Proceedings of the Entomological Society of Washington

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to JULY 2000 NO. 3
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Tat

“PROCEEDINGS

of the

ENTOMOLOGICAL SOCIETY

VOL;; 102

CONTENTS

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

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

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

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

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

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

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

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

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

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

(Continued on back cover)

THE

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v

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

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

ALESSANDRA R. P. BAPTISTA AND WAYNE N. MATHIS

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

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

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

Oceanian, Oriental, Palearctic

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

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

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

482

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

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

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

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

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

NEOTROPICAL REGION

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

KEY TO NEW WORLD SPECIES OF CYAMOPS

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

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

C. halteratus Sabrosky

— Wing at least partially infuscate

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

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

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

tion of hindtibia darkened; postpronotum

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

VOLUME 102, NUMBER 3

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

C. americus Baptista and Mathis

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

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

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

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

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

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

C. neotropicus Hennig

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

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

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

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

483

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

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

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

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

484

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

Cyamops funkae.

Abbreviations: aed.

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

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

Thorax: Dorsocentral setulae typically

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

6th-7th st.

Hi! Mtg Mi,

WAAL),
Say \f 44
As My
NO
>

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

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

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

VOLUME 102, NUMBER 3

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

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

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

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

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

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

485

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

Cyamops sp. 1
(Figs. 5—6)

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

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

Cyamops sp. 2
(Fig. 7)

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

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

AFROTROPICAL REGION

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

486 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

i

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

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

°°

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

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

2; hindfemur mostly yellow, infuscate apically;

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

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

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

VOLUME 102, NUMBER 3

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

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

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

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

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

487

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

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

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

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

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

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

488 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

Distribution.—Cyamops nigeriensis is
known only from Nigeria.

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

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

VOLUME 102, NUMBER 3

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

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

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

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

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

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

489

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

Female: Unknown.

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

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

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

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

AUSTRALASIAN/OCEANIAN REGION

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

490

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

KEY TO AUSTRALIAN/OCEANIAN SPECIES OF

. Wing hyaline (Australia) .

CYAMOPS

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

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

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

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

wide, not constricted medially; 1st costal ratio

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

setae; basal scutellar seta about % length of

apicalsSetalg sa. tc < Ree he ee eee ee 4

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

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

. Mesofacial plate without setae; tibia and tar-

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

tose; vertex dull microtomentose (Ponape) . . .

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

Comb on ventral margin of midcoxa present

(Australia) C. pectinatus Khoo

. Peristomal setae on mesofacial plate (Austra-

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

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

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

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

ally (American Samoa)

C. samoensis, new species

- Legs mostly yellowish; midfemur yellow;

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

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

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

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

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

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

seta about % of apical seta. Legs mostly yel-

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

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

VOLUME 102, NUMBER 3 491

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

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

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

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

492

16

Figs. 14-16.

COR ses eS
urs
am A)

lS

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

oN ‘ (HY

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

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

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

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

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

VOLUME 102, NUMBER 3

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

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

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

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

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

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

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

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

493

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

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

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

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

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

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

494

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

Etymology.—The species epithet, fiji, is

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

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

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

VOLUME 102, NUMBER 3

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

Description.—Adult ¢ length 1.7—1.8

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

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

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

495

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

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

Thorax: Legs with femora mostly brown.

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

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

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

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

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

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

496 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

23

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

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

VOLUME 102, NUMBER 3

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

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

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

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

497

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

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

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

Cyamops pectinatus Khoo

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

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

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

*‘Ponape”’ species complex
(Fig. 23)

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

498

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

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

Cyamops sp. 3

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

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

ORIENTAL REGION

KEY TO ORIENTAL SPECIES OF CYAMOPS

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

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

. Basal scutellar seta very thin, about % that of

apical seta (Thailand) C. kaplanae, new species

— Basal scutellar seta well developed, about %—

%4 of apical seta

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

N

, of wing usu-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ally hyaline on anterior basal portion, other-

wise brown; right surstylus subrectangular,

much smaller than left surstylus (Fig. 17)

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

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

ally mostly infuscate dividing subapical white

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

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

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

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

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

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

VOLUME 102, NUMBER 3

499

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

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

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

500

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

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

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

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

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

VOLUME 102, NUMBER 3

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

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

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

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

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

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

501

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

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

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

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

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

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

502

aed.apod.

29

aed.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

VOLUME 102, NUMBER 3

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

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

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

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

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

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

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

503

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

Cyamops kaplanae. 20, Segments 5-7,

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

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

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

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

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

504

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

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

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

Thorax: Wing: Hyaline

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

PALEARCTIC REGION

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

Cyamops sp. 4

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

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

PHYLOGENETIC CONSIDERATIONS

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

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

VOLUME 102, NUMBER 3

505

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

C. banvaneue.

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

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

ACKNOWLEDGMENTS

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

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

LITERATURE CITED

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

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

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

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

506

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

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

America North of Mexico. United States. Depart-

ment of Agriculture, Agriculture Handbook No.

276, iv + 1696 pp.

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

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

Diptera of the Afrotropical Region. British Mu-

seum (Natural History), London, 1437 pp.

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

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

W. EVAN BRASWELL AND JAMES R. OTT

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

reprint requests

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

tribution data for D. ampla in central Texas.

Key Words:

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

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

gregarious behavior, aposematic coloration, group defense

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

METHODS

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

508

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

VOLUME 102, NUMBER 3

RESULTS AND DISCUSSION

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

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

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

509

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

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

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

510 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

O §0 100 MILES
i

lentes Ihe
hatched) in Texas.

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

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

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

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

VOLUME 102, NUMBER 3

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

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

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

511

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

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

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

512 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

VOLUME 102, NUMBER 3

Nn
—_
W

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

yellow background color with longitudinal black bands).

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

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

20

15

10

Larval Body Length (mm)

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

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

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

picted with shaded symbols.

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

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

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

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

VOLUME 102, NUMBER 3

10

Frequency

0 100

200

ails

400 500

300
Mean Dispersal Distance/aggregation (cm)

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

host plants under field conditions.

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

SUMMARY

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

516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

LOO

75

Percent of Leaves Remaining
N Nn
Nn =)

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

30
2215,
20
1s

10

Percent of Larvae Which Dispersed

0 25 50 a5 100
Percent of Leaves Remaining

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

VOLUME 102, NUMBER 3

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

ACKNOWLEDGMENTS

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

517

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

LITERATURE CITED

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

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

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

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

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

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

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

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

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

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

1911. Descriptions of the larvae of some

Mexican Lepidoptera. Proceedings of the Ento-

mological Society of Washington 13: 227-232.

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

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

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

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

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

518 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

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

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

RICHARD D. GOEDEN

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

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

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

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

520

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

MATERIALS AND METHODS

The present study was based in large part
on dissections of flower heads of Machaer-
anthera canescens (Pursh) A. Gray collect-
ed discontinuously during 1988-1997
mainly from the following three locations
in the northern section of the San Bernar-
dino Nat. Forest, SW San Bernardino Co.:
Onyx Peak at 2700-m elevation, Caribou
Creek along Van Duzen Canyon Road at
2120 m, and N shore Big Bear Lake at 2020
m. One-liter samples of excised, immature
and mature flower heads containing eggs,
larvae, and puparia were transported in
cold-chests in an air-conditioned vehicle to
the laboratory and stored under refrigera-
tion for subsequent dissection, photogra-
phy, description, and measurement. Six
eggs, six first-, 12 second-, and 13 third-
instar larvae, and five puparia dissected
from flower heads were preserved in 70%
EtOH for scanning electron microscopy
(SEM). Additional prepuparia and puparia
were placed in separate, glass shell vials
stoppered with absorbant cotton and held in
humidity chambers at room temperature for
adult and parasitoid emergence. Specimens
for SEM were hydrated to distilled water in
a decreasing series of acidulated EtOH.
They were osmicated for 24 h, dehydrated
through an increasing series of acidulated
EtOH and two, 1-h immersions in hexa-
methyldisilazane (HMDS), mounted on
stubs, sputter-coated with a gold-palladium
alloy, and studied and photographed with a
Philips XL-30 scanning electron micro-
scope in the Institute of Geophysics and

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Planetary Physics, University of California,
Riverside.

Most adults reared from isolated puparia
were individually caged in 850-ml, clear-
plastic, screened-top cages with a cotton
wick and basal water reservoir and provi-
sioned with a strip of paper toweling im-
pregnated with yeast hydrolyzate and su-
crose. These cages were used for studies of
longevity and sexual maturation in the in-
sectary of the Department of Entomology,
University of California, Riverside, at 25 +
1°C, and 14/10 (L/D) photoperiod. Six pairs
of virgin males and females obtained from
emergence cages also were held in each of
six separate, clear-plastic, petri dishes pro-
visioned with a flattened, water-moistened
pad of absorbant cotton spotted with honey
(Headrick and Goeden 1994) for observa-
tions of their courtship and copulation be-
havior.

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

RESULTS AND DISCUSSION
Taxonomy

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

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

VOLUME 102, NUMBER 3

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

Acc V Spot Magn
100kV 3.0 _1940x

AccV Spot Magn
100kV30 1940x

Det WD Exp
SEU Allon

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

heretofore have not been described or fig-
ured.

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

37

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

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

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

522 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

AccY SpotMagn D
10.0 kV 1.0 272

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

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

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

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

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

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

VOLUME 102, NUMBER 3

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

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

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

523

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

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

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

524 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

AccV SpotMagn Det WD Exp H——{ 200um

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

10.0 kV 1.0

Fig: 3:

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

WD Exp

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

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

9-integumental petal; (D) anterior spiracle.

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

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

VOLUME 102, NUMBER 3

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

525

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

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

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

526 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

Sa
BS Exp
as

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

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

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

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

0,.0-KV.1.0.- 2738x__SE—-A

VOLUME 102, NUMBER 3

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

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

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

527

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

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

DISTRIBUTION AND Hosts

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

528

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

100kV 10 23x E 39:7. 27

nies
7%
4

x

(Ae yg PY id
Me AAA hee)

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

BIOLOGY

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

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

VOLUME 102, NUMBER 3

¥
4
'

4
iv

'
Ny)
\i
\"

7

Fig. 6.

529

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

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

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

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

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

530

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

VOLUME 102, NUMBER 3

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

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

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

531

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

ACKNOWLEDGMENTS

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

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

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

ROBERT W. SITES AND MICHAEL R. WILLIG

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

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

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

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

534

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

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

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

MATERIALS AND METHODS

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

VOLUME 102, NUMBER 3

535

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

AWS
(10,10)

NEVADA: Lincoln Co.
Ash Warm Springs

3750 ft. elev.; 36°C

21 July 1992; CL 2711

J. T. and D. A. Polhemus

COL
(10,10)

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

J. T. Polhemus

EEL
(8,10)

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

R. K. Allen

HCF
(0,1)

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

9-TX-1965

E. J. Allen

HCF
(10,8)

IDAHO: Owhyee Co. (18)
nr. Bruneau

10-X-1975

A. D. Allen

IDA
(10,10)

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

T7S ROE sect. 26

22 Sept. 1988

R. S. Zack, coll.

INY
(10,10)

CALIFORNIA: Inyo Co.
Shoshone Hot Spring

CL 2904

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

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

27 August 1989

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

23 Sept. 1988; 12°C

coll: R. W. Sites

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

7 mi. W. Hwy 93

30°C; CL2898

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

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

28 Aug 1991; R. S. Zack and

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

16-X-1982

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

J. T. Polhemus

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

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

Janes AL
in analyses.

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

RESULTS

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

VOLUME 102, NUMBER 3

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

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

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

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

3)3)7/

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

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

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

538 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

Character PCl PeZ PC3 PC4

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

% variance (total) 0.922 0.026 0.016 0.008

% variance (shape) —_ 0.335 0.205 0.104

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

DISCUSSION

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

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

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

VOLUME 102, NUMBER 3 539

Males

UTA

SHA

—

o PC3

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

PC2

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

540

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

VOLUME 102, NUMBER 3

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

ACKNOWLEDGMENTS

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

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102(3), 2000, pp. 542-548

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

GEORGE A. FOSTER AND WAYNE N. MATHIS

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

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

Tethina and a diagnosis for the senior synonyms are provided.

Key Words:

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

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

Diptera, Tethinidae, Tethina, neotropics

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

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

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

KEY TO NEOTROPICAL SPECIES OF TETHINA

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

N

VOLUME 102, NUMBER 3

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

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

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

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

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

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

Tethina insulans Curran

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

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

543

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

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

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

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

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

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

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

544

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

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

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

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

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

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

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

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

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

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

545

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

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

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

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

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

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

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

546

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

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

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

VOLUME 102, NUMBER 3 547

a= hypandrium

,

Sa th

Mat

7 Pag
‘ raat
i can]
1 x OE
wk
fetes

Sebel ges -— hypandrial apodeme

—— aedeagal apodeme

SN \ 3
“SS ventral lobe A
surstylus

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

548

ACKNOWLEDGMENTS

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

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

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

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

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

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

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

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

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

asitoids, lady beetles, Diaspidiotus ancylus

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

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

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

550

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

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

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

METHODS

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

VOLUME 102, NUMBER 3

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

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

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

551

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

RESULTS

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

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

2nd m&f

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

crawler peak

Ist m&f

crawler peak

v

CS Se es eee Se Sener |

J F M A M J

Fig. 1.

A S O N D J

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

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

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

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

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

‘emerged. We found a few adult males on

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

VOLUME 102, NUMBER 3

500
400

+ SE)

07 11

May June

1000
800

No. of Crawlers/Tape (mean

13 26 9 25 9

May June

27 10 25 11

1.3 18 4
July

553

a) Chatsworth

30°16 28 18 7 23
July Aug. Sept. Oct.

15 28 14 6

Aug. Sept. Oct.

Dates in 1998

Fig. 2.

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

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

season.

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

guishable from second-instar females at that
time.

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

554

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

Percent of Population in each Instar

Dates in 1998

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

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

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

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

VOLUME 102, NUMBER 3

es
a

ee "eo
ee
ee"

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

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

35/5)

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

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

556 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

yea
we

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

VOLUME 102, NUMBER 3

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

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

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

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

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

S5y/

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

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

558

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

DISCUSSION

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

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

VOLUME 102, NUMBER 3

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

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

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

559

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

ACKNOWLEDGMENTS

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

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

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

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

STEPHEN D. GAIMARI AND MICHAEL E. IRWIN

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

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

each species is diagnosed.

Key Words:

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

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

phylogeny, revision, morphology, cladistics

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

METHODS

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

For each species, at least one male ab-

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

i

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

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

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

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

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

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

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

VOLUME 102, NUMBER 3

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

DESCRIPTIONS AND REDESCRIPTIONS OF
SPECIES

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

Ozodiceromyia Bigot

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

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

563

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

Ozodiceromyia mexicana-group

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

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

564

Table 2.

AMNH
ANIC
ANSP
ASUT
BMNH
BPBM
BYUC
CASC
CDFA

CICESE

CMNH
CNCI
CSUC

CUIC
DEIC
DENH
EBCC

EMEC
EMUS
ESUW

FMNH
FSCA
IEXA
INBC
INHS
IZAS
IZAV

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

List of codons used for repositories of specimens.

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

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

Academy of Natural Sciences, Philadelphia, Pennsylvania.

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

The Natural History Museum, London, United Kingdom.

Bernice P. Bishop Museum, Honolulu, Hawaii.

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

California Academy of Sciences, San Francisco, California.

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

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

Carnegie Museum of Natural History, Pittsburgh, Pennsylvania.

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

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

Cornell University Insect Collection, Ithaca, New York.

Deutsches Entomologisches Institut, Eberswalde, Germany.

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

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

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

Entomological Museum, Utah State University, Logan, Utah.

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

Field Museum of Natural History, Chicago, Illinois.

Florida State Collection of Arthropods, Gainesville, Florida.

Instituto de Ecologia, Xalapa, Vera Cruz, Mexico.

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

Illinois Natural History Survey, Champaign, Illinois.

Institute of Zoology, Academica Sinica, Beijing, China.

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

Kansas State University Insect Collection, Manhattan, Kansas.

Kagoshima University Insect Collection, Kagoshima, Japan.

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

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

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

Museo Entomologico, Universidad de Chile, Santiago, Chile.

Muséum d’ Histoire Naturelle, Genéve, Switzerland.

Muséum National d’Histoire Naturelle, Paris, France.

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

Montana State University Entomological Collection, Bozeman, Montana.

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

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

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

Naturhistoriska Riksmuseet, Stockholm, Sweden.

Natal Museum, Peitermaritzburg, Natal, South Africa.

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

Nevada State Department of Agriculture, Reno, Nevada.

New York State Museum, Albany, New York.

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

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

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

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

Stephen D. Gaimari private collection.

VOLUME 102, NUMBER 3

Table 2. Continued.

565

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

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

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

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

University of Arizona Insect Collection, Tucson, Arizona.

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

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

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

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

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

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

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

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

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

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

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

566

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

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

Male.—Body length: 8.5 mm.

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

VOLUME 102, NUMBER 3

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

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

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

567

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

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

Wing: 7.4 mm long.

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

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

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

568

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

VOLUME 102, NUMBER 3

569

Fig. 1.

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

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

Known distribution for Ozodiceromyia argentifera.

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

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

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

570

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

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

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

Male.—Body length: 7.6 mm.

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

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

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

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

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

VOLUME 102, NUMBER 3

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

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

571

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

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

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

Wing: 7.26 mm long.

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

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

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

572

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

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

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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

VOLUME 102, NUMBER 3

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

573

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

574

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Szerlip, J. A. Powell, 13-VIII-1974, at light
(1 36, MEI 052175, KSUC); Santa Cruz
County, K. Roever, 3-VIII-1959, black light
trap (1 2, MEI 038886, UAIC; 1 2, MEI
038887, IZAS); Canelo, G. D. Butler, 3-
VIII-1965 (1 2, MEI 038901, BPBM); Ca-
nelo Hills, Parker Cyn, R. L. Westcott, 30-
VII-1979 (1 3, MEI 079463, CNCI); Pena
Blanca, Werner, Olson, 11-VIII-1983 (1 2,
MEI 076929, UAIC); Sycamore Canyon,
near Ruby, K. Roever, 2-VIII-1959, light
trap (1 2, MEI 038876, UNAM; 1 2°, MEI
038885, INBC; 1 2, MEI 038900, UNSM)
V. L. Versterby, 6-IX-1963 (1 @, MEI
038880, MEIC); Patagonia Mountains, F
G. Werner, G. D. Butler, 8- VITI-1955, west-
ern slope (1 6, MEI 038867, SDGC), 9-
VIII-1966, western slope (1 2, MEI
038918, MEIC), G. D. Butler, E G. Werner,
9-VIII-1955, western slope (1 ¢6, MEI
038906, QCAZ; 1 3, MEI 038907, ANSP;
1 3, MEI 038908, SDGC; 1 6, MEI
038914, UAIC; 1 36, MEI 038917, UNSM),
Mount Washington, 1676.4 m, L. G. Be-
zark, R. A. Cunningham, D. E. Russell, 12-
VIII-1991 to 13-VIII-1991, Hg vapor and
UV blacklight (1 2, MEI 038115, WSUC);
Pajarito Mountains, R. H. Arnett Jr, E.
VanTassell, 6-VIII-1961 (1 2, MEI 038784,
FSCA); Santa Rita Mountains, Madera
Canyon, 3-VIII-1977 (1 36, MEI 052167,
SDGC), A. J. Gilbert, N. J. Smith, 11-VIII-
1981 (1 2, MEI 038119, SDGC; 1 2, MEI
038120, MEUC), D. K. Faulkner, 9-VIII-
1978 to 20-VIII-1978 (1 2, MEI 052168,
ZMAS), D. J. Knull, J. N. Knull, 1-VIII-
1972 (2 Y, MEI 052223, TAUD, Da
Faulkner, 20-VIII-1979 (1 2, MEI 052165,
BMNH), E. M. Fisher, 7-VIII-1962 to 9-
VIII-1962 (1 2, MEI 030943, CMNH; 1 °,
MEI 030929, CNCI; | 2, MEI 031008,
EMUS; 1 6, MEI 030942, KUIC), G. R.
Ballmer, K. Brown, 31-VII-1965 (1 3, MEI
038860, 1 2, MEI 038859, MEIC; 1 °,
MEI 038808, NHRM), L. G. Bezark, G. M.
Nishida, C. Kitayama, B. Tilden, 24-VIII-
1975, UV light (1 6, MEI 041768, NHRM;
1 3, MEI 041773, UMRM; 1 2, MEI
041770, MEIC), L. M. Martin, 14-VIII-

VOLUME 102, NUMBER 3

1949 (1 2, MEI 030936, MRSN), 15-VIII-
1949 (1 36, MEI 030934, LACM; 1 2, MEI
030933, CNCI; 1 2, MEI 030935, UMSP),
16-VIII-1949 (1 3, MEI 030941, LACM),
31-VII-1947 (1 2, MEI 030939, INHS), M.
A. Cazier, R. Schrammel, C. Vaurie, P. Vau-
ne. 13-Vil-1952.(1. 2 MEP, 038781;
AMNB), P. H. Sullivan, 31-VIII-1970 (1 @,
MEI 038856, KUIC), R. L. Westcott, 13-
VII-1964 (1 6, MEI 079570, WFBM; 1 °,
MEI 079569, FSCA), W. A. McDonald, 17-
VIII-1955 to 20-VIII-1955 (1 ¢, MEI
038902, LACM), 1,402.08 to 1,706.88 m,
4-VIII-1975 (1 3, MEI 076916, CICESE;
1 3, MEI 076918, FMNH; 2 6, MEI
076919, 076915, 1 2, MEI 076920, UAIC;
1, 22; MEL 076940, ‘-ZMHB:.«1. 2... MEI
O76917. -EBCE: a1. SieMEL 076914,
NHMW), 1,487.424 m, J. G. Franclemont,
20-VII-1959 (1 °°, MEI 038810, CUIC), L.
M. Martin, 20-VIII-1946 to 29-VIII-1946
(1 2, MEI 031062, EMEC; 1 2, MEI
030938, CNCD, V. L. Versterby, 8-IX-1963
(1 2, MEI 038894, QCAZ; 1 2, MEI
038895, MEIC), 12-VIII-1963 (1 6, MEI
038896, MUSM; | 36, MEI 038897, ANIC;
1 6, MEI 038898, IZAV), 21-VII-1963 (1
36, MEI 038881, MHNG), 22-IX-1963 (1
?, MEI 038883, UMRM), 31-VII-1963 (1
36, MEI 038882, MEUC), 1,524 m, K. W.
Brown, P. Petrulis, 3-[X-1971, UV light (1
6, MEI 033978, 1 @%, MEI 033979,
PMNH), 1,554.48 m, D. R. Davis, 10-VII-
1964 to 26-VII-1964 (1 3, MEI 052152,
SEMC; 1 2, MEI 052153, NMSA), Bog
Springs, 1,706.88 m, E. Lindquist, 6-VIII-
1973 (1 36, MEI 052231, CNCI); County
unknown, Floricera, Sta Rita Mts., Olson,
Burme, Frank, 30-VII-1980 (1 ¢, MEI
076938, UAIC); Garces, N. Banks, VIII (1
2, MEI 038864, MCZC) ; New Mexico, Hi-
dalgo County, 1,645.92 m, Skeleton Can-
yon, G. R. Ballmer, 12-VIII-1965 (1 ¢,
MEI 038892, USNM); 56.35 km E of
Douglas, Arizona, C. W. Sabrosky, 22-IX-
1965 (1 2, MEI 038857, USNM); Coro-
nado National Forest, Peloncillo Moun-
tains, 1.4 km W National Forest Boundary,
Black C.C.C. Dam Road, NFR63, 1661.16

S/S

m, M. A. Metz, 28-VI-1997, malaise trap
(2 36, MEI 103430, 103426, SDGC; 2 6,
MEI 103431, 103432, MEIC; 1 6, MEI
103433, OSEC; 1 3d, MEI 103427, ZMUC),
4.0 km W National Forest Boundary Black
C.C.C. Dam Road, NFR63, 1,682.496 m,
M. A. Metz, 28-VI-1997, malaise, 9 m
Fock’s (1 ¢, MEI 103428, INHS; 1 6, MEI
103429, ZMAS; 1 6, MEI 103424, TAUI;
1 3d, MEI 103425, MNHN).

Diagnosis.—The scape in this species is
setose, but most of the setae are short, with
few longer ones. The lower frons bulges
only slightly, with silver pruinescence lat-
erally, extending down along parafacial.
The face below the antenna also has silver
pruinescence. In males, the notum is dense-
ly covered with erect and appressed pale or
gold pile. In females, the notum and ab-
dominal tergites are covered with short,
fine, appressed brown and white pile. In the
male genitalia, the setae of the epandrium
and gonocoxites are orange. The subepan-
drial plate is attached to the epandrium at
the posterolateral corners, and the sclero-
tized portion of the subepandrial plate is V-
shaped posteriorly. The gonocoxites lack a
suture along the midline separating the two
lateral halves. The outer gonocoxal process
is widened distally, with a small patch of
setae on the dorsolateral surface. The ven-
tral gonocoxal process of the gonocoxites is
present, elongated and flattened; there are
no clumped setae at its base. The base of
the ventral lobes is U-shaped and broad be-
tween the two halves. The parameral sheath
of the aedeagus is smooth dorsally. The
ventral apodeme lacks a ventral keel. The
distiphallus is swollen basally, and is re-
curved and parallel with the ventral apo-
deme at the tip. In the female terminalia,
the furca is not notched posteriorly, and
lacks a posterolateral peg. The anterolateral
furcal prongs are separated by a distance
subequal to the greatest width of the furca.
The common spermathecal duct is broad
basally, at half the width of the entire furca;
the duct tapers quickly.

Rip:

Autapomorphy.—Gonocoxal setae pale
(character 26, state 0).

Distribution.—The known distribution
for this species is found in Fig. 2. Most of
the localities fall within an elevational
range of 1,000—2,000 m, although 1 2 was
collected near 500 m. The distribution en-
compasses parts of the following biogeo-
graphical regions: the Provincia Xer6fila
Mexicana, the northern part of the Provin-
cia Mesoamericana de Montafia, and the
montane regions of the southeastern quarter
of Arizona and southwestern New Mexico.

Etymology.—Patronym for Todd P. Li-
vdahl, professor of Biology at Clark Uni-
versity, Worcester, Massachusetts, alma
mater of SDG; to be treated as a noun in
apposition.

Biology.—This species is most active in
July and August, with a range from the end
of June (4 d<d) through September. This
species seems to be commonly collected at
UV and other light sources at night.

Ozodiceromyia mexicana Bigot
(Figs. 9-10, 16, 21, 25, 30-31, 34-35,
41)

Ozodiceromyia mexicana Bigot 1890: 321.

Type locality: Mexico. Type: HT? in

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Bae

Known distribution for Ozodiceromyia livdahli.

BMNH. Bigot 1890: 323 (key ref.);
Krober 1912: 211 (repr. orig. descr.),
1913: 8 (cat. cit.); Cole 1923: 19—20 (an-
not. listing); Gaimari 1998: 124-125 (lit.
list), 157-159 (in phylogeny).

Ozodiceromya (sic) mexicana: Bigot 1890:
321 (orig. descr.); Irwin and Lyneborg
198la: 196 (illust.: coxa), 199 (illust.: an-
tenna), 254 (illust.: ¢ genit.), 257 (list-
ing).

Ozodiceronyma (sic) mexicana: Godman
1901: 378 (listing).

Euphycus setosus Kréber 1912: 211. Type
locality: Mexico. Type: LTd (designated
herein) in NHMW. Irwin and Lyneborg
1981a: 257 (prop. syn.).

Ozodiceromya (sic) setosa: Irwin and Ly-
neborg 198la: 257 (comb. change).

Female holotype.—The HT?® is in poor
condition, i.e., the abdomen and thorax are
completely covered with orange fungus.
Fortunately, the head is mounted separately
and is fungus-free. The head is distinct
enough to associate this specimen with a
recognizable species. The following de-
scription deals only with the characteristics
that can be seen in the specimen. Following
this, the full redescription of the recognized

VOLUME 102, NUMBER 3

species comes from the LT¢ of the junior
synonym, Euphycus setosus Krober.

Body length: 7.1 mm.

Head: 1.13 mm long, 1.73 mm wide,
1.28 mm high. Distance between eyes at
level of anterior ocellus 0.33 mm; at anten-
nal level 1.01 mm; at genal level 0.96 mm.
Scape 0.59 mm long, 0.12 mm wide; with
long, fine and thick, black setae, evenly dis-
tributed over entire scape (including median
surface). Pedicel 0.18 mm long, barrel-
shaped, setose (including median surface).
Antennal base to nearest edge of eye 0.20
mm. Frons bulging 0.17 mm beyond eye in
lateral view. Parafacial lacking pile; silver
pruinescence in thin line along edge of eye;
face and parafacial otherwise glabrous,
shiny black. Frons shiny black; lower frons
globose and bulging to antennal insertion;
upper frons distinct from lower frons, bulg-
ing. Genal pile short and black.

Legs: Foreleg dark brown, except fore-
tibia paler and lacking setae on proximal,
dorsal surface. Midleg dark brown, except
midtibia orange basally.

Wing: 5.85 mm long.

Male.—Lectotype of Euphycus setosus
Kroéber, to represent the species Ozodicer-
omyia mexicana.

Body length: 6.9 mm.

Head (Fig. 9): 0.90 mm long, 1.59 mm
wide, 1.35 mm high. Distance between eyes
at antennal level 0.83 mm; at genal level
1.01 mm. Antenna dark brown. Scape 0.71
mm long, 0.18 mm wide; with long, fine
and thick, black setae, evenly distributed
over entire scape (including median sur-
face). Pedicel 0.20 mm long, barrel-shaped,
setose (including median surface). First fla-
gellomere 0.60 mm long, 0.15 mm wide;
with short setae covering basal two-thirds.
Stylus 0.09 mm long, inserted subapically.
Antennal base to nearest edge of eye 0.26
mm. Frons globose, bulging 0.18 mm be-
yond eye in lateral view. Parafacial lacking
pile, with silver pruinescence only along
eye edge; pruinescence extends dorsally in
thin line along eye margin halfway up
frons; face and parafacial otherwise gla-

577

brous, shiny black. Frons otherwise shiny
black; with long, fine, black setae (up to
0.38 mm long); upper frons bare. Genal pile
short, darkened. Palpal pile white basally,
dark brown distally. Postgenal pile white;
occipital pile black; occiput with silver
pruinescence only along edge of eye, and
with several black setae. Median occipital
sclerite flattened; glabrous, shiny black; up-
per edge not rounded. Postocular setae
black, arranged in single, transverse row.
Ocellar tubercle shiny black, with fine, for-
ward-directed, black setae.

Thorax: Scutum and scutellum with
erect black pile sparsely interspersed with
appressed gold pile. Scutum 2.08 mm long,
1.70 mm wide; ground color black; with sil-
ver-grey pruinescence dorsally, lacking
pruinescence laterally; median vitta diffuse
bronze; dc vittae absent; dc setae absent [or
1 pair present]. Scutellum with silver prui-
nescence, reduced pruinescence anteriorly.
Halter yellow. Katatergite with dense, long,
orange pile. Anepimeron, katepimeron, and
meron lacking pile, and with reduced silver
pruinescence, appearing as vertical brown
stripe from wing base to between second
and third coxae. Anepisternum with silver
pruinescence, and with white pile. Katepis-
ternum with silver pruinescence, with white
pile only on vertical crest along middle of
pleurite. Prepimeron with silver pruinesc-
ence, lacking pile.

Legs: Coxae with silver pruinescence,
posterior surfaces less so. Posterior surface
of hindcoxa with long, white pile (Fig. 16).
Femora dark brown. Fore- and midfemora
with appressed, scale-like, white pile, and
with long, erect, recumbent, white and
black pile. Hindfemur with appressed,
scale-like brown and white pile. Tibiae with
short, erect, black setae; proximal, dorsal
surfaces with sparser setal covering, or
nearly bare. Foretibia orange basally, be-
coming dark brown distally; clavate distal-
ly. Midtibia orange, becoming darkened at
distal tip. Hindtibia orange basally, becom-
ing dark brown distally. Foretarsus dark
brown. Mid- and hindtarsi dark brown ex-

578

cept for basal two tarsomeres orange at
their bases.

Wing: 6.24 mm long. Basal costal lobe
with setae arranged in line along outer
edge, extending into outer row of costal se-
tae; second line of setae along base of basal
costal lobe, extending into inner row of cos-
tal setae. Entire membrane slightly dark-
ened; veins and membrane orange basally,
brown distally.

Abdomen: Tergite 1 with erect and re-
cumbent gold pile, except dark brown me-
dioposteriorly; remaining tergites with erect
and recumbent, white pile; dorsally, with
silver pruinescence; laterally, lacking prui-
nescence (showing dark brown ground col-
or), except tergite 1 and posterior edge of
tergite 2 with silver pruinescence. Sternites
with silver pruinescence only along anterior
edge of sternite 2.

Terminalia (MEI 052240): Sternite 8
with black setae along posterior edge,
which is emarginate medially. Tergite 8
dumbell-shaped, with black setae along
posterior edge. Epandrium (Fig. 25) 0.24
mm long, 0.63 mm wide at widest point;
orange; emarginate anteriorly; dorsal sur-
face with white setae on posterior two-
thirds; lateral edges parallel; posterolateral
corners extended posteriorly. Subepandrial
plate (Fig. 25) unattached to epandrium;
sclerotized portion reduced, divided into
two small plates. Sclerotized portion of cer-
ci 0.23 mm long; extending posteriorly be-
yond posterolateral corners of epandrium;
subequal in length to ventral epandrial
sclerite. Gonocoxites (Figs. 30-31) 0.77
mm wide; orange; with white setae, up to
0.50 mm long; fusion 0.35 mm long at mid-
line, with suture. Inner gonocoxal process
flanged, knob with several white setae. Out-
er gonocoxal process flange-like, 0.23 mm
long; bare. Gonocoxal apodeme entirely
within anterior edge of gonocoxite; lacking
sclerotized bridge to parameral sheath of
phallus. Ventral gonocoxal process absent,
lateral part of corresponding edge with
dense patch of fine orange setae, up to 0.30
mm long. Ventral lobes distinct, fused ba-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

sally by thin, transparent membrane. Gon-
ostylus expanded ventrally into large lobe;
with dorsobasal lobe; lacking subapical, lat-
eral spur. Dorsal apodemes (Fig. 34) of ae-
deagus parallel; subequal in length to ven-
tral apodeme; parameral sheath with dis-
tinct transverse wrinkles dorsally. Ventral
apodeme thin, of uniform width; lacking
ventral keel. Ejaculatory apodeme 0.35 mm
long; stick-like, slightly expanded distally.
Lateral ejaculatory process a complete ring
dorsally, but notched; set into aedeagus;
lightly sclerotized. Distiphallus long, re-
curved; distally parallel with ventral apo-
deme (Fig. 35).

Female (MEI 052163).—Similar to 6
except as follows: body length 8.6 mm.

Head (Fig. 10): 1.10 mm long, 1.76 mm
wide, 1.28 mm high. Distance between eyes
at level of anterior ocellus 0.32 mm; at an-
tennal level 0.83 mm; at genal level 0.95
mm. Scape 0.81 mm long, 0.15 mm wide.
First flagellomere 0.71 mm long, 0.15 mm
wide. Antennal base to nearest edge of eye
0.32 mm. Frons shiny black; lower frons
globose and bulging to antennal insertion,
with fine, black setae (up to 0.12 mm long)
in patch above antennal base; upper frons
bulging and distinct from lower frons.

Thorax: Scutum and scutellum with re-
cumbent black pile and appressed gold pile.
Scutum 2.08 mm long, 1.80 mm wide. Ane-
pisternum with reduced silver pruinescence;
pile of anepisternum and katepisternum
sparse, short, white.

Wing: 6.12 mm long.

Abdomen: Tergites and sternites lacking
pruinescence (showing dark brown ground
color); tergites with appressed and semi-ap-
pressed gold pile.

Terminalia (paratype, MEI 070426) (Fig.
41): Furca 0.53 mm long, 0.38 mm wide;
anterior edge not sclerotized; anterolateral
prongs dorsoventrally flattened, separation
subequal to greatest width of furca; pos-
terolaterally with small, ventromedially ori-
ented peg; posterior edge notched. Furcal
bulla not sclerotized; posteriorly with in-
dented cavity. Gonopore basal to furcal bul-

VOLUME 102, NUMBER 3

la, originating within posterior cavity. Com-
mon spermathecal duct 1.14 mm long.
Spermathecal ducts originate from clean tri-
furcation at base of central sac duct; central
sac duct absent. Central sac 0.89 mm long,
0.30 mm wide. Spermatheca 0.15 mm long,
0.15 mm wide; rounded distally, but basal
edge flattened.

Type materials—HOLOTYPE 2°: (MEI
103180) top label: Mexic[o]; with the fol-
lowing handwritten, second label (note, this
is a standard provisional label of Bigot
which he never updated): n. genus, Ozodi-
ceromyia, J. Bigot, O. mexicana 2, n. sp.
Inedict. [= unedited, unpublished], Quincy
9b, 1888 J. Bigot, Mexique. This specimen,
which is from BMNH, is in poor condition
and is entirely covered with orange fungus,
except for the head, which is mounted sep-
arately on the same pin.

LECTOTYPE ¢ (here designated to fix
the current interpretation of this name and
to ensure stability and uniformity in its fu-
ture interpretation) of Euphycus setosus:
(MEI 084117) with the following labels:
Bilimek, Mexico, 1871, Guadalupe/Cotype
(pink label)/Phycus setosus Krob., det.
Kréber 1911/Lectotypus 6, Euphycus se-
tosus Kroéber, designated in 1998 by SD
Gaimari et ME Irwin (red label). This
pinned specimen is in very good condition,
and is deposited in NHMW. PARALEC-
TOTYPES: Bilimek, Mexico, 1871/Type
(pink label)/Phycus setosus Krob., det.
Kréber 1911/Paralectotypus 2, Euphycus
setosus Krober, designated in 1998 by SD
Gaimari et ME Irwin (red label) (pinned °,
MEI 084118, NHMW); Mexico/Cotype
(pink label)/Phycus setosus Krob., det.
Kréber 1911/Paralectotypus 6, Euphycus
setosus Kroéber, designated in 1998 by SD
Gaimari et ME Irwin (red label) (2 pinned
33, MEI 084115, 084116, NHMW); Type
(pink label)/Mexico, Sta. Fe, 1871/Type
No. 24183, U.S.N.M. (red label)/Euphycus
setosus Kroéb., Kréber det. 1911/Paratype
only, det. WwWirth/Ozodiceromyia mexi-
cana Bigot, det WWirth/Paralectotypus <6,
Euphycus setosus Kréber, designated in

7/8)

1998 by SD Gaimari et ME Irwin (red la-
bel) (pinned 3, MEI 032957, USNM).
Materials examined.—MEXICO: Coa-
huila, 30.6 km SE of Saltillo, Highway 57,
2,194.56 m, C. O’Brien, L. O’Brien, G.
Wibmer, 12-IX-1982 (1 2, MEI 052245,
MCZC); Distrito Federal, TX-1898 (2 &,
MEI 032959, 032960, USNM); Mexico
Citys? R.) Muller iG. <d3MEE032958;
USNM), W. G. Downs, 14-X-1951 (¢ and
2, MEI 038775, 038774, AMNH); Pedre-
gal de San Angel, H. Perez, 27-IX-1969 (1
2, MEI 084201, KSUC); Tizapan (1 °,
MEI 052178, AMNH; 2 6, MEI 084194,
084195, UNAM); Guerrero, 12.9 km E of
Taxco, 1,828.8 m, R. H. Painter, E. M.
Painter, 15-[X-1963 (1 6, MEI 052188,
VAIC); 22 km NE Telolopan, 1,530 m, J.
A. Powell, J. A. Chemsak, 16-[X-1982 (1
2, MEI 081567, EMEC); Hidalgo, 16.1 km
S of Zimapan, 1,889.76 m, J. A. Powell,
28-IX-1975 (1 3, MEI 052247, SDGC);
Tepeapulco, G. E. Bohart, W. J. Hanson,
18-IX-1974 (1 36, MEI 007797, EMUS; 1
6, MEI 007795, FSCA; 1 3, MEI 007796,
UAIC; 1 2, MEI 007798, EBCC); Jalisco,
6.4 km W Mazamilta, 2,072.64 m, R. E
Smith, 16-X-1950 (1 2, MEI 052182,
ZMHB); Mexico, C. W. Johnson coll. (1 6,
MEI 032961, USNM; 1 3, MEI 052180,
MCZC); Amecameca, R. Dreisbach, K.
Dreisbach, 25-[X-1957 (1 6, MEI 052192,
MEIC; 1 2, MEI 052181, INHS); 45.1 km
SE of San Juan del Rio, G. E. Bohart, W.
J. Hanson, 31-VIII-1974 (1 6, MEI
007800, EMUS); Michoacan, 4.8 km W of
Morelia, 1,859.28 m, R. H. Painter, E. M.
Painter, 21-X-1963 (1 do, MEI 052185,
KUIC), 1,950.72 m, R. H. Painter, E. M.
Painter, 20-[X-1963 (1 6, MEI 052190,
KUIC; 1 36, MEI 052191, FSCA; 1 2°, MEI
090233, SDGC); 6 km S of Ocampo,
Lag.[una] Verde, A. L. Norrbom, 6-X-1991,
observed waving forelegs acting like a
wasp (1 2, MEI 052163, USNM); 11.3 km
E of Quiroga, 2,346.96 m, R. H. Painter, E.
M. Painter, 21-I[X-1963 (1 ¢, MEI 052176,
KSUC); 14.5 km W of Ciudad Hidalgo,
2,194.56 m, R. H. Painter, E. M. Painter,

580

19-IX-1963 (1 2, MEI 052177, KSUC);
14.5 km W of Morelia, 2,072.64 m, J. A.
Powell, J. A. Chemsak, T. Eichlin, T. P.
Friedlander, 9-X-1975 (3 3, MEI 052236,
0522387052239, CASC:'1 6, MEI 052237;
ZMAS); 20.9 km N of Morelia, 1,981.2 m,,
R. H. Painter, E. M. Painter, 22-[X-1963 (1
2, MEI 107043, MEIC); N of Patzcuaro, 3
km E of Zintzuntzan, 2,100 m, T. Griswold,
26-X=1987 (1 2, METI. 052229; EMUS);
SW of Patzcuaro, 3 km E of Zirahuén,
2,000 m, T. Griswold, 27-X-1987 (1 6,
MEI 052227, WFBM; 1 2, MEI 052228,
EMUS); Morelos, 19 km E Cuernavaca,
Canyon del Lobo, E. M. Fisher, 15-X-1986
(6 3, MEI 070420, 070421, 070423,
070424, 070425, 070427, 1 ¢, MEI
070419, CDFA; 1 6, MEI 070422, 1 2,
MEI 070428, SDGC; 1 3, MEI 070429,
ZMUC; 1 6, MEI 070430, MNHN; 1 <4,
MEI 070431, INHS; 1 2, MEI 070426,
MEIC; 1 2, MEI 070432, CNCI); Canon
del Lobos, M. Rodriguez, 19-X-1985 (1 6,
MEI 084184, CICESE); Tepoztlan, 1,700
m, C. D. Michener, R. Murillo, J. M. La-
bougle, 13-XI-1980 (1 36, MEI 052220,
SEMC); Nuevo Leon, 4.8 km E of Galeana
Jct., 1,828.8 m, J. A. Chemsak, J. Powell,
A. Michelbacher, M. M. Michelbacher, 15-
IX-1976 (1 36, MEI 081532, EMEC; 1 9,
MEI 081530, NHMW; 1 ¢, MEI 081531,
SDGC); 6.4 km W Iturbide, 1,676.4 m, J.
A. Chemsak, 22-IX-1976, at light (1 6,
MEI 052240, MEIC; 1 2, MEI 052241,
CASC), J. A. Chemsak, J. A. Powell, 13-
IX-1976 to 14-IX-1976, at light (1 6, MEI
052197, EMEC); 8.1 km E of Galeana Jct.,
J. A. Powell, J. A. Chemsak, 16-[X-1976 to
17-IX-1976 (1 3, MEI 052202, MNHN; 1
36, MEI 052204, AMNH; 1 6, MEI
052205.) SEMC; 1 (dé; SME 052206;
BMNH), J. A. Powell, J. A. Chemsak, A.
E. Michelbacher, M. M. Michelbacher, 16-
IX-1976 to 17-IX-1976 (10 o, MEI
052196, 052199, 052200, 081551, 081550,
081549, 081548, 081547, 081538, 081537,
1 ¢&, MEI 081539, EMEC; 1 6, MEI
052201, UCRC; 1 3, MEI 052198, EBCC;
1 2, MEI 081536, ZMUC); 11.3 km W of

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Iturbe, 1,767.84 m, R. H. Painter, E. M.
Painter, 25-[X-1963 (1 6, MEI 052187,
KSUC); 14.5 km S of junction 60 to Dr.
Arroyo, R. Turnbow, 22-X-1979 (1 36, MEI
052174, UGCA); 16.1 km E of San Rob-
erto, 2,133.6 m, 15-[X-1976 (1 36, MEI
052203, MZSP); 16.1 km NW Provedencia,
1828.8 m, J. A. Chemsak, J. Powell, A.
Michelbacher, M. Michelbacher, 27-IX-
1976 (6 and 2 in copula, MEI 081553,
081552, EMEC); 16.1 km N Provedencia,
J. Powell, J. A. Chemsak, A. Michelbacher,
M. M. Michelbacher, 25-[X-1976 (1 6,
MEI 081534, EMEC); 22.5 km N San Juan-
ito, 2,194.56 m, J. A. Chemsak, J. Powell,
A. Michelbacher, M. M. Michelbacher, 22-
IX-1976 (2 3, MEI 081528, 081529,
EMEC); 66 km S of Saltillo, 1,889.76 m,
7-IX-1962, U. Kans. Mex. Exped., on flow-
ers of Encelia farinosa (2 6, MEI 052184,
052186, KSUC); 66 km SE of Saltillo,
1,889.76 m, N. Marston, 7-[X-1962 (1 6,
MEI 052184, KSUC); 70.8 km SE of Sal-
tillo, G. E. Bohart, W. J. Hanson, 30-VIII-
1974 (1 3, MEI 007803, EMUS); probably
Nuevo Leon, Guadalupe, [D.] Bilimek,
1871 (1 3, 109316, NHMW); Oaxaca, 11.3
km SE of Nochixtlan, 2,133.6 m, J. A.
Powell, J. A. Chemsak, T. Eichlin, T. P.
Friedlander, 7-X-1975 (2 6, MEI 052242,
052243, CASC; 1 6, MEI 052244, DEIC;
1 2, MEI 052235, MZSP); 14.5 km NE
San Jose del Estado, 2,590.8 m, R. E Smith,
25-X-1966 (1 36, MEI 081533, EMEC);
San Luis Potosi, 31 km SE San Luis Potosi,
2,250 m, E. Ramirez, 17-X-1996 (1 3, MEI
103423, NHMW; 1 6, MEI 103422,
EBCC; 1 6,-MEI 103420, ZMHB:) ives
MEI 103419, UCRC), 18-X-1996 (1 ¢,
MEI 103403, SDGC; 1 3, MEI 103421,
CNCI); Veracruz, Perote, San Juan del
Monte, A. Cordoba, 14-X-1989 to 15-X-
1989 (1 36, MEI 080258, 1 2, MEI 080259,
IEXA; 1 2, MEI 080260, BMNH); Zaca-
tecas, 3.2 km S of Luis Moya, 1,920.24 m,
R. H. Painter, E. M. Painter, 3-[X-1962 (1
36, MEI 052189, INBC); (unknown state),
Carr. la para cuautla km 2; 15 Nov. 1978,
coll: E. Olvara (1 6, MEI 084199, UCDC;

VOLUME 102, NUMBER 3

1 3, MEI 084202, UNAM), 19 Nov. 1978
(1 36, MEI 084200, DEIC); Cuamauaca
moz, km 57.5 Autopista, 15 Nov. 1978, J.
C. Medina (1 6, MEI 084188, UCDC); San
Lucas PUZ., Matorrol espinoso [= spiny
bush], 21-[X-89, A. Exuihua M. (1 2°, MEI
079915, UNAM).

Diagnosis.—The scape in this species is
densely setose, with long setae. The lower
frons is globose and distinctly bulging, is
shiny black, lacking pruinescence, and has
only a thin line of silver pruinescence along
the eye margin extending down along the
parafacial. The face below the antenna is
shiny black, lacking pruinescence. In males,
the notum is covered with erect black pile
interspersed with appressed gold pile. In fe-
males, the notum and abdominal tergites are
covered with short, fine, appressed gold
pile. In the male genitalia, the setae of the
epandrium and gonocoxites are orange. The
subepandrial plate is attached to the epan-
drium only through a thinly membranous
connection laterally. The sclerotized portion
of the subepandrial plate is not V-shaped
posteriorly, but is reduced to two smaller
lateral plates. The gonocoxites have a su-
ture along the midline separating the two
lateral halves. The outer gonocoxal process
tapers evenly towards apex and is bare of
setae. The ventral gonocoxal process of the
gonocoxites is absent; there is a distinct,
dense clump of orange setae on the poster-
oventral edge of the gonocoxite. The base
of the ventral lobes is V-shaped and narrow
between the two halves. The parameral
Sheath of the aedeagus has distinct, trans-
verse wrinkles dorsally. The ventral apo-
deme lacks a ventral keel. The distiphallus
is evenly tapered to apex, lacking any
swelling, and is recurved and parallel with
the ventral apodeme at the tip. In the female
terminalia, the furca is notched posteriorly,
and ventromedially oriented pegs are pres-
ent on the posterolateral portion of furca.
The anterolateral furcal prongs are separat-
ed by a distance subequal to the greatest
width of the furca. The common sperma-

581

thecal duct is not widened basally, and ta-
pers normally.

Autapomorphies.—Lower frons globose
and bulging to antennal insertion (character
9), posterolateral surface of hindcoxa pilose
(character 14), parameral sheath with trans-
verse wrinkless dorsally (character 17),
subepandrial plate attached to epandrium
laterally only (character 24), posterolateral
corners of epandrium held by separate,
sclerotized membrane, (character 25), pres-
ence of distinct internal keel where gono-
coxites fused (character 27), posterior edge
of furca with distinct notch (character 36),
medially directed peg present on postero-
lateral furcal margin (character 37).

Distribution.—The known distribution
for this species is found in Fig. 3. Nearly
all of the specimens were collected at ele-
vations ranging from 1,500—2,600 m. The
distribution encompasses parts of the fol-
lowing biogeographical regions: the Prov-
incia Xerdfila Mexicana, and the Provincia
Mesoamericana de Montafia.

Biology.—This species is most active in
September and October, with several spec-
imens collected in late August and in the
middle of November. Two males were col-
lected on flowers of Encelia farinosa A.
Gray (Asteraceae), and one female was ob-
served in life waving her forelegs in appar-
ent mimicry of the antennae of a sphecoid
wasp.

Ozodiceromyia parargentifera Gaimari
and Irwin, new species
(Figs. 36-37, 40)

Male.—Body length: 7.0 mm.

Head: 1.04 mm long, 2.00 mm wide,
1.58 mm high. Distance between eyes at an-
tennal level 0.62 mm; at genal level 0.83
mm. Antenna brown, except basal half of
scape orange. Scape 0.95 mm long, 0.15
mm wide; with short, fine, black setae,
evenly distributed over entire scape (includ-
ing median surface), and few larger setae.
Pedicel 0.18 mm long, barrel-shaped, setose
(including median surface). First flagello-
mere 0.96 mm long, 0.15 mm wide; with

Fig. 3:

short setae covering basal two-thirds. Stylus
0.09 mm long, inserted subapically. Anten-
nal base to nearest edge of eye 0.15 mm.
Frons bulging 0.09 mm beyond eye in lat-
eral view. Face below antenna with silver
pruinescence. Parafacial lacking pile, with
silver pruinescence extending to antennal
base and dorsally along eye margin halfway
up frons. Frons otherwise shiny black; with
few, short, fine, black setae (up to 0.17 mm
long) dorsolateral to antennal base; remain-
der of frons bare. Genal pile short, dark-
ened. Palpal pile white basally, dark brown
distally. Postgenal and occipital pile white;
occiput with silver pruinescence only along
edge of eye, and with several black setae.
Median occipital sclerite flattened; gla-
brous, shiny black; upper edge not rounded.
Postocular setae black, arranged in single,
transverse row. Ocellar tubercle with silver
pruinescence, and fine, forward-directed,
black setae.

Thorax: Scutum and scutellum with fine,
erect black pile. Scutum 2.22 mm long,
1.53 mm wide; ground color black; with sil-
ver-grey pruinescence dorsally, lacking
pruinescence laterally; median vitta diffuse
bronze; dc vittae absent; 1 pair dc setae.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Known distribution for Ozodiceromyia mexicana.

Scutellum with silver pruinescence, reduced
pruinescence anteriorly. Halter brown. Ka-
tatergite with dense, long, white pile. Ane-
pimeron, katepimeron, and meron lacking
pile, and with reduced silver pruinescence,
appearing as vertical brown stripe from
wing base to between second and third cox-
ae. Anepisternum with silver pruinescence,
and with white pile. Katepisternum with sil-
ver pruinescence, with white pile only on
vertical crest along middle of pleurite. Pre-
pimeron with silver pruinescence, lacking
pile.

Legs: Coxae with silver pruinescence,
posterior surfaces less so. Posterior surface
of hindcoxa lacking pile. Forefemur dark
brown; dorsally with appressed, scale-like
white pile, and erect, white and brown pile.
Foretibia with short, erect, black setae;
proximal, dorsal surface with sparser setal
covering; orange on basal half, becoming
dark brown distally; clavate distally. Fore-
tarsus dark brown. Mid- and hindlegs bro-
ken off.

Wing: 5.88 mm long. Basal costal lobe
with setae arranged in line along outer
edge, extending into outer row of costal se-
tae; second line of setae along base of basal

VOLUME 102, NUMBER 3

costal lobe, extending into inner row of cos-
tal setae. Entire membrane slightly dark-
ened; veins and membrane orange basally,
darker distally.

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

Terminalia: Sternite 8 with fine black se-
tae restricted to posterior edge, which is
emarginate medially. Tergite 8 dumbell-
shaped, with fine black setae restricted to
posterior edge. Epandrium 0.18 mm long,
0.68 mm wide at widest point; orange;
emarginate anteriorly; dorsal surface with
black setae on posterior half; lateral edges
parallel; posterolateral corners extended
and pointed posteriorly. Subepandrial plate
attached to posterolateral and posterior edg-
es; sclerotized portion V-shaped posteriorly.
Sclerotized portion of cerci 0.20 mm long;
extending posteriorly slightly beyond pos-
terolateral corners of epandrium; subequal
in length to ventral epandrial sclerite. Gon-
ocoxites 0.72 mm wide; orange; with black
setae, up to 0.47 mm long; fusion 0.36 mm
long at midline, lacking suture. Inner gon-
ocoxal process flanged; knob with several
black setae. Outer gonocoxal process
flange-like, 0.33 mm long; with short, fine
setae along dorsal edge. Gonocoxal apode-
me entirely within anterior edge of gono-
coxite; lacking sclerotized bridge to para-
meral sheath of phallus. Ventral gonocoxal
process 0.27 mm long, flanged; bare. Ven-
tral lobes distinct, fused basally by thin,
transparent membrane. Gonostylus expand-
ed ventrally into large lobe; with dorsobasal
lobe; with subapical, lateral spur. Dorsal
apodemes (Fig. 36) of aedeagus parallel;
shorter than ventral apodeme; parameral
sheath smooth dorsally. Ventral apodeme
uniformly wide; with ventral, longitudinal
keel. Ejaculatory apodeme 0.30 mm long;
stick-like, expanded distally and bilobed.

583

Lateral ejaculatory process a complete ring
dorsally, but notched; set into aedeagus;
lightly sclerotized. Distiphallus swollen ba-
sally, long, undulating, recurved; distally
perpendicular to ventral apodeme (Fig. 37).

Female.—Similar to ¢ except as follows:
body length 9.6 mm.

Head: 1.22 mm long, 2.28 mm wide,
1.59 mm high. Distance between eyes at
level of anterior ocellus 0.50 mm; at anten-
nal level 0.95 mm; at genal level 1.04 mm.
Antenna brown. Antennal base to nearest
edge of eye 0.30 mm. Parafacial pruinesc-
ence ends at antennal level. Frons shiny
black; lower frons bulging, with short, fine,
black setae (up to 0.09 mm long) dorsolat-
eral to antennal base; upper frons bulging
and wrinkled, distinct from lower frons.
Median occipital sclerite and upper edge
rounded; transverse row of black setae
across median occipital sclerite in addition
to row of postocular setae.

Thorax: Scutum and scutellum with
erect and recumbent, short, black pile [one
paratype also has short, fine, appressed gold
pile]. Scutum 2.58 mm long, 1.76 mm
wide. Pile of anepisternum and katepister-
num short, white; densest on upper half of
anepisternum.

Legs: Foretibia dark brown [or orange
basally]. Mid- and hindfemora dark brown.
Mid- and hindtibiae light brown. Mid- and
hindtarsi dark brown, except for basal two
tarsomeres mostly orange.

Wing: 7.20 mm long.

Abdomen: Tergites with short, recum-
bent, white pile. Tergites and sternites with
reduced silver pruinescence (showing dark
brown ground color), except posterolateral
edges of basal tergites with silver pruinesc-
ence.

Terminalia (paratype, MEI 027022):
Furca (Fig. 40) 0.53 mm long, 0.32 mm
wide; anterior edge not sclerotized; antero-
lateral prongs dorsoventrally flattened, sep-
aration narrower than greatest width of fur-
ca. Furcal bulla not sclerotized; posteriorly
with indented cavity. Gonopore basal to
furcal bulla, originating within posterior

584 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

cavity. Common spermathecal duct basally
0.11 mm wide, tapering to 0.05 mm within
0.33 mm distance from gonopore; 0.62 mm
long. Spermathecal ducts originate from
clean trifurcation with central sac duct; cen-
tral sac duct wider in basal diameter than
spermathecal ducts. Central sac duct 0.45
mm long. Central sac 0.68 mm long, 0.41
mm wide. Spermatheca 0.17 mm in diam-
eter; rounded, but basal edge slightly flat-
tened.

Type materials—HOLOTYPE 36 (MEI
027017) with the following label: MEXI-
CO, Guerrero, 4 mi. W of Chilpancingo,
July 15, 1984, Carroll, Schaffner, Friedlan-
der. This pinned specimen, which is depos-
ited in CASC (with permission of TAMU),
is in good condition except for its missing
mid- and hindlegs.

Materials examined.—PARATYPES.
MEXICO: same label as holotype (1 &,
MEI 027022, SDGC); Michoacan, 9.3 km
SE Quiroga, 25-VIII-1991, W. FE Barr, on
roadside vegetation, 2,653 m (1 2, MEI
079906, WFBM); Puebla, 4.4 mi. SW [San
Francisco] Acatepec, 9-VII-1981, Bogar,
Schaffner, Friedlander (allotype 2, MEI
027003, TAMU).

Diagnosis.—This species is much like
Ozodiceromyia argentifera and Ozodicero-
myia livdahli. The scape in this species is
setose, but most of the setae are short, with
few longer ones. The lower frons bulges
only slightly, with silver pruinescence lat-
erally, extending down along parafacial.
The face below the antenna also has silver
pruinescence. In males, the notum has some
fine, erect, black setae (the holotype, the
only known male, appears rubbed dorsally,
and so presence or condition of appressed
pile is unknown). In females, the notum has
short, fine, erect and appressed black pile,
and occasionally appressed gold pile. In the
male genitalia, the setae of the epandrium
and gonocoxites are black. The subepandri-
al plate is attached to the epandrium at the
posterolateral corners, and the sclerotized
portion of the subepandrial plate is V-
shaped posteriorly. The gonocoxites lack a

suture along the midline separating the two
lateral halves. The outer gonocoxal process
is widened distally, with a small patch of
setae on the dorsolateral surface. The ven-
tral gonocoxal process of the gonocoxites is
present, elongated and flattened; there are
no clumped setae at its base. The base of
the ventral lobes is U-shaped and broad be-
tween the two halves. The parameral sheath
of the aedeagus is smooth dorsally. The
ventral apodeme has a ventral keel. The dis-
tiphallus is swollen basally, and is perpen-
dicular to the ventral apodeme throughout.
In the female terminalia, the furca is not
notched posteriorly, and lacks a posterolat-
eral peg. The anterolateral furcal prongs are
separated by a distance narrower than the
greatest width of the furca. The common
spermathecal duct is broad basally, at half
the width of the entire furca; the duct tapers
quickly.

Autapomorphies.—Halter brown (char-
acter 15), ventral apodeme longer than dor-
sal apodemes (character 18), ventral surface
of ventral apodeme with longitudinal keel
(character 19), distiphallus perpendicular to
dorsal apodemes (character 23, state 0), an-
terior prongs of furca narrowly separated
(character 38).

Distribution.—The known distribution
for this species is found in Fig. 4. This spe-
cies is found in southern Mexico, in the
states of Puebla, Michoacan, and Guerrero.
The type locality, in Guerrero, is part of the
northern slope foothills of the Sierra Madre
de Sur range, at an elevation over 1,500 m.
The remaining localities are all above this
elevation, as high as 2,600 m. The distri-
bution encompasses parts of the following
biogeographical regions: the Provincia Me-
soamericana de Montana, and the Provincia
Pacifica.

Etymology.—Gr., para: beside, near +
“argentifera,’ referring to the similarity
with Ozodiceromyia argentifera (Krober);
to be treated as a noun in apposition.

Biology.—Adults are active at least
through the months of July and August.

VOLUME 102, NUMBER 3

585

Fig. 4.

KEY TO SPECIES

A comprehensive key to species of Ozo-
diceromyia is not presented herein. This
will be included in the larger revision of the
entire genus in progress by the authors. The
current key is a continuation of Gaimari and
Irwin’s (2000) key to world genera of Cy-
clotelini. An alternative is the key to the
therevid genera of North America found in
Irwin and Lyneborg (198la, b). The first
couplet below separates members of the

zodiceromyia signatipennis

Ozodiceromyia costalis

2 4 10 11 21 29 31 32 35

1 5 6 7 8 15 16 20 23 33 34 39

Ripe:

3 11 22 26 28 29 30 35 40 41

Known distribution for Ozodiceromyia parargentifera.

Ozodiceromyia mexicana-group from. all
other members of the genus. Note that the
male genitalic characters in the key are usu-
ally visible without dissection. The female
genitalic character in couplet 3 requires dis-
section, but should not be necessary for
routine identification.

il. Antenna (Figs. 7-10) longer than head.
Scape setose throughout. Pedicel barrel-
shaped, longer than wide, setose throughout.
First flagellomere setose over basal two-

9 14 17 24 25 27 36 37

zodiceromyia mexicana
15 18 19 23 38
zodiceromyia parargentifera

zodiceromyia argentifera

Ozodiceromyia livdahli

Single most parsimonious cladogram for the species of the Ozodiceromyia mexicana-group, showing

character state changes under ACCTRAN character optimization. Characters are numbered as in the text; hash
marks are as follows: black = forward change with no homoplasy; white = change with homoplasy.

586

O. signatipennis

O. costalis

O. mexicana

O. parargentifera

O. argentifera

O. livdahli

Single most parsimonious cladogram for

Fig. 6.
the species of the Ozodiceromyia mexicana-group,
with Bremer support indices indicated above each
branch and post-successively reweighted, rescaled Bre-
mer support indices below each branch.

thirds. Wing membrane and veins orange, at
least basally; distal third of wing often dark-
GneGenwithudankeneGuvielisS mei elie eect
SiS Saris |e eee Ozodiceromyia mexicana-group
= Antenna (Fig. 11) shorter than head, rarely
as long as head. Scape usually bare on me-
dian surface. Pedicel as long as wide, usu-
ally bare on median surface. First flagello-
mere setose at most in basal third. Wing
membranes hyaline or banded, but never or-
ange; veins dark, rarely orange; distal third
of wing not darkened relative to basal two-
thirds other Ozodiceromyia
2(1). Scape densely setose, with long setae, lon-
ger than those of first flagellomere (Figs. 9—
10). Lower frons globose, distinctly bulging,
and shiny black, with silver pruinescence
only along eye margin (Figs. 9-10). Face
beneath antenna shiny black, lacking prui-
nescence (Figs. 9-10). Male: notum with
erect black pile; posterior surface of hind-
coxa with long, white pile (Fig. 16); outer
gonocoxal process tapering evenly, bare
(Fig. 31); ventral gonocoxal process of gen-
italia absent, dense clump of orange setae
on corresponding edge (Fig. 30)
Ozodiceromyia mexicana
— Scape setose, but with short setae, with few
long setae interspersed (Figs. 7-8). Lower
frons not globose, only slightly bulging; sil-
ver pruinescence present dorsolaterally to
antennal base (Figs. 7-8). Face beneath an-
tenna with silver pruinescence (Figs. 7-8).
Male: pile of notum variable, but usually
both erect and appressed pile present; pos-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

terior surface of hindcoxa lacking pile (Fig.
14); outer gonocoxal process widened dis-
tally, with small patch of short setae on dor-
solateral surface (Fig. 27); ventral gonocox-
al process present, long and flattened, with
no densely clumped setae (Fig. 26)
. Halter brown in both sexes. Male: notum
with erect black pile; setae of gonocoxites
and epandrium black; distiphallus perpen-
dicular to ventral apodeme throughout
length (Fig. 37). Female: distance between
anterolateral furcal prongs distinctly nar-
rowed anteriorly (Fig. 40)
waite ay fe ae WA gee Ozodiceromyia parargentifera
- Halter yellow in both sexes. Male: notum
with both erect and appressed pale or gold
pile; setae of gonocoxites and epandrium
black or orange; distiphallus recurved be-
fore apex to become parallel with ventral
apodeme (Fig. 33). Female: distance be-
tween anterolateral furcal prongs not nar-
rowed anteriorly, subequal to widest part of
furca (Fig. 39)
4(3). Male: setae of gonocoxites and epandrium
black (Figs. 23, 26—27). Female: appressed
pile on abdominal tergites gold

Ozodiceromyia argentifera
— Male: setae of gonocoxites and epandrium
pale or orange (Fig. 28). Female: appressed
pile on abdominal tergites brown and white
SACRE es ah ace Recker Ozodiceromyia livdahli

CHARACTER DESCRIPTIONS

This section is devoted to description of
morphology and taxon distribution of states
of the 41 characters used in this analysis.
Of these, 11 are taken from the head, 5
from the thorax, 19 from the male termin-
alia, and 6 from the female terminalia.
Characters described and discussed by Gai-
mari and Irwin (2000) are referred to the
appropriate character number therein for
description, although state distributions are
discussed herein. All characters are binary.
Character distributions are discussed rela-
tive to the presented cladogram (Fig. 5).

HEAD
I. Antennal length

0 shorter than or subequal to head
length
1 longer than head
The states for this character are described
as character 1 in Gaimari and Irwin (2000).

VOLUME 102, NUMBER 3 587

Figs. 7-13. Heads, ¢ (7, 9, 11), 2 (8, 10); 2, antennal first flagellomere (12), tip only (13). (7-8) Ozodi-
ceromyia argentifera (MEI 038790, 052183). (9-10, 13) Ozodiceromyia mexicana (MEI 052192, 107043,
070426). (11-12) Ozodiceromyia signatipennis (MEI 044750, 056973). Measure bars, 0.2 mm.

In the current analysis, an elongated anten- 2. Medial surface of scape
na (Fig. 7) is autapomorphic for the mexi-

cana-group, and is found in no other known 0 without setae
Ozodiceromyia. Members of several other 1 setose

cycloteline genera, including Cyclotelus In nearly all Ozodiceromyia, the scape is
Walker, also share this state. bare on the median surface (Fig. 11). Only

588

in the mexicana-group and the outgroup
taxon Ozodiceromyia costalis (and several
undescribed, putatively related species) is
this surface setose (Fig. 9).

3. Setae of scape

0 longer than those of first flagellomere
1 most are shorter than or equal to
those of first flagellomere

The setae of the scape are longer than those
of the first flagellomere (Fig. 9), plesiom-
orphically, as in both outgroup taxa and
Ozodiceromyia mexicana. As a Synapomor-
phy for the clade of Ozodiceromyia parar-
gentifera + Ozodiceromyia argentifera +
Ozodiceromyia livdahli, the setae of the
scape are shorter than or equal to those of
the first flagellomere, at most with several
longer setae interspersed among the short
setae (Fig. 7).

4. Medial surface of pedicel

Q without setae

lL Ssetose
The states for this character are described
as character 3 in Gaimari and Irwin (2000).
The apomorphically setose medial surface
of the pedicel (Fig. 9) is present in all mem-
bers of the mexicana-group, and in the out-
group taxon Ozodiceromyia costalis.

5. Setae of first flagellomere

QO basal only, or bare

1 covering more than basal half
In nearly all Ozodiceromyia, including both
outgroup taxa, the setae of the first flagel-
lomere are restricted to the base (Fig. 11).
Only as an autapomorphy for the mexicana-
group do these setae cover more than the
basal half of the first flagellomere (Fig. 9).

6. Setae of first flagellomere

OQ less dense ventrally, or bare

1 fully surrounding with equal density
Plesiomorphically, the ventral surace of the
first flagellomere is bare, or only sparsely
setose (Fig. 11), as in nearly all Ozodicer-
omyia including both outgroup taxa. Only
as an autapomorphy for the mexicana-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

group do setae fully surround the first fla-
gellomere with equal density (Fig. 9).

7. Antennal style

QO originates near tip, but extended
beyond first flagellomere distally
1 distinctly subapical, and not extended
beyond tip of first flagellomere
Plesiomorphically, the antennal style origi-
nates very close to the apex of the first fla-
gellomere, but slightly subapical and ven-
tral. Despite this, the tip of the style extends
distally beyond the tip of the first flagellom-
ere (Fig. 12). As an autapomorphy for the
mexicana-group, the antennal style origi-
nates distinctly subapically on the first fla-
gellomere within a ventral pit, with the sty-
lar tip not reaching the apex of the first fla-
gellomere (Fig. 13).

8. First stylar segment

0 as long as wide

1 wider than long, and ring-like
Plesiomorphically, the first stylar segment
is as long as wide (Fig. 12). As an auta-
pomorphy for the mexicana-group, this seg-
ment is reduced to a small, flattened ring
set within a ventral pit (Fig. 13).

9. Lower frons at antennal insertion

O normal, not globose

1 globose, bulging
At the antennal insertion, the lower frons
bulges slightly in most Ozodiceromyia (Fig.
7). As an autapomorphy for Ozodiceromyia
mexicana, the lower frons is distinctly bulg-
ing and globose (Figs. 9-10).

10. Setae of 3 frons

O scattered

1 in patches or absent
In many Ozodiceromyia, including the out-
group taxon Ozodiceromyia signatipennis,
the setae on the male frons are scattered
(Fig. 11). In the mexicana-group and Ozo-
diceromyia costalis, these setae are absent
or reduced to small patches located dorso-
laterally from the antennal base (Fig. 7).

VOLUME 102, NUMBER 3

589

Figs. 14-16.
of wing, @, left is basal (15); hindcoxa, d, left is anterior (16). (14-15) Ozodiceromyia argentifera (MEI
037789, 052183); arrow (14) points to katepisternum. (16) Ozodiceromyia mexicana (052192). Measure
bars, 0.2 mm.

11. Face directly below antenna

QO with full pruinescence
1 shiny, with little or no pruinescence
(usually black)
For most Ozodiceromyia, the face below
the antenna is fully covered with silver

Thorax and associated structures: lateral thorax, d, left is anterior (14); basal costal lobe

pruinescence (Fig. 8). In Ozodiceromyia
costalis (and several undescribed, putative-
ly related species) and Ozodiceromyia mex-
icana, the face completely lacks pruinesc-
ence, or any other vestiture, exposing the
black, shiny cuticle (Fig. 9).

590 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 17-25. 4d genitalic and pregenitalic structures: 8th sternite (17) and tergite (18); gonostylus (19-20);
ventral lobe (21—22); epandrium, split view (left, dorsal; right, ventral) (23, 25), dorsal view (24). (17-19, 23)
Ozodiceromyia argentifera (MEI 037789); arrow (23) points to subepandrial plate. (20, 22) Ozodiceromyia
signatipennis (MEI 044750). (21, 25) Ozodiceromyia mexicana (MEI 052240); arrow (21) points to pilose,
secondary distal lobe. (24) Ozodiceromyia livdahli (MEI 038822). Measure bars, 0.2 mm.

VOLUME 102, NUMBER 3

THORAX
12. S notal vestiture

OQ without decumbent pile

1 with thick decumbent pile
Although decumbent pile on the male no-
tum is present in other members of this ge-
nus, and other therevids, its absence is con-
sidered plesiomorphic at this level of anal-
ysis. The presence of thick, decumbent pile
is a synapomorphy for Ozodiceromyia ar-
gentifera + Ozodiceromyia livdahli.

13. Fine, erect setae of 3 notum

O some black

1 entirely pale or gold, none black
All taxa studied have fine, erect setae on
the male notum. Plesiomorphically, most of
these setae are black, although some pale
setae may be interspersed. As a synapo-
morphy for Ozodiceromyia argentifera +
Ozodiceromyia livdahli, all of these fine,
erect, notal setae are pale or gold. Note, this
does not include the notal macrosetae.

14. Posterolateral surface of hindcoxa

0 lacking pile

1 pilose
Plesiomorphically, the posterolateral sur-
face of the hindcoxa is bare (Fig. 14). As
an autapomorphy for Ozodiceromyia mexi-
cana, this surface has long, thin pile (Fig.
16). This state is also present in certain oth-
er groups within Ozodiceromyia.

15. Halter color

QO brown

1 yellow
In most species of Ozodiceromyia, the hal-
ter is brown. In several groups, including
the mexicana-group, yellow halter is syna-
pomorphic. The only exception is the
brown halter found in Ozodiceromyia par-
argentifera, which is considered an auta-
pomorphy for the species.

16. Wing color

O clear, or with darkened patches
1 cloudy yellow or orange

591

Although banded wings are known in Ozo-
diceromyia, clear wings or those with dark-
ened patches along certain wing veins are
plesiomorphic at this level of analysis. Only
in the mexicana-group is the wing entirely
cloudy yellow or orange, becoming dark-
ened in the distal third. This is considered
an autapomorphy for the mexicana-group.

MALE TERMINALIA
17. Parameral sheath texture

OQ smooth dorsally

1 with transverse wrinkles dorsally
Plesiomorphically, the parameral sheath of
the aedeagus is smooth dorsally (Fig. 32).
As an autapomorphy for Ozodiceromyia
mexicana, the parameral sheath has distinct
transverse wrinkles dorsally (Fig. 34).

18. Relative lengths of dorsal and ventral
apodemes

QO subequal, or dorsal apodeme longer
1 ventral apodeme longer
The states for this character are described
as character 37 in Gaimari and Irwin
(2000). The longer ventral apodeme (Fig.
37) is autapomorphic for Ozodiceromyia
parargentifera.

19. Ventral surface of ventral apodeme

O without longitudinal keel

1 with longitudinal keel
Most species of Ozodiceromyia lack a ven-
tral longitudinal keel on the ventral apode-
me. A longitudinal keel is present (Fig. 37)
as an autapomorphy for Ozodiceromyia
parargentifera.

20. Distiphallus length

0 shorter than dorsal apodemes
1 as long or longer than dorsal apo-
demes

At this level of analysis, having a disti-
phallus shorter than the dorsal apodemes is
considered plesiomorphic (Fig. 38). As an
autapomorphy for the mexicana-group, the
distiphallus is distinctly longer than the dor-
sal apodemes (Fig. 33).

592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 102, NUMBER 3

21. Dorsal part of basal distiphallus

QO with median groove

1 without median groove, rounded
A median, longitudinal groove is present on
the dorsal part of the basal distiphallus in
the outgroup taxon Ozodiceromyia signati-
pennis. This groove is lacking, and the dis-
tiphallus is entirely rounded above in the
mexicana-group and Ozodiceromyia costal-
is.

22. Basal portion of distiphallus

Q tapering evenly

1 swollen
Plesiomorphically, the basal portion of the
distiphallus is not swollen and tapers grad-
ually and evenly to the apex (Fig. 35). As
a synapomorphy for Ozodiceromyia parar-
gentifera + Ozodiceromyia argentifera +
Ozodiceromyia livdahli, this part of the dis-
tiphallus is distinctly swollen (Figs. 33, 37).

23. Orientation of distiphallus at apex

Q perpendicular to dorsal apodemes

1 parallel with dorsal apodemes
In both outgroup taxa, the distiphallus is
perpendicular to the dorsal apodemes
throughout, not recurving anteriorly or
posteriorly. As a synapomorphy for mem-
bers of the mexicana-group, the distiphal-
lus recurves anteriorly before its apex, be-
coming parallel with the dorsal apodemes
(Figs. 33, 35). As an autapomorphy for
Ozodiceromyia parargentifera, the disti-
phallus loses its curve and becomes per-
pendicular with the dorsal apodemes (Fig.
37); as in state 0:

24. Subepandrial plate attachment to
epandrium

Q laterally and posteriorly
1 laterally only

ee

Figs. 26-31.

595

Plesiomorphically, the subepandrial plate is
solidly attached to the epandrium both lat-
erally and posterolaterally (Fig. 23). As an
autapomorphy for Ozodiceromyia mexi-
cana, the subepandrial plate is greatly re-
duced, and is only attached to the epan-
drium through a lateral membranous con-
nection (Fig. 25).

25. Posterolateral corners of epandrium

O held by subepandrial plate

1 held by sclerotized membrane
The posterolateral corners of the epandrium
are held together by the subepandrial plate
(Fig. 23), plesiomorphically. As an auta-
pomorphy for Ozodiceromyia mexicana,
the corners are held by a separate, sclero-
tized membrane (Fig. 25), apparently not
associated with the subepandrial plate.

26. Gonocoxal setae color

O mostly pale

1 all black
Although this is variable throughout Ozo-
diceromyia, pale gonocoxal setae are ple-
siomorphic at this level of analysis. Under
the accelerated transformation character op-
timization model, presence of all black se-
tae (Fig. 27) is synapomorphic for the clade
of Ozodiceromyia parargentifera + Ozo-
diceromyia argentifera + Ozodiceromyia
livdahli, with a subsequent autapomorphic
change to pale setae in Ozodiceromyia li-
vdahli (Fig. 28)

27. Distinct internal keel where
gonocoxites fused

QO absent

1 present
At this level of analysis, the lack of a dis-
tinct internal keel (visible externally as a
groove) where the gonocoxites are fused

3d gonocoxites and associated structures, split view (left, ventral; right, dorsal) (26, 30), lateral

view, gonostylus removed, left is dorsal (27), lateral view, left is dorsal (28-29, 31). (26-27) Ozodiceromyia
argentifera (MEI 037789); arrow (26) points to ventral gonocoxal process, arrow (27) points to inner gonocoxal
process. (28) Ozodiceromyia livdahli (MEI 038822); arrow points to outer gonocoxal process. (29) Ozodicero-
myia signatipennis (MEI 044750). (30-31) Ozodiceromyia mexicana (MEI 052240). Measure bars, 0.2 mm.

594 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 32-38. Aedeagal complex: dorsal view (32, 34, 36); lateral view (33, 35, 37-38). (32-33) Ozodicer-
omyia argentifera (MEI 037789); arrow (32) points to lateral ejaculatory process, arrow (33) points to disti-
phallus. (34-35) Ozodiceromyia mexicana (MEI 052240); arrow (34) points to ejaculatory apodeme, arrow (35)
points to dorsal apodeme. (36-37) Ozodiceromyia parargentifera (MEI 027017); arrow (37) points to ventral
apodeme. (38) Ozodiceromyia signatipennis (MEI 044750). Measure bar, 0.2 mm.

VOLUME 102, NUMBER 3

(Fig. 26) is plesiomorphic. As an autapo-
morphy for Ozodiceromyia mexicana, this
distinct line of fusion is present (Fig. 30).

28. Ventral gonocoxal process

Q absent or only small fold of edge

1 elongated and flanged
Plesiomorphically, the ventral gonocoxal
process is absent, or is only a small fold of
the edge (Fig. 30). As a synapomorphy for
the clade of Ozodiceromyia parargentifera
+ Ozodiceromyia argentifera + Ozodicer-
omyia livdahli, the ventral gonocoxal pro-
cess is present, elongated, and flanged (Fig.
26).

29. Setae of ventral gonocoxal process
(including corresponding edge if absent)

QO absent
1 distinctly different
gonocoxal setae

Plesiomorphically, setae are absent on the
ventral gonocoxal process (Fig. 26), or on
the corresponding edge if absent. In Ozo-
diceromyia mexicana and Ozodiceromyia
costalis, the setae are distinctly different
from the remaining gonocoxal setae (Fig.
30).

from other

30. Setae of outer gonocoxal process

Q absent

1 present
The outer gonocoxal process is bare of se-
tae (Fig. 31) plesiomorphically. As a syna-
pomorphy for the clade of Ozodiceromyia
parargentifera + Ozodiceromyia argenti-
fera + Ozodiceromyia livdahli, there is a
small patch of short setae on the dorsolat-
eral portion of the outer gonocoxal process
(Fig. 27).

31. Outer and ventral gonocoxal
processes

O fused

1 unfused
The outer and ventral gonocoxal processes
are plesiomorphically fused (Fig. 29), as in
the outgroup taxon Ozodiceromyia signati-
pennis. These processes are separate and

595

unfused (Figs. 27, 35) in the mexicana-
group and in Ozodiceromyia costalis.

32. Projection on gonocoxal apodeme for
articulation with aedeagus

QO absent

1 present
There is no sclerotized connection between
the gonocoxal apodeme and the aedeagus
in any taxon in this study. Ozodiceromyia

signatipennis completely lacks any projec-

tions on the gonocoxal apodeme. However,
in the mexicana-group and in Ozodicero-
myia costalis, there is a medially directed
projection of the gonocoxal apodeme (Fig.
30) that provides a site for articulation with
the aedeagus.

33. Distal part of ventral lobe

OQ bare, or with fine, short pile on outer
edge

1 with pilose distal lobe
Most Ozodiceromyia, including both out-
group taxa, lack a secondary distal lobe on
the ventral lobe (Fig. 22). Only as an au-
tapomorphy for the mexicana-group is a
finely pilose, secondary lobe present distal-
ly on the ventral lobe (Fig. 21).

34. Lobe of gonostylus

Q expanded ventrally into small lobe

1 lobe greatly expanded ventrally
In the more primitive Ozodiceromyia (see
Gaimari 1998), the gonostylus is strap-like,
with no lobe ventrally. More advanced
members of the genus have a small lobe on
the ventral part of the gonostylus (Fig. 20),
which is the plesiomorphic state at this lev-
el of analysis. As an autapomorphy for the
mexicana-group, this lobe is greatly en-
larged (Fig. 19).

35. Subapical spur of gonostylus

OQ present

1 absent
The states of this character are described
as character 56 in Gaimari and Irwin
(2000). The subapical spur of the gono-
stylus is absent in both Ozodiceromyia

596

mexicana and the outgroup taxon Ozodi-

ceromyia costalis. Although this character

is quite variable in the genus, the change
to state 0 appears to be a synapomorphy
for the clade of Ozodiceromyia parargen-
tifera + Ozodiceromyia argentifera +

Ozodiceromyia livdahli.
FEMALE TERMINALIA

36. Posterior edge of furca

Q rounded

1 with notch or bend
Plesiomorphically, the posterior edge of the
furca is rounded normally, with no notches
or bends (Fig. 39). As an autapomorphy for
Ozodiceromyia mexicana, a distinct, ante-
riorly directed notch is present (Fig. 41).

37. Medioventral edge of posterolateral
furca

QO smooth

1 with small process
Plesiomorphically, there are no medially di-
rected pegs on the inner furcal margin (Fig.
39). As an autapomorphy for Ozodicero-
myia mexicana, there is a small process, or
peg, on the inner ventral edge of the furca
on the posterolateral margin (Fig. 41).

38. Space between anterior prongs of
furca

0 subequal to greatest furcal width
1 distinctly narrower than greatest
furcal width

The space between the anterior prongs of
the furca is approximately the same width
as the greatest furcal width (Fig. 39), ple-
siomorphically. As an autapomorphy for
Ozodiceromyia parargentifera, the width
between the anterior prongs of the furca is
distinctly narrowed (Fig. 40).

39. Posterior part of furcal bulla

QO normal, not indented
1 indented as cavity to house common
spermathecal duct
Plesiomorphically, the posterior part of the
furcal bulla us rounded normally. As an au-
tapomorphy for the mexicana-group, this

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

posterior margin is distinctly indented as a
cavity to house the common spermathecal
duct (Fig. 39).

40. Basal part of common spermathecal
duct

QO normal width, more or less even
throughout length

1 distinctly widened, tapering quickly
The basal part of the common spermathecal
duct is plesiomorphically of normal width,
more or less even throughout its length,
only gradually tapering (Fig. 41). As a syn-
apomorphy for the clade of Ozodiceromyia
parargentifera + Ozodiceromyia argenti-
fera + Ozodiceromyia livdahli, the base is
distinctly enlarged and widened, tapering
quickly to normal width (Fig. 39).

41. Individual spermathecal duct origin

Q from base of central sac

1 from common spermathecal duct
Plesiomorphically, the spermathecal ducts
originates from the base of the central sac
(Fig. 41). As a synapomorphy for the clade
of Ozodiceromyia parargentifera + Ozo-
diceromyia argentifera + Ozodiceromyia
livdahli, the spermathecal ducts originate
on the common spermathecal duct, with a
short duct then leading to the central sac
(Fig. 39).

RESULTS AND DISCUSSION

Cladistic analysis.—The analysis pro-
duced a single most parsimonious tree (Fig.
5) of length 47, Consistency Index (Kluge
and Farris 1969) of 0.87 (0.80 excluding
autapomorphies), Retention Index (Farris
1989) of 0.81, and Rescaled Consistency
Index (Farris 1989) of 0.71. The topology
remained stable through successive weight-
ing (Farris 1969; Carpenter 1988, 1994),
implemented by PAUP by reweighting all
characters on a base weight of 1,000 by the
maximum values of their Rescaled Consis-
tency Indices, leveling out at a single iter-
ation. After this single iteration, the tree sta-
tistics above were all 0.97 or higher. Be-
cause the expected amount of homoplasy

VOLUME 102, NUMBER 3

S27)

Figs. 39-41. @ reproductive structures: furca (dorsal view) through central sac and basal part of sper-
mathecal ducts (both spermathecal ducts truncated basally) (39); furca (dorsal view) with truncated common
spermathecal duct (40); furca (dorsal view) through central sac and spermathecae (41). (39) Ozodiceromyia
argentifera (MEI 052230); arrow points to central sac. (40) Ozodiceromyia parargentifera (MEI 027022);
arrow points to furcal bulla. (41) Ozodiceromyia mexicana (MEI 070426); arrow points to spermatheca.

Measure bars, 0.2 mm.

increases with the number of taxa in a par-
simony analysis, the regression equation of
Sanderson and Donoghue (1989) was used
to calculate the expected value for the Con-

sistency Index. The calculated value of 0.78
for 6 taxa is lower than the value indicated
in the current analysis, excluding autapo-
morphies, so the amount of homoplasy is

598

less than expected for an analysis with this
number of taxa under the 1989 model. The
relatively high Retention Index indicates
that a high proportion of the potential syn-
apomorphies in the data matrix are present
as homologies on the cladogram.

Bremer support (aka, decay analysis)
may reflect the robustness of certain ele-
ments of the resulting cladograms (Bremer
1988, Donoghue et al. 1992, Kallersj6 et al.
1992). A decay analysis entails calculating
the difference in tree length between the
most parsimonious tree and the shortest tree
which lacks each nodal group. This reflects
how much additional evidence, in the form
of characters supporting an alternative
grouping, would be necessary to overthrow
a clade in the most parsimonious hypothe-
sis. Using AUTODECAY (Eriksson 1997,
currently in version 2.9.9) in combination
with PAUP, Bremer support indices were
calculated for each nodal group in the clad-
ogram. Following Gaimari and Irwin
(2000), Bremer support indices were also
calculated using the successively reweight-
ed data, and were rescaled per the proce-
dure outlined by Bremer (1994). The re-
sulting Bremer support indices for each
node are displayed on the cladogram in Fig.
6. The Bremer support values are an indi-
cation of the number of characters in op-
position to the current topology that would
be necessary to break down the individual
nodes of the tree. Justification for excluding
perturbation (e.g., bootstrap, jackknife) and
permutation (PTP, T-PTP) based methods to
assess branch support is provided by Gai-
mari and Irwin (2000).

Classification.—The internal classifica-
tion of the Therevidae has only recently be-
gun to receive attention. Not until recently
(Irwin and Lyneborg 1981a) was the family
formally divided into two subfamilies,
Therevinae and Phycinae. More recently,
Gaimari and Irwin (2000) defined and pro-
posed a phylogenetic classification for gen-
era of Cyclotelini, including Ozodicero-
myia, and Gaimari (1998) defined putative

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 3. Classification for the species of the Ozo-
diceromyia mexicana-group.

Cyclotelini Gaimari and Irwin, 2000
Ozodiceromyia Bigot, 1890
Ozodiceromyia **mexicana-group’’
O. mexicana Bigot, 1890
O. parargentifera, n. sp.
O. argentifera (Krober 1929)
O. livdahli, n. sp.

species-groups within the genus, including
the mexicana-group as defined herein.

According to the cladogram, the Bremer
support index of 7 additional steps for the
clade, and the rescaled Bremer support in-
dex of 13 additional steps, the Ozodicero-
myia mexicana-group appears well support-
ed by autapomorphies. This hypothesis will
be most thoroughly tested in the phyloge-
netic analysis of the entire genus underway.
The classification in Table 3 is presented as
a phyletic sequence of taxa (Nelson 1969,
1972, 1973) within this small group of Ozo-
diceromyia. The group will be classified
within the larger context of the entire genus
with the pending revisionary work under-
way.

ACKNOWLEDGMENTS

Heartfelt thanks are extended to Jill D.
Mullett for her superb illustrations; Gail E.
Kampmeier (INHS) (and our cadre of lab
assistants, including Mona Abou-Batnin,
Jessica Adams, Jamie Bender, Jessica Bick-
ham, Amanda Buck, Shelly Cook, Andrea
Gavurnik, Mary Grabowski, Dyanna Greg-
ory, Traci Gustafson, Sarah Kastelic, Sandy
Kawanaka, Erin Leslie, Leslie Marsh, Rob-
ert Pollok, Ann Ramsey, Stephanie Stroth-
off, Johnson Zeledon, and Anjun Zhou) for
making this research much easier through
databasing; Martin Hauser (INHS), Kevin
C. Holston (INHS), the late Ellis G.
MacLeod (University of Illinois), Mark A.
Metz (INHS), Evert I. Schlinger (Schlinger
Foundation), Donald W. Webb (INHS), Bri-
an M. Wiegmann (North Carolina State
University), Shaun L. Winterton (UQIC),

VOLUME 102, NUMBER 3

Long-Long Yang (North Carolina State
University), and David K. Yeates (Univer-
sity of Queensland), for many fruitful dis-
cussions about systematic theory, therevid
and asiloid evolution and characters, and
for numerous shared experiences in the
field. We also thank Neal L. Evenhuis
(BPBM) and E Christian Thompson (Sys-
tematic Entomology Lab, USDA) for pro-
viding helpful nomenclatural and _ biblio-
graphic insight, Adrian Pont (Oxford Uni-
versity) for deciphering the handwritten la-
bel of J. Bigot, and Mark A. Metz for
translating German primary literature. In
addition, we wish to thank the following
curators for loans of type materials and for
their help during the travels of SDG study-
ing type materials: John E. Chainey
(BMNH); Loic Matile (MNHN), Ruth Con-
treras-Lichtenberg (NHMW), and Vadim F
Zaitsev and Vitali N. Tanasijtshuk (ZMAS);
and for loans of specimens (including
types) we thank the following curators, in
order by collection: David A. Grimaldi
(AMNH), Richard W. Baumann (BYUC),
Paul H. Arnaud and Norman D. Penny
(CASC), Eric M. Fisher (CDFA), Chen W.
Young (CMNH), Jeffrey M. Cummings
(CNCI), E. Richard Hoebeke (CUIC), Don-
ald S. Chandler and John FEF Burger
(DENH), Enrique Ramirez (EBCC), Cheryl
B. Barr, John A. Chemsak, and Jerry Pow-
ell (EMEC), Wilford J. Hanson (EMUS),
Howard V. Weems, Jr. and Gary J. Steck
(FSCA), Vicente Hernandez-Ortiz (IEXA),
Donald W. Webb (INHS), H. Derrick
Blocker (KSUC), Brian V. Brown (LACM),
Philip D. Perkins (MCZC), Michael Ivie
(MTEC), Don C. Arnold (OSEC), Norman
E Johnson (OSUC), Raymond J. Pupedis
(PMNH), David Faulkner (SDMC), Robert
W. Brooks (SEMC), Wade C. Sherbrooke
(SWRS), Edward G. Riley (TAMU), Carl
A. Olson (UAIC), Steven L. Heydon and
Lynn S. Kimsey (UCDC), Saul I. Frommer
(UCRC), Cecil L. Smith (UGCA), Atilano
Contreras-Ramos and Harry Brailovsky
(UNAM), FE Christian Thompson (USNM),
Frank W. Merickel (WFBM), Richard S.

599

Zack (WSUC), H. Wendt (ZMHB), and
Leif Lyneborg and Verner Michelsen
(ZMUC). The distributional maps were
generated using VERSAMAP, version 2.04,
by Charles H. Culberson. We also extend
our thanks to FE Christian Thompson, for his
critical review of the manuscript, and help-
ful suggestions. Special thanks are due to
our wives, Helen G. Gaimari and Bonnie J.
Irwin, for their unending patience and un-
derstanding. This study was supported by
NSF DEB-PEET grant # 95-21925 to MEI,
in addition to funding provided from the
Schlinger Foundation and a graduate re-
search assistantship to SDG through the I-
linois Natural History Survey.

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PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 601-608

REVIEW OF THE ASIAN SAWFLY GENUS ANISOARTHRA CAMERON
(HYMENOPTERA: TENTHREDINIDAE)

V. VASU, DAviID R. SMITH, AND MALKIAT S. SAINI

(VV, MSS) Department of Zoology, Punjabi University, Patiala-147 002, India; (DRS)
Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department
of Agriculture, % National Museum of Natural History, Washington, D.C. 20560-0168,
U.S.A. (e-mail: dsmith @sel.barc.usda.gov)

Abstract.—Anisoarthra Cameron includes three species, A. coerulea Cameron from
India and Sri Lanka, A. diascoreae (Rohwer), n. comb., from India, and A. birmanica
(Malaise), n. comb., from Burma and India (new record). The species are described and
illustrated, and a key is provided. Anisoarthra diascoreae feeds on yam, Dioscorea sp.

(Dioscoreaceae).

Key Words:
oclia

Species of Anisoarthra are relatively
large, mostly metallic bluish-black or vio-
laceous sawflies of the subfamily Blenno-
campinae. A characteristic, shared with sev-
eral other genera of the subfamily, is the
comblike, four- or five-toothed tarsal claws.
The three species of the genus occur only
in southern Asia—Burma, India, and Sri
Lanka. Yam, Dioscorea sp., is a host plant
for Anisoarthra diascoreae (Rohwer). A
species in a related genus in the Blenno-
campinae, Senoclidia purpurata (Smith), is
known to be a pest of yam in Papua New
Guinea (Szent-Ivany 1974). Since yam is an
important food plant in the tropics and is
distributed by commerce, it is important to
know and be able to recognize its potential
pests.

Here we redescribe the species of Ani-
soarthra and give a key for their separation.

The National Museum of Natural Histo-
ry, Smithsonian Institution, Washington,
D.C., is designated as USNM; Zoological
Department, Punjabi University, as ZDPU;
and Canadian National Collection, Ottawa,
as CNC.

India, Sri Lanka, Burma, Dioscorea, Dioscoreaceae, Blennocampinae, Sen-

Anisoarthra Cameron

Anisoarthra Cameron 1876: 461. Type spe-
cies: Anisoarthra coerulea Cameron.
Designated by Rohwer 1911.

Senoclia Cameron 1877: 88 (unnecessary
new name for Anisoarthra Cameron).—
Rohwer 1921: 105 (new species; key to
species).—Malaise 1937: 50 (separation
from Neoclia Malaise; new species from
Burma; key to species).—Benson 1938:
367 (in Senocliini).—Malaise 1964: 20
(Anisoarthra not preoccupied and is cor-
rect name for the genus).

Description.—Antenna covered with ap-
pressed hairs; scape about 2 longer than
broad, pedicel as long as broad, 3rd and 4th
segments subequal in length, each a little
dilated at apices; 5th segment dilated at
apex and little longer than 4th; 6th segment
about % length of 5th, dilated at apex; 7th
segment shorter than 6th; apical two seg-
ments each slightly shorter than 7th (Fig.
2). Clypeus smooth, truncate or with slight-
ly rounded anterior margin; malar space lin-

602

ear to less than half diameter of front ocel-
lus; genal carina absent. Epicnemium ab-
sent. Mesoscutellum raised, conspicuous,
smooth and shining. Forewing (Fig. 1) with
vein M and Rs+M meeting before attaining
Sc+R; vein 2A+3A furcate or curved up
at its apex, furcation sometimes faint; 4 cu-
bital cells. Hindwing (Fig. 1) with one mid-
dle cell; petiole of anal cell as long as cell.
Tarsal claws comblike, with 4—5 teeth, if
with 4 teeth, then acute basal lobe present
(Figs. 3-5). Male penis valve with long api-
cal filament (Fig. 12).

Remarks.—The comblike tarsal claws
are characteristic of several other genera of
Blennocampinae, such as Neoclia Malaise
and Brykella Malaise, but Anisoarthra may
be separated by the venation of the fore-
wing (vein M and RS+M meeting before
attaining Sc+R; vein 2A+3A curved up or
furcate at its apex) and hindwing (petiole of
anal cell about half as long as cell), absence
of a genal carina, and absence of an epic-
nemium. The long apical filament of the
male penis valve does not occur in other
known Asian Blennocampinae and appears
unique to Anisoarthra.

Much of the literature pertaining to this
genus has been under the name Senoclia
Cameron, an unnecessary replacement
name for Anisoarthra. Cameron (1876)
originally included two species, A. coerulea
and A cyanella from Ceylon. Though the
type of Anisoarthra cyanella is probably
lost, strong indications are that it belongs to
Senoclidea Rohwer and that it is not from
Ceylon as given in the original description
(see discussion by Smith 1982). In fact, hid-
den in a footnote, Cameron (1877) stated
“Tam told by Mr. FE Smith that the locality
for Senoclia cyanella (l.c. p. 462) is New
Gunea, and not Ceylon.’’ Consequently, we
do not include A. cyanella. Rohwer (1921)
added two species from India, one of which
was synonymized with A. coerulea by
Smith (1982), and gave a key to three spe-
cies. Malaise (1937) added a species from
Burma and gave a key to species. Malaise

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

did not examine types and based separation
of Rohwer’s species on the literature.

Benson (1938) placed Anisoarthra (men-
tioned as Senoclia) in the tribe Senocliini
of the Blennocampinae. The tribe included
a small group of Austro-Oriental genera
(Kampongia Malaise, Neoclia Malaise, Ne-
sotomostethus Rohwer, and Anisoarthra)
separated from other Blennocampinae by
the point of fusion between veins M and Ir-
m of the hindwing which reaches to or very
close to Rs, vein M of the forewing arising
from vein Rs+M. Nesotomostethus is sep-
arated from Anisoarthra by the three-
toothed tarsal claws with a basal lobe and
presence of an epicnemium. Kompongia is
separated by the two-toothed tarsal claws
with a basal lobe and cell M absent in the
hindwing.

KEY TO SPECIES OF ANISOARTHRA

1. Forewing darkly, uniformly infuscated; legs
black; tarsal claws with 4 teeth and a basal lobe
(Fig. 3) A. coerulea Cameron

— Forewing not completely infuscated, approxi-

mately basal half hyaline; bases of tibiae or
most of tibiae and extreme apices of femora
with a whitish or brownish spot; tarsal claws
with 4 or 5 teeth (Figs. 4—5)
. Tarsal claws with 4 distinct teeth and a distinct
triangular basal lobe (Fig. 4); supraclypeal area
flat; labrum black; lateral furrows on head par-
allel; postocellar area broader than long, as 4:
3; about % or less of bases of tibiae whitish to
brownish; foretibial spur furcate at apex; male
hypopygium truncate with slight notch at cen-
tenes. SEO aca ae A. diascoreae (Rohwer)

— Tarsal claws with 5 distinct teeth, basal lobe
indistinct, rounded (Fig. 5); supraclypeal area
triangularly roundly raised; labrum brownish to
white; lateral furrows diverging posteriorly;
postocellar area broader than long, as 3:2; basal
% or foretibia, midtibia except extreme apex,
and basal % hind tibia white; foretibial spur
simple; male hypopygium convex, rounded . .
bags nee ees ee A. birmanica (Malaise)

i)

Anisoarthra coerulea Cameron
(Figs. 3, 8)

Anisoarthra coerulea Cameron 1876: 462
(2, 6).—Smith 1982: 188, figs. 1, 9 13
(2, 6, Sri Lanka; syn.: bilanga Rohwer).

Senoclia coerulea: Kirby 1882: 181, pl. 8,

VOLUME 102, NUMBER 3

603

Figs. 1-5.
3, A. coerulea. 4, A. diascoraea. 5, A. birminaca.

fig. 21; Dalla Torre 1894: 186; Rohwer
1921: 105 (in key).—Malaise 1937: 52
(in key; as caerulea).
Senoclia bilanga Rohwer 1921: 106 (2, 3).
Senoclia caerulea var. bilanga: Malaise
19372 D2.

Female.—Length, 9.0—-11.0 mm. Bluish
black with metallic hue. Forewing darkly,
uniformly infuscated; hindwing lightly in-
fuscated; veins and stigma dark brown.

Antennal length shorter than abdomen,
2X head width. Clypeus with anterior mar-
gin truncate to slightly rounded; labrum
broader than long, as 2:1, with deflexed and
roundly pointed anterior margin; supracly-
peal area triangularly raised; inner margins
of eyes converging below with lower inter-
ocular distance to interocular distance at
level of front ocellus to eye length as 4:5:

1, Anisoarthra diascoraea, forewing and hindwing. 2, A. diascoraea, antenna. 3—5, Tarsal claws.

4; malar space linear; head in lateral view
with frontal area almost on same level as
eyes and supraantennal tubercles moderate,
sloping backwards and confluent with low
frontal ridges; median fovea prominent,
ditchlike on anterior half and shallow pos-
teriorly, reaching median ocellus; postocel-
lar area convex, broader than long, as 3:2;
temple lateral to lateral ocellus flat; post-,
inter-, and circumocellar furrows distinct;
lateral furrows distinct, deep, parallel and
abruptly ending well before posterior mar-
gin of head; head from above narrowing be-
hind eyes; postocellar, ocello-occipital, ocu-
lo-ocellar, and oculo-occipital distances
subequal. Mesoscutellum convex, append-
age not carinated or grooved; distances be-
tween cenchri to distance between tegulae
as 1:5. Tarsal claws comblike with 4 dis-

604

tinct teeth and triangular basal lobe (Fig. 3);
foretibial spur furcate at apex; hind basitar-
sus longer than following 3 segments com-
bined, as 7:5; hind tibial spurs subequal in
length, length of inner tibial spur to apical
width of hind tibia to outer apical tibial spur
length as 3:2:2. Lancet similar to Fig. 6,
with 22 serrulae. Sheath short, in lateral
view with dorsal margin bent down apically
(Fig. 8).

Head with sparse, minute, irregular punc-
tures, surface shining; mesonotum and me-
sepisternum with minute, scattered punc-
tures, surface shining with general oily lus-
ter; mesoscutellum and appendage impunc-
tate, surface polished; abdomen impunctate,
surface subshining. Body covered with
mixed metallic blue and silvery pubes-
cence.

Male.—Length, 9.0—10.0 mm. Similar to
female. Hypopygium truncate with slight
notch at center. Genitalia similar to Figs.
12.

Types.—The lectotype 2 of A. coerulea,
designated by Smith 1982, is in The Natural
History Museum, London, labeled *““Type,
H.T.”; “B.M. Type Hym. 1.363”; ““B.M.
Type, Hym. Anisoarthra coerulea (Camer-
ometsro) > Ceylon’: ~Kby-p: Sai, 215”

The holotype of Senoclia bilanga, a &,
is in the USNM, labeled ‘“‘Kollegal, 2,000
feet (about 606 meters), Coimbatoire, S. In-
dia, 1-[X-17, Ramakrishna.”’

Specimens examined.—INDIA: Tamil
Nadu, Kollegal (Coimbatoire), 600 m,
25.9.1917; Kollegal, 2,000 ft., 1-[X-17, Ra-
makrishna, coll. (d allotype of S. bilanga,
USNM); Nilgiri Hills, Kallar, 1,250 ft.,
South India, Oct. 1955, PS. Nathan. SRI
LANKA: North Central Province; North
Western Province; Central Province (see
Smith 1982 for records).

Distribution.—India (Tamil Nadu); Sri
Lanka.

Host.—Unknown.

Remarks.—The uniformly infuscated
wings, entirely bluish black legs, short an-
tennal length (2 times the head width), and
4-toothed tarsal claws with a basal lobe will

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

separate this species from both other spe-
cies of Anisoarthra. The female lancet and
male genitalia are ver similar to those illus-
trated for S. diascoreae (Figs. 6, 11-12).

Rohwer (1921) distinguished bilanga be-
cause of differences he observed in the
male. The females he had were identical to
A. coerulea. The characters he used were
the uniformly infuscate wings and the head
more distinctly punctured and with stronger
antennal furrows. We do not see these dif-
ferences in the males examined.

Anisoarthra diascoreae (Rohwer), new
combination
(Figs. ly, 24,6; 95 1l=12)

Senoclia diascoreae Rohwer 1921: 105 (f,
m).—Malaise 1937: 51 (in key).

Female.—Length, 10.0—11.5 mm. Bluish
black with metallic hue; basal ¥% or less of
tibiae and small spot on apices of femora
brownish to whitish. Wings darkly infus-
cated beyond proximal end of cell 1M in
forewing and hindwing; rest of wings hya-
line; veins and stigma dark brown to black.

Antennal length equal to abdomen
length, subincrassinate in middle, 2.4
head width; clypeus truncate; labrum broad-
er than long, as 2:1, with deflexed roundly
pointed anterior margin; supraclypeal area
almost flat; inner margins of eyes converg-
ing below, lower interocular distance to in-
terocular distance at front ocellus to eye
length as 7:9:8; malar space about half di-
ameter of front ocellus; head in lateral view
with frontal area though prominently raised
still slightly below level of eyes and su-
praantennal tubercles slightly indicated and
confluent with roundly raised frontal ridges;
median fovea prominent, ditchlike on an-
terior half and shallow posteriorly, reaching
median ocellus; postocellar area convex,
broader than long, as 4:3; temples lateral to
lateral ocelli depressed, post-, inter-, and
circumocellar furrows distinct; lateral fur-
rows distinct, deep, parallel and abruptly
ending well before posterior margin of
head; head from above parallel behind eyes;

VOLUME 102, NUMBER 3

Figs. 6-12. 6, Lancet of Anisoarthra diascoreae. 7, Lancet of A. birmanica. 8, Female sawsheath of A.
coerulea. 9, Female sawsheath of A. diascoreae. 10, Female sawsheath of A. birmanica. 11, Ventral view of
genital capsule of A. diascoraea. 12, Lateral view of penis valve of A. diascoraea.

606

postocellar, ocello-occipital, oculo-ocellar,
and oculo-occipital distances as 4:7:7:5.
Mesoscutellum subconvex with faintly in-
dicated lateral carina; mesoscutellar ap-
pendage neither carinated or grooved; dis-
tance between cenchri to distance between
tegulae as 2:9. Tarsal claws with 4 distinct
teeth and triangular basal lobe (Fig. 4); for-
etibial spur furcate at apex; hind basitarsus
longer than 3 following segments com-
bined, as 5:4; length of inner tibial spur to
apical width of hind tibia to outer apical
tibial spur length as 5:3:4. Lancet as in Fig.
6, with 22 serrulae. Sheath in lateral view
long wit dorsal margin slightly curved up-
ward (Fig. 9).

Head with shallow and scattered punc-
tures, surface shining, mesonotum with mi-
nute, scattered punctures, surface shining;
mesoscutellum with conspicuous punctures
on posterior slope, surface polished; mesos-
cutellar appendage impunctate, polished;
mesepisternun and mesosternum punctured
like mesonotum, surface shining with gen-
eral oily luster; abdomen shining with some
scattered and minute punctures. Body cov-
ered with mixed metallic blue and silvery
pubescence.

Male.—Length, 9.0—10.0 mm. Similar to
female except antennal segments 7—9 and
extreme apices of femora brownish. Malar
space linear; postocellar area broader than
long, as 3:2. Hypopygium truncate, with
shallow notch at center. Genitalia as in Figs.
11-12.

Types.—The holotype 2 is in the
USNM, labeled “‘N. Malabar, Taliparamba,
India’; “‘on pepper vine June ‘18.’ Allo-
type and paratypes in USNM; paratypes in
the Zoological Survey of India, Calcutta.

Specimens examined.—INDIA: Kerala,
Poonnudi Range (Trivandrum), 3,000 ft,
May 1972; South Canara Dt., Kollar Ghat,
3000 ft., 18-21-IX-18, T.V.R. (1 ¢ paratype,
USNM); N. Malabar, Taliparamba, July—
Aug. 1918, PS. Nathan (¢ allotype,
USNM), same data, 16-26-[X-18, Ramak-
rishna, coll. (1 6 paratype, USNM); Am-
matti, Coorg, S. India, May 1951, BS. Na-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

than; Nilgiri Hills, S. India, 3,500 ft., May
1950, P.S. Nathan.

Distribution.—India (Kerala).

Host.—Rohwer (1921) stated adults were
bred ‘‘on a creeper, Diascorea,’’ probably a
misspelling for Dioscorea sp. (Dioscorea-
ceae). An adult was collected “‘on pepper
vine”’ (? black pepper, Piper nigrum L. [Pi-
peraceae]).

Remarks.—The tarsal claws with 4 teeth
and a distinct triangular basal lobe, whitish
to brownish bases of tibiae and extreme api-
ces of femora, hyaline basal half of fore-
wing, furcate foretibial spur, postocellar
area broader than long, and antennal length
2.4 times head width, will separate this spe-
cies from other species of Anisoarthra.

Anisoarthra birmanica (Malaise), new
combination
(Figs..3,; 7710)

Senoclia birmanica Malaise 1937: 51 (f).

Female.—Length, 12.0-13.0 mm. Blu-
ish-black with metallic hue antenna, la-
brum, extreme apex of forefemur, basal half
of foretibia, midtibia except extreme apex,
and basal ¥ of hind tibia whitish to light
brownish. Forewing and hindwing deeply
infuscated beyond proximal end of cell 1 M
and infuscation linearly extending to base
of wings covering entire anal cell; veins and
stigma dark brown to black.

Antennal length equal to abdomen
length; subincrassinate in middle, 3.2
head width. Clypeus truncate; labrum
broader than long as 2:1 deflexed and
roundly pointed anteriorly; supraclypeal
area subtriangularly roundly raised; inner
margins of eyes converging below, lower
interocular distance to interocular distance
at level of front ocellus to eye length as 3:4:
3; malar space linear; head in lateral view
with frontal area almost at level of eyes and
supraantennal tubercles significant with
steep posterior slope and meeting low lying
frontal ridges; median fovea prominent on
anterior half and shallow posteriorly, reach-
ing median ocellus; frontal area anterior to

VOLUME 102, NUMBER 3

median ocellus conspicuously depressed,
thus forming humplike raised projection be-
tween depression and median fovea; pos-
tocellar area convex, broader than long, as
3:2; temples lateral to lateral ocelli subcon-
vex; post-, inter-, and circumocellar furrows
distinct; lateral furrows quite distinct, deep,
diverging backwards and abruptly ending
well before hind margin of head; head from
above parallel behind eyes; postocellar,
ocello-occipital, oculo-ocellar, and oculo-
occipital distances as 4:4:5:4. Mesoscutel-
lum subconvex, its appendage neither cari-
nated nor grooved; distance between cen-
chri to distance between tegulae as 1:5; tar-
sal claws comblike with 5 teeth and
indistinct rounded basal lobe (Fig. 5); for-
etibial spur simple, hind basitarsus longer
than following 3 segments combined, as 5:
4; length of inner tibial spur to apical width
of hindtibia to outer apical tibial spur length
as 5:3:4. Lancet as in Fig. 7, with 21 ser-
rulae; sheath in lateral view long with dor-
sal margin slightly curved up apically (Fig.
10).

Head with dense, irregularly spaced,
prominent punctures and surface between
punctures microsculptured, surface shining;
mesonotum with dense, irregular, minute
punctures, surface shining; mesoscutellum
with distinct, irregular, scattered punctures,
more conspicuous on its posterior slope,
surface polished; mesoscutellar appendage
impunctate, polished; mesepisternum and
mesosternum with few, scattered micro-
punctures, surface shining with general oily
luster; abdomen impunctate, shining. Body
covered with mixed blackish and metallic
blue pubescence.

Male.—Length, 10.0—11.0 mm. Similar
to female. Hypopygium truncate at apex.
Genitalia similar to Figs. 11—12.

Types.—Described from 2 ¢, deposited
in the Naturhistoriska Riksmuseet, Stock-
holm, Sweden. From Burma (Taungdo at
the south end of Inle Lake in southern Shan
States, alt. 900 m, 19.I1X.1934) (Malaise
1937).

Specimens examined.—INDIA: Assam,

607

Shillong, 1,450 m, 7.6.1943 (ZDPU; As-
sam, Kameng, Bokhar, 28-V-61, 2,500’
(CNC).

Distribution.—Burma; India (Megha-
laya).

Host.—Unknown.

Remarks.—The mostly whitish to
brownish tibiae, whitish extreme apices of
the femora, hyaline basal half of forewing
and hindwing, triangularly raised supracly-
peal area, 5-toothed tarsal claws with and
indistinct rounded basal lobe, white labrum,
and simple foretibial spurs will separate this
species from other species of Anisoarthra.
We did not examine the types. Malaise’s
(1937) description is sufficient to character-
ize this species. A series of three females
and three males from Kameng, Assam as-
sociate the sexes.

ACKNOWLEDGMENTS

Financial assistance rendered by USS.
PL-480 in collaboration with ICAR is ac-
knowledged with thanks. We are grateful to
Henri Goulet, Agriculture and Agri-Food
Canada, Ottawa (Canadian National Collec-
tion), for allowing examination of speci-
mens under his care and for reviewing the
manuscript. We also thank the following for
review of the manuscript: A. S. Jensen and
E. E. Grissell, Systematic Entomology Lab-
oratory, USDA, Beltsville, MD, and Wash-
ington, DC, respectively. Cathy Anderson,
Systematic Entomology Laboratory, helped
arrange the plates.

LITERATURE CITED

Benson, R. B. 1938. On the classification of sawflies
(Hymenoptera: Symphyta). Transactions of the
Royal Entomological Society of London 87: 353-—
384.

Cameron, P. 1876. VIII. Description of new genera and
species of Tenthredinidae and Siricidae chiefly
from the East India, in the collection of the British
Museum. Transactions of the Entomological So-
ciety of London, pp. 459-471.

. 1877. VII. Descriptions of new genera and
species of East Indian Tenthredinidae. Transac-
tions of the Entomological Society of London, pp.
87-92.

Dalla Torre, C. G. 1894. Catalogus Hymenopterorum,

608 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Vol. 1. Tenthredinidae incl. Uroceridae (Phyllo-
phaga & Xylophaga). Lipsiae, 459 pp.

Kirby, W.E 1882. List of Hymenoptera in the British
Museum, Vol. 1. London, 450 pp.

Malaise, R. 1937. New Tenthredinidae mainly from the
Paris Museum. Revue Frangaise d’Entomologie 4:
43-53.

. 1964. New genera and species of the subfam-
ily Blennocampinae (Hymenoptera, Tenthredini-
dae). Entomologisk Tidskrift 85: 20-39.

Rohwer, S. A. 1911. Technical papers on miscella-
neous forest insects II. The genotypes of the saw-

flies or woodwasps of the superfamily Tenthredi-

noidea. United States Bureau of Entomology

Technical Bulletin 20, pp. 69-109.

. 1921. Notes on sawflies, with descriptions of
new genera and species. Proceedings of the Unit-
ed States National Museum 59: 83-109.

Smith, D. R. 1982. Symphyta (Hymenoptera) of Sri
Lanka. Proceedings of the Entomological Society
of Washington 85: 117—127.

Szent-Ivany, J. J. H. 1974. Insect pests of yam (Dios-
corea ssp.) in Papua New Guinea. Folia Ento-
mologica Hungarica (Series Nova) 28: 205-210.

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 609-612

CHESPIRITOS, A NEW GENUS OF LIMOSININAE
(DIPTERA: SPHAEROCERIDAE) FROM COSTA RICA

S. A. MARSHALL

Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada
N1G 2W1 (e-mail: smarshall @evbhort.uoguelph.ca)

Abstract.—Chespiritos sindecimus new genus, new species, is described from speci-
mens taken in decaying bromeliads in Costa Rica. The relationship of Chespiritos to other

Limosininae is discussed.

Key Words:

Chespiritos sindecimus new genus, new
species, was collected around the leaf-bases
of wind-downed bromeliads at 2,400 m in
the Rio Macho reserve, Costa Rica. Most
of the bromeliads examined were found to
contain specimens of this new species, as
well as specimens of an unidentified Prer-
ogramma Spuletr.

Terminology in this paper follows Mar-
shall and Langstaff (1998). Terms used for
the same structures in other recent papers
on Sphaeroceridae are indicated in paren-
theses.

Chespiritos Marshall, new genus

Type species.—Chespiritos sindecimus,
new species.

Diagnostic characters and similar gen-
era.—Chespiritos stands out as a distinct
genus because of its broad head with a large
and exposed clypeus, two pairs of large in-
terfrontal bristles, six scutellar bristles, dis-
tinctive male genitalia, and the complete
lack of a tenth tergite in the female. The
most closely related genus is probably
Sclerocoelus Marshall, with the broad lu-
nule and well-developed intra-alar bristles
providing some evidence for this relation-
ship. Chespiritos, however, lacks the elab-
orately sclerotized male genital pouch,

Diptera, Sphaeroceridae, Costa Rica, taxonomy

complex subcercus, and broad alula which
define Sclerocoelus (Marshall 1995), and
Sclerocoelus lacks the basal scutellar bris-
tles, two interfrontal bristles, highly derived
phallus, and totally reduced female tergite
10 (epiproct of authors) which appear to de-
fine Chespiritos. Similar small basal scutel-
lar bristles do appear elsewhere in the
Sphaeroceridae (some species of the Pulli-
mosina subgenus Dahlimosina Rohacek,
and three species of Spelobia Spuler, for ex-
ample). The female tergite 10 of other Li-
mosininae is occasionally reduced (as in
some Sclerocoelus), and is occasionally ab-
sent (as in some Pterogramma with greatly
reduced cerci), but the complete absence of
tergite 10 in combination with the presence
of well-developed cerci is a distinctive at-
tribute of Chespiritos.

Chespiritos sindecimus Marshall,
new species
(Figs. 1-8)

Description.—Length from base of an-
tenna to wing tip 3.0 mm, general color
brown, tibiae and tarsi luteous. Head strik-
ingly broad, frontal width between eyes 3X
height of interfrontal area; clypeus large,
dark and exposed. Frons with two equal in-

610

? Surstylus

l

Sternite 5

Sternite 8

Sternite 7 7

Figs. 1-8.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Cercus
Tergite 8 6

Tergite 7

Chespiritos sindecimus. 1-3 and 5, Male abdominal structures. 1, terminalia, left lateral. 2, ter-

minalia, posterior. 3, aedeagus and associated structures. 5, sternites 5-7. 4, Right wing of male. 6-8. Female
abdominal structures. 6, tergites 7-8 and cerci. 7, spermathecae. 8, sternites 7—10 and cerci.

terfrontal bristles and a small lower setula;
two large orbital bristles; inner occipital
bristle large, postvertical and postocellar
bristles very small. Orbit, interfrontal stripe
and ocellar triangle with indistinct silver
pollinosity. Palpus clavate, setulose, with

only weak apical hairs and a single preap-
ical ventral bristle. First flagellomere flat-
tened laterally, rounded apically; arista aris-
ing dorsolaterally, length twice head height,
aristal hairs long (5X aristal width at mid-
length). Prosternum bare. Thorax heavily

VOLUME 102, NUMBER 3

pollinose; 2 strong postsutural dorsocentral
bristles separated by 6 rows of acrostichal
setae; outer postpronotal bristle strong, both
presutural and postsutural intra-alar bristles
strong. Scutellum long, with 2 pairs of long
marginal bristles and a pair of small basal
bristles. Dorsal surface of mid tibia with 5
bristles proximally (3 anterodorsal and 2
posterodorsal; the posterodorsal bristles
small and sometimes difficult to distinguish
from posterodorsal setulae) and 5 bristles
distally (2 anterodorsal, 2 dorsal, 1 poster-
odorsal). Male mid tibia with an apicoven-
tral bristle, an anteroventral bristle near
middle, and a double row of short stout
ventral bristles on distal half, base of mid
femur with corresponding stout bristles; fe-
male mid tibia with a mid ventral and an
apical ventral bristle. Hind tibia with a
small anteroventral bristle at apex. Wing
(Fig. 4) long, third costal sector 0.8 length
of second, vein R,,; gently sinuate; costa
extending very far (at least 10 vein-widths)
beyond apex of vein R,,,;; distance between
crossveins dm-cu and r-m 3% as long as
dm-cu, both M,,, and CuA, extending be-
yond discal cell as pigmented processes; al-
ula narrow.

Female abdomen: Tergite 7 with a pale
posteromedial notch; tergite 8 pale but con-
vex posteromedially, laterally expanded and
with differentiated posteroventral parts; ter-
gite 10 absent (Fig. 6). Cercus short, strong-
ly tapered, entirely setulose, with 3 small
outer bristles, a long straight apical bristle
and a curved inner preapical bristle. Ster-
nite 7 with a pale posteromedial notch; ster-
nite 8 reduced to a small, medially pale,
transverse sclerite with two stout bristles on
each half; sternite 10 reduced to two small
plates, one under each cercus (Fig. 7). Area
between sternite 8 and sternite 10 with a
large, hyaline vaginal sclerite with a prom-
inent posteromedial process. Spermathecae
(3) large, spherical, transversely wrinkled,
with long conical necks and short scleroti-
sed parts of ducts (Fig. 8). Pleural mem-
brane wrinkled and densely setose.

Male abdomen: Syntergite 1+2 twice as

611

long as tergite 3, middle part pale almost to
hind margin. Sternite 5 small but complex,
with two pairs of prominent posterior lobes,
outer pair long-setose and inner pair bare
and blade-like. Middle part of sternite 5
dark, very short, with a long deflexed (bent
upwards) posteromedial lobe ending in two
small, bristle-bearing lobes (Fig. 5). Ster-
nite 6 with a narrow ventral part anterior to
a distinct (but medially membranous) gen-
ital pouch; a distinct ring sclerite in right
membrane beside genital pouch. Epandrium
uniformly long-setose, subanal plate broad,
weakly bilobed ventrally, subepandrial
sclerite broad and arching slightly above
level of subanal plate; each half of subanal
plate with a single long bristle (Fig. 2). Sur-
stylus leaf-like, with a row of stout bristles
on anterior edge and a basal patch of fine
bristles (Fig. 1). Hypandrium stout, deeply
cleft posteriorly and with an elongate apo-
deme anteriorly, anterior arms long and
fused both with apodeme and epandrium,
no median posterior part but with small
lobe connecting anterior arm to postgonite,
pregonite (suspensory sclerite) small and
closely appressed to postgonite. Postgonite
(paramere, gonostylus) dark, flattened, apex
serrate with a posteroapical lobe (Fig. 3).
Basiphallus stout, wedge-like; distiphallus
heavily sclerotised, tubular basally and with
two detached distal dorsal lobes and a spat-
ulate ventral lobe.

Type material.—Holotype (d¢, INBio)
and 9 paratypes (2 2, 2 d, INBio; 2 2, 3
6 GUELPH): COSTA RICA. Cartago, Rio
Macho, La Esperanza, El Guarco (8 km
from km 61 Interamerican Highway) 2,400
m, LN-188200, 549800, Hoja Tapanti, in
bromeliads on ground, 1.vi.1998, S. A.
Marshall. Holotype and two paratype @
with two large mites each.

Etymology.—The genus is named after a
well-known truck stop on the Interamerican
Highway near the type locality; the gender
is masculine. The specific name refers to
the lack of tergite ten on the female abdo-
men.

Comments.—The type series of Chespir-

612

itos sindecimus was collected as part of an
INBio (Instituto Nacional de Biodiversidad,
Costa Rica) sponsored field trip for partic-
ipants in a planning workshop for a major
biodiversity inventory project in Costa
Rica, and this description can be viewed as
an initial contribution to that inventory.

ACKNOWLEDGMENTS

Thanks to INBio (Instituto Nacional de
Biodiversidad, Costa Rica), and especially
dipterist Manuel Zumbado, for making it

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

possible for me to collect this and many
other interesting Sphaeroceridae in Costa
Rica.

LITERATURE CITED

Marshall, S. A. 1995. Sclerocoelus and Druciatus, new
genera of New World Sphaeroceridae (Diptera:
Sphaeroceridae: Limosininae). Insecta Mundi 9:
283-289.

Marshall, S. A. and R. Langstaff. 1998. A revision of
the New World Opacifrons Duda (Diptera,
Sphaeroceridae, Limosininae). Contributions in
Science 474: 1-27.

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 613-624

A REVIEW OF THE FLEA GENERA HECTOPSYLLA FRAUENFELD AND
RHYNCHOPSYLLUS HALLER (SIPHONAPTERA: PULICIDAE)

MICHAEL W. HASTRITER AND EUSTORGIO MENDEZ

(MWH) Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM,
P.O. Box 20200, Provo, Utah 84602-0200, U.S.A. (e-mail: hastritermw @ sprintmail.com);
(EM) Instituto Conmemorativo Gorgas de Estudios de la Salud, Apartado Postal 6991,

Zona 5, Panama, Republica de Panama

Abstract.—Rhynchopsyllus, a monotypic genus, is proposed as a new junior synonym
of Hectopsylla. A key for the 12 species of Hectopsylla is provided with annotation of
host preferences and geographical distribution of Hectopsylla pulex (Haller).

Key Words:

Rhynchopsyllus pulex described by Hall-
er (1880) has been a source of taxonomic
confusion. Schreiter and Shannon (1927)
and Traub and Gammons (1950) erected
Maxilliopsylla lilloi and Rhynchopsyllus
megastigmata, respectively, from females
only. Tipton and Mendez (1966) subse-
quently described the male of R. megastig-
mata from Panama (in the presence of ac-
companying females). Later Méndez (1977)
reported associated males and females from
Colombia to be the same as R. megastig-
mata from Panama, but considered them
conspecific with R. pulex. Jordan (1939)
synonymized M. lilloi with R. pulex. Jordan
and Rothschild (1906), Dalla Torre (1924),
and Pinto (1930) each reported the species
as Hectopsylla (based only on females). In
addition, Anduze et al. (1947) and Cova
Garcia and Tallaferro (1959) questioned the
status of Rhynchopsyllus because of simi-
larities with Hectopsylla. If one uses the
characters described by Hopkins and Roths-
child (1953), Panamanian and Colombian
males reported by Tipton and Méndez
(1966) and Méndez (1977) clearly belong
to Hectopsylla. This paper provides obser-
vations to support synonymizing the mono-

Hectopsylla, Rhynchopsyllus, Siphonaptera, Pulicidae, key

typic genus Rhynchopsyllus with Hectop-
sylla.

Mammalian synonymies follow those of
Wilson and Reeder (1993) and avian syn-
onymies are those described in Peters
(1934, 1940), Deignan et al. (1964) and
Sibley and Monroe (1990). The deposito-
ries of material examined for this study are
annotated after host/locality data as: The
Natural History Museum, London
(BMNH), Field Muséum of Natural Histo-
ry, Chicago (FMNH), J.C. Beaucournu Col-
lection (JCB), Museum d’ Histoire Naturel-
le, Belgium (Md’HN), Michael W. Hastriter
Collection (MWH), Robert E. Lewis Col-
lection (REL), and National Museum of
Natural History, Smithsonian Institution,
Washington D.C. (USNM).

Genus Hectopsylla Frauenfeld

Hectopsylla Frauenfeld 1860: 464. Type
species (by monotypy): Hectopsylla psit-
taci Frauenfeld.

Rhynchopsyllus Haller 1880: 72. Type spe-
cies (by monotypy): Rhynchopsyllus pu-
lex Haller. New synonymy.

Maxilliopsylla Schreiter and Shannon 1927:
6. Type species (by monotypy): Maxil-

614

liopsylla lilloi Schreiter and Shannon.
Synoymized by Jordan 1939: 303.

The diagnostic features distinguishing
Rhynchopsyllus from Hectopsylla have
been based only on females of the former.
Jordan (1934) and Hopkins and Rothschild
(1953) based separation of these two genera
by the morphology of the spermatheca and
maxilla. Jordan (1934) specifies the differ-
ences as ‘a long, narrow, pointed and
curved maxilla and a conical projection at
the orifice of the spermatheca”’ in Rhyn-
chopsyllus, whereas the maxilla of Hectop-
sylla is ‘“‘short and broadly triangular and
the orifice of the spermatheca is flush with
the surface, not projecting on a cone”’.
More recent descriptions of the male of R.
pulex by Tipton and Méndez (1966) and
two additional species (H. gracilis Mahnert,
1982 and H. pascuali Beaucournu and A\l-
cover, 1989) provide evidence that would
suggest Rhynchopsyllus is synonymous
with Hectopsylla. Hectopsylla gracilis and
H. pascuali also have sharply pointed max-
illae that are directed caudad. Both species
clearly belong to Hectopsylla. Comparison
of these three species illustrates the simi-
larity of their maxillae (Figs. 19-21). Rhyn-
chopsyllus pulex is the only species among
the two genera, which bears a broad conical
projection at the orifice of the duct of the
spermatheca. Although the spermatheca is
morphologically distinct from that of all
species of Hectopsylla (except H. stomis to
which it is quite similar) the authors con-
sider the conical structure relevant as a spe-
cies distinction only.

Both genera share a common sessile par-
asitic mode of life (primarily in the fe-
males). Modifications for a sessile mode of
life shared by females of both genera in-
clude extreme serration and sometimes ex-
treme elongation of the lacinia, angular
frons, compression of thoracic segments,
partial covering of the anterior portion of
antennal fossa, and expansibility of inter-
segmental abdominal membranes (not true-
ly neosomic). The expansibility of interseg-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

mental abdominal membranes is demon-
strated most dramatically among females of
R. pulex, H. knighti, and H. psittaci (the
only species within the two genera with
preference for volant hosts). This is most
extreme in R. pulex and is accompanied by
autoseverence of appendages (a feature of
species of Tunga). Evidence of autosever-
ence in R. pulex includes catabolic scarring
of remaining coxae and trochanters, absent
in all species of Hectopsylla examined.

Neither males of Rhynchopsyllus, nor
Hectopsylla have developed adaptations to
accomodate a sessile mode of life (shorter
mouthparts, fewer serrations on lacinia, ten-
dency for rounded frons, and inexpansible
abdomens as demonstrated by examination
of mounted male specimens of Hectopsylla
previously attached to hosts). The feeding
and copulatory behavior of Rhynchopsyllus
or Hectopsylla males is virtually unknown.
Their lack of abundance on hosts (none for
Rhynchopsyllus) would suggest that feeding
is minimal (if at all) and the occasional col-
lection of male specimens of Hectopsylla
from a host likely occurs while they are
seeking females for mating.

The clasper of Rhynchopsyllus is closely
allied with that of Hectopsylla, possessing
a posteriorly projecting manubrium termed
manubrium 2 by Hopkins and Rothschild
(1953). The pattern of the aedeagus of
Rhynchopsyllus is also similar to those of
all species of Hectopsylla. The sclerotized
inner tube (S.I.T.) is nearly identical, being
extremely long and narrow with a small
basal capsule to guide the short penis rod
into the S.I.T. (Figs. 5—6, 8). Dorsal to the
capsule and running obliquely parallel to
the S.I.T. is a heavily sclerotized structure
referred to as the crescent sclerite by Tipton
and Méndez (1966) in their description of
R. pulex males. This pair of sclerites is pre-
sent in all Hectopsylla species, as well as
R. pulex, but is not present in Tunga, or
Echidnophaga (the most closely allied gen-
era). Since the structure is dorsal and dis-
tinct from the defined capsule of the aedea-
gus (and not the roof of the capsule = cres-

VOLUME 102, NUMBER 3

615

Table 1. A summary of specimens examined during this study illustrating host preferences within the genus

Hectopsylla.

Non-Chiropteran Host

Flea species Chiropteran Mammal Aves Unknown

H. broscus 10) 1/13! 0) 0)
H. coniger 0 1/7 0) 0)
H. cypha 0 5/74 0 0)
H. eskeyi 0 5/40 0 10)
H. gemina 0) 18/54 0) 8/6
H. gracilis 0) 2/9 0) 0)
H. knighti 0) 0) 0/1 0)
H. pascuali 0) 1/2 0) 10)
H. psittaci O 6) 5/61 0/5
H. pulex 0/34 0/3 O 3/3
H. stomis 0) WAS) 0/5 0/4
H. suarezi 0) 5/6 0) 0/1

0/0 = number of males/number of females.

cent sclerite), it is not the crescent sclerite.
These sclerites are herein referred to as the
dorsal armature (D.A.) (Figs. 5—6). The
apex of the D.A. of specimens of all species
appears fused or at least contiguous with
the S.I.T. (Fig. 6) (the position of the D.A.
as illustrated in Fig. 5:is atypical of most
specimens examined). Based on the com-
mon morphological features discussed, the
authors consider the monotypic genus
Rhynchopsyllus a junior synonym of Hec-
topsylla.

Hectopsylla pulex (Haller)
(Figs. 6—7, 19)

Rhynchopsyllus pulex Haller 1880: 72.

Rhynchopsylla_ pulex: Taschenberg 1880:
56.

Hectopsylla psittaci Baker 1904: 375, 434.

Hectopsylla pulex: Jordan and Rothschild
19065.59;.63.

Rhynchopsyllus pulex: Cunha 1914: 172.

Hectopsylla pulex: Dalla Torre 1924: 20.

Maxilliopsylla lilloi Schreiter and Shannon
1927: 6 (type species of Maxilliopsylla,
by monotypy).

Rhynchopsyllus pulex: Ewing 1929: 158.

Hectopsylla pulex: Pinto 1930: 332.

Rhynchopsyllus pulex: Jordan 1934: 19;
Jordan 1939: 303 (synonymy of M. /il-
loi); Guimaraes 1940: 219; Fuller 1942:

44; Anduze et al. 1947: 1-10; Macchia-
vello 1948: 15; Augustson and Ryan
1948: 111.

Rhynchopsyllus megastigmata Traub and
Gammons 1950: 271; Hopkins and
Rothschild 1953: 68.

Rhynchopsyllus pulex: Hopkins and Roths-
child 1953: 66; Johnson 1957: 237; Bar-
rera and Diaz-Ungria 1957: 174.

Hectopsylla pulex: Cova Garcia and Tallaf-
erro, 1/9592 35.

Rhynchopsyllus megastigmata: Tipton and
Méndez 1966: 296.

Rhynchopsyllus pulex: Tamsitt and Fox
1970: 1093; Tipton and Machado-Allison
1972: 4; Méndez 1977: 164 (synonymy
of R. megastigmata).

Material examined.—ARGENTINA:
Buenos Aires, 1 2, Ex: Nyctinomus bras-
iliensis = Tadarida brasiliensis (1. Geof-
froy) (BMNH). BOLIVIA: S. Cruz de la
Sierra, 2 2, Ex: Molossus obscurus =
Molossus molossus (Pallas) (BMNH);
Magdalena, Dept. Beni, 2 2, Ex: Eumops
bonariensis (Peters), 12 Nov 1966; Mag-
dalena, Dept. Beni, 1 2, Ex: Noctilio la-
bialis = Noctilio albiventris Desmarest,
20 Aug 1964 (USNM). BRAZIL: 1
(paratype), Ex: Molossus sp.; Lagoa San-
ta, 1 2, Ex: Eumops perotis (Schinz), 2
Feb 1916 (BMNH); Lagoa Santa, 2 °,

616

Ex: E. perotis, 2 Jan 1944 (USNM); Pe-
tropolis, 1 2, Ex: “‘ear of bat’’; Parana
Prov., 3 2°, Ex: Histiotus velatus (1. Geof-
frey) (BMHH). COLUMBIA: Anserman-
uevo, Dept. Valle, 4 2, Ex: M. molossus,
20 Jul 1967; Cali, Dept. Valle, 2 2, Ex:
Molossus major = M. molossus, 22 Nov
1962; Cali, Dept. Valle, 12, Ex: M. mo-
lossus, Jun 1967; Cartago, Dept. Valle, 2
2, Ex: Molossus bondae J.A. Allen, 10
Oct 1967 (USNM). ECUADOR: Albitig-
ua, elev. 1,000 m, 3 2, Ex: Rhipidomys
leucodactylus (Tschudi); Gualquiza, elev.
760 m, 12, Ex: ‘“‘bat’?. PANAMA: Pa-
cora, 2d, Ex: “bat guano’’, 22 Jun 1961
and 27 Feb 1962 (USNM); Pacora, 1 6,
ie oeExe bat guano’, 20, 22 Jun 1961:
Pacora, 2 °, Ex: “‘bat guano’’, 12, 14 Feb
1962; (MWH). PERU: Rio Chinchao,
Dept. Huanuco, | 2, Ex: Molossus ob-
scurus = M. molossus (BMNH); Quince
Mil, Cuzco Prov., 1°, Ex: T. braziliensis,
19 Jun 1950 (USNM). TRINIDAD: San-
tissimas il! “2 Exe’ “bat-4-27 Jul i915
(USNM). UNITED STATES: Frio Cave,
Uvalde County, Texas, 1 2, Ex: Tadarida
mexicana = T. brasiliensis, 30 Mar 1955
(USNM). VENEZUELA: Esteban,
Djiques, 2 2, Ex: M. obscurus; Merida,
1 2, Ex: Vespertilio fuscus = Eptesicus
fuscus (Beauvois) (BMNH); 3 km S and
46 km W Caracas, Dto. Federal, 2 2, Ex:
M. obscurus = M. molossus, 19, 20 Aug
1966 (USNM).

Remarks.—Hectopsylla pulex, one of
12 species of Hectopsylla, occurs primar-
ily on bats of the family Molossidae and
to a lesser extent on Vespertilionidae. The
remaining species parasitize either Aves
(HM. knighti, H. psittaci) or non-Chirop-
teran mammals (Caviidae: H. cypha, H.
eskeyi, H. gemini, H. suarezi; Muridae:
H. pascuali, H. gracilis; Mustelidae: H.
broscus, H. coniger; and Chinchillidae:
H. stomis) (Table 1). The known country
records of Hectopsylla pulex, its host spe-
cies and host synonymies are listed in Ta-
ble 2. The occurrence of a single female
of H. pulex on Zonotrichia pileata =

Nw

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Zonotrichia capensis (Miller) (reported
in Cunha 1914) and three females ex-
amined from the Murid rodent R. leuco-
dactylus are likely accidental associa-
tions.

Little is known about the life cycle of
H. pulex. Females have been collected
frequently on chiropteran hosts while
males have been collected only from bat
guano associated with species of Molos-
sus in Colombia and Panama.

KEY TO SPECIES OF HECTOPSYLLA

Maxilla rounded at apex (Fig. 11) (male un-
known) knighti
Maxilla broadly angular to sharply pointed at
apex (Bigs: 1222) oie cs eae a ee 2
Segment V of all tarsi with seven or eight
(usually eight) lateral planter bristles; tarsal
claws with a prominent basal tooth; metepi-
meron of female without a process on dorso-
posterior margin of metepimeron psittaci
Segment V of all tarsi with at most six pairs
of lateral plantar bristles; tarsal claws without
a prominent basal tooth; metepimeron of fe-
male nearly always with a process at dorso-

posterior margin (Fig. 2) (except pulex) ... 3
Malet. fie afeiiege & 2c A eee 4
Femalé;..¢ cbs ssieacite a betel th eee 13

P! of clasper narrower at apex than at base
(Figs. 3, 7)
P! of clasper broader at apex than at base (Fig.
A) verses ak suyex genie hit ee Ee eee 7
Tarsal segments V with six pairs of lateral
plantar bristles; median dorsal lobe of aedea-
gus absent or vestigial ............. broscus
Tarsal segments V with five or fewer pairs of
lateral plantar bristles on tarsi; median dorsal
lobe well developed (Fig. 5)
P? and P® of clasper pincer-like (Fig. 4); distal
arm of st. [IX divided into several lobes; fifth
tarsal segments with four pairs of lateral plan-
tar"bristles "iss seg. a ee ee © oe stomis
P? and P® not pincer-like (Fig. 7); distal arm
of st. [IX composed of a single lobe; fifth tar-
sal segments with five pairs of lateral plantar
DTIStleS yas cece meee gaeys ok ee eee pulex
Median dorsal lobe of aedeagus poorly de-
veloped, at most discernible
Median dorsal lobe well developed (Fig. 5),
distinctly visible
Median lobe (L?) of distal portion of st. IX
enlarged with concavity at ventral margin;
apical margin of P! of clasper straight; P* not
bifurcate apically gracilis
Ventral margin of L? lacking concavity; apical

VOLUME 102, NUMBER 3

617

Table 2. Distribution and host-parasite relationships of Hectopsylla pulex.

Country

Host species!

Flea sex

Argentina

Myotis nigricans (Schinz)

2

Nyctinomus brasiliensis = Tadarida brasiliensis (1. Geot- 2

froy)

Promops perotis = Eumops perotis (Schinz)
Zonotrichia pileata = Zonotrichia capensis? (Miiller)
Eumops bonariensis (Peters)

Bolivia

Molossus obscurus = Molossus molossus (Pallas)
Noctilio labialis = Noctilio albiventris Desmarest

Brazil E. perotis

Histriotus velatus = Histiotus velatus (11. Geoffroy)
Molossus bondae J. A. Allen

Molossus obscurus = M. molossus

Molossus rufus = Molossus ater E. Geoffroy
Nyctinomus brasiliensis

Colombia Molossus bondae

Molossus major = M. molossus
Molossus molossus major = M. molossus

M. molossus

Molossus obscurus obscurus = M. molossus
Noctilio labialis = Noctilio albiventris

‘““Bat’?

Rhipidomys leucodactylus* (Tschudi)

E. perotis

Geoffroy)
Histiotus sp.

Molossus obscurus = M. molossus

Tadarida brasiliensis
Salata

E. perotis

Tadarida mexicana =

Trinidad
United States

Venezuela

T. brasiliensis
Nyctinomus macrotis = Nyctinomops macrotis (Gray)

Tadarida yucatanica = Nyctinomops laticaudatus (E.

T. brasiliensis
M. major = M. molossus

M. obscurus = M. molossus

Myotis nigricans

Vespertilio fuscus = Eptesicus fuscus (Beauvois)

+O +0

Oy Oy
40)4+0) 40 FO GO" — 40) +0) +O 40) 40) 4OT4O 407410 40 40 tORtO tO

Oy
+O

+O +O +O +0 +0 +0 +0 +0 +0 +0

' Host synonymy after Wilson and Reeder (1993).
? Only known record occurring on a bird.

3 Only known record occurring on a non-Chiropteran mammal.

margin of P! somewhat convex; P2 strongly
bifurcated into two dentate projections
pascuali
9. Dorso-posterior margin of metepimeron with
a caudally directed process (Fig. 2)
— Dorso-posterior margin of metepimeron en-
tirewlackinoyprocess oe Sood se. 4s] - 10
10. Sclerotized inner tube (S.I.T.) with a sharp
thorn-like spine at ventral midpoint (Fig. 8);
apex of ventral lobe (L*) of distal portion of
st. [IX pointed; fifth tarsal segments with three
pairs of lateral plantar bristles (some with
three and four on same metatarsus) . gemina
— §S.LT. without thorn-like spine at ventral mid-

. coniger

point, if with tubercle or protuberance then
rounded and blunt (Fig. 5); apex of L* round-
ed; fifth tarsal segments with either four or

five pairs of lateral plantar bristles ...... 11
. L? with lateral patch of long thin setae; usu-
ally four pairs of lateral plantar bristles on
fiithytarsalfscoments ee ae een ie cypha
L? without lateral patch of setae, but with ver-
tical line of setae; four or five pairs of lateral
plantar bristles on fifth tarsal segments .. . 12
. Fifth tarsal segments with four pairs of lateral
plantar/bristles® ets. <5. a ee oe ae eskeyi
Fifth tarsal sements with five pairs of lateral
platitarsbristles, . "2088 Pee 2 eee suarezi

618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

9 10

Figs. 1-10. 1, Head, female H. broscus. 2, Metepimeron, H. eskeyi (female paratype). 3, P' of clasper, H.
stomis. 4, Clasper, H. eskeyi (holotype). 5, terminal portion of aedeagus, H. eskeyi (holotype), 6, Terminal prtion
of aedeagus, H. pulex. 7, Clasper, H. pulex. 8, Sclerotized inner tube, H. gemina. 9, Metatarsal claw, H. pascuali
(allotype). 10, Metatarsal claw, H. gracilis (female paratype). Abbreviations: CR. = crochet; C.S. = crescent
sclerite; D.A. = dorsal armature; L.L. = lateral lobe; L.S. = lateral sclerite; M.D.L. = medium dorsal lobe; P',
P2 and P? = processes of clasper; PS.C. = pseudocrochet; S.I.T. = sclerotized inner tube. Scale = 100p.

VOLUME 102, NUMBER 3

13. Metepimeron with a dorso-posterior projec-
tion (Fig. 2)
— Metepimeron without a dorso-posterior pro-
jection pulex
14. Posterior margin of occiput with a well de-
finedwlobe (bisa) tees see ce wea ene eee 15
— Posterior margin of occiput without a well de-

fined lobe (though slightly indicated in eskeyi)

15. Tergum II with three setae per side and t. VII
with two or three setae per side; frons with
convex angle midway between oral angle and
internal incrassation (Fig. 1); five or six (usu-
ally six) lateral plantar bristles on fifth tarsal
SECMENtS Aaesissloticbs CG.cs ae a Sew broscus

— Tergum II with one or two setae per side and
t. VII with one per side; frons with angular
projection immediately below internal incras-
sation; three lateral plantar bristles on fifth
(RICA SSSNETNIG 5 poo ea pom co ooo oc coniger

16. Sclerotized rim of sensilial plate anterior to
sensilial pits greater than twice width of dou-
blesowsOl pits ef cei Pi sns agls: Sacp tego ee stomis

— Sclerotized rim of sensilian plate anterior to
sensilial pits at most as wide as double row
of pits

17. Fifth tarsal segments with three pairs of lat-
eral plantar bristles; ventral apical margin of
t. VIII with tooth-like projection

gemina

— Fifth tarsal segments with either four or five
pairs of lateral plantar bristles; ventral apical
margin of t. VIII without tooth-like projection

18. Fifth tarsal segments with five pairs of lateral
plantar bristles (occasional specimens may
have four and five on the same _ tarsus)

Pe Neher b ot Picacho oats a aeaee Shad 2-3 ketaars suarezi

— Fifth tarsal segments with four pairs of lateral
Plantarbnistlesse ee serene ee ae 19

19. Dorsal margin of metepimeron heavily scler-
otized, sclerotization extending to apex of
process which is markedly turned down; usu-
ally three setae on metepimeron ....... cypha

— Dorsal margin of metepimeron not noticeably
sclerotized (Fig. 2); usually four setae on me-

He PIMELOME foe eee ob a ea wen meets S 20

20. Hilla of spermatheca nearly as wide through-
out as width of bulga; duct of spermatheca
connecting at cribriform area of bulga on ven-

Teal MAT OWN wma. tte cis acc fies Sees ioe eskeyi

— Hilla of spermatheca much narrower than
width of bulga, with marked narrowing from
base of bulga to apex of hilla; duct of sper-
matheca connecting at cribriform area of bul-

ga on ventral apical margin ........... 21
21. Base of metatarsal claw with small cleft, or
CHHAUNS (CS ex OD) A) Gi Stu au 6 Blond ote. 0, Sto oe pascuali

619

— Base of metatarsal claw without cleft, or sinus
(Fig. 10) gracilis

OTHER MATERIAL EXAMINED

Hectopsylla broscus Jordan and
Rothschild 1906
(Fig. 12)

Material examined.—ARGENTINA:
Central Pampa, 1 2? (lectotype), Ex: Co-
nepatus humboldtii Gray; Central Pampa, 5
2 (paralectotypes), Ex: C. humboldtii; San
Rafael, Mendoza Prov., 1 6, 1 2, Ex: Co-
nepatus suffocans = C. chinga (Molina);
Don Roberto, San Luis Prov., 1 2, Ex: Co-
nepatus chinque = C. chinga, 14 Jun 1962;
‘Southern Argentina,’ 5 2, Ex: Zaedyus
pichiy (Desmarest), 30 Jun 1962 (BMNH).

Remarks.—All records of this species are
restricted to Argentina and skunks of the
genus Conepatus appear to be the preferred
host.

Hectopsylla coniger Jordan and Rothschild
1906
(Fig. 17)

Material examined.—BOLIVA: Pampa
Olliga, 1 ¢ (lectotype), Ex: Conepatus ar-
equipae = C. chinga, 19 Oct 1901; Pampa
Olliga, 1 2 (lectoallotype), 6 2 (paralec-
totypes), Ex: C. chinga, 19 Oct 1901
(BMNB).

Hectopsylla cypha Jordan 1942
(Fig. 14)

Material examined.—ARGENTINA: Las
Catitas, Mendoza Prov., 9 2 (paratypes),
Ex: Octomys barrerae = Tympanoctomys
barrerae (Lawrence), Jul 1939; Mendoza, 1
2, Ex: Microcavia australis (1. Geoffroy
and d’Orbigny), 6 Aug 1959; Puesto “‘La
Carpa,’’ Mendoza, 5 2, Ex: Graomys gri-
seoflavus griseoflavus = Graomys griseo-
flavus (Waterhouse), 12, 14 Jul 1959; Pues-
to ‘“‘La Carpa,’”’ Mendoza, 2 2, Ex: M. aus-
tralis, 13, 15 Jul 1959; San Rafael, Men-
doza Prov., 1 2 (neoallotype), 2 ¢, 52 @
(paratypes), Ex: M. australis, Apr—Jul
1939; Santa Rosa, Mendoza Prov., 2 6, 4
2 (paratypes), Ex: M. australis, Jul 1939

620 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

19 20 21 22

Figs. 11-22. First segment of maxillary palpus, maxilla, and ventral margin of head (genal lobe); male is
positioned directly above female for each species. 11, H. knighti (female holotype). 12, H. broscus (female
lectotype). 13, H. stomis (female holotype). 14, H. cypha (paratypes). 15, H. gemina. 16, H. suarezi. 17, H.
coniger (lectotype and allolectotype). 18, H. eskeyi (holotype and paratype). 19, H. pulex. 20, H. pascuali
(holotype and paratype). 21, H. gracilis (holotype and paratype). 22, H. psittaci. Scale = 100w.

VOLUME 102, NUMBER 3

(BMNH). Zapallar, La Rioja Prov., 1 ¢, Ex:
*“Viscacha”’ Lagostomus sp. Brookes, 8 Jan
1933 (USNM).

Remarks.—Mountain cavies (M. austral-
is), which are restricted to Argentina and
southern Chile, are the preferred host for
this flea. The geographic distribution of T.
barrerae is also restricted to Argentina and
it is rarely collected. Hectopsylla cypha oc-
curring on other hosts that have much
broader geographical distributions (Lagos-
tomus sp. and G. griseoflavus) are likely ac-
cidental associations.

Hectopsylla eskeyi Jordan 1933
(Figs. 2, 4—5, 18)

Material examined.—BOLIVIA: Pucara,
elev. 2,400 m, 1 2, Ex: G. griseoflavus, 28
Sep 1954; Samaipata, Dept. Santa Cruz,
elev. 1,650 m, 1 ¢, Ex: G. griseoflavus, 28
Jan 1955; Serrano, Dept. Chuquisaca, elev.
2,160 m, 1 2, Ex: Hesperomys muriculus
= Calomys callosus (Rengger), 24 Jul 1955
(BMNH). ECUADOR: Riobamba, Chim-
borazo Prov., elev. 2,800 m, 1 @°, Ex: Rat-
tus rattus (Linn.), 12 Jun 1956; Sanjapam-
ba, Tungurahua Prov., 6 2, Ex: Cavia por-
cellus (Linn.), 1956 (BNMH). PERU:
Huancabamba, elev. 1,960 m, 4 6,9 2, Ex:
Cavia cobaya (domestic) = C. porcellus,
17, 19 May 1956; Lima, 1¢ (holotype), 1
2 (Neoallotype), 1 ¢, 18 @ (paratypes),
Ex: “rats,” 1930; Yura, Arequipa Prov.,
elev. 8,200 ft, 1 2, Ex: Cavia musteloides
= Galea musteloides Meyen, 8 Aug 1939
(BMNH); Yura, Arequipa Prov., elev. 8,200
ft, 1 2, Ex: G. musteloides (REL).

Remarks.—Hectopsylla eskeyi occurs in
the higher Andean valleys on a variety of
hosts and little can be said of its host spec-
ificity.

Hectopsylla gemina Jordan 1939
(Figs. 8, 15)

Material examined.—ARGENTINA:
Mendoza, | ¢, 4 2, Ex: M. australis, 9 Aug
1959; Las Catitas, Mendoza Prov., 1 3, 1
2, Ex: Octomys barrerae = T. barrerae; La
Paz. Mendoza\Prov., 3° 6, 232 ,°Ex: M.

621

australis, Feb-Mar 1939; Puesto “‘La Car-
pa,’ Mendoza Prov., elev. 600 m, 1 6, 6
2, Ex: M. australis, 13-17 Jul 1959; Puesto
“Pugin,” Algarrobito, Mendoza Prov., 620
m, 4 6, 7 @, Ex: M. australis, 24-31 Jul
1959; San Rafael, Mendoza Prov., 86, 6 @,
Mar—May 1939; Santa Rosa, Mendoza
Prov., 66, 7 2 (paratypes), Ex: M. aus-
tralis, Jul & Feb 1939; Fortin Uno, Rio Ne-
gro Prov., 1 6 (holotype), 1 2 (paratype),
Ex: M. australis, Aug 1937; near General
Roca, Rio Negro Valley, Rio Negro Prov.,
1 36,2 2, Ex: Ctenomys sp. Blainville, 25
Mar 1952 (BMNH). Nacunan, Mendoza
Prov., 1 2, Ex: Akodon sp. Meyen, 15 Mar
1983 (REL). General Acha, La Pampa
Prov., 1 36, Ex: M. a. australis, 20 May
1936; La Cristina, 1 2, Ex: Galea sp. Mey-
en, 26 Oct 1966 (USNM).

Remarks.—All records of this species are
restricted to Argentina, the preferred host
being mountain cavies (M. australis). Al\-
though this flea has not been associated
with H. cypha, it parasitizes the same host
species in the same general geographic re-
gion.

Hectopsylla gracilis Mahnert 1982
(Figs. 10, 21)

Material examined.—ARGENTINA:
Puesto “‘La Carpa,’’ Mendoza Prov., elev.
600 m, 1 2, Ex: Canis familiaris Linn., 17
Jul 1959; Puesto “‘La Carpa,’’ Mendoza
Prov., elev. 600 m, 6 2, Ex: G. griseoflavus
ssp., 14 Jul 1959 (BMNH). Puerto Madryn,
Chubut Prov., 1 d (holotype), Ex: Elig-
modontia morgani Allen, 23 Apr 1978;
Puerto Madryn, Chubut Prov., 1 6, 1
(paratypes), Ex: E. morgani, 5 Apr 1978
(Md’HN); Puerto Madryn, Chubut Prov., 1
2, Ex: E. morgani, 24 Apr 1978 (REL).

Remarks.—Specimens from Mendoza
Province (BMNH) were not recognized as
a H. gracilis until Mahnert’s description in
1982. The geographic range of the preferred
host, E. morgani, is limited to Argentina
and adjacent southern Chile (Wilson and
Reeder 1993).

Hectopsylla knighti Traub and Gammons
1950
(Fig. 11)

Material examined.—-México Michoa-
can State, Municipality of Tancitaro, Tan-
citami, 1 2 (holotype), Ex: “‘head of
swift,” May 1940 (FMNBH).

Remarks.—Additional observations of
‘‘swifts’’ in the region of Michoacan, Méx-
ico, are needed to find the males of this
species and further elucidate its taxonomic
status.

Hectopsylla pascuali Beaucournu and
Alcover 1990
(Figs. 9, 20)

Material examined.—ARGENTINA: Rio
Chapelco, Neuquén Province, 1d (holo-
type), 1 @ (allotype), 1 2 (paratype), Ex:
Chelemys macronyx (Thomas), 9 Dec 1987
(JCB).

Hectopsylla psittaci Frauenfeld 1860
(Fig. 22)

Material examined.—ARGENTINA: 1 2
(lectotype), 2 2 (paralectotypes), Ex: Strix
pelate = Tyto alba (Scopoli); Buenos Aires,
2 2, Ex: “owl,” 1913; Chivilcoy, Buenos
Aires Province, 2 9, Ex: “‘hen,’’ 2 Feb
1936 (BMNH); Las Roses, (B.A.), 1 2, Ex:
‘“‘Paloma”’; Zapallar, La Rioja Prov., 3 °,
Oct 1933 (USNM). BRAZIL: Progne, Rio
de Janeiro State, 2 2 (USNM). CHILE: St.
Jogo (Santiago de Chile), 2 2 (syntypes),
Ex: Cyarzolyscus patagonius or Enicogna-
thus leptorhynchus (King) (BMNH); San-
tiago, 1 2, 11 Sep 1951; Santiago, 2 2, Ex:
“turtle dove’? (USNM). HOLLAND: den
Haag, 22 ¢, Ex: Phasianus sp. (L.), Aug
1926; Rotterdam, 4 2, Ex: Gallus domes-
ticus = Gallus gallus (L.), July 1936
(BMNH); den Haag, 3 2, Ex: Phasianus
sp., Aug 1926 (USNM). ENGLAND: Lon-
don (Zoological Gardens), 3 9, Ex: Citto-
cincla macrura (sic) = Copsychus mala-
baricus (Swinhoe); London (Zoological
Gardens), 3 6, Ex: ‘“‘birds in western avi-
ary’; London (Zoological Gardens) 3 °,

2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Ex: Cittocincla macrura (sic) or Copsychus
saularis (L.) (BMNH); London (Zoological
Gardens), 2 2, Ex: ‘‘Cittocincla (sic) = Kit-
tocincla (Gould) or Copsychus’”> (USNM).
PERU: Ex Ribeyro;sLimas lsd. 2697 Ex
‘“‘“chickens’’; Hacienda Las Vegas, 30 km
from Barranca City, 1 2, Ex: swallow “‘va-
riety called Santa Rosita,’ 1947 (BMNH);
Cuzcon, Ocongata River, elev. 3,500—4,000
m, 2 2, Ex: Merganetta leucogenis turneri
= Merganetta armata Gould, 26 Jul 1949;
Hacienda Ceapano, Ocongate, Cuzco Prov.,
1 2, Ex: Colaptes rupicola puna = Colap-
tes rupicola d’Orbigney, 14 Aug 1950
(USNM). UNITED STATES: Oceanside,
San Diego County, California, 1 ?, Ex:
Bubo virginianus pacificus = Bubo virgi-
nianus (Gmelin), 7 Jun 1942 (BMNH);
Oceanside, San Diego County, California, 1
2, Ex: B. v. pacificus, 7 Apr 1942 (USNM);
Alameda (6.5 km w. Newark), California, 1
36, 4 2, Ex: cliff swallow nests, 15 Jan
1981 (REL).

Remarks.—Hectopsylla psittaci, a pri-
mary parasite of birds, is recorded from di-
verse areas. However, it is native to the
southern cone of South America with pop-
ulations dispersed to the southwestern Unit-
ed States by owls and other migratory bird
species. Schwan et al. (1983) reported large
numbers of this species from the nests of
Cliff Swallows (Hirundo pyrrhonota Vieil-
lot) and Black Phoebes [Sayornis nigricans
(Swainson)] in southern California. The
species likely occurs through Central Amer-
ica as well.

Hectopsylla stomis Jordan 1925
(Figs. 3, 13)

Material examined.—ARGENTINA:
Mariano, Buenos Aires Prov., 1 2 (holo-
type), Ex: “‘bird’’, Dec 1912; Canada Mar-
iano, Buenos Aires Prov., 4 2 (paratypes),
Ex: ‘‘birds,’’ 30 Dec 1912; Bahia Blanca,
Euenos Aires Prov., 19 2 (paratypes), Ex:
Mephitis sp. E. Geoffroy and G. Cuvier
(sic), only Conepatus spp. Gray occur in
Argentina, 15 Jan 1911; Pique, Buenos Ai-
res, 1 3, 4 2, Ex: Lagostomus maximus

VOLUME 102, NUMBER 3

(Desmarest), 5 May 1928; El Quebrachal,
Salta: Proy., elev: 620 m; 2 9, Ex: L. mmax-
imus, 30 Jul 1940; El Ojito, Santiago del
Estero Prov., elev. 620 m, 1 2, Ex: L. max-
imus, 19 Jul 1940; La Paz, Mendoza Prov.,
elev, 620'm:"6" 6, 22 2) Ex) maximus, 1
Mar 1939; Puesta “‘pugin’”’ Algarrobito,
Mendoza Prov., elev. 620 m, 4 ¢, Ex: L.
maximus, 25 Jul 1959; San Rafael, Men-
doza Prov., 1 2, Ex: M. australis, 20 Apr
1939 (BMNH). Chasic6, Buenos Aires
Prov., 1 2, Ex: L. m. maximus, May 1968;
Patagonia, 4 2, Ex: ‘‘Viscacha’’ Lagosto-
mus sp., Sep 1962; Victoria, La Pampa
Broviil): SsoBxeeRattus sp.y29 Jum-1935
(USNM).

Remarks.—Lagostomus maximus, occur-
ring in southern Paraguay, and in northern
and central Argentina is the primary host
for this flea. Hectopsylla stomis has been
collected throughout the year.

Hectopsylla suarezi C. Fox 1929
(Fig. 16)

Material examined.—ECUADOR: Gua-
moti, Chimborazo Prov., 1 6, Ex: “‘guinea
pig’; Guamoti, Chimborazo Prov., 1 2, Ex:
‘“suinea pig,” 8 Aug 1927. PERU: near
Hauncabamba, Dept. Piura, 4 6, 4 @, Ex:
Cavia aperea Erxleben (domesticated),
1946/47; near Hauncabamba, Dept. Piura, 1
°, Ex: ‘‘among human clothes’? (BMNH).
Near Hauncabamba, Dept. Piura, 1 @
(USNM).

Remarks.—This flea is associated with
domestic guinea pigs associated with hu-
man habitations. Astute collecting may
yield many more specimens than are cur-
rently known, although the wide use of
DDT in human dwellings for controlling
the vectors of Chagas’ disease (conenose
bugs) and malaria (mosquitoes) may ex-
plain the absence of these fleas in collec-
tions after the mid-1940s.

ACKNOWLEDGMENTS

The authors express their appreciation to
Theresa Howard, The Natural History Mu-
seum, London; Nancy Adams, National

623

Museum of Natural History, Washington,
D.C.; Robert E. Lewis, Iowa State Univer-
sity, Ames; Jean-Claude Beaucournu, La-
boratoire de Parasitologie Medicale,
Rennes, France; and personnel from the
Field Museum of Natural History, Chicago,
and the Muséum d’Histoire Naturelle, Bel-
gium, for the loan of specimens.

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PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 625-633

THE PUPAE OF THREE SPECIES IN MINOMYIA THEOBALD FROM NEPAL
AND A KEY TO THE KNOWN PUPAE OF THE GENUS IN THE ORIENTAL
AND AUSTRALASIAN REGIONS
(DIPTERA: CULICIDAE)

RICHARD E DARSIE, JR.

Florida Medical Entomology Laboratory, University of Florida, 200 9" Street, SE, Vero
Beach, FL 32962, U.S.A. (e-mail: rfd@ gnv.ifas.ufl.edu)

Abstract.—Four species of the genus Mimomyia are known from Nepal, Mi. chamber-
laini, Mi. luzonensis, Mi. hybrida and Mi. intermedia. Complete descriptions and illustra-
tions of the first three species are provided. A key to the known pupae of Mimomyia in
the Oriental and Australasian regions is included.

Key Words: Mimomyia, pupae, Nepal

Four species of the genus Mimomyia
Theobald are known from Nepal. The first
to be reported was Mi. (Etorleptiomyia) lu-
zonensis (Ludlow) by Pradhan and Darsie
(1989); then Mi. (Mimomyia) chamberlaini
Ludlow and Mi. (Mim.) hybrida (Leicester)
by Darsie and Pradhan (1990); and most re-
cently Mi. (Mim.) intermedia Barraud, a
single female in the Nepal collection by
Darsie et al. (1992). Immature stages of the
latter are unknown. One of the purposes of
this study is to give a complete description
of the pupae of the former three species.

Due to the size of the trumpets and forms
of the paddles of the three pupae being de-
scribed, only these structures have been il-
lustrated and/or briefly described by Bar-
raud (1934), Mattingly (1957, 1971), Del-
finado (1966, Mi. chamberlaini and Mi. lu-
zonensis)) and Baisas (1974, Mi. luzonensis
only). Chen and Lien (1956) described the
pupae of Mi. luzonensis and Mi. chamber-
laini, as Mi. metallica (Leicester) in some-
what more detail; however, no complete de-
scription and illustration has previously
been made. Mimomyia chamberlaini and
Mi. metallica were considered by Mattingly

(1957) as the same polymorphic species,
and Knight and Stone (1977) listed Mi. me-
tallica as an infrasubspecies of Mi. cham-
berlaini. However, Lee et al. (1988) have
raised Mi. metallica to a subspecies of Mi.
chamberlaini.

The genus Mimomyia consists presently
of three subgenera and 45 species. Of those,
32 are in Subsaharan Africa, mostly in the
subgenera Mimomyia Theobald and Ingra-
mia Edwards, while 14 occur in the Ori-
ental and Australasian regions, 7 in the sub-
genus Mimomyia Theobald, 4 in the sub-
genus Etorleptiomyia Theobald, and 3 in
the subgenus /ngramia (Knight and Stone
1977, Knight 1978, White 1974). Pupae of
10 species, including the three treated here,
have been adequately described and are in-
cluded in the following key (Belkin 1962;
Knight and Chamberlain 1948; Mattingly
HOSA ao).

METHODS AND MATERIALS

The methods are as described by Darsie
(1998). Morphological nomenclature fol-
lows Harbach and Knight (1980). The pu-
pae of Mi. chamberlaini, collected in Nepal,

626

are in the author’s collection at the Florida
Medical Entomology Laboratory, Vero
Beach, FL, USA. For the other two species,
no pupae from Nepal are available and
those from Thailand, Malaysia, and the
Philippines were borrowed from the Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, DC. Abbre-
viations used in the descriptions are br
meaning branches and Le and Pe meaning
larval and pupal exuviae.

DESCRIPTIONS

Mimomyia (Mimomyia) chamberlaini
Ludlow
(Fig. 1)

Pupa.—Position and size of setae as fig-
ured, range and modal number of branches
in Table 1. Abdomen 3.01—4.21, * 3.86
mm, paddle 1.07—1.26, x 1.14 mm.

Cephalothorax: Striped and mottled with
darker brown pattern, those on mesothorac-
ic wing match adult wing venation. Setae
1,3,5-CT very long and stout, usually dou-
ble; 8-CT long, mostly with 5 br (4—6);
trumpet very long, 1.53—2.09, x 1.96 mm,
index 12.7—17.0, * 15.6, pinna in apical
0.16—0.23 * 19.0, with normal opening for
water surface respiration, tracheoid part
0.72-0.75 x 0.73 of total length.

Abdomen: Segments I-III medium
brown, IV—VIII with sublateral dark brown
stripes. Seta 1-I subdendritic float seta with
13—38 br; 2,3-I long, single; 1-II long, rath-
er stout, with 5-10 br; 1-III-VII and 5-IV-
VII stout, with short and long br, 1-III,1V
with 6—12 br, 1-V,VI with 4—11 br, 1-VI
with 4—6 br, 3—VII absent; 5-IV-VI with 4—
11 br, 5-VII with 3—7 br; 6-III-VI long, sin-
gle, seldom double; 6-VII ventral, short,
usually double (1—4); 9-VIII attached to a
rather pointed, apicolateral process, short,
usually 3, 4 br (1—4); female genital lobes
with rather pointed prolongation apicolater-
ally, with medium-sized spicules.

Paddle: uniformly brown, except for large
white area laterally in basal 0.58—0.74, some
pupae (70%) also with smaller white area

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

medially; index 1.75—2.18, x 1.99; midrib
extending to apex; outer margin with large
spine-like spicules in apical 0.2—0.44, tiny
spicules on apex in Nepal specimens, bare
point apically in Thai specimens; inner mar-
gin with similar large spicules in apical
0.12—0.22; seta 1-P short, single.

Material examined.—NEPAL: Lumbini
District, Shivapur, IX-13-92, 1 2 LePe, IX-
16-92, 1 2 LePe, ex Eichornia pond (Dar-
sie and Courtney). MALAYSIA: Perak,
Tanjong Tualang ER., 1968, 3 2 Pe, 2 d
Pe (811-15, 102, 104, 105, 113). PHILIP-
PINES: Mindoro Island, San Jose, I[-25-45,
1 3 LePe, ex Pistia marsh (E.S. Ross).

Mimomyia (Etorleptiomyia) luzonensis
Ludlow
(Fig. 2)

Pupa.—Position and size of setae as fig-
ured, range and modal number of branches
in Table 2. Abdomen 3.61—3.87, x 3.72 mm,
paddle 1.0—1.04, x 1.03 mm.

Cephalothorax: Uniformly light brown,
setae 1,3-CT long, single, 3-CT stout; 5-CT
medium long, usually double; 6-CT light in
color, very long and stout, single; 8-CT un-
usually stout, single; trumpet extremely
long, length 2.78—3.28, x 3.03 mm; index
25.9-35.4 xX 32.4; tracheoid 2.14—2.63, x
2.36 and pinna 0.13—0.16 x 0.148 of total
length.

Abdomen: Uniformly light tan; Seta 1-I
medium long, single; 2,3-I long to very
long, rather stout, single; 1-II long, stout,
double, sometimes single; 1-III-VI long,
stout, aciculate, usually 4 br, 1-VII very
long, triple; 3-VII absent; 5-IV-VI long,
stout, aciculate, usually triple, 5-VII 4,5 br;
6-II-VI long, single; 6-VII long, 4—6 br; 9-
II-VII short, larger than usual, single; 9-
VIII medium long or long, aciculate, 7—11
br; female genital lobe produced apicolater-
ally, with many large spines.

Paddle: light tan, linear, index 4.68—5.35
x 4.98, midrib extending to apex, outer mar-
gin with spicules coarse in apical 0.5, finer
basally; inner margin with coarse spicules
to near base, seta 1-P short, thin, single.

VOLUME 102, NUMBER 3 627

Fig. 1. Pupa of Mimomyia (Mim.) chamberlaini. A, Cephalothorax. B, Metanotum and abdomen. C, Lateral
process of female genital lobe. D, Enlarged spicules on border of paddle, upper from Nepal, lower from Thailand.
Abbreviations: Bu = external buttress; CT = cephalothorax, GL = genital lobe, Mr = midrib. Pa = paddle, T
= respiratory trumpet; scale in mm.

628

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Setal branching of Mimomyia chamberlaini pupa.
Abdominal Segments
Cephalo-

Seta thorax I I Ul IV Vv VI Vil Vill
0) — — 1 | l 1 l

| 24 (2)! 13-38(?) 6-10(6) 7-12(10) 6-11(8) 411(7) 48 (8) 4-6 (?) —

2 1-3 (2) 1 1 ] 1 1 1 —

3 24(2) 1 ILC) 1-5 (2) 2-6 (3) 1-3 (2) 1-4 (3) — —

4 1-5 (3) 3-5 (4) 2-6 (3) 1 1-5 (1) 1-3 (1) 1-2 (1) 1-4 (1)
5) 1-5 (4) 1-3 (1) 2-8 (4) 3-6 (4) 4-10 (?) 5-11 (9) 4-10(?) 3-7 (4) —

Gj Il 1-4 (2) 14 (1) 1 1 1251) 2 C1) 1-4 (2) —

7 1 I) (G))) 1EORCL) l PG) 1e27 1) E22 (GL) als) —

8 4-6 (5) 2 (Cy) = 1-3 (1) 1-3 (1) 1-3 (1) 1-3 (1) 1-4 (3) —

9 14(@) — ] 1 1 1 1 1 14 (3)
10 2-4 (4) — — 1 2 (CL) 12) 1 1-3 (1) _—
11 1 (1) == = 1 1 1 1 1,2 (1) —
WA NY) = = = = = — 1 —
14 -— _— = 1 1 1 1 1 1

' Range followed in parenthesis by the mode.

Material examined.—THAILAND, Chon-
buri Province, Bang Lamung, Khao Mai
Keao, X-8-63, 2 2, 3 6, Pe, ex pond on
trackway (Kol, Vuth, Chmnong; Nepal:
Sunsari District, Tarahara XI-25-87, 1 &,
SPP, RFD; Sindhuli Garhi District, Bardi-
agoth, IV-28-90, 2 2, SPP.

Mimomyia (Mimomyia) hybrida
(Leicester)
(Fig. 3)

Pupa.—Position and size of setae as fig-
ured, range and modal number of branches
in Table 3. Abdomen 2.92-—3.47, x 3.27 mm,
paddle 0.78—0.87, « 0.83 mm.

Cephalothorax: Mottled with darker
brown pattern, seta 1,3-CT very long, usu-
ally triple; 5-CT very long, 3-5 br; 6-CT
very long, rather stout, single; 8-CT very
long, with 4-6 br; trumpet length 1.46—1.92,
x 1.64 mm, index 9.0—-13.2, * 11.1; trach-
eoid 0.82—0.1.05, * 0.93 and pinna 0.16—
0.24 x 0.20 of total length, pinna split into
two spinulose processes, adapted for pierc-
ing plant tissue.

Abdomen: Uniformly light tan; seta 1-I
float seta, 11—16 br; stout; 1-II long, rather
stout, with 4—7 br; seta 1-ITI-VI with thin,
long or very long br, 1-[I with 4—9 br, 1-
IV-VI with 2—5 br, 1-VII long, single, sel-

dom double; 2,3-I long, single, 2-I stout; 3-
VII absent; 5-IV-VI with thin, moderately
long or long br, 5-I[V with 3-6 br, 5-V with
4 or 5 br, 5-VI with 2-4 br; 6-I-VI medium
long, single, 6-VI rarely double; 6-VII ven-
tral, short, single or double; 9-VIII short,
usually double, when single, aciculate; fe-
male genital lobe with rounded projections
apicolaterally, with numerous small spicules.

Paddle: ovoid, entirely light brown, bi-
lobate apically, median lobe larger, index
1.64—-1.92, * 1.78 outer margin with coarse
spicules in apical 0.20—0.29, inner margin
with coarse spicules in apical 0.20—0.29,
seta 1-P absent.

Material examined.—THAILAND, Chieng-
mai Province, Chiengmai, IX-30-52, 1 @, 1
3d, LePe, ex pond (M300-A, M-300-1) (M.
Rattanopradith); Nonthburi Province, Pak
Kert, Ko Kret, IV-17-64, 1 92, LePe, ex
ditch (Prajim). Nepal: Sunsari District, Tar-
ahara, IX-X-85, 2 92; Morang District,
Khanar, [X-X-85, 2 2; Sonapur, IX-X-85 2
? (Burgess, unpublished data).

KEY TO THE KNOWN PUPAE OF THE GENUS
MIMOMYIA IN THE ORIENTAL AND
AUSTRALASIAN REGIONS

Partially adapted from Mattingly (1957)

VOLUME 102, NUMBER 3 629

at

ee
————e

seed FEE

i

Fig. 2. Pupa of Mimomyia (Eto.) luzonensis. A, Cephalothorax. B, Metanotum and abdomen. For explana-
tions of abbreviations, see Fig. 1.

Ike Seta 9-VIII very large with about 8 or more with numerous branches; trumpet < 20
br, paddle at least 4X greatest width; seta 1- width at middle (subg. Mimomyia) ...... 7
I small, single or bifid; trumpet length at 2(1). Paddle length at least 10X greatest width;
least 20X width at middle ............ 2, seta 1,5-VII single or bifid; trumpet with

= Seta 9-VIII small, with at most 3 br; paddle tracheoid spiculose (subg. Ingramia) .... 3

less than 3X greatest width; seta 1-I large — Paddle length at most 8X greatest width;

630

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Setal branching of Mimomyia luzonensis pupa.
Abdominal Segments
Cco)h2l> een ee

Seta thorax I Ul i IV Vv VI Vil Vill

0) — oo 1 l 1 ] !

| 1 l e252) 3,4 (4) 3,4 (4) 4,5 (4) 4-8 (5) 3 —

2 1 l 1 ] 1 l os

3 l 1 2h) 1-3 (3) 1-3 (2) — a

4 1-3 (2)! 2-6 (2) l—3 (1) 14 (1) 2,3 (2) E22) 1-3 (1) 1-3 (2) 1-4 (1)

5) 2 ] 2) 2,3 (2) 3 3 3-5 (3) 4,5 (4) —

6 1 2G) I l l 1 1 4-6 (4) a

7/ os (1) 2 (UY) 1,2 (1) 1-3 (2) 1-3 (1) Si) 122) =

8 1 _ -~ Nez) l 1,2 (1) 1,2 (1) 1-3 (2) —

9 1-3 (2) 1 l 1 1 1 l 1 7-10 (10)
10 2 ()) _ — NG) 1,2 (2) 2s) 1-5 (1) 1-3 (1) —
11 1 — — 2 ()) 1 1 1 - =
12 iL? () —_— — — — — — -- --
14 a — — l 1 1 ] 1 1

' Range of branching followed in parenthesis by the mode.

seta 1,5-VII with at least 3 br, tracheoid of
trumpet smooth (subg. Eforleptiomyia) ... 4

. Seta 5-II,III about 0.75 length of seta 3-II-
Ill fusca (Leicester)

— Seta 5-IJ,HI 0.35 or less length of seta 3-
II deguzmanae (Mattingly)

4(2). Inner margin of paddle with small spinules
in basal 0.5.4) ee ase es thee ees rel 5

— Inner margin of paddle with prominent
SPINES a: ool coe GPM Me oe ae dep e ui « 6

. Seta 6-II, UI, VII longer than following ter-

gum bougainvillensis (Belkin)
— Seta 6-II, Il, VII shorter than following ter-

gum elegans (Taylor)
. Setae 2,3-I subequal; seta 8-II present.....
solomonis (Belkin)
— Seta 2-I 2.0 length of 3-I; seta 8-II absent
luzonensis (Ludlow)
. Trumpet with 2 pointed processes apically

eae eater amir ears hares, a ey Mut Mo eee ae 8
— Trumpet without 2 pointed processes at apex
Sebiek Maine seh. Sette ey sek eee coe ere bs Y
8(7). Setae 1,5-IV-VI with some branches longer
than following tergum; paddle emarginate
apicallive. cs ocr Sees hybrida (Leicester)
— Setae 1,5-IV-VI shorter than following ter-
gum; paddle rounded apically ..........
se ace esi Sieh Stussy = gurneyi (Belkin)
9(7). Lateral pale area in basal 0.58—0.74 of pad-

dle, 0.5 of total length chamberlaini Ludlow
— Pale area in distal 0.5 of paddle, 0.3 of total
length or less aurea (Leicester)

COMPARATIVE MORPHOLOGY OF SUBGENERA
OF MIMOMYIA BASED ON PUPAL
CHARACTERS

Based on evidence found in the descrip-
tions of the pupae of 19 species in the sub-

genus Ingramia, five species in the subge-
nus Etorleptiomyia, and 9 in the subgenus
Mimomyia., the latter appears to be the
most primitive. That is because it has seta
8-CT a normal branched seta, seta 1-I a
dendritic float seta, and paddle with indices
of 1.6—2.18, similar to most other mosquito
pupae. On the other hand, the subgenera
Etorleptiomyia and Ingramia have seta 8-
CT stout, almost spike-like in most species,
seta 1-I a single seta, and the paddles linear
in shape with indices of 4.5—20. All sub-
genera have long tracheoid respiratory
trumpets; however, those of Mimomyia
have indices of varying from, 9.0—17.0,
while those of the other two subgenera have
indices of 25—50. Contrarily, seta 9-VIII is
greatly reduced in the subgenus Mimomyia,
with 2—5 branches, while it is fully devel-
oped in the other two with 8—22 branches.
All in all, pupal evidence supports the ev-
olutility of Etorleptiomyia and Ingramia at
a later time than Mimomyia. These obser-
vations also consider the work of Grjebine
(1986) on the Ficalbiini of Madagascar.

ACKNOWLEDGMENTS

The author is indebted to the National Geo-
graphic Society for partial support of this
study; to T. Gaffigan, Walter Reed Biosys-

VOLUME 102, NUMBER 3 631

Bole re

Fig. 3. Pupa of Mimomyia (Mim.) hybrida. A, Cephalothorax. B, Metanotum and abdomen. For explanation
of abbreviations, see Fig. 1.

632

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 3. Setal branching of Mimomyia hybrida pupa.
Abdominal Segments
Cephalo-

Seta thorax I II Il IV Vv VI Vil Vill
0) = l 1 1 l l

1-3 (3)! 11-16 (?) 4-7 (5) 4-9 (5) 2-5 (3) 2-5 (4) 2-4 (4) 2 (A) --
2 Z ] 1 1 1 1,2 (1) —
3) 14 (3) | 152, (Lb) 1-3 (1) DS(Z) 1—3 (2) — —
4 1 (3) 2-4 (3) 1-4 (2) 3,4 (3) 2 2 2 (2) 2 (CU) 1-3 (1)
5 3-6 (5) 1 2-356) Le (QD) 3-6 (?) 4,5 (4) 2-4 (3) 12 (2) —
6 1 i ] 1 1 2a) 2) —
7 1-3 @) 1 It 127 Gls) 2, (ly) 14 (1) 1 1
8 4-6 (5) - — 2 {) 2 (Cy) i174 (Ab) 12 (Ul) 2b) ooo
5 1-3 (1) 1 1 1 1 1 1 1 1 25@)
10 2,3 (2) — — 1 1 125) 1 1 —
11 il22 (ID) = — 1 1 l 1 1 -=
12 il (U) _ — — —- — -—- — —
14 _ — — 1 1 1 1 1 1

' Range followed in parenthesis by the mode.

tematics Unit, Smithsonian Institution, for
the loan of specimens, to B. Bower-Dennis
for the illustrations, and to Drs. J. Rey and
C. Lord for reviewing the manuscript. This
is Florida Agricultural Experiment Station
Journal Series No. RO7078.

LITERATURE CITED

Baisas, E E. 1974. The mosquito fauna of Subic Bay
Naval Reservation, Republic of the Philippines.
U.S. Navy, Headquarters First Medical Service
Wing Technical Report 72-2, 1—170.

Barraud, P. J. 1934. Family Culicidae. Tribes Megar-
hinini and Culicini. The fauna of British India,
including Ceylon and Burma. Diptera. Vol. V, 463
Pp-

Belkin, J. N. 1962. The mosquitoes of the South Pa-
cific (Diptera, Culicidae). I, Il, University of Cal-
ifornia Press, 608 pp., 412 figs.

Chen, H. and C. Lien. 1956. Mosquitoes of genus Fi-
calbia in Taiwan (Formosa), China. Journal of the
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Darsie, R. F, Jr. 1998. Descriptions of the pupae of six
species of Armigeres Theobald, subgenus Leices-
teria Theobald (Diptera: Culicidae) from Nepal.
Proceedings of the Entomological Society of
Washington 100: 234-246.

Darsie, R. FE, Jr. and S. P. Pradhan. 1990. The mos-
quitoes of Nepal their identification, distribution
and biology. Mosquito Systematics 22: 69-130.

Darsie, R. F, Jr., S. P. Pradhan, and R. G. Vaidya. 1992.
Notes on the mosquitoes of Nepal I. New species
records from 1991 collections. Mosquito System-
atics 24: 23-28.

Delfinado, M. D. 1966. The culicine mosquitoes of the
Philippines, tribe Culicini (Diptera, Culicidae).
Memoirs of the American Entomological Institute
7: 1-252.

Grjebine, A. 1986. Insectes Diptéres Culicidae Culi-
cinae Ficalbiini. Muséum National d’ Histoire Na-
turelle, Paris, Faune de Madagascar No. 68, 1—
441.

Harbach, R. E. and K. L. Knight. 1980. Taxonomists’
glossary of mosquito anatomy. Marlton, Plexus
Publishing, Inc. 413 pp.

Knight, K. L. 1978. Supplement to a catalog of the
mosquitoes of the world. Thomas Say Foundation,
Supplement to Volume VI.

Knight, K. L. and R. W. Chamberlain. 1948. A new
nomenclature for the chaetotaxy of the mosquito
pupa, based on a comparative study of the genera
(Diptera: Culicidae). Proceedings of the Helmin-
thological Society of Washington 15: 1-18.

Knight, K. L. and A. Stone. 1977. A catalog of the
mosquitoes of the World. Thomas Say Foundation
6: 1-611.

Lee, D. J.. M. M. Hicks, M. L. Debenham, M. Grif-
fiths, J. H. Bryan, and E. N. Marks. 1988. The
culicidae of the Australasian Region. Vol. 10, En-
tomological Monograph 2, Canberra.

Mattingly, P. E 1957. The culicine mosquitoes of the
Indomalayan Area Part I. Genus Ficalbia Theo-
bald. British Museum of Natural History, pp. 1—
6l.

. 1971. Contributions to the mosquito fauna of
Southeast Asia. XII. Illustrated keys to the genera
of mosquitoes (Diptera, Culicidae). Contributions
of the American Entomological Institute 7(4): 1—
84.

VOLUME 102, NUMBER 3

Pradhan, S. P. and R. E Darsie, Jr. 1989. New mosquito
records for Nepal. Journal of the American Mos-
quito Control Association 5; 21—24.

White, G. B. 1974. Priority of Ingramia Edwards,

633

1912, over Ravenalites Doucet, 1957, as the name
for a subgenus of Mimomyia Theobald, 1903
(Diptera: Culicidae). Mosquito Systematics 6:
239-242.

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 634-642

A STUDY OF THE GENUS FALCOSYNTRETUS TOBIAS FROM THE NEW
WORLD WITH FIVE NEW SPECIES AND A KEY TO KNOWN SPECIES
(HYMENOPTERA: BRACONIDAE: EUPHORINAE)

JENO PAPP AND ScoTT R. SHAW

(JP) Department of Zoology, Hungarian Natural History Museum, pf. 137, H-1431,
Budapest HUNGARY; (SRS) Scott R. Shaw, Insect Museum, % Department of Renew-
able Resources, University of Wyoming, Laramie, WY 82071-3354, U.S.A. (e-mail:

braconid@uwyo.edu)

Abstract.—Five new species of Falcosyntretus Tobias from North and Central America
are described and illustrated: F. complanatus, F. falcoi, F. fallax, F. muesebecki, and F.
transversus. A key for the six known New World species of Falcosyntretus is provided.

Key Words:

The genus Falcosyntretus Tobias was
erected based on one Asian species, F. fal-
cifer (Tobias 1965). Although a species
now assigned to this genus, F. venustus,
was described by Muesebeck (1936), the
presence of this genus in the New World
was not recognized until that species was
reclassified by Shaw (1985). Shaw (1985)
also established the monophyly of Falco-
syntretus on the basis of three synapomor-
phies: the propodeum being mostly smooth
and polished, the petiolate first metasomal
tergum being smooth and polished, and the
curved ovipositor which is as long or longer
than the first metasomal tergum. Shaw
(1985) defined the tribe Syntretini to in-
clude Falcosyntretus and five other genera
sharing several synapomorphies including
cleft tarsal claws, forewing vein M+CU ab-
sent, and the petiolate first metasomal seg-
ment being fused ventrally. Tobias (1986)
indicated that the European species Syntre-
tus xanthocephalus Marshall should be
transferred to Falcosyntretus. Most recent-
ly, Papp (1992) described F. elabsus from
Korea.

As far as known, all members of the tribe

Falcosyntretus, Euphorinae, parasitoids

Syntretini have distinctive cleft tarsal claws
(Figs. 13-16) and are koinobiont endopar-
asitoids of adult Hymenoptera, including
bees and ichneumonids (Shenefelt 1969,
Shaw 1988). This peculiar life style may
partly account for their rarity in collections,
because few attempts are made to rear par-
asitoids from hosts such as adult bumble
bees or adult ichneumonids. Specimens can
be identified as Falcosyntretus using the
key provided by Shaw (1997). A generic
diagnosis was provided by Shaw (1985).
Morphological terminology follows that of
Sharkey and Wharton (1997). Types are de-
posited at the University of Wyoming, Lar-
amie (UWL); the Hungarian Natural His-
tory Museum, Budapest (HNHM); the Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C.
(USNM); the Canadian National Collection,
Ottawa (CNC); and the Zoological Muse-
um, Lund (ZML).

The root of the generic name, falco,
means ‘‘hawk’’ in Latin and refers to the
falcate (curved) ovipositor of the type spe-
cies. However, this character can be mis-
leading for generic recognition because it is

VOLUME 102, NUMBER 3

clear from examination of females of sev-
eral species that the ovipositor is very flex-
ible, and its appearance varies depending on
its position at death. It may appear curved
or straight, or somewhat shorter or longer
depending on its position or degree or ex-
sertion at death. A more useful character for
generic recognition is the mostly smooth
propodeum that lacks a carinate areola
(Figs. 9-12).

KEY TO FEMALES OF THE NEw WorLD
SPECIES OF FALCOSYNTRETUS

l. First metasomal tergite distinctly narrowest
medially, giving the appearance of a con-
striction near the spiracles (Fig. 1); body
size usually smaller than 3mm; antenna
short, with 17—23 antennomeres ........ D,

- First metasomal tergite about as broad me-

dially as basally, or just barely narrower, but

not appearing contricted at middle (Figs. 2—

4); body size usually larger than 3mm, an-

tenna longer, with more than 23 antenno-

meres

Occipital carina effaced dorsally (Fig. 5);

vein cu-a of hind wing absent; vertex of

head with a broad dark brown to black band

that meets compound eye margins com-

Plete lyme we asks res eo Seortes seat marr ee Oe

Spas Falcosyntretus transversus, new species

- Occipital carina complete dorsally (as in
Fig. 6); vein cu-a of hind wing present; ver-
tex of head sometimes with dark markings
around ocelli, but never meeting margins of
compound eyes
a bear's Falcosyntretus muesebecki, new species

3(1). First metasomal tergite distinctly broader
posteriorly, surface entirely smooth (Figs.
2—4); propodeum entirely smooth (Figs. 9—

10) or with scattered, extremely faint rugae
or pitting postero-medially (Fig. 11) ..... 4

- First metasomal tergite about evenly broad
over its entire length, not greatly broader
posteriorly than anteriorly or medially, sur-
face smooth but with two small longitudi-
nally striate areas latero-medially (Fig. 4);
propodeum with median line distinctly ru-
gose (Fig. 12)
Peery os Falcosyntretus falcoi, new species

4(3). Posterior margin of median ocellus about
even with anterior margins of lateral ocelli
(Fig. 7); propodeum entirely smooth..... 5

~ Posterior margin of median ocellus distinct-
ly anterior to anterior margins of lateral
ocelli (Figs. 6, 8); propodeum variable,
sometimes smooth but often with faint ir-

2(1)

635

regular rugae or pitting postero-medially
(Fig. 11) .. Falcosyntretus fallax, new species
5(4). Mesoscutum entirely black; propodeum
with a complete semi-circular carina bor-
dering the junction with the metasoma (Fig.
9) .. Falcosyntretus complanatus, new species
— Mesoscutum yellowish orange, with some
black markings; propodeum with a distinct
break medially in the carina bordering the
junction with the metasoma (Fig. 10) ......
ete Falcosyntretus venustus (Muesebeck)

Falcosyntretus complanatus Papp and
Shaw, new species
(Figs. 2, 9)

Description of holotype female.—Body
length 4 mm. Head in dorsal view 1.6X
broader than long, eye somewhat protrud-
ing and longer than temple, temple rounded
close behind eye. Ocelli large and forming
a low triangle, distance between median
and lateral ocelli shorter than greatest di-
ameter of an ocellus, posterior border of
median ocellus even with anterior border of
lateral ocelli. Antenna just shorter than
body and with 30 antennomeres, first fla-
gellomere 1.4 as long as second flagel-
lomere, further flagellomeres proximo-dis-
tally 2.0—2.2X as long as broad. Mesosoma
in lateral view 1.4X as long as high. Pter-
ostigma 3.5X as long as wide. Hind wing
vein cu-a present basally. Hind femur 5.3 X
as long as broad. Hind basitarsus as long as
tarsomeres 2—3 combined. Metasoma some-
what longer than head and mesosoma com-
bined. Proximal 0.5 of petiole flattened, its
hind width 1.4 greater than basal width,
petiole itself as long as tarsomeres 1—2
combined. Tergites long, second tergite
slightly wider posteriorly than long medi-
ally, third tergite quadrate or as long as
wide behind. Ovipositor sheath as long as
hind tarsomeres 1—2 combined.

Color: Body color mostly golden yellow,
darker dorsally. Antenna, longitudinal
streak of vertex brownish black. Mesosoma
dorsally (mesonotum, scutellum, metano-
tum and propodeum) and petiole black. Ter-
gites dark brown to black. Legs mostly yel-
low, coxae and trochanters whitish, hind fe-
mur apically and tarsomeres dark brown.

636 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-4. First metasomal tergites of Falcosyntretus species, dorsal view. 1, F. muesebecki, 300X. 2, F.
complanatus, 210. 3, F. venustus, 220. 4, F. falcoi, 200X.

VOLUME 102, NUMBER 3

Figs. 5-8.
carina. 6, F. fallax, dorsal view, 200.7, F. complanatus, lateral view, 220. 8, F. muesebecki, dorsal view, 200.

Wings subhyaline, pterostigma and vena-
tion dark brown.

Variation, paratype females.—Body 4.0—
4.2 mm long. Head in dorsal view 1.48—
1.54 broader than long. Antenna with 25—
31 antennomeres. First flagellomere 1.3—
1.4X< as long as second flagellomere. Pter-
ostigma 3.4—3.8X as long as wide. Hind fe-
mur 4.1—5.0 as long as broad. Hind femur
apically and tarsomeres dark brown to
black.

Material examined.—Holotype ?: COS-
TA RICA: Puntarenas, San Vito, Estac.
Biol. Las Alturas, 1,500 m, ex. Malaise,
January 1992 (P. Hanson) (UWL). Para-
types: same data as holotype, 1 2 (UWL);
same data except December 1991, 1 &
(UWL); same data except June 1992, 2 ¢
(UWL); same data except March 1992, 1 @
(UWL); same data except May 1992, 1 @
(UWL); San José, Cerro de la Muerte, 26
km. N. San Isidro, ex. Malaise, 2,100 m,

637

Heads of Falcosyntretus species. 5, F. transversus, dorso-lateral view, 285, note effaced occipital

February—May 1991 (P. Hanson), 1
(HNHM).

Comments.—The form of the petiolate
first metasomal tergum is distinctive in this
species; it is broad and flat through its basal
half, and is barely or not at all constricted
at the middle (Fig. 2). Falcosyntretus com-
planatus is most similar to F. venustus
(Muesebeck) but can be distinguished by
the presence of a complete semi-circular ca-
rina bordering the junction of the metasoma
to the propodeum (Fig. 9) and by the me-
soscutum being entirely black.

Etymology.—The species name com-
planatus (= flattened) refers to the dorsally
flattened petiole.

Falcosyntretus falcoi Papp and Shaw,
new species
(Figs. 4, 12-13)

Description of holotype female.—Body
length 4 mm. Head in dorsal view 1.6

638

broader than long, eye somewhat protrud-
ing and longer than temple, temple rounded
close behind eye. Ocelli large and forming
a low triangle, distance between median
and lateral ocelli shorter than greatest di-
ameter of an ocellus, posterior border of
median ocellus even with anterior border of
lateral ocelli. Antenna just shorter than
body and with 30 antennomeres, first fla-
gellomere 1.4X as long as second flagel-
lomere, further flagellomeres proximo-dis-
tally 2.0—2.2 as long as broad. Mesosoma
in lateral view 1.4 as long as high. Pter-
ostigma 3.5X as long as wide. Hind wing
vein cu-a present basally. Hind femur 5.3 X
as long as broad. Hind basitarsus as long as
tarsomeres 2—3 combined. Propodeum
mostly smooth except median line distincty
rugulose. Metasoma somewhat longer than
head and mesosoma combined. broad and
flat through its entire length, parallel-sided
with no median constriction, and entirely
smooth except for two small longitudinally
striate areas latero-medially. Hind width of
petiole barely broader than anterior width.
Entire petiole as long as tarsomeres 1—2
combined. Tergites long, second tergite
slightly wider posteriorly than long medi-
ally, third tergite quadrate or as long as
wide behind. Ovipositor sheath as long as
hind tarsomeres 1—2 combined.

Color: Body color mostly golden yellow,
darker dorsally. Antenna, longitudinal
streak of vertex brownish black. Mesosoma
dorsally (mesonotum, scutellum, metano-
tum and propodeum) and petiole black. Ter-
gites dark brown to black. Legs mostly yel-
low, coxae and trochanters whitish, hind fe-
mur apically and tarsomeres dark brown to
black. Wings subhyaline, pterostigma and
venation dark brown.

Variation, paratype female.—Essentially
as in holotype female.

Material examined.—Holotype 2: MEX-
ICO: Chiapas, San Cristobal de las Casas,
7,000 ft., 7 June 1969, Malaise trap, (CNC).
Paratypes: UNITED STATES: 1 @, North
Carolina, Swain Co., Smokemont, 16 July
1977, Malaise trap, North Carolina Depart-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ment of Agriculture (USNM). HONDU-
RAS: 1 @, Olancho, Parque Nacional La
Muralla, 15 km N La Union, 15.07N,
86.45W, April 1995, R. Cave, Malaise trap
in high elevation rain forest, (ZML).

Comments.—The form of the petiolate
first metasomal tergum is very distinctive
in this species: it is broad and flat through
its entire length, parallel-sided with no me-
dian constriction, and entirely smooth ex-
cept for two small longitudinally striate ar-
eas latero-medially (Fig. 4). It is most sim-
ilar to F. complanatus but can be distin-
guished by the mentioned petiolar
characters, and also differs by having a dis-
tinctly rugulose median area on the propo-
deum (Fig. 12). The wings of this species
were illustrated in Marsh et al. (1987, fig.
288).

Etymology.—This species is named in
honor of Guido Falco, the owner of Pen-
guin’s restaurant in Santo Domingo de He-
redia, Costa Rica.

Falcosyntretus fallax Papp and Shaw,
new species
(Figs. 6, 11, 14)

Description of holotype female.—Body 3
mm long. Head in dorsal view 1.7 broader
than long, eye somewhat shorter than tem-
ple, temple not rounded close behind eye.
Ocelli forming a high triangle, distance be-
tween median and lateral ocelli as long as
greatest diameter of an ocellus, posterior
border of median ocellus distinctly anterad
of anterior border of lateral ocelli. Antenna
as long as body and with 27 antennomeres,
first flagellomere 1.3X as long as second
flagellomere, flagellomeres proximo-distal-
ly 1.6—2.0X as long as broad. Mesosoma in
lateral view 1.4X as long as high. Fore
wing as long as body. Pterostigma 3.6X as
long as wide. Hind wing vein cu-a present
basally. Hind femur 4.2 as long as broad.
Hind basitarsus as long as tarsomeres 2—4
combined. Metasoma as long as head and
mesosoma combined. Petiole 1.5 broader
posteriorly than basally, petiole itself as
long as middle tarsomeres 1—2 and half of

VOLUME 102, NUMBER 3

Figs. 9-12.
230X. 11, F. fallax, 110X. 12, F. falcoi, 200.

tarsomere 3. Tergites 2—3 transverse, sec-
ond tergite 1.4 and third tergite 2.6 as
wide posteriorly as long medially. Ovipos-
itor sheath as long as hind basitarsus and
half of second tarsomere combined.

Color: Ground color of body mostly yel-
low. Wide longitudinal streak of vertex
brownish black. Mesosoma above (meson-
otum, scutellum, metanotum and propo-
deum) and petiole black. Tergites brown.
Legs yellow, except coxae and trochanters
whitish. Wings subhyaline, pterostigma
brownish opaque.

Variation, paratype females.—Similar to
the holotype except body length 2.6—3.1
mm, usually 2.0—3.0 mm, long; head in
dorsal view 1.57—-1.67X, usually 1.58—
1.6X, as broad as long; antenna about as
long as body and with 25-28, usually 27—
28, antennomeres; pterostigma 3.3—3.7X,
usually 3.5—3.7X, as long as wide; hind fe-
mur 4.1—5.1 X, usually 4.4—4.7X, as long as

639

oan

s

Propodea of Falcosyntretus species, dorsal views. 9, F. complanatus, 210. 10, F. venustus,

broad medially; scutellum varying from
yellow to brown; tergites 2—7 varying from
brown to nearly black, pterostigma varying
from pale brown to yellowish brown.
Material examined.—Holotype 92:
UNITED STATES: Wyoming, Albany
County, Medicine Bow National Forest, 5
miles SW Lincoln Monument, mixed co-
nifer/aspen forest, taken with Malaise trap,
20-26 August 1990. Paratypes: 1 2, same
data as holotype; 5 2 with same locality
data as holotype except collected (1 each)
on following dates: 23-38 July 1990, 28
July—2 August 1990, 13-20 August 1990,
15-19 July 1991, and 29 July—5 August
1991; 1 2, same data except meadow east
of highway, Malaise trap, 13—20 August
1990; 2 2, same data except mixed forest
near sagebrush, Malaise trap, 28 August
1990; 1 2, same data except meadow near
mixed forest, 20—27 August 1991, swept
and aspirated, S. R. Shaw; 3 2, same data

640

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 13-16. Cleft tarsal claws of Falcosyntretus species. 13, F. falcoi, 1,500X. 14, F. fallax, 800. 15, F.
transversus, 1,500X. 16, F. venustus, 1,500.

except Happy Jack Recreation Area, mixed
forest near sagebrush, Malaise trap, 13—20
August 1990. CANADA: 2 2, Quebec,
Summit King Mountain, 26 June and 4 July
1977, M. Sanborne; 1 2°, New Brunswick,
Kouchibouguac National Park, 9 August
1977, S. J. Miller, code 5800B; 1 3, same
locality except 18 July 1977, G. A. Cald-
erwood, code 5649G. Holotype and 8 par-
atypes from Wyoming are deposited at
UWL, four paratypes from Wyoming at
HNHM, and the remaining (Canadian) par-
atypes at CNC.

Comments.—The new species, F. fallax,
is somewhat similar to F. venustus (Mue-
sebeck), but can be distinguished by the fol-
lowing characters. In F. fallax the posterior
border of the median ocellus is set distinctly
in front of the anterior margins of the lateral
ocelli (Fig. 6), the mesonotum is entirely
black, the propodeum often has some irreg-
ular rugose sculpture posteromedially (Fig.

11), and the junction of the propodeum and
metasoma is bordered by a semicircular ca-
rina. In F. venustus the median ocellus is
barely in front of the lateral ocelli, the me-
sonotum has at least some yellow markings
anteriorly and is often extensively marked
with yellow, the propodeum is entirely
smooth posteromedially, and the carina bor-
dering the propodeal/metasomal junction is
broken medially to form two separate short
carinae (Fig. 10). The wings and propo-
deum of this species were illustrated in
Shaw (1997, figs. 19 and 58).

Etymology.—The species name fallax (=
false) refers to the deceptive features of this
species that might cause it to be confused
with F. venustus.

Falcosyntretus muesebecki Papp and
Shaw, new species
(Figs. 1; 8)
Description of holotype female.—Body
2.7 mm long. Head in dorsal view 1.6X

VOLUME 102, NUMBER 3

broader than long, eye as long as temple,
temple rounded close behind eye. Ocelli
small and forming a high triangle, distance
between median and lateral ocelli equal to
greatest diameter of lateral ocellus, hind
margin of median ocellus distinctly before
anterior borders of lateral pair of ocelli. An-
tenna about as long as body and with 22
antennomeres, first flagellomere 1.2 as
long as second flagellomere, flagellomeres
proximo-distally 1.8—2.2 as long as broad.
Metasoma shorter than head and mesosoma
combined. First metasomal tergum as long
as fore femur. Tergites 2—3 quadrate, a bit
shorter medially than broad behind. Ovi-
positor sheath as long as middle basitarsus
and second tarsomere combined.

Color: Ground color of body yellow, ex-
cept flagellum, ocellar triangle, scutellum,
metanotum, propodeum, and dorsum of me-
tasoma mostly brownish black. Pterostigma
opaque yellowish brown.

Variation, paratype females.—Similar to
holotype except body 2.7—2.8 mm long.
Eye somewhat protruding and a bit longer
than temple. Antenna about one-quarter
shorter than body and with 19 antennom-
eres, first flagellomere slightly longer than
second flagellomere. Pterostigma 2.8 as
long as wide. Hind femur 3.8 as long as
broad medially. Ovipositor sheath some-
times almost as long as hind basitarsus.
Ocellar field brown to black, sometimes lat-
eral pair of spots on mesonotum, metano-
tum and propodeum brownish black. Hind
half of petiole and tergites 4—7 sometimes
dark brown, tergites 2—3 sometimes yellow.

Variation, paratype males.—Similar to
female except body 2.3—3.0 mm long. Head
in dorsal view 1.65—1.7X as broad as long.
Antenna with 17—23 antennomeres. Pter-
ostigma 2.75—2.8X as long as wide. Hind
femur 3.5—3.6X as long as broad. Mesoso-
ma and metasoma brown to brownish black,
pronotum yellow, tergites 2+3 yellow to
brown. First metasomal tergum gradually
wider apically, not constricted medially as
in female.

Material |= examined.—Holotype 2:

641

UNITED STATES: Georgia, Forsyth, 5—10
June 1971, ex. Malaise trap, EF T. Naumann
(CNC). Paratypes: 1 2, same data as ho-
lotype (CNC); 1 female, Michigan, Onton-
ogan County, 18 June 1960, R. and K.
Dreisbach; 1 36, New Mexico, Catron
County, 8 mi. S.E. Luna, 7,500 ft, 9-14
July51979;°Ss and JeuPeck- (EN@) 1) 2
(HNHM), 3 36 (USNM), North Carolina,
Alleghany County, Doughton Park, Malaise
trap operated by North Carolina Depart-
ment of Agriculture, 25 July 1977; 1 °,
CANADA: Newfoundland, South Branch,
July 1973, Malaise trap, Heinrich (CNC).

Comments.—The new species, F. mue-
sebecki is most similar to F. transversus
new species, which also has a small body
length (less than 3 mm) and strongly con-
stricted first metasomal tergum (see Fig. 1).
Falcosyntretus muesebecki can be distin-
guished from F. transversus by the pres-
ence of the occipital carina dorsally, the
presence of hind wing vein cu-a, and less
extensive black markings on the vertex
(black blotch not reaching compound eyes).

Etymology.—This new species is dedi-
cated to the late Carl E W. Muesebeck
(1894-1987), well-known specialist of the
parasitoid wasps and the first reviser of the
euphorine braconids of the Nearctic Region
(see Muesebeck 1936).

Falcosyntretus transversus Papp and
Shaw, new species
(Big2 5,15)

Description of holotype female.—Body
2.2 mm long. Head in dorsal view 1.6X
broader than long, eye somewhat shorter
than temple, temple rounded continuously,
occipital carina effaced. Ocelli small and
forming a high triangle, distance between
median and lateral ocelli longer than great-
est diameter of lateral ocellus, hind border
of median ocellus distinctly anteriad of an-
terior borders of lateral ocelli. Antenna
somewhat shorter than body and with 21
antennomeres, first flagellomere hardly lon-
ger than second flagellomere, flagellomeres
proximo-distally 1.8—2 as long as broad.

642

Mesosoma in lateral view 1.4X as long as
high. Pterostigma 3.3X as long as wide.
Hind wing vein cu-a absent. Hind femur
4.4X as long as broad, hind basitarsus as
long as tarsomeres 2-3 plus half of tarso-
mere 4. Metasoma somewhat longer than
mesosoma. Petiole half as long as hind tib-
ia. Tergites 2—3 transverse, second tergite
1.6—-1.7X and third tergite 1.8—2 as broad
as long medially. Ovipositor sheath also
half as long as hind tibia.

Color: Ground color of body yellow.
Transverse streak on vertex dark brown.
Three spots on mesonotum, scutellum, me-
tanotum, propodeum and tergites brownish
yelow to brown. Legs yellow, coxae and
trochanters whitish yellow. Pterostigma
opaque yellow.

Variation, partype females.—Similar to
the holotype except body 2.1mm _ long.
Head in dorsal view 1.5 as broad as long.
Antennae with 20—21 antennomeres. Pter-
ostigma 3X as long as wide. Hind femur
4.5 as long as broad.

Material examined.—Holotype @&:
UNITED STATES: Wyoming, Albany
County Medicine Bow National Forest, 2
miles N on road No. 705, willow bog, taken
with Malaise trap, 19—23 July 1991 (UWL).
Paratypes: 1 2, same data as holotype ex-
cept 9-15 July 1991 (UWL). CANADA: 1
2, Manitoba, Riding Mountain National
Park, Dead Ox Creek, hardwood forest, 400
m, 28 June 1979, Mason (CNC).

Comments.—The new species, F. trans-
versus, is distinctive by the absence of the
occipital carina and the transverse dark
brown streak on upper part of head meeting
the eyes.

Etymology.—The species name “‘trans-
versus” refers to the transverse brown
streak above on the head.

ACKNOWLEDGMENTS

We thank Dr. David Smith (Systematic
Entomology Laboratory, USDA, Washing-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ton, D.C.) and Dr. John Huber (Canadian
National Collection, Ottawa) for providing
loans of specimens. Ms. Teresa Williams
(Western Research Institute, Laramie) as-
sisted with environmental scanning electron
microscopy. Ms. Norma Hosher assisted
with word-processing.

LITERATURE CITED

Marsh, P. M., S. R. Shaw, and R. A. Wharton. 1987.
An identification manual for the North American
genera of the family Braconidae (Hymenoptera).
Memoirs of the Entomological Society of Wash-
ington, No. 13, 98 pp.

Muesebeck, C. E W. 1936. The genera of parasitic
wasps of the braconid subfamily Euphorinae, with
a review of the Nearctic species. United States
Department of Agriculture Miscellaneous Publi-
cations 241: 1—38.

Papp, J. 1992. Braconidae (Hymenoptera) from Korea,
XIV. Acta Zoologica Hungarica 38(1—2): 63-73.

Sharkey, M. J. and R. A. Wharton, 1997. Morphology
and terminology, pp. 19-37. Wharton, R. A., P.
M. Marsh, and M. J. Sharkey, eds., Manual of the
New World Genera of the Family Braconidae
(Hymenoptera), Special Publication of the Inter-
national Society of Hymenopterists, Number 1,
Washington, D.C., 439 pp.

Shaw, S. R. 1985. A phylogenetic study of the subfam-
ilies Meteorinae and Euphorinae (Hymenoptera:
Braconidae). Entomography 3: 277-370.

. 1988. Euphorine phylogeny: the evolution of

diversity in host utilization by parasitoid wasps

(Hymenoptera: Braconidae). Ecological Entomol-

ogy 13:323-335.

. 1997. Subfamily Euphorinae, pp. 234—254. In
Wharton, R. A., PR. M. Marsh, and M. J. Sharkey,
eds., Manual of the New World Genera of the
Family Braconidae (Hymenoptera), Special Pub-
lication of the International Society of Hymen-
opterists, Number 1, Washington, D.C., 439 pp.

Shenefelt, R. D. 1969. Hymenopterorum Catalogus,
Braconidae 1, Euphorinae. W. Junk, The Hague.
176 pp.

Tobias, V. I. 1965. Generic groupings and evolution of
parasitic Hymenoptera of the subfamily Euphori-
nae (Hymenoptera: Braconidae), I. Entomological
Obozrrenie. 44: 841—865 (in Russian). [Transla-
tion in: Entomological Review Washington 44:
494-508. ]

. 1986. Subfamily Euphorinae, pp. 181—250.Jn

Medvedev, G. S., ed., Identification Keys for In-

sects of the European Part of the U.S.S.R., Vol.

III. Hymenoptera. Part 4. Nauka Publishing

House, Leningrad (in Russian).

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 643-651

PREY SELECTION AND DIURNAL ACTIVITY OF
HOLCOCEPHALA OCULATA (F.) (DIPTERA: ASILIDAE) IN
COSTA RICA

Louis M. LAPIERRE

Department of Organismic Biology, Ecology, and Evolution, University of California,
Los Angeles, CA 90095-1606, U.S.A. (e-mail: louis @ucla.edu)

Abstract.—Holcocephala oculata (F) is an abundant small robber fly in open areas at
La Selva Biological Station, Costa Rica, and was studied during August 1995. The ma-
jority (92%) of prey taken by H. oculata belong to the orders Hymenoptera, Diptera, and
Coleoptera, with the remaining belonging, in order of decreasing frequency, to Hemiptera,
Thysanoptera, Homoptera, Araneida, Strepsiptera, and Psocoptera. A comparison of sticky
trap samples of flying insects and robber fly prey items reveal differences in the proportion
of orders represented. Either H. oculata prefers particular orders of prey, or sticky traps
are inadequate in sampling insect faunas. Cannibalism was not observed for H. oculata.
Flies are more numerous in sunny areas at the beginning and end of the day. Flies are
rarely observed in exposed areas on clear days when incident radiation is high. Areas
shaded from direct incident radiation through the day show little change in fly numbers.
Courtship and copulation, described herein, generally occur after 1200. Holcocephala
oculata perches less than a meter off the ground, on average, with grass species tending
to be the substrate most often chosen. No evidence for invertebrate predation on H. oculata
was found, although several suitable predaceous arthropods co-occur with the robber fly
and at least one, the ponerine ant Ectatomma ruidum Roger, readily attacked robber flies
during feeding trials.

Key Words: Asilidae, Holcocephala oculata, robber fly, prey selection, courtship, diurnal

activity, La Selva Biological Station, Costa Rica

Robber flies are among the larger and
more visible members of the Diptera but are

item externally. In addition, some members
of this group have courtship displays that

poorly understood when it comes to details
of their ecology and behavior. A number of
temperate zone Asilidae have been studied
in detail (Lavigne and Holland 1969; Hes-
penheide and Rubke 1977; Hespenheide
1978, 1989; Weeks and Hespenheide 1985),
but only a few tropical species (Fisher
1983; Shelly 1984a, 1988; Fisher and Hes-
penheide 1992). These flies typically cap-
ture their prey by waiting at a perch, inter-
cepting the prey in mid-flight, and then re-
turning to the perch and consuming the

usually involve characteristic aerial maneu-
vers on the part of the male (Fisher and
Hespenheide 1982, 1992).

Holcocephala oculata (F.) (Dasypogoni-
nae: Danilini; Fig. 1) is the smallest (mean
length 5.20 mm; n = 10) of four species of
Holcocephala that are known to occur at La
Selva Biological Station, Costa Rica (E.
Fisher and H. A. Hespenheide, unpub-
lished). Adults perch on low-growing veg-
etation in open areas. Here I describe the
prey selection and diurnal activity of H.

644

Fig. 1.
perch behavior.

Holcocephala oculata showing typical

oculata and test the hypothesis that net in-
cident radiation affects robber fly densities
more than ambient temperature or time of
day.

MATERIALS AND METHODS

The data were collected during 10 sepa-
rate days between 9-29 August 1995 at the
Organization for Tropical Studies’ La Selva
Biological Station which is located at the
confluence of the Rio Puerto Viejo and Rio
Sarapiqui, Heredia Province, Costa Rica
(McDade et al. 1994).

I chose two 50 m transects along weedy
margins bordering mono- and polyculture
plots within the “‘Huertos”’ project, an on-
going study on the sustainability of soil fer-
tility in reconstructed tropical ecosystems
(J. Ewel, personal communication). One
transect was continually exposed to the sun
from dawn until dusk (“‘sunny”’?) whereas
the other was shaded most of the time

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(‘“‘shady’’). In addition to the transects,
neighboring weedy margins within the
Huertos project were used for additional be-
havioral and ecological observations on H.
oculata. The weedy margins are cut at three
to six month intervals and maintained at a
height of less than three meters, and include
several species of grasses, the most com-
mon being Paspalum conjugatum Berg.,
Digitaria sp., and Cynodon dactylon (L.)
Pers. In addition, sedges (Cyperus spp.),
and a diverse assemblage of plant species
adapted to disturbed areas are common
within the margins, as are fallen stems and
leaves from adjacent tree plots.

At least five additional asilid species
(three Holcocephala, one Atractia, one
Mallophora) also co-occur with H. oculata
in the weedy margins, and many species of
predaceous arthropods may be found there
as well (personal observation). The biology
of one other species (H. affinis (Bellardi))
has been studied at La Selva and is gener-
ally similar to that of H. oculata (H. A.
Hespenheide, unpublished).

To determine the diversity of prey taken
by H. oculata, | captured feeding robber
flies in the sunny transect during the 10
days and collected their prey items. Prey
were collected by placing a plastic vial over
a robber fly disturbing it so that it released
the prey item into the vial, after which the
robber fly was released. Preliminary cap-
tures revealed no detrimental effects due to
this method of removing prey from the rob-
ber flies; released individuals remained in
the transect and were often seen with a new
prey item moments later. This method of
prey removal has been used by others (see
Hespenheide and Rubke 1977; Hespenheide
1978, 1989; Shelley 1984a, 1988). I record-
ed the time at which the prey item was col-
lected in order to observe any temporal pat-
tern in prey choice. Prey were not sampled
from robber flies in the shady transect.

To gather information on the diversity of
available prey items, I placed two sticky
traps (21.5 X 28 cm transparencies coated
with Tanglefoot®) on poles 1 m above the

VOLUME 102, NUMBER 3

ground in the sunny transect during study
days 8—10 while the robber flies were being
sampled. Only arthropods measuring up to
4.3 mm (the maximum prey size collected
from H. oculata) were considered in the
analysis of the sticky traps. I identified prey
and sticky trap samples to order. To deter-
mine if H. oculata exhibits a preference for
certain prey over others or is behaving op-
portunistically, I performed two analyses to
compare the actual robber fly prey with the
available prey gathered from sticky traps.
In one analysis I compared a subsample of
the actual prey collected during the days the
sticky trap samples were being employed
(days 8—10). In a second analysis I included
the entire actual prey sample collected dur-
ing the 10 sampling days.

To determine the daily cycle of H. ocu-
lata, 1 censused flies on all 10 days dividing
each into five two-hour periods beginning
at 0800 h and ending at 1600 h. I conducted
a census at the beginning of each of the five
periods. Flies in the shady transect were
simply counted while those in the sunny
transect were counted and assigned to one
of four activities: Feeding, copulating,
courting, or other. The activity “‘other”’ in-
cluded flies that were perched but not feed-
ing and those that were flying between
perches. During the intervening time peri-
ods, I kept out of the transects to minimize
my disturbing the flies.

Prior observations on the daily cycle of
H. oculata indicated that individuals avoid-
ed areas exposed to direct sunlight during
the hottest times of the day, especially on
clear days. I gathered data on ambient tem-
perature and incident radiation to determine
possible influences on the daily cycle of H.
oculata. My hypothesis was that net radia-
tion was more important than temperature
or time of day in determining robber fly
density in a given area; the null hypothesis
was that there was no difference among the
three independent variables in determining
robber fly density. Net radiation was mea-
sured with a net radiometer monitored by
the Huertos project. Ambient temperature

645

was measured at a single fixed location in
the shade between the two transects.

To describe the courtship behavior of H.
oculata, I surveyed the transects during the
study periods and observed courting pairs.
I recorded the time and noted whether the
outcome of the courting attempt was suc-
cessful or unsuccessful in initiating copu-
lation.

To determine whether or not H. oculata
exhibits a preference for type or height of
perch, I walked the weedy margins in and
around the transects and recorded the
choice of perch substrate, location on sub-
strate, and height of perch above ground for
all H. oculata encountered. The location
was defined as precisely where on the sub-
strate the fly chose to perch. Above ground
height referred to the shortest distance be-
tween the fly’s position on the substrate and
the ground.

For all analyses, data on mean values are
evaluated with the non-parametric Wilcox-
on sign rank test, and contingency tables
employing the G-test or Fisher’s exact test
are used to compare percentile data.

The arthropod predators most often ob-
served in the transects foraging on sub-
strates used by H. oculata were Ectatomma
ruidum Roger, an aggressive ponerine ant,
and orb-weaving spiders (Araneidae). I pre-
formed feeding trials to determine the pal-
atability of this fly to E. ruidum with robber
flies disabled by having one wing removed.
Orb-weavers construct webs in the weedy
margins at the level of H. oculata’s perch
heights. Because prey items accumulate in
these webs, information on the spider’s prey
choice is readily available. I carefully re-
moved the silk from entrapped prey to es-
tablish whether H. oculata was among
those preyed upon by the spiders. Other ar-
thropod predators observed included jump-
ing spiders (Salticidae), which have been
observed to take robber flies (H. A. Hes-
penheide personal communication), and ti-
ger beetles (Cicindelidae). However, I ob-
served so few individuals (n = 4 and n =

646

2, respectively) that I am unable to evaluate
their potential as robber fly predators.

RESULTS AND DISCUSSION

I collected nine orders of arthropod prey
from H. oculata (Table 1). Among these the
Hymenoptera, Diptera, and Coleoptera to-
gether account for 92% of the total prey
sample. The most frequent prey at the study
site were ant reproductives (28.5%). The
average prey size was 1.48 mm (n = 379)
with a range of 0.4 mm (nematocerous Dip-
tera) to 4.3 mm (Coleoptera). Hymenoptera,
Diptera, and Coleoptera were also found to
dominate prey samples of Holcocephala
fusca Bromley and H. abdominalis (Say) in
the eastern United States (Dennis 1979,
Scarbrough 1982). However, H. calva
(Loew), also from the eastern United States
and while showing a similar preference for
Hymenoptera and Diptera, differs from oth-
er Holcocephala studied to date by prefer-
ing Hemiptera, Homoptera, and Psocoptera
over Coleoptera (Scarbrough 1982). The
predator to prey size ratio for H. oculata is
3.51 (mean robber fly length 5.20 mm).
This is identical to that determined for H.
fusca (3.5, mean robber fly length 5.9 mm;
Dennis 1979), and intermediate between H.
abdominalis and H. calva (3.4 and 3.7, re-
spectively, mean robber fly lengths 5.81
mm and 7.25 mm, respectively; Scarbrough
1982). Compared to other asilid genera, the
predator to prey size ratio for H. oculata is
close to that reported for the similarly sized
robber fly Nannocyrtopogon neoculatus
Wilcox and Martin (3.66, mean robber fly
length 6.62 mm) in the southwestern United
States (Hespenheide1978), but differs from
that reported for the much larger Sticho-
pogon trifasciatus (Say) (2.23, mean robber
fly length 13.00 mm), also from the south-
western United States (Weeks and Hespen-
heide 1985). No incidence of cannibalism
by H. oculata was observed.

Factors that contribute to prey choice for
particular asilid species likely include the
relative difficulty in handling prey as well
as the availability of prey in space and time

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(Lavigne and Holland 1969, Hespenheide
and Rubke 1977). For H. oculata, compar-
isons of available prey from sticky traps
with actual prey collected during the same
period (days 8—10) and actual prey from the
remaining days (days 1—7) show dissimilar
proportions with respect to some prey or-
ders (Fig. 2). For days 8-10, Hymenoptera,
Diptera, and Coleoptera are the predomi-
nant orders of actual prey of H. oculata as
well as of sticky trap samples at 90% and
83%, respectively. In general, there is no
significant difference between the two sub-
samples of actual prey. This could indicate
that H. oculata is consistent in prey choice,
or that the relative availability of prey
changed little over the course of the study.
The higher proportion of formicid Hyme-
noptera among prey could be the result of
robber flies preferring this group. The high-
er proportion of non-nematocerous Diptera
in the sticky trap samples could be due to
robber flies not favoring this group, perhaps
because they are generally too difficult to
capture. These results are in contrast to
those found by Shelly (1984a) for Atractia
marginata Osten Sacken in Panama where
non-nematocerous Diptera composed a
lower proportion of prey relative to what
was available from sticky traps, and Hy-
menoptera were insufficiently represented
in either sample. Alternatively, the differ-
ences observed between robber fly prey and
sticky trap samples may be an artifact of
the limited ability of sticky traps to ade-
quately sample them, and of sticky traps
themselves being more attractive to some
taxa than others (Shelly 1984b, 1988; Men-
sah 1996).

Size comparisons among and between
actual and available prey also reveal signif-
icant differences (Table 1). In general, there
is no difference (P > .8; Wilcoxon rank
sum test) between the average size prey tak-
en by H. oculata and the average size prey
available to it. Differences emerge, how-
ever, when comparisons are made between
individual prey orders. These results indi-
cate that H. oculata prefers significantly

VOLUME 102, NUMBER 3 647

0.3

[| Actual prey (days 8-10)

at 0.25 WM Available prey (days 8-10)
: FF] Actual prey (days 1-7)
=) 02
2
jor
0.15 7
=
£
=
SOx a
e a
= b
0.05
b aba
Thy Hom

S
~)
n

©
Nn

Proportion of H. oculata

0.25

0800 1000 1200 1400 1600

Time (hours)

Figs. 2-3. 2, Proportions of actual and potential prey of H. oculata: available prey from sticky traps (days
8-10; closed bars), actual prey from H. oculata (days 8-10; open bars), and remaining actual prey (days 1-7;
cross-hatched bars). Relationships among taxa that are significantly different at P < .05 (G-test) are represented
by different letters. Prey taxa are defined as follows: Hym-F ants; Hym-NE non-ant Hymenoptera; Dip-N,
nematocerous Diptera; Dip-NN, non-nematocerous Diptera; Col, Coleoptera; Hem, Hemiptera; Thy, Thysanop-
tera; Hom, Homoptera. 3, Proportion of H. oculata involved in various activities at each census period. Values
for each census period are summed over the 10 study days. CP = copulating, CT = courting, F = feeding, O
= other (i.e., perched without prey or flying).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

648

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VOLUME 102, NUMBER 3

smaller non-nematocerous Diptera prey
than the average size available to it, and
significantly larger Coleoptera. H. oculata
chooses significantly larger formicid Hy-
menoptera than the overall average size of
prey captured, and smaller non-formicid
Hymenoptera and Araneida. This apparent
size preference has also been observed in
other robber flies (Hespenheide and Rubke
1977, Hespenheide 1978, Shelly 1984a)

The proportion of flies involved in the
four activities (copulation, courtship, feed-
ing, or other) at different times of the day
is shown in Fig. 3. Copulation and court-
ship tend to peak in mid- to late afternoon
and the proportion of flies feeding generally
increases through the day. A linear regres-
sion shows a strong inverse relationship be-
tween fly dens'ties and net radiation (Fig.
4; P < .000’ r*? = .54) and a weaker in-
verse relauunship between fly densities and
ambient nperature (P < .0001; r° = .34),
althougn u.ese factors are correlated (P <
.0OO1; r2 = .67).

Differences in robber fly behavior be-
tween the shady and sunny transects further
support the hypothesis that net radiation is
the primary influence. The cycle of average
daily robber fly density in the sunny and
shady transects is shown in Fig. 5. During
the study period, robber flies in the shady
transect exhibited no significant change in
average hourly density throughout the day
Whereas those in the sunny transect de-
creased significantly at midday compared to
dawn or dusk. By 1400 h direct sunlight
had entered the shady transect due to the
sun’s position at that hour, and the affect of
this is represented in Fig. 5 as the period in
the shady transect during which the fewest
flies were counted. Flies in the shady tran-
sect tended to congregate in shady patches
when sunlight entered the area, or relocate
out of the transect. In the sunny transect,
the few flies that chose to remain when net
radiation was highest were found perched
under grass blades, positioned in such a
way so that the blade blocked the sun’s
rays. These observations indicate that the

649

H. oculata density

0 200 400 600

Net radiation (watts/m2)

800

Mean H. oculata density

0800

1000 1200 1400 1600

Time (hrs.)

Figs. 4—5S. 4, Linear relationship between H. oculata
density and net radiation: as net radiation increases fly
density decreases (P < .0001; r°? = .54). 5, The daily
cycle of H. oculata in the sunny transect (open circles)
and shady transect (closed circles). Data are shown as
the mean density, + SE, for each of five time periods
for 10 separate days. Different letters represent signif-
icant differences among times within transects at P <
.O5 (Wilcoxon sign rank test); letters a—c refer to com-
parisons within the sunny transect, and letters d and e
to comparisons within the shady transect.

flies may be responding primarily to inci-
dent radiation. Robber flies in general are
believed to relocate to more shaded perches
presumably to maintain a more “normal”

650

body temperature (Lavigne and Dennis
1973).

Fourteen pairs of H. oculata involved in
courtship were observed in detail during
this study with five resulting in the male
joining the female in copulo. Males locate
females by slowly cruising through the hab-
itat close to the ground, pausing briefly to
inspect perch sites for females. When a
perched female is located, the male hovers
several centimeters behind and slightly be-
low the perched female and begins to fly
upwards and make physical contact with
her. Each time the male makes contact, the
female’s wings are pushed apart; possibly
to provide him with an exposed abdomen
to hold on to for leverage in initiating cop-
ulation. These collisions happen, on aver-
age, at about seven second intervals be-
tween which the male resumes the hovering
position, rubbing his tarsae together prior to
the next attempt. Once in copulo the pair
remains attached, tail to tail with the female
perched and the male dangling upside
down, for an average of 11 minutes (n =
5). The female will often relocate to another
perch, towing the male behind her. The nine
unsuccessful courtship attempts ended with
the female, apparently uninterested in
courting the male, flying far enough away
such that he lost track of her. The copula-
tory position I describe here for H. oculata
is similar to that of H. fusca in North Amer-
ica (Dennis 1979).

Among the 80 H. oculata observed dur-
ing the study of perch preference, most
chose grasses (90%) over woody branch
tips, herbs, and sedges in and around the
study area, with a mean + SE perch height
of 45.5 + 1.7 cm. The grass species and
their order of perch preference included P.
conjugatum (39%), Digitaria sp. (24%), C.
dactylon (18%), and undetermined blades
(19%). Among the major parts of an indi-
vidual grass, 60% of robber flies preferred
the tips of spikes, 28% the main part of the
blade, 9% the tip of the blade, and 3% the
stem of the spike. This clear preference for
grasses, and for P. conjugatum in particular,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

may be an artifact of their relative abun-
dance being higher in these margins, which
appears to be the case. Alternatively, H.
oculata exhibits a species-specific perch be-
havior (see Fig. 1) like many asilids (Fisher
and Hespenheide 1982, 1992), and grasses
may present the optimal substrate for this.

Ectatomma ruidum seized the robber flies
immediately in all 10 of the feeding trials.
I conclude from this that, given the oppor-
tunity under non-experimental conditions,
the ant would prey upon H. oculata.

Of 62 prey items removed from 18 orb-
weaver webs, 47 were ant reproductives,
nine were flies, and six were beetles. Inter-
estingly, these orders also dominated the
prey sampled from Micrathena schreibersi
(Perty) webs in Panama (see Shelly 1984b).
None of the orb-weaver’s prey in my study
were robber flies. The absence of robber
flies as orb-weaver prey might be a result
of robber flies cruising through the habitat
at a slow enough rate as to evade becoming
ensnared. I observed one case in which a
robber fly collided with a web and imme-
diately flew around it. Non-nematocerous
Diptera are strong fliers and as a group have
been found to be disproportionately repre-
sented in webs of M. schreibersi (Shelly
1984b), and this may likely be due to their
ability to both avoid and escape becoming
caught. This may explain the absence of H.
oculata in orb-weaver webs during my
study.

ACKNOWLEDGMENTS

I am grateful to Seth Bigalow, Ankila
Hiremath, and John J. Ewel for allowing me
to work in the study site of the Huertos pro-
ject (NSF grant DEB-9318403) and for pro-
viding the data on net radiation. I am also
grateful to the principal investigators and
staff of the Arthropods of La Selva project
(John T. Longino, Robert K. Colwell, Dan-
ilo Brenes Madrigal, Ronald Vargas Castro,
Maylin Paniagua Guerrero, and Nelci
Oconitrillo Miranda; NSF grant DEB-
9706976) for allowing me to use the facil-
ities contained there. I thank Grant L. Gen-

VOLUME 102, NUMBER 3

try for first introducing me to H. oculata,
and Henry A. Hespenheide and A. G. Scar-
brough for reviewing the manuscript and
offering valuable comments. Eric Fisher
provided the final determination for H. ocu-
lata. Funding support during this project
was supplied by the Vice-Chancellor for
Research at UCLA, Kumar Patel, and the
Organization for Tropical Studies (OTS).

LITERATURE CITED

Dennis, D. S. 1979. Ethology of Holcocephala fusca
in Virginia (Diptera: Asilidae). Proceedings of the
Entomological Society of Washington 81(3): 366—
378.

Fisher, E. M. 1983. Pilica formidolosa (mosca asesina,
robber fly), pp. 755-758. In Janzen, D. H., ed.,
Costa Rican Natural History. The University of
Chicago Press, Chicago, 816 pp.

Fisher, E. M. and H. A. Hespenheide. 1982. Taxonomy
and ethology of a new Central American species
of robber fly in the genus Glaphyropyga (Diptera:
Asilidae). Proceedings of the Entomological So-
ciety of Washington 84(4): 716-725.

1992. Taxonomy and biology of Central
American robber flies with an illustrated key to
genera (Diptera: Asilidae), pp. 611—632. /n Quin-
tero, D. and A. Aiello, eds., Insects of Panama and
Mesoamerica: Selected Studies. Oxford Universi-
ty Press, New York, New York, 692 pp.

Hespenheide, H. A. 1978. Prey, predatory and court-
ship behavior of Nannocrytopogon neoculatus
Wilcox and Martin (Diptera: Asilidae). Journal of
the Kansas Entomological Society 51(3): 449—
456.

. 1989. Notes on the biology of Efferia wilcoxi

(Bromley) (Diptera: Asilidae). The Pan-Pacific

Entomologist 65(4): 375-380.

651

Hespenheide, H. A. and M. A. Rubke. 1977. Prey,
predatory behavior, and the daily cycle of Holo-
pogon wilcoxi Martin (Diptera: Asilidae). Pan-Pa-
cific Entomologist 53(4): 277-285.

Lavigne, R. J. and E R. Holland. 1969. Comparative
behavior of eleven species of Wyoming robber
flies (Diptera: Asilidae). University of Wyoming
Agricultural Experimental Station Science Mono-
graph 18: 1-61.

Lavigne, R. J. and D. S. Dennis. 1975. Ethology of
Efferia frewingi with a description of the eggs
(Diptera: Asilidae). Annals of the Entomological
Society of America 68: 992-996.

McDade, L. A., K. S. Bawa, H. A. Hespenheide and
G. S. Hartshorn, eds. 1994. La Selva: Ecology and
Natural History of a Neotropical Rain Forest. The
University of Chicago Press, Chicago, 486 pp.

Mensah, R. K. 1996. Evaluation of coloured sticky
traps for monitoring populations of Austroasca
vividigrisea (Paoli) (Hemiptera: Cicadellidae) on
cotton farms. Australian Journal of Entomology
35(4): 349-353.

Scarbrough, A. G. 1982. Coexistence in two species
of Holcocephala (Diptera: Asilidae) in a Mary-
land habitat: Predatory behavior. Proceedings of
the Entomological Society of Washington 84(2):
349-365.

Shelly, T. E. 1984a. Prey selection by Atractia mar-
ginata. Proceedings of the Entomological Society
of Washington 86(1): 120-126.

. 1984b. Prey selection by the neotropical spi-

der Micrathena schreibersi with notes on web-site

tenacity. Proceedings of the Entomological Soci-

ety of Washington 86(3): 493-502.

. 1988. Relative abundance of day-flying in-
sects in treefall gaps vs. shaded understory in a
neotropical forest. Biotropica 20(2): 114-119.

Weeks, L. and H. A. Hespenheide. 1985. Predatory
and mating behavior of Stichopogon (Diptera: As-
ilidae) in Arizona. Pan-Pacific Entomologist
61(2): 95-104.

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 652-677

A REVISION OF THE SHORE-FLY GENUS LEMNAPHILA CRESSON
(DIPTERA: EPHYDRIDAE)

WAYNE N. MATHIS AND JAMES E EDMISTON

(WNM) Department of Entomology, National Museum of Natural History, Smithsonian
Institution, Washington, DC 20560-0169, U.S.A. (e-mail: mathis.wayne@nmnh.si.edu);
(JFE) Director, Institute for Franciscan Environmental Studies, Quincy University, Quincy,
IL 62301, U.S.A. (e-mail: edmisja@quincy.edu)

Abstract.—Species of the shore-fly genus Lemnaphila Cresson are revised, including
the description of two new species: L. grossoae (Argentina. Tucuman: San Miguel de
Tucuman) and L. longicera (Peru. Loreto: Iquitos). Lemnaphila is known thus far only
from the New World, where there are now five species. All species with known biologies
are miners in various species of Lemnaceae. Although Lemnaphila is clearly a monophy-
letic clade based on several synapomorphies, its relationship to other genera of the tribe
Hydrelliini is unresolved.

Key Words: Revision, Diptera, Ephydridae, shore flies, Lemnaphila, L. longicera, L. gros-

soae, New World

Specimens of Lemnaphila Cresson are
among the smallest shore flies, with adult
flies often being less than one mm in length.
Their small size corresponds well with, and
is apparently an adaptation to, the mining
behavior of the larval instars in the tiny
thalli of the duckweed family Lemnaceae,
especially the genus Lemna L. Their small
size evidently has been a deterrent to their
study, as specimens are seldom collected
and are poorly represented in collections,
and very few studies that concern the genus
have been published. Duckweed, moreover,
is often an overlooked component of aquat-
ic environments, and studies pertaining to
the rearing and identification of phytopha-
gous insects associated with species of
Lemnaceae are uncommon (Scotland 1940,
Buckingham 1989). Our experience in the
field, however, indicates that specimens of
Lemnaphila are frequently numerous when
the duckweed habitat is adequately sam-

pled. Indeed, recent collecting has revealed
a fifth species from Peru, a sixth species
from Argentina, and also major extensions
in the distributions of the other species. Ad-
ditional studies on their natural history, in-
cluding fresh larval and puparial material of
L. scotlandae Cresson, were conducted in
Ohio. Describing and documenting these
discoveries are the primary objectives for
this revision. Further information on the
natural history of L. scotlandae will be pub-
lished separately (Edmiston and Foote, in
preparation).

Cresson (1933) first described Lemna-
phila as a monotypic genus, being known
only from L. scotlandae Cresson. Miss
Minnie B. Scotland, after whom this spe-
cies was named, discovered the rather mi-
nute adults of L. scotlandae on Lemna that
was growing in a fish hatchery in Ithaca,
New York. Miss Scotland conducted studies
on this species and published three papers

VOLUME 102, NUMBER 3

(1934, 1939, 1940) on its natural history.
The genus remained monotypic until Lizar-
ralde de Grosso (1977, 1978) described
three additional species, all based on spec-
imens collected in the neotropics. In the
first paper, Lizarralde de Grosso (1977) de-
scribed two of the new species: L. neotro-
pica (Argentina (La Plata), Panama, and Ja-
maica) and L. wirthi (Ecuador). Lizarralde
de Grosso (1978) described the third spe-
cies, L. lilloana, from specimens collected
in Argentina (Tucuman). In the second pa-
per, she also described the immature stages
of L. lilloana and L. neotropica and report-
ed the occurrence of L. wirthi in Argentina
(Tucuman). Lizarralde de Grosso (1980)
published a third paper in which the larvae
of Lemnaphila are distinguished from other
shore flies.

Other than catalog entries (Wirth 1965,
Mathis and Zatwarnicki 1995), new state
records (Deonier and Regensburg 1978,
Steinly et al. 1987), and records of parasa-
toids (Marsh 1979, Muesebeck 1979), few
published records on Lemnaphila are avail-
able. Johannsen (1935) included Lemnaphi-
la in his generic key to the immature stages
of shore flies, as did Courtney et al. (1996)
and Merritt et al. (1996). The latter two pa-
pers also summarized known information
on the biology of the larvae. Wirth and
Stone (1956) and Wirth et al. (1987) in-
cluded Lemnaphila in their keys to the
shore-fly genera of the Nearctic Region. In
a synopsis of shore-fly biology, Foote
(1995) also reviewed the biology of Lem-
naphila.

The descriptive terminology, with the ex-
ceptions noted in Mathis (1986) and Mathis
and Zatwarnicki (1990a), follows that pub-
lished in the Manual of Nearctic Diptera
(McAlpine 1981). Because specimens are
small, usually less than 3.5 mm in length,
study and illustration of the male terminalia
required use of a compound microscope.
Although we followed the terminology for
most structures of the male terminalia that
other workers in Ephydridae have used (see
references in Mathis 1986; Mathis and Za-

653

twarnicki 1990a, 1990b), Zatwarnicki
(1996) now uses alternative terms (medan-
drium, transandrium) that are based on the
‘“‘hinge”’ hypothesis for the origin of the er-
emoneuran hypopygium. The terminology
for structures of the male terminalia is pro-
vided directly on Figs. 4—6 and is not re-
peated for comparable illustrations of other
species.

Dissections of male genitalia were per-
formed using the method of Clausen and
Cook (1971) and Grimaldi (1987). External
morphology and internal genitalic features
were drawn using a drawing tube on a Wild
M20EB compound and a MS dissecting mi-
croscope. The species descriptions are com-
posite and are not based solely on the ho-
lotypes. Two head and two venational ratios
used in the descriptions are defined below
(all ratios are based on measurements from
a minimum of five specimens). All mea-
surements were made from pinned speci-
mens.

Eye ratio is the eye width divided by the
eye height.

Gena-to-eye ratio is the genal height
measured at the maximum eye height di-
vided by the eye height.

Costal vein ratio is the straight line dis-
tance between the apices of R,,; and R,,./
distance between the apices of R, and R,,;.

M vein ratio is the straight line distance
along vein M between crossveins (dm-cu
and r-m)/distance apicad of dm-cu.

The phylogenetic analysis was performed
with the assistance of Hennig86©, a com-
puterized algorithm that produces clado-
grams by parsimony. Before analysis, char-
acter data were arranged in transformation
series and then polarized, primarily using
outgroup procedures.

Rearing.—All field and laboratory work
was carried out in northeastern Ohio near
the city of Kent in Portage County. Labo-
ratory rearings, providing life cycle data,
were carried out in an environmental cham-
ber programmed to give a photoperiod of
16L:8D and a temperature variation be-
tween 22-—32° C. Adults were collected

654

from a marsh near the Kent State University
stadium and placed in plastic tumblers with
approximately 100 ml of pond water. Lem-
na thalli (approximately 25) were placed in
each tumbler. The tumbler was covered
with cheese cloth held in place by a rubber
band.

Immature descriptions.—Specimens col-
lected from the field or reared in the labo-
ratory were preserved in 70% ethanol or
double mounted and dried. External and in-
ternal morphologic structures were drawn
using an ocular grid on a Zeiss compound
microscope. Continuous characters were
measured using a calibrated ocular micro-
meter attached to a Zeiss compound micro-
scope. Descriptive terminology of immature
stages follows McAlpine (1981). The ten-
toropharyngeal sclerite length and width are
the maximum straight line distances in lat-
eral view from the anterior tip to the dorsal
cornu posterior tip (A-PDC), from the an-
terior tip to the ventral cornu posterior tip
(A-VDC), and from the dorsal cornu pos-
terior tip to the ventral cornu posterior tip
(PDC-VDC).

Fecundity.—Females collected from the
field were isolated in individual tumblers.
The number of eggs laid on Lemna thalli
were counted on each of four days. Each
day, the female was removed from the tum-
bler and placed in a fresh preparation with
25 new Lemna thalli.

Life History.—Eggs deposited on the
Lemna thalli were placed individually in 20
ml plastic vials. Eggs were observed each
day for hatching, and individual flies were
observed each day for molting between in-
stars, pupariation, and adult emergence.

Specimens.—Although most specimens
for this study are in the National Museum
of Natural History, Smithsonian Institution,
Washington, D.C. (USNM), we also studied
numerous specimens that are deposited in
the American Museum of Natural History
(AMNH), New York; the Academy of Nat-
ural Sciences of Philadelphia (ANSP),
Pennsylvania; The Natural History Muse-
um (BMNH), London, England; Canadian

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

National Collection (CNC), Ottawa, Ontar-
io; Cornell University (CU), Ithaca, New
York; Guelph University (GUE), Guelph,
Ontario; personal collection of James FE Ed-
miston (JEPC); Instituto Miguel Lillo
(IML), Tucuman, Argentina; Ohio State
University (OHSU), Columbus; Universi-
dad de La Plata, La Plata, Argentina
(UNLP) and Washington State University
(WSU).

SYSTEMATICS

We follow Deonier’s (1995) tribal char-
acterization of Hydrelliini and the generic
composition of the tribe that Mathis and Za-
twarnicki (1995) published. Deonier (1995)
published a key to the three genera, includ-
ing Cavatorella, which he described in the
same paper. Deonier’s concept of Lemna-
phila, as reflected in the key, was based pri-
marily on L scotlandae, the type species,
but with the addition of other species, the
characterization of the genus has been ad-
justed as indicated in the following revised
key to genera.

KkyY TO GENERA OF HYDRELLIINI
ROBINEAU-DESVOIDY

(modified from Deonier 1995)

1. Posterior notopleural seta inserted near dorsal
angle; prescutellar acrostichal seta lacking;
ocellar setae lacking; anepisternum bearing a
single seta along posterior margin; lacking an
intrapostalar setula ....... Lemnaphila Cresson

— Posterior notopleural seta inserted at same lev-

el as anterior seta near ventral margin; pres-
cutellar acrostichal seta present; ocellar setae
usually present; anepisternum usually bearing
1 large seta and 1-2 smaller setae along pos-
terior margin; usually bearing 1 intrapostalar
setula
. Wing apex broadly rounded; vein R,,; long,
making costal section II much longer than sec-
tion III; interfacetal setulae conspicuous and
dense; male lacking ventroclinate processes
from dorsad of fused surstyli.............

Hydrellia Robineau-Desvoidy

— Wing apex narrowly rounded, ellipsoidal; vein
R,,; short, making costal section II subequal
to section III; costa bearing several interfrac-
tural setae; interfacetal setulae inconspicuous,
sparse; male with ventroclinate styliform pro-

Nw

VOLUME 102, NUMBER 3

cesses projected from dorsad of fused surstyli
Peay See hs Sh sO ecient Cavatorella Deonier

Genus Lemnaphila Cresson

Lemnaphila Cresson, 1933:229. Type spe-
cies: Lemnaphila scotlandae Cresson,
1933, monotypy; 1944:175—176 [review,
tribal relationships].-Scotland, 1934:291
[biology, larval miner in Lemna]; 1939:
713-718 [biology, parasatoids]; 1940:
322-324 [review of biology, figs. of egg,
puparium, adults].—Johanssen, 1935:48—
51 [larval key].—Wirth, 1965:745 [Ne-
arctic catalog].—Wirth and Stone, 1956:
465, 469 [key to genus, leaf mining hab-
it]|—Lizarralde de Grosso, 1977:159-—
164 [review]; 1980:55—60 [larval key];
1989:54—55 [review, Argentina fauna].
Wirth et al., 1987:1035, 1041 [fig. of
head, key to genus].—Mathis and Za-
twarnicki, 1995:96—97 [world catalog].—
Courtney et al., 1996:508 [larval key to
genus].—Merritt et al., 1996:545 [habitat,
habits, trophic relationships].

Diagnosis.—Lemnaphila is a genus of
the tribe Hydrelliini, subfamily Hydrelli-
inae, that is similar and closely related to
Hydrellia Robineau-Desvoidy and Cavato-
rella. Lemnaphila, which are minute to
small shore flies (length 0.83—1.50 mm), is
distinguished from these and other shore-fly
genera by the following combination of
characters:

Head: Frons much wider than long,
sometimes with distinct mesofrons and par-
afrons; mesofrons broadly reaching anterior
margin of frons, vestiture variable. Ocellar

and paravertical setae lacking; pseudopos- |

tocellar setae well developed, proclinate
and slightly divergent. Pedicel lacking a
large, dorsoapical seta; arista with 5—9 long
rays along dorsum. Face in lateral view ver-
tically straight or shallowly concave and
with ventral third slightly angled anteriorly;
ventral facial margin at most shallowly
arched; facial seta 1 near ventrolateral mar-
gin, sometimes with a few setulae, gener-
ally weakly developed. Eye comparatively
large, especially in lateral view, bearing nu-

655

merous interfacetal setulae. Gena very
short, gena-to-eye ratio less than 0.15; lack-
ing a prominent genal seta. Clypeus not
generally exposed.

Thorax: Chaetotaxy: dorsocentral setae 2
(0+2 or with anterior seta on transverse su-
ture), posterior dorsocentral seta displaced
laterally (this seta has sometimes been la-
beled as the intra-alar seta); lacking pres-
cutellar acrostichal setae, other acrostichal
setae variable, either lacking or with a few
setulae in 2 anterior rows; presutural supra-
alar seta present but often reduced; lacking
postpronotal, postsutural supra-alar, and in-
trapostalar setae; 1 postalar seta; 2 lateral
scutellar setae (lacking a smaller setula be-
tween 2 larger setae), basal seta shorter than
apical seta; 2 notopleural setae, anterior seta
usually shorter, posterior seta inserted near
dorsal angle of notopleuron; 1 anepisternal
seta along posterior margin; katepisternal
seta variable, lacking, reduced, or well de-
veloped. Wing: generally hyaline or very
faintly infuscate; costal vein long, extended
to vein M; vein R,,, short, with costal sec-
tion II equal to section IJ or much shorter
than section III; lacking greatly elongate in-
terfractural setulae other than apical seta.
Legs lacking dorsoerect setae on dorsal sur-
face.

Abdomen: Dorsum comprising tergites
1-5, with tergites 1-2 somewhat fused;
sternite 1 greatly reduced or lacking, 2—5
present; 5th sternite of male comparatively
well developed and much larger than ster-
nites 2—4, bearing setulae, shape variable
with species. Male terminalia: Epandrium
weakly developed, incomplete dorsally with
a membranous gap between lateral exten-
sions, lateral extensions as a fused lateral
phalange at base of cerci; cerci well devel-
oped, fused with median margin of epan-
drium; surstylus greatly reduced as a small,
usually narrow process from ventral margin
of fused epandrium/cerci; subepandrial
plate well developed, simple, arched; ae-
deagus well developed, sometimes angulate
in lateral view; aedeagal apodeme narrow
to very broadly developed, keel variously

656

developed, usually enlarged; postgonite
well developed, usually bearing setulae
near midlength; pregonite apparently lack-
ing; hypandrium simple, lateral apices ori-
ented posteriorly, shallowly arched or nar-
rowly V-shaped.

Discussion.—As noted in the generic di-
agnosis, we place Lemnaphila in the tribe
Hydrelliini, subfamily Hydrelliinae, where
it is closely related to Hydrellia and Ca-
vatorella. Although this tribal assignment
and the generic associations are now ap-
parent, these relationships were not always
recognized and there was some confusion
that led to a rather circular history. When
Cresson (1933) described Lemnaphila he
placed the genus “. . . in the tribe Hydrel-
liae of the subfamily Notiphilinae, near the
genus Nostima.’’ Cresson based his generic
placement on adult characters (setulose
eyes, antennal shape, and the arrangement
and number of dorsocentral setae). Cresson
(1944), however, then removed WNostima
Coquillett, Philygria Stenhammar, and
Lemnaphila from the tribe Hydrelliini and
created a new tribe, Hydrinini (unavailable,
based on a junior homonym, = Philygriini
Lizarralde de Grosso), for these three gen-
era. Despite removing Lemnaphila from a
close association with Hydrellia, Cresson
(1944:176) continued to recognize that
Lemnaphila has ‘‘. . . the appearance of a
small Hydrellia.”’ The tribe Philygriini was
then characterized primarily by the elevated
insertion of the posterior notopleural seta
and the reduced setation. Wirth (1965) fol-
lowed Cresson’s suprageneric arrangement
for the Tribe Philygriini in his catalog of
North American Ephydridae. Dahl (1959),
in his review of Scandinavian Ephydridae,
removed Nostima and Philygria from the
subfamily Notiphilinae (= Hydrelliinae)
and placed these genera along with Hyadi-
na Haliday in the subfamily ‘“‘Hydrininae”’
(unavailable, based on a junior homonym,
= Ilytheinae Cresson), which Dahl de-
scribed in the same paper. Mathis and Za-
twarnicki (1990b) also recognized Philygri-
ini but with only two genera: Philygria and

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Nostima. They suggested that Lemnaphila
was better placed in the tribe Hydrelliini
(Subfamily Hydrelliinae) near the genus
Hydrellia based especially on characters of
the male terminalia and larvae, and in sub-
sequent publications (Zatwarnicki 1992,
Deonier 1995, and Mathis and Zatwarnicki
1995), Lemnaphila was placed in the Hy-
drelliini, along with Hydrellia and Cavato-
rella.

Cresson (1933, 1944) first noted that
adults of Lemnaphila are similar to Ay-
drellia, and a close relationship between
these two genera is strongly supported by
larval characters as well. Johannsen (1935)
was able to distinguish between larvae of
Lemnaphila and Hydrellia only on the basis
of larval size and the preference Lemna-
phila has for feeding on Lemna. Hydrellia
is generally characterized by its stem- and
leaf-mining habits, and Johannsen observed
that the mouthparts of Lemnaphila and Hy-
drellia were identical except for size. Hen-
nig (1943) also proposed a close relation-
ship between larvae of Lemnaphila and Hy-
drellia. Lizarralde de Grosso (1977, 1978)
described additional Neotropical species of
Lemnaphila, including larval descriptions,
and indicated that the Neotropical larvae of
Hydrellia and Lemnaphila are very similar
and closely related.

Specimens of Lemnaphila are not only
tiny but they are highly derived, in part and
presumably as an adaptation to the mining
habit in Lemnaceae. The structures of the
male terminalia also reflect their derived
condition. The surstylus, for example, is
highly modified as a much reduced struc-
ture at the ventral margin of the fused epan-
drium/cercus. The reduced epandrium that
is fused to the cercus and the enlarged cerci
are also evidence of the derived status.

Key TO SPECIES OF LEMNAPHILA CRESSON

1. Scape, pedicel, and basal half of 1st flagellom-
ere yellow; length of Ist flagellomere twice its
height; face in profile straight; length of outer
vertical seta less than half length of inner ver-
tical seta. Katepisternum whitish yellow, con-
colorous with forecoxa

VOLUME 102, NUMBER 3

L. longicera, new species
— Antenna entirely dark brown; length of Ist fla-
gellomere only slightly longer than its height;
face shallowly concave in lateral view with
ventral third slightly angled forward; length of
outer vertical seta about 3/4 length of inner
vertical seta. Katepisternum brown .,...... 2
. Fronto-orbital setulae greatly reduced, at most
hairlike, none well developed; mesofrons sub-
shiny, distinct from densely microtomentose
parafrons. Scutellum lacking microtomentum
or sparsely microtomentose, not appearing vel-
vety; katepisternal seta reduced, hairlike .....
L. scotlandae Cresson
— At least | fronto-orbital seta well developed,
anterior setula proclinate, posterior seta much
longer, length subequal to outer vertical seta,
lateroclinate, parallel to outer vertical seta; me-
sofrons and parafrons densely microtomentose,
appearing velvety, or with small area immedi-
ately anterior of ocellar triangle bare (L. /il-
loana), shiny. Scutellum moderately to densely
microtomentose, brownish black to black,
sometimes appearing velvety; katepisternal
seta well developed, length subequal to poste-
MOmAnepIStCHnaleScla weer een eka eMenen ens one 3
3. Ventral third of face densely microtomentose,
blackish brown, concolorous with dorsal por-
tion of face; coxae, femora, and tibiae yellow
PCR e ee, tier L. wirthi Lizarralde de Grosso
— Ventral third of face densely microtomentose,
silvery white, distinctly contrasted with black-
ish brown dorsal portion; femora brown, oth-
erwise legs yellow
4. Area immediately laterad and anteriad of ocel-
lar triangle silvery white, concolorous with
fronto-orbit immediately anterior and posterior
of lateroclinate fronto-orbital seta ..........
L. grossoae, new species
— Area immediately laterad and anteriad of ocel-
lar triangle dark brown, similar to coloration of
OcCellaratian leew ccs. HHS ee eects & ele 5
5. Mesofrons and parafrons densely microtomen-
tose, appearing velvety blackish brown; pos-
terior fronto-orbit extensively white to silvery
Witte 5 6 G6 bE L. neotropica Lizarralde de Grosso
— Mesofrons just anterior of ocellar triangle bare,
shiny, blackish brown, contrasted with densely
microtomentose parafrons; posterior fronto-or-
bit with very small area white ............
L. lilloana Lizarralde de Grosso

Nw

Lemnaphila longicera Mathis and
Edmiston, new species
(Figs. 1-3)

Diagnosis.—This species is distinguished
from congeners by the following combina-

657

tion of characters: Scape, pedicel, and basal
half of Ist flagellomere yellow; length of
Ist flagellomere twice height; face in profile
straight; face mostly uniformly microto-
mentose and colored, coloration varies de-
pending on angle of view, except for nar-
row, vertical stripe; length of outer vertical
seta less than half inner vertical seta; scu-
tellum densely microtomentose, appearing
velvety, dark brown; katepisternum whitish
yellow, concolorous with forecoxa; legs en-
tirely yellow, with coxae whitish yellow
and apical half of tibiae slightly brownish
yellow.

Description.—Minute to small shore
flies, length 0.90—1.35 mm.

Head: Frons with distinct, subshiny me-
sofrons; parafrons densely microtomentose,
dark brown, appearing velvety. Fronto-or-
bital setae very weakly developed and/or
lacking, at most with 1-2 hairlike setulae;
length of outer vertical seta less than half
inner vertical seta, outer seta curved later-
ally, inner seta very shallowly curved, near-
ly vertical; ocelli arranged in equilateral tri-
angle, or nearly so. Scape, pedicel, and bas-
al half of Ist flagellomere yellow, apical
half of Ist flagellomere brown; length of Ist
flagellomere twice height, apex bluntly
rounded; arista with 8—9 long, dorsal rays.
Face in profile straight at anterior margin;
varying in color from different angles, from
lateral view mostly yellowish with brown-
ish yellow area at midheight, from anterior
view whitish gray, sometimes slightly dark-
er medially at midheight, and with keel-like
brown vertical stripe. Eye ratio 0.68—0.71.
Gena-to-eye ratio 0.10—0.13. Maxillary pal-
pus yellow; mouthparts mostly yellow to
grayish yellow.

Thorax: Mostly dark brown; scutum dark
brown with moderately dense whitish gray
microtomentum; scutellum with distal %
densely microtomentose, dark brown, ap-
pearing velvety, basal % whitish gray; no-
topleuron, area of scutum immediately pos-
teriad of notopleuron densely microtomen-
tose, dark brown, appearing velvety; anepis-
ternum and anepimeron concolorous, dark

658

Figs. 1-3.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Lemnaphila longicera. 1, Male terminalia (epandrium, cercus, surstylus), lateral view. 2, Internal

male terminalia (aedeagus, aedeagal apodeme, postgonite, hypandrium, and subepandrial plate), ventral view. 3,

Fifth sternite, ventral view.

brown but lighter than notopleuron; katepis-
ternum, meron, and coxae concolorous,
whitish yellow. Chaetotaxy: anterior dorso-
central seta inserted near level of transverse
suture; acrostichal setae lacking; basal scu-
tellar seta about % length of apical seta; ka-
tepisternal seta lacking. Wing: length 1.08—
1.20 mm; width 0.45—0.49 mm; vein R,,;
shallowly sinuous, shallowly curved toward
posterior margin, than shallowly recurved,

bowed anteriorly; costal vein ratio 1.0—1.07,
length of costal section II about equal to sec-
tion III; M vein ratio 0.42—0.47. Halter knob
bright yellow; stem brownish yellow. Legs:
femora whitish yellow to yellow basally, be-
coming more yellow apically; tibiae yellow,
becoming slightly darker apically; foretarsi
brownish yellow to brown, mid- and hind-
tarsus slightly lighter than foretarsus, more
yellowish.

VOLUME 102, NUMBER 3

Abdomen: Dark brown, male Sth tergite
subshiny; 5th sternite of male (Fig. 3) well
developed, narrowly heart shaped, longer
than wide, tapered gradually to narrowly
rounded anterior margin, posterior margin
deeply emarginate. Male terminalia (Figs.
1—2): Epandrium (Fig. 1) reduced to nar-
row, fused, lateral phalange at basolateral
margin of cercus; cercus (Fig. 1) greatly en-
larged, projected posteriorly, bearing 4—5
large setulae at anteroventral margin; su-
bepandrial plate simple, arched, connected
ventrally with postgonite; aedeagus in lat-
eral view (Fig. 1) with moderately broad
base tapered irregularly thereafter to narrow
process, apex narrowly rounded; aedeagal
apodeme with ventral keel comparatively
long and narrow in lateral view (Fig. 1),
slightly asymmetrical; postgonite prominent
with rounded apex, bearing setulae about %4
distance from connection with subepandrial
plate; hypandrium V-shaped in ventral view
(Fig. 2):

Type Material—The holotype male is la-
beled ““PERU. Loreto: Iquitos [,] 16Feb1984
[,] Wayne N. Mathis/sHOLOTYPE Lemna-
phila longicera 6 W.N. Mathis & Edmiston
USNM [red; species name, gender, and au-
thors handwritten].’” The holotype is double
mounted (minuten in a block of plastic), is
in excellent condition, and is deposited in
the USNM. Twenty-two paratypes (8 d, 14
2; USNM) bear the same label data as the
holotype.

Distribution.—WNeotropical: Peru (Lore-
to).

Etymology.—The species epithet, longi-
cera, alludes to the elongate antenna that,
among other characters, distinguishes this
species.

Remarks.—Although this species clear-
ly belongs in Lemnaphila, it is very dis-
tinct structurally (see diagnosis and key)
and apparently represents a lineage that
has undergone considerable anagenetic
evolution. We know nothing about the nat-
ural history of the immature stages but
suggest that they will be found to be min-
ers in Lemnaceae.

659

Lemnaphila grossoae Mathis and
Edmiston, new species

Diagnosis.—This species is distinguished
from congeners by the following combina-
tion of characters: Antenna entirely dark
brown; length of Ist flagellomere only
slightly longer than height; face in lateral
view shallowly concave; ventral third of
face densely microtomentose, silvery white,
distinctly contrasted with blackish brown
dorsal portion; mesofrons densely microto-
mentose, area immediately laterad and an-
teriad of coellar triangle silvery white, oth-
erwise brown to blackish brown; fronto-or-
bit anteriad and posteriad of long laterocli-
nate fronto-orbital seta silvery white,
otherwise velvety blackish brown; scutel-
lum sparsely microtomentose, appearing
subshiny to faintly dull, brown; katepister-
num brown; femora brown; tibiae and most
tarsomeres yellow.

Description.—Small shore flies, length
1.30 mm.

Head: Frons entirely microtomentose, bi-
colored, brown and silvery white; parafrons
and anterior half of fronto-orbit concolo-
rous, densely microtomentose, brownish
black, appearing velvety, area immediately
laterad and anteriad of ocellar triangle sil-
very white; posterior fronto-orbit (from
base of outer vertical seta to base of later-
oclinate fronto-orbital seta) with silvery
white microtomentum, concolorous with
area laterad and anteriad of ocellar triangle.
One fronto-orbital seta well developed,
length subequal to outer vertical seta, ori-
entation postero-obliquely lateroclinate to
lateroclinate; other fronto-orbital setae an-
terior of large seta, at best weakly devel-
oped, hairlike; length of outer vertical seta
about *% that of inner vertical seta, outer
vertical seta distinctly curved laterally, in-
ner vertical seta very shallowly curved,
nearly vertical; ocelli arranged in isosceles
triangle, with distance between posterior
ocelli greater than between either posterior
ocellus and anterior ocellus. Antenna en-
tirely dark brown; length of Ist flagellom-

660

ere only slightly longer than height; arista
with 6—7 long, dorsal rays. Face shallowly
concave in lateral view; ventral third of face
densely microtomentose, silvery white, dis-
tinctly contrasted with blackish brown dor-
sal portion; dorsal portion of face with very
shallow carina, mostly faintly grayish black
except for black area on either side of mid-
line just dorsad of silvery white microto-
mentum. Eye ratio 0.70. Gena-to-eye ratio
0.19. Maxillary palpus blackish brown;
mouthparts generally brownish black.

Thorax: Mostly blackish brown; scutum
subshiny, sparsely microtomentose; scutel-
lum with moderately dense microtomen-
tum, brownish black, somewhat dull, not
velvety appearing like frons except partially
along sides; pleuron mostly blackish brown
to black except most of anepisternum ap-
pearing silvery gray to silvery white. Chae-
totaxy: anterior dorsocentral seta inserted
behind level of transverse suture; acrosti-
chal setulae sparse anteriorly, in 2 rows,
lacking posteriorly; basal scutellar seta
about % length of apical seta; katepisternal
seta well developed, length subequal to pos-
terior anepisternal seta. Wing: length 1.30
mm; width 0.50 mm; vein R,,; shallowly
bowed anteriorly; costal vein ratio 1.25;
length of costal section II about half of sec-
tion III; M vein ratio 0.55. Halter knob
bright yellow; stem blackish brown. Legs:
Femora mostly brown, apices yellowish;
tibiae and most tarsomeres yellow, 5th tar-
somere brown.

Abdomen: Tergites very sparsely micro-
tomentose to bare, blackish brown.

Type Material.—The holotype female is
labeled ““ARGENTINA San Miguel de
TUCUMAN 2-XII-1976 Col. L. de Grosso
[handwritten]/Lemnaphila lillolana det. M.
Lizarralde de Grosso/PARATIPO [yellow]/
COLECCION INST.-FUND M. LILLO
(4000)—S.M. TUCUMAN TUCUMAN—
ARGENTINA [yellow]/HOLOTYPE Lem-
naphila grossoae 2 W.N. Mathis & Edmis-
ton USNM [red; species name, gender, and
authors handwritten].’’ The holotype is
double mounted (glued to a large triangle),

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

is in good condition (glue partially obscur-
ing tarsi), and is deposited in the IML.

Distribution.—WNeotropical: Argentina
(Tucuman).

Etymology.—The species epithet, gros-
soae, is a patronym to honor and recognize
the numerous contributions of Dr. Mercedes
Lizarralde de Grosso to the study of shore
flies, Lemnaphila in particular.

Remarks.—This species was misidenti-
fied as L. lillolana but the pattern of micro-
tomentum and coloration of the head, es-
pecially the frons, differs from that species
as noted in the diagnosis. In addition to the
holotype female, we dissected the male al-
lotype/paratype of L. lillolana and discov-
ered that it is likewise not conspecific with
L. lillolana. Our basis for this conclusion is
comparison of the structures of the male
terminalia with those that Lizarralde de
Grosso (1978:15, figs. 6-13) published.
Thus, this male, which lacks its head, may
be conspecific with this species.

Lemnaphila lillolana Lizarralde de Grosso

Lemnaphila lillolana Lizarralde de Grosso,
1978:14-17 [immature stages; host
plants: Spirodela biperforata Kock, Wol-
fiella lingulata (Hegelm.), Salvinia mini-
ma Baker (Lemnaceae)]; 1989:55 [re-
view, Argentina fauna].-Mathis and Za-
twarnicki, 1995:96 [world catalog].

Diagnosis.—This species is distinguished
from congeners by the following combina-
tion of characters: Antenna entirely dark
brown; length of Ist flagellomere only
slightly longer than height; face in lateral
view shallowly concave; ventral third of
face densely microtomentose, silvery white,
distinctly contrasted with blackish brown
dorsal portion; mesofrons just anterior of
ocellar triangle bare, shiny, blackish brown,
contrasted distinctly with densely microto-
mentose parafrons; posterior fronto-orbit
with very small area white; scutellum
densely microtomentose, appearing velvety,
dark brown; katepisternum brown; femora
brown; tibiae and most tarsomeres yellow.

VOLUME 102, NUMBER 3

Description.—Small shore flies, length
1.40—1.50 mm.

Head: Frons with mesofrons except for
ocellar triangle bare of microtomentum,
shiny, blackish brown; parafrons, and an-
terior half of fronto-orbit concolorous,
densely microtomentose, brownish black,
appearing velvety; posterior fronto-orbit
(from base of outer vertical seta to base of
lateroclinate fronto-orbital seta) with whit-
ish gray microtomentum. One fronto-orbital
seta well developed, length subequal to out-
er vertical seta, orientation postero-oblique-
ly lateroclinate to lateroclinate; other fron-
to-orbital setae anterior of large seta, at best
weakly developed, hairlike; length of outer
vertical seta about % that of inner vertical
seta, outer vertical seta distinctly curved
laterally, inner vertical seta very shallowly
curved, nearly vertical; ocelli arranged in
isosceles triangle, with distance between
posterior ocelli greater than between either
posterior ocellus and anterior ocellus. An-
tenna entirely dark brown; length of 1st fla-
gellomere only slightly longer than height;
arista with 7—8 long, dorsal rays. Face shal-
lowly concave in lateral view; ventral third
of face densely microtomentose, silvery
white, distinctly contrasted with blackish
brown dorsal portion; dorsal portion of face
with very shallow carina, mostly faintly
grayish black except for black area on ei-
ther side of midline just dorsad of silvery
white microtomentum. Eye ratio 0.75—0.77.
Gena-to-eye ratio 0.1 1—0.12. Maxillary pal-
pus blackish brown; mouthparts generally
brownish black.

Thorax: Mostly blackish brown; scutum
subshiny, sparsely microtomentose; scutel-
lum with moderately dense microtomen-
tum, brownish black, dull, not velvety ap-
pearing like frons; pleuron mostly blackish
brown to black except dorsal half of ane-
pisternum from some angles appearing sil-
very gray. Chaetotaxy: anterior dorsocen-
tral seta inserted behind level of transverse
suture; acrostichal setulae sparse anteriorly,
in 2 rows, lacking posteriorly; basal scutel-
lar seta about % length of apical seta; ka-

661

tepisternal seta well developed, length sub-
equal to posterior anepisternal seta. Wing:
length 1.45-1.60 mm; width 0.60—0.74
mm; vein R,,; shallowly bowed anteriorly;
costal vein ratio 1.30—1.33; length of costal
section II about half of section III; M vein
ratio 0.49—0.50. Halter knob bright yellow;
stem blackish brown. Legs: Femora mostly
brown, apices yellowish; tibiae and most
tarsomeres yellow, 5th tarsomere brown.

Abdomen: Dark brown.

Type Material.—The holotype female
(not a male as indicated in the original de-
scription) is labeled “‘ARGENTINA[.] San
Miguel de TUCUMAN[,] 2.XII.1976 [2
Dec 1976] Col. L. de Grosso [handwritten,
black margin]/Lemnaphila Lilloana Lizar-
ralde de Grosso det. Lizarralde de Grosso
[handwritten, black marginJ/HOLOTIPO
[red]J/COLECCION INST. - FUND. M.
LILLO (4000) - S. M. TUCUMAN TUC-
UMAN - ARGENTINA [yellow].”’ The ho-
lotype is double mounted (glued to mod-
erately large paper triangle), is in fair con-
dition (specimen slightly teneral, eyes and
face partially collapsed, right wing with
posterior margin folded back on itself), and
is deposited in the IML.

Other Specimens Examined.—ARGEN-
TINA. Tucumdn: San Miguel de Tucuman,
2 Dec 1976, M. Lizarralde de Grosso (6 9;
paratypes; IML).

Distribution.—Neotropical: Argentina
(Tucuman).

Remarks.—This species is structurally
very similar to L. neotropica and is obvi-
ously closely related. The only difference
that we have discovered in external struc-
tures is the shiny mesofrons with the ex-
ception of the ocellar triangle (L. neotro-
pica has a completely microtomentose, vel-
vety, blackish brown mesofrons). The other
differences that Lizarralde de Grosso noted
(1978:17) are variable and are within the
ranges of either species. No males of this
species were available for description and
illustration.

662

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 4-7.

Lemnaphila neotropica. 4, Male terminalia (epandrium, cercus, surstylus, subepandrial plate,

postgonite), posterior view. 5, Same, lateral view. 6, Internal male terminalia, lateral view. 7, Fifth sternite and

hypandrium, ventral view.

Lemnaphila neotropica Lizarralde de
Grosso
(Figs. 4—7)

Lemnaphila neotropica Lizarralde de Gros-
so, 1977:160; 1978:17-18 [biology, im-
mature stages, host plants: Lemna minima
Philippi, Hydromystia stolonifera,; para-
site: Hexacola sp. (Cynipidae)]; 1989:55
[review, Argentina fauna].—Mathis and
Zatwarnicki, 1995:96—97 [world catalog].

Diagnosis.—This species is distinguished
from congeners by the following combina-
tion of characters: Antenna entirely dark
brown; length of Ist flagellomere only
slightly longer than height; face in lateral
view shallowly concave; ventral third of
face densely microtomentose, silvery white,

distinctly contrasted with blackish brown
dorsal portion; mesofrons and parafrons en-
tirely densely microtomentose, appearing
velvety blackish brown; scutellum densely
microtomentose, appearing velvety, dark
brown; katepisternum brown; femora
brown; tibiae and most tarsomeres yellow.

Description.—Minute to small shore
flies, length 0.90—1.30 mm.

Head: Frons with mesofrons, parafrons,
and anterior half of fronto-orbit concolo-
rous, densely microtomentose, brownish
black, appearing velvety; posterior fronto-
orbit (from base of outer vertical seta to
base of lateroclinate fronto-orbital seta)
with whitish gray microtomentum. One
fronto-orbital seta well developed, length
subequal to outer vertical seta, orientation

VOLUME 102, NUMBER 3

postero-obliquely lateroclinate to laterocli-
nate; other fronto-orbital setae anterior of
large seta, at best weakly developed, hair-
like; length of outer vertical seta about %
that of inner vertical seta, outer vertical seta
distinctly curved laterally, inner vertical
seta very shallowly curved, nearly vertical;
ocelli arranged in isosceles triangle, with
distance between posterior ocelli greater
than between either posterior ocellus and
anterior ocellus. Antenna entirely dark
brown; length of Ist flagellomere only
slightly longer than height; arista with 6—7
long, dorsal rays. Face shallowly concave
in lateral view; ventral third of face densely
microtomentose, silvery white, distinctly
contrasted with blackish brown dorsal por-
tion; dorsal portion of face with very shal-
low carina, mostly faintly grayish black ex-
cept for black area on either side of midline
just dorsad of silvery white microtomen-
tum. Eye ratio 0.75—0.77. Gena-to-eye ratio
0.11—0.12. Maxillary palpus blackish
brown; mouthparts generally brownish
black.

Thorax: Mostly brownish black to black;
scutum subshiny, sparsely microtomentose;
scutellum with moderately dense microto-
mentum, brownish black, dull, not velvety
appearing like frons; pleuron mostly
brownish black to black except dorsal half
of anepisternum from some angles appear-
ing silvery gray. Chaetotaxy: anterior dor-
socentral seta inserted behind level of trans-
verse suture; acrostichal setulae sparse an-
teriorly, in 2 rows, lacking posteriorly; bas-
al scutellar seta about % length of apical
seta; katepisternal seta well developed,
length subequal to posterior anepisternal
seta. Wing: length 0.80—1.30 mm; width
0.45—0.65 mm; vein R,,; shallowly bowed
anteriorly; costal vein ratio 1.60—1.69;
length of costal section II slightly less than
half of section III; M vein ratio 0.46—0.55.
Halter knob bright yellow; stem blackish
brown. Legs: Femora mostly brown, apices
yellowish; tibiae and most tarsomeres yel-
low, 5th tarsomere brown.

Abdomen: Dark brown, male 5th tergite

663

subshiny; 5th sternite of male in ventral
view (Fig. 7) much wider than long, broad-
ly and shallowly U-shaped, bearing numer-
ous setulae along posterior margin, these
also in membrane between sternite and hy-
pandrium. Male terminalia (Figs. 4—5):
Epandrium comparatively well developed
as a wide phalange at base of cerci; cerci
(Figs. 4—5) well developed, irregularly
hemispherical around posteromedial mar-
gin, narrowed at dorsal and ventral angles,
lateral margin nearly straight in lateral
view, shallowly concave in posterior view;
surstylus a short process, only slightly lon-
ger than wide, bearing numerous setulae,
these longer apically; subepandrial plate
bandlike in lateral view (Fig. 6), distinctly
arched in posterior view (Fig. 4); aedeagus
very well developed, angulate in lateral
view, robust, especially basiphallus; aedea-
gal apodeme greatly enlarged, especially
expanded keel, keel slightly asymmetrical
in lateral view, fanlike; postgonite promi-
nent, heavily sclerotized, base in ventral
view rectangular with a narrowed, apical,
arched process extended from ventromedial
portion of base, apical process parallel sid-
ed, apex narrowly rounded; pregonite ap-
parently lacking; hypandrium (Fig. 7) in
ventral view a broad, arched band, posterior
Margin concave, anterior margin convex,
with a short, tonguelike, medial process.

Type Material.—The holotype male is la-
beled “‘[Argentina. Buenas Aires:] LA
PLATA 18/10/71 [18 Oct 1971] COL.
[Loidcono-] DIAZ [handwritten; black sub-
margin|/HOLOTIPO [red]/$/MUSEO DE
LA PLATA LEMNAPHILA NEOTROPI-
CA L. DE GROSSO Det LIZARRALDE
DE GROSSO [species name and author
handwritten; black submargin].”’> The ho-
lotype is double mounted (glued to a large
paper triangle), is in fair condition (abdo-
men missing, some setae misoriented), and
is deposited in the UNLP.

Other Specimens Examined.—ARGEN-
TINA. Buenos Aires: La Plata, 18 Oct 1971,
Loidcono-Diaz (1 6, 1 2; paratypes;
UNLP).

664 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 8-10. Lemnaphila scotlandae. 8, Male terminalia (epandrium, cercus, surstylus, aedeagus, aedeagal
apodeme, subepandrial plate, postgonite), lateral view. 9, Internal male terminalia (surstylus shown on right side
only, postgonite shown on left side only), dorsal view. 10, Fifth sternite, ventral view.

DOMINICAN REPUBLIC. La Vega: Jar-
abacoa (1-2 km S; 19°06.9'N, 70°37'W;
520 m), 14-17 May 1998, D. and W. N.
Mathis (11 6, 4 2; USNM).

GRENADA. St. George: Point Salines
Airport (W end; 12°00.3'N, 61°47.7'W), 12
Sep 719979 WiN- Mathis: (7.3659 bh @s
USNM).

JAMAICA. St. Ann: Runaway Bay
(stream bed), Feb 1969, W. W. Wirth (1 od;
paratype; USNM).

PANAMA. Aguadulce Cocle, 25 Sep
1951, ES. Blanton (1 6; paratype, USNM).

Distribution.—Neotropical: Argentina
(Buenos Aires, Chaco, Tucuman), Panama,
West Indies (Dominican Republic, Grenada,
Jamaica).

Remarks.—This species and L. wirthi are
very similar and obviously closely related.
This relationship is established by the fol-
lowing synapomorphies: 1. Aedeagus large,
prominent, angulate; 2. Sth sternite of male
much wider than long and bearing numer-
ous setulae along posterior, concave mar-
gin.

Lemnaphila scotlandae Cresson
(Figs. 8-21)

Lemnaphila scotlandae Cresson, 1933:229
[USA. New York: Tompkins County, Ith-

aca, Fall Creek; HT 3, ANSP (6501)];
1944:176 [tribal assignment].—Johan-
nsen, 1935:50—51 [immature stages].—
Scotland, 1934:291 [biology, larval min-
er in Lemna]; 1939:713-—717 [host plant:
Lemna minor L. (Lemnaceae); parasit-
oids: Opius lemnaphilae Muesebeck
(Braconidae), Trichopria angustipennis
Muesebeck, 7. paludis Muesebeck (Dia-
priidae)]; 1940:322—324 [immature stag-
es, host plants and parasitoids].—Mue-
sebeck, 1939:58—62 [descriptions of par-
asitoids]; 1979:1146—-1147 [catalog of
parasitoids, Trichopria angustipennis and
T. paludis|.—Wirth, 1965:745 [Nearctic
catalog].—Deonier and Regensburg,
1978:155 [records for Ohio].—Marsh,
1979:209 [catalog of parasitoid, Opius
lemnaphilae|.—Ferrar, 1987:169 [sum-
mary of biology].—Steinly et al., 1987:
166, 168 [records for Illinois, discus-
sion].—Buckingham, 1989:219-221
[parasitoids: Opius lemnaphilae Muese-
beck (Braconidae), Trichopria angusti-
pennis Muesebeck, T. paludis Muesebeck
(Diapriidae); records for Florida; review
of biology].—Mansor and Buckingham,
1989:115-118 [host range].—Foote,
1995:430 [review of biology].—Mathis

VOLUME 102, NUMBER 3

and Zatwarnicki, 1995:97 [world cata-
log].

Diagnosis.—This species is distinguished
from congeners by the following combina-
tion of characters: Antenna entirely dark
brown; length of Ist flagellomere only
slightly longer than height; mesofrons sub-
shiny, distinct from densely microtomento-
se parafrons; face in lateral view shallowly
concave; ventral third of face densely mi-
crotomentose, appearing velvety, dark
brown to golden yellow, depending on an-
gle; scutellum lacking dense microtomen-
tum, gray, not appearing velvety; katepis-
ternum brown; legs brown.

Description.—Minute to very small
shore flies, length 0.83—1.2 mm; yellowish
brown, dark brown, to black with silvery
gray and yellowish silver microtomentum.

Head: Frons generally moderately dense-
ly invested with silvery whitish gray micro-
tomentum; mesofrons from ocellar triangle
anteriad moderately densely silvery whitish
gray microtomentose, subshiny; parafrons
brown; fronto-orbit generally concolorous
with mesofrons or slightly more brownish.
Occiput brown with yellowish silver micro-
tomentum; outer vertical seta lateroclinate;
length of outer vertical seta about %4 inner
vertical seta; inner vertical seta parallel to
slightly convergent; fronto-orbital setae mi-
nute, hairlike; pseudopostocellar setae
prominent, proclinate and somewhat diver-
gent; paravertical setae absent; ocellar setae
absent; postocullar setae absent; ocelli ar-
ranged in isosceles triangle, with distance
between posterior ocelli much greater than
between either posterior ocellus and ante-
rior ocellus. Antenna entirely dark brown;
length of Ist flagellomere only slightly lon-
ger than its height; arista bearing 5—7 rays
along dorsum. Facial background colora-
tion shiny dark brown with sparse silvery
gray microtomentum. Oral margin slightly
protruded dorsally. Face with transverse
crease at about ventral third, shallowly con-
cave in lateral view, ventral third slightly
angled forward, densely microtomentose,

665

dark brown. Parafacial band shiny dark
brown. Eye ratio 0.78—0.83. Gena-to-eye
ratio 0.14—0.17. Gena and postgena brown
with sparse silvery yellow microtomentum;
parafacial setae minute; gena bearing small
black setulae. Maxillary palpus dark brown
with prominent setae; prementum shiny
dark brown, bulbous.

Thorax: Mesonotum brown with silvery
yellow microtomentum; scutellum lacking
microtomentum or microtomentum sparse,
not appearing velvety; scutum length 0.29—
0.43 mm; scutellar length 0.10—0.17 mm;
subscutellum brown with yellowish silver
microtomentum; anepisternum with dorsal
half brown with dense silvery gray micro-
tomentum, ventral half with sparse silvery
and golden microtomentum; katepisternum
brown with sparse silvery golden microto-
mentum; anatergite brown with yellowish
silver microtomentum. Chaetotaxy: anterior
dorsocentral seta inserted near level of
transverse suture; acrostichal setulae lack-
ing; scutellar setae with anterolateral setae
3%44 length of posteromedial setae and me-
diolateral setulae; katepisternal seta weakly
developed, hairlike, much shorter than pos-
terior anepisternal seta. Wing: length 1.02—
1.44 mm; width 0.42—0.61 mm; vein R,,;
shallowly bowed anteriorly; costal vein ra-
tio 1.76—1.85, with length of costal section
II about half costal section III; M vein ratio
0.46—0.55. Halter knob bright yellow; stem
blackish brown. Legs: yellowish brown to
brown; tarsomeres 1—4 yellowish brown,
darker distally; tarsomere 5 brown.

Abdomen: Tergite 3 length, males 0.13-—
0.17 mm, females 0.14—0.20 mm; tergite 4
length, males 0.18—0.20 mm, females 0.18—
0.20 mm. Background coloration brown to
dark brown. Tergites 1—5 covered with
sparse yellow microtomentum; 5th sternite
slightly longer than wide, tapered slightly
anteriorly, bearing numerous setulae. Male
Terminalia (Figs. 8—9): Epandrium much
reduced, as a small, lateral phalange at base
of fused cercus; cercus comparatively great-
ly enlarged, height nearly equal to width of
5th sternite of male; subepandrial plate an

666

Figs. 11-16.
ventral view. 13, Micropylar end of egg, ventral view. 14, Third-instar larva, cephalic end, lateroblique view.
15, Same, antennae. 16, Same, posterior end, ventral view.

arched, narrow, bandlike plate; surstylus
(Fig. 8) a long, narrow, parallel-sided pro-
cess bearing 2 apical setulae; aedeagus
cornucopialike in lateral view (Fig. 8), with
broad base, tapered to narrow apex that is
hooked; aedeagal apodeme with keel com-
paratively long and narrow in lateral view,
essentially symmetrical; subepandrial plate
forming rounded arch, connected ventrally
with postgonite; postgonite (Fig. 8) promi-
nent with a posterior and anterior process;
posterior gonal process prominent with api-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Scanning electron micrographs of Lemnaphila scotlandae. 11, Egg, dorsal view. 12, Same,

cal % parallel-sided, apex rounded, ventral-
ly projected setula about *% distance from
connection with subepandrial plate; anterior
gonal process in lateral view (Fig. 8) a
small, narrow, tapered process about %
length of surstylus, bearing apical setulae;
hypandrium (Fig. 10) narrowly V-shaped.
Type Material.—The holotype male is la-
beled ‘‘d/Fall Creek, Ithaca, N[ew].
Y[ork]./M. B. Scotland VII 25—46 [sic; Jul
25—26],1933/TYPE No. 6501 Lemnaphila
SCOTLANDAE E.T. Cresson, Jr, [red;

VOLUME 102, NUMBER 3

number and species name handwritten].”’
The holotype is double mounted (glued to
a pin in a rectangular block of cork), is in
excellent condition (slightly teneral), and is
deposited in the ANSP (6501).

Other Specimens Examined.—NEARC-
TIC. CANADA. ONTARIO. Guelph, 12 Jan
1976, J. M. Cumming (14 6, 16 2; GUE);
16 Jan 1976, J. M. Cumming (5 2°; GUE);
8 Aug 1978, W. A. Attwater (3 36,5 2;
GUE); Limehouse, 1 Sep 1974, R. E.
Roughley (1 3, 1 2; GUE); 2 Sep 1974,
RES Roughley (1 6,5. 2: CNC, GUE); 6
Oct 1974, R. E. Roughley (on Lemna minor
in Black Creek), (1 2; GUE); Ottawa, 1967,
J. G. Chillcott (2 36, 2 2; CNC); Wainfleet
Bog (8 km S Welland), 14—28 Sep 1987,
A. Stirling (2 6; WSU).

UNITED STATES. FLORIDA. Alachua:
Gainesville, 13—31 Oct 1986 (Lemna val-
diviana: pools at Division of Plant Indus-
try), G. R. Buckingham (4 6, 14 &, 17;
JEPC); Pierce’s Homestead (Malaise trap;
S9-TI1OS-RI8E), 11 May 1974, W. H.
Pierce (1 2; USNM). Broward: Ft. Lauder-
sale (Lemna valdiviana), 14 Nov 1988, G.
R. Buckingham (1 6; USNM). Putnam:
Rodman Reservoir (Lemna minor), 6 Oct
1987, G. R. Buckingham (1 6, 1 Q&;
USNM). Sumter: Lake Okahumpka (Lemna
valdiviana), 22 Oct 1987, G. R. Bucking-
ham (1 2; USNM). MICHIGAN. Clinton:
Bath, 10 Oct 1965, (1 2; USNM). NEW
YORK. Tompkins: Ithaca, Fall Creek, 25—
26 Jul 1933, M. B. Scotland (12 6, 11 &;
allotype; AMNH, ANSP, BMNH, CU,
USNM); Ithaca, Sep 1933, M. B. Scotland
(7 6, 11 2; CU, USNM). OHIO. Portage:
Kent (near Kent State University stadium),
23 May—4 Aug 1986, 1987, J. Edmiston
Mi2ad, 12625 JEPC):Kent.24 Jun=2°Sep
1987 (lab reared), J. Edmiston (9 3, 14 @;
JEPC). Franklin: Columbus (Mirror Lake),
23 Aug 1919, W. C. Kraatz (1 6; OHSU).
Geauga: LaDue Reservoir (41°22.2'N,
80°13.4'W; duckweed), 12 Sep 1976, B. A.
Steinly (32 36, 21 2; USNM).

Distribution.—WNearctic: Canada (ON),
USA (FL, IL, MI, NY, OH).

667

Natural History.—Adult L. scotlandae
are commonly found on or flying over the
surface of floating duckweed thalli. The
adults utilize the Lemna for food, leaving
characteristic elongate gouges on the sur-
face of the thallus. Seasonal distribution
and overwintering stages are unknown;
adults, however, have been collected in late
November, suggesting that adults are able
to survive the cold season like other north-
temperate Ephydridae. Adults collected
from the field and reared in the laboratory
lived between 2—21 days (n = 13).

Females lay eggs on the margin of a
Lemna thallus, usually one egg per thallus.
Females collected from the field and reared
in the laboratory laid up to 41 eggs per day,
averaging 21—26 eggs per day (n = 11) dur-
ing their first four days in the laboratory.
One female laid 334 eggs during 20 days.

Eggs hatch in 4—6 days (x = 4.21; n =
10), after which the first-instar larvae begin
feeding on thallus tissue. Burrowing
through the duckweed, the first-instar lar-
vae feed for 1-3 days (x = 1.92; n = 10)
before molting; second-instar larvae feed
for 1-4 days (x = 2.62; n = 10) before
molting; third-instar larvae feed for 2—7
days (x = 4.85; n = 10) before pupariation
occurs within the hollowed-out duckweed
thallus. The pupal period ranged between
6-8 days (x = 7.15 days; n = 10).

Although the immatures and adults of
this species are very tiny, three hymenop-
terous parasatoids, as noted in the synony-
my, have been reared from puparia (Mue-
sebeck 1939, Scotland 1939, Buckingham
1987).

DESCRIPTION OF IMMATURE STAGES

Egg (Figs. 11—12).—Length 0.35-0.37
mm (x = 0.36; n = 12); width 0.13—0.17
(x = 0.16; n = 12); ovoid, ends bluntly
rounded; micropylar end porous; 4 longi-
tudinal ridges dorsally with parallel rows of
minute pores between ridges; flattened ven-
trally with medioventral rows of minute
pores and laterally with parallel rows of mi-
nute pores. Chorion light brown.

668

Mature third-instar larva (Figs. 20-
21).—Length 1.11—1.34 (x = 1.24; n = 5);
width 0.22—0.32 (x = 0.28; n = 5). ver-
miform, 12 segmented, white. Anterior end
blunt with ventrally projected mandibles,
posterior end with medioventrally projected
spiracles. Integument covered with minute
setulae. Cephalic segment (segment 1,
pseudocephalic segment): antenna 2 seg-
mented with rounded socket, proximal an-
tennal segment minute, circular; distal an-
tennal segment ovoid, greater than 10X
length of proximal segment, lateroventrally
with rounded sensory pits.

Segment 2 (prothoracic segment): ven-
trally and laterally with rows of comblike
setae, projected dorsomedially over cephal-
ic segment. Segment 3: longest segment, 1
¥% length of segment 2 with rows of comb-
like setae. Segments 3—11 similar, rounded,
ventral surface with creeping welts; welts
of segments 3 and 4 small; welts of seg-
ments 5—11 large. Segment 12 with perianal
pad ventrally and posterior spiracles ventro-
medially; perianal pad bilobed, each lobe
ovoid; posterior spiracle with cylindrical
base and conical projection; conical projec-
tion approximately twice as long as cylin-
drical base and tapered to point.

Cephalopharyngeal skeleton (Figs. 17—
19): A-PDC 0.20-0.21 mm (n = 7), A-
VDC 0.18—-0.20 mm (n = 7), PDC-VDC
0.08 mm (n = 7). Mandibles fused dorsally,
anteriorly pointed, posteriorly broadened.
Hypopharyngeal sclerite minute, posterior-
ly rounded, anteriorly slightly indented.
Tentoropharyngeal_ sclerite anteriorly
roundly pointed with anteroventral projec-
tion, dorsal cornu posteriorly sharply point-
ed, ventral cornu posteriorly broadened.
First- and second-instar larvae similar. Dis-
tinguishable by size of tentoropharyngeal
sclerite.

Puparium (Fig. 21).—Length 1.17—1.36
(x = 1.28; n = 13); width 0.53-—0.60 (x =
0.56; n = 13); width broadest posterome-
dially in ventral view, tapering at each end;
anterior end slightly curved ventrally,
broadly rounded with remnants of cephal-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

opharyngeal skeleton clearly visible; pos-
terior end slightly curved ventrally, spiracle
remnants dark, perianal pad remnants visi-
ble; margins of puparium smooth. Color
uniformly pale brown except for black ce-
phalopharyngeal skeleton remnants.
Remarks.—This species is commonly
called the Lemna fly (Scotland 1934, 1939,
1940) or the duckweed miner, and the
adults and immature stages live entirely in
and around duckweed (Lemnaceae). Miss
Minnie B. Scotland (1933), after whom the
species was named, first reared L. scotlan-
dae from duckweed (Lemna minor L.) that
was collected in a fish hatchery at Fall
Creek near Ithaca, New York. The species
is now known to occur from southern Can-
ada to northern Florida and west to Illinois.

Lemnaphila wirthi Lizarralde de Grosso
(Figs. 22—26)

Lemnaphila wirthi Lizarralde de Grosso,
1977:163; 1978:18-19 [review, descrip-
tion of male terminalia]; 1989:55 [re-
view, Argentina fauna].—Mathis and Za-
twarnicki, 1995:97 [world catalog].

Diagnosis.—This species is distinguished
from congeners by the following combina-
tion of characters: Antenna entirely dark
brown; length of Ist flagellomere only
slightly longer than height; face in lateral
view shallowly concave; ventral third of
face densely microtomentose, blackish
brown, concolorous with dorsal portion of
face; scutellum densely microtomentose,
appearing velvety, dark brown; katepister-
num brown; coxae, femora, and tibiae yel-
low.

Description.—Minute to very small
shore flies, length 0.95—1.20 mm.

Head: Frons with mesofrons, parafrons,
and anterior half of fronto-orbit concolo-
rous, densely microtomentose, brownish
black, appearing velvety; posterior fronto-
orbit (from base of outer vertical seta to
base of lateroclinate fronto-orbital seta)
with whitish gray microtomentum. One
fronto-orbital seta well developed, length

VOLUME 102, NUMBER 3 669

19

20

Figs. 17-21. Lemnaphila scotlandae. 17, Cephalopharyngeal skeleton, first-instar larva. 18, Same, second-
instar larve. 19, Same, third-instar larva. 20, Third-instar larva, lateral view. 21, Puparium, ventral view.

subequal to outer vertical seta, orientation like; length of outer vertical seta about 7
postero-obliquely lateroclinate to laterocli- that of inner vertical seta, outer vertical seta
nate; other fronto-orbital setae anterior of distinctly curved laterally, inner vertical
large seta, at best weakly developed, hair- seta very shallowly curved, nearly vertical;

670 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 22-26. Lemnaphila wirthi. 22, Male terminalia (epandrium, cercus, surstylus, aedeagus, aedeagal apo-
deme), posterior view. 23, Same (with addition of postgonite), lateral view. 24, Internal male terminalia, lateral
view. 25, Postgonite and subepandrial plate, ventral lateral view. 26, Fifth sternite and hypandrium, ventral view.

ocelli arranged in isosceles triangle, with
distance between posterior ocelli greater
than between either posterior ocellus and
anterior ocellus. Antenna entirely dark

brown; length of Ist flagellomere only
slightly longer than height; arista with 6-7
long, dorsal rays. Face with shallowly con-
cave in lateral view; ventral third of face

VOLUME 102, NUMBER 3

densely microtomentose, concolorous with
frons; dorsal portion of face with very shal-
low carina, mostly black except shiny black
carina. Eye ratio 0.68—0.74. Gena-to-eye ra-
tio 0.09—0.11. Maxillary palpus blackish
brown; mouthparts generally brownish
black.

Thorax: Mostly brownish black to black;
scutum subshiny, sparsely microtomentose;
scutellum with dense microtomentum,
brownish black, not shiny, but velvety ap-
pearing like frons; pleuron mostly brownish
black to black except dorsal half of anepis-
ternum from some angles appearing silvery
gray. Chaetotaxy: anterior dorsocentral seta
inserted behind level of transverse suture;
acrostichal setulae sparse anteriorly, in 2
rows, lacking posteriorly; basal scutellar
seta about % length of apical seta; katepis-
ternal seta well developed, length subequal
to posterior anepisternal seta. Wing: length
1.08—1.24 mm; width 0.45—0.57 mm; vein
R,,; mostly straight to very shallowly
bowed anteriorly; costal vein ratio 1.5—
1.73; length of costal section II about half
of section III; M vein ratio 0.44-0.49. Hal-
ter knob bright yellow; stem brownish yel-
low. Legs: yellow except for brown 5th tar-
someres.

Abdomen: Dark brown, male 5th tergite
subshiny; 5th sternite of male (Fig. 26)
broadly and shallowly U-shaped, thickly
formed, bearing numerous setulae along
posterior margin and on membrane imme-
diately adjacent. Male terminalia (Figs. 22—
24): Epandrium (Figs. 22—23) comparative-
ly well developed as a phalange at ventro-
basal angle of cercus; cerci very well de-
veloped, in lateral view (Fig. 23) irregularly
hemispherical around posteromedial mar-
gin, basolateral margin even, nearly straight
in lateral view, shallowly concave in pos-
terior view; surstylus (Figs. 22—23) long,
slender, mostly parallel sided, apical % well
sclerotized, bearing several setulae along
length, apex rounded; subepandrial plate
(Figs. 24—25) evenly arched, slender, band-
like in lateral view (Fig. 24); aedeagus an-
gulate in lateral view (Fig. 24), basiphallus

671

in lateral view somewhat quadrate with dis-
tiphallus extended from ventral portion of
basiphallus, distiphallus long, slender, shal-
lowly sinuous, apex recurved; aedeagal
apodeme (Fig. 24) enlarged, in lateral view
semioval with keel prominent, wide; post-
gonite in lateral view long and conspicu-
ously sinuous, base moderately slender, api-
cal % more slender, apex acutely pointed,
apices cruciate in ventral view (Fig. 25);
hypandrium in ventral view (Fig. 26) nar-
rowly V-shaped with arms slightly flared
apically, base wide and bluntly rounded.

Type Material.—The holotype female is
labeled ““ECUADOR. Manabi[:] Estero
Balsa 9 Sept 1955/Collr.Levi-Castillo/Lem-
naphila 2 n sp. det WWirth °60 [?]/LEM-
NAPHILA WIRTHI L. DE GROSSO
[handwritten]./HOLOTYPO [red].’’ The
holotype is double mounted (glued to a pa-
per triangle), is in good condition (right
wing in glue, some setae broken), and is
deposited in the USNM.

Other Specimens Examined.—NEO-
TROPICAL. DOMINICAN REPUBLIC. La
Vega: Jarabacoa (1-2 km S; 19°06.9'N,
70°37'W; 520 m), 8-21 May 1995, 1998,
W. N. Mathis (296, 132; USNM).

Distribution.—Neotropical: Argentina
(Tucuman), Ecuador, West Indies (Domini-
can Republic).

Remarks.—The specimens from the Do-
minican Republic are a major extension in
the known range of this species, and em-
phasizes the apparent lack of collecting this
species and Lemnaphila generally. We now
know this species from Argentina in the
south to Ecuador and the Dominican Re-
public in the north.

PHYLOGENETIC CONSIDERATIONS

Lemnaphila, as discussed previously (ge-
neric diagnosis and discussion), is similar
and related to Hydrellia and Cavatorella,
and these three genera comprise the tribe
Hydrelliini in the subfamily Hydrelliinae.
The tribe Hydrelliini is distinguished from
other tribes of Hydrelliinae by the follow-
ing characters: specimens usually dull and

672

densely microtomentose (some species sec-
ondarily sparsely microtomentose, subshi-
ny, blackish); ocellar setae seldom as strong
as pseudopostocellar setae, usually much
weaker; eye bearing numerous short, inter-
facetal, fluted setulae; postsutural supra-alar
seta usually short, not longer than posterior
notopleural seta; posterior notopleural seta
at same level as anterior seta (independently
and secondarily inserted above level of an-
terior seta in Lemnaphila, as in Atissini);
costa extended to vein M; midtibiae lacking
dorsal, spinelike setae.

Within Hydrelliini, the three included
genera are distinguished from each other by
the characters noted in the generic key (p.
654). The relationships among these genera,
however, are unresolved and remain prob-
lematic. For example, although Lemnaphila
and its included species are a well-corrob-
orated, monophyletic lineage (see below),
this lineage may eventually be found to be
an included lineage within Hydrellia, i.e.,
Hydrellia lacks characterization by syna-
pomorphies and its monophyly is unsub-
stantiated. Cavatorella may likekwise be an
included lineage within Hydrellia, perhaps
closely related to Lemnaphila.

Synapomorphies that conclusively estab-
lish the monophyly of Lemnaphila are: 1.
Ocellar setae lacking (also lacking in some
species of Hydrellia); 2. Prescutellar acros-
tichal seta lacking; 3. Vein R,,, short with
costal section II shorter than section III (a
shortened R,,, also occurs in some species
of Hydrellia and in Cavatorella); 4. Poste-
rior notopleural seta inserted near dorsal (su-
tural) angle, clearly higher than the level of
the anterior seta; 5. Anepisternum bearing a
single seta along the posterior margin; 6.
Epandrium weakly developed, incomplete
dorsally, with membranous gap between lat-
eral extensions, these extensions are as nar-
row, lateral phalanges at base of cerci; 7.
Cerci fused with medial margin of epan-
drium; 8. Surstylus reduced to a simple,
small process at the ventral margin of the
fused epandrium/cerci complex; 9. Larvae
mining the thalli of Lemnaceae (Hydrellia

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

albilabris (Meigen) from the Old World also
mines the thalli of Lemna (Mathis and Za-
twarnicki 1995)). The mining habit of H. al-
bilabris in Lemna apparently arose indepen-
dently. We have examined specimens of H.
albilabris to verify that they are not closely
related to Lemnaphila.

In the presentation on species-level rela-
tionships that follows, the characters used
in the analysis are noted first. Each char-
acter is immediately followed by a discus-
sion to explain its states and to provide per-
spective and any qualifying comments
about that character. After presentation of
the information on character evidence, an
hypothesis of the cladistic relationships is
presented and briefly discussed. The clad-
ogram (Fig. 27) is the primary mode to con-
vey relationships, and the discussion is to
supplement the cladogram and is intended
only to complement the latter. In the dis-
cussion of character data, a “0” indicates
the state of the outgroup; a “1” or “2” in-
dicate the respectively more derived states.
All multistate characters (1, 4, 8, 11, 14, 18,
20) were treated as nonadditive characters
(—), and characters 5, 6, and 9, which are
autapomorphies for L. longicera, were
made inactive (]) for the analysis, such as
calculation of the consistency index. The
numbers used in the presentation are the
same as those on the cladogram (Fig. 27),
and the sequence is the same as noted in
the character matrix (Table 1).

The other two genera of Hydrