ccscceeeee\ereecessneceseeees 47] 
LATTIN, JOHN D.— Physopleurella floridana Blatchley, 1925, a synonym of Physopleurella 

mundula (White, 1977) (Hemiptera: Cimicoidea: Anthocoridae) ........................2022 200 460 


LATTIN, JOHN D.— Dufouriellus ater (Puton), Macrotrachelia nigronitens (Stal), and Xylo- 
coris (Arrostelus) flavipes (Reuter) (Hemiptera: Cimicoidea: Anthocoridae): First records 


hr @O naa tS A Wy aa TDS VAT S os crossictsasseavsee beter sxe le love ios SLLE ee EVE eke SITET SGT STE Jo iS LAS Tee 466 
LATTIN, JOHN D.— Scoloposcelis pulchella (Zetterstedt, 1838) in North America (Hemiptera: 
IAM OCOTIGAE)) ercrevove che esters lore eestor se Ee eC OAC aeT ae eee Pen eree 729 


LATTIN, JOHN D.— Scoloposcelis discalis Van Duzee, 1914, a synonym of Anthocoris gal- 
actinus Fieber, 1837, and Xylocoris umbrinus Van Duzee, 1921, a synonym of Piezostethus 


californicus Reuter 1884 (Hemiptera: Heteroptera: Anthocoridae) ...........................225. 971 
LAVIGNE, ROBERT J. and STEPHEN BULLINGTON—Notes on the ethology of Bohartia 

martini Adisiomarto and Wood (Diptera: Asilidae) in Wyoming, USA ....................202+- 968 
MAS YAIUINSeel RUEDARIEE OROLDOIME eee eee eee eee eee cee eee eee eee 235 
McCAFFERTY, W. P. and H. M. BARBER-JAMES—Tanzaniops, replacement name for certain 

Athican Bactidae (Ephemeroptera) ieee eeeeeeeeee ae eee eee eee eee eee eee Eee eee eee eee eee 238 
McC AEEE RIS Wa P——SeevAGOBUSSEUKE May cere erence ree eeeeeee eee ene e eee 474 
MIG CANS SE IEE WW, 12 See WN COIS, JWI IMI, caocoosoonccncdnccsccanesGbGabsondocabausceensooe 734 
Mc CAREERS Wa P——ScewACOBUS SE URE avis eee eee reer eee eee eee eee eee eee ee 979 


NELDER, MARK P., WILL K. REEVES, JAMES A. KORECKI, and LANCE A. DURDEN— 

Fleas of the genus Ceratophyllus (Siphonaptera: Ceratophyllidae) in the southeastern United 

Rl eS US eee coe Sot tires 8c GUC nace Aaa aera Aanaaideicl qe deer ah An deni aacccrbe yacdtauna a 471 
REEVES) WIE Ke SeeiNEBRDE Ra MARKS Pier tee eee eee eee eee eee eee eee eee ee eete 471 


VOLUME 107, NUMBER 4 995 


RUEDA, LEOPOLDO M., YA-JUN MA, GUANG-HONG SONG, and QI GAO—Notes on 
the distribution of Anopheles (Anopheles) sinensis Wiedemann (Diptera: Culicidae) in China 


and the status of some Anopheles Hyrcanus group type specimens from China ............... 235 
SITIES), INO BIE Wi See WACOBIWISS, ILLUMI IMI, cacccasossnnoaocncconnodonncsnocns0oseeannsonnoser 734 
SIMIAN SL, IDANAID) IR-—=Syee SIMO), IDVARIRIEIN| .coccoscoanconsosnnsdncadcdcasocsondoacsboossosoosnaneooe 463 
SONG, GUANG-HONG—See RUEDA, LEOPOLDO M. ............... 0.0... ccc ceeeeeeeceeeeeeees 235 
VARNADO, WENDY C., JEROME GODDARD, and BRUCE A. HARRISON-——New state 

record of Culex coronator Dyar and Knab (Diptera: Culicidae) from Mississipppi ........... 476 
WOODLEY, NORMAN E.—Distributional notes on Bertamyia notata (Loew), including the 

first report of the family Platypezidae from the Caribbean (Diptera: Platypezidae) ........... 731 

BOooK REVIEWS 
BICKLEY, WILLIAM E.— I/dentification and Geographical Distribution of the Mosquitoes of 

North America, North of Mexico [Second Edition], by Richard E Darsie, Jr, and Ronald A. 

AV VT De eevee ctayes ee ree torch cyavclcittesnrstatare Mitt da vie cman ate sO Se re MOM" APTN ION op CSE ey ene diy ONE te 737 
DAVIS, DONALD R.— Lepidoptera, Moths and Butterflies, Volume 2, Niels P. Kristensen, 

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UTION LIBRARIES 


wi il 


176 5922 


CONTENTS 
(Continued from front cover) 
MAWDSLEY, JONATHAN R.—Additional historic records of Cicindela dorsalis Say and 
Cicindela puritana Horn (Coleoptera: Carabidae: Cicindelini) from the Chesapeake Bay 
RERION WA ac MA UN kat AR ai ett ist ae OR ras eeUaa Sey A lata otal Datstegy iE Sn ls Se a aRtat ete 


METLEVSKI, JAN—Contribution to the taxonomy and faunistics of the genus Meropleon Dyar 
(Vepidoptera Noctuidae) CF Neh aun Ma tecayabyler aie. ote clnicte fe letolll agit alalel ta al ataeca Sanne tae he Astana oe 


NAVARRETE-HEREDIA, JOSE LUIS—A new species of Phanolinus Sharp (Coleoptera: 
Staphylinidae) with a key and comments for Mexican species ................00ecseeeeeee eens 


PINKAEW, NANTASAK, ANGSUMARN CHANDRAPATYA, and RICHARD L. BROWN— 
Two new species and a new record of Eucoenogenes Meyrick (Lepidoptera: Tortricidae) from 
Thailand with a discussion of characters defining the genus .................-.e eee cece eceeeee 


POINAR, GEORGE, JR. and ALEX E. BROWN—New Aphidoidea (Hemiptera: Sternorrhyncha) 
IN Busmesecamben yy F MAN sks Nis eM nek Ri AM Me i Dea TIAA eet sc 


RAZOWSKI, JOZEF and JOHN W. BROWN—Review of Oregocerata Razowski (Lepidoptera: 
Tortricidae: Euliini) with descriptions of four new Species ............ 02. eee ee eee ee eee cece 


SCARBROUGH, AUBREY G. and JESSICA E. COSTANTINO—The genus Ommatius 
Wiedemann, Dilatipennis species group (Diptera: Asilidae) ...................-.. eee eee eee eee 


SMITH, DAVID R.—Review of the Aulacidae (Hymenoptera) of Chile and adjacent Argentina .. 


SMITH, DAVID R. and NATHAN M. SCHIFF—A new western Nearctic species of Calameuta 
Konow, (Hymenopteray Cephidac) aaa a ciety Aaa etciate cau stee eiahe sae ka eo elors erate te ote ave eee 


TOGASHI, ICHIJI—A new species of the genus Conaspidia Konow (Hymenoptera: 
Tenthredinidae) from Japan, with a key to the Japanese species ...................-2-0-0-- eee 


TOGASHI, ICHIJI—An additional species of the genus Masaakia Takeuchi (Hymenoptera: 
Tenthredinidae)) froma apann Waa wale ceca sonic rien eebs Cal aabstnye ust oii, (ret 2k G7 oy eee an Ra 


WHEELER, A. G., JR. and E. RICHARD HOEBEKE—Livilla variegata (L6w) (Hemiptera: 
Sternorrhyncha: Psyllidae) new to North America, with records of three other Palearctic 
psyilidssnew to-Newfoundl aml iyo sere ig eis uate ia ae ees aS eda 2 paints ee 


NOTES 


BAUMGARDNER, DAVID E.—New and additional records of mayflies (Ephemeroptera) from 
the southwestern United States, including a new country record ...................-0-00eee eee 


CARROLL, J. F. and T. L. CYR—A note on the densities of Ixodes scapularis (Acari: Ixodidae) 
and white-tailed deer on the campus of the National Institute of Standards and Technology, 
Micarpy beam IS Aired esa GRU UL SSR RAD CRIA Sg a ZA oe toe aera a aa 


JACOBUS, LUKE M. and W. P. McCAFFERT Y—4pobaetis futilis (McDunnough), a new com- 
binationin: Nearctic Bactidac (Ephemeroptena)) seen oa eee eee cee arene eae 


LATTIN, JOHN D.—Scoloposcelis discalis Van Duzee, 1914, a synonym of Anthocoris galacti- 
nus Fieber, 1837, and Xylocoris umbrinus Van Duzee, 1921, a synonym of Piezostethus cal- 
ifornicus Reuter 1884 (Hemiptera: Heteroptera: Anthocoridae) ...................2...00e eee 


LAVIGNE, ROBERT J. and STEPHEN BULLINGTON—Notes on the ethology of Bohartia mar- 
tini Adisiomarto and Wood (Diptera: Asilidae) in Wyoming, USA .................0..2-.e eee 


BOOK REVIEWS 


MAWDSLEY, JONATHAN R.—4 Field Guide and Identification Manual for Florida and 
Fasiern.Uss: Nicer Beetlesjoy Paul Mi. Choate. aengan nea: eaiee ye eee cine eee eee ee 


SCHAEFER, CARL W—The Insects: Structure, Function and Biodiversity, by Dunston P. 
VaXi0010) RO CMR RS REY BNR IN NTU RENAL VG CMe on Ta EE DA ee Se 


MISCELLANEOUS 
Sacicty Meets ers eek CAN I ESA NO OR Ue A UI ee ee ne 
Table of Contents, Volume 107 


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808 


812 


887 


869 


835 


903 


789 
820 


864 


846 


914 


941 


977 


973 


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O7t 


968 


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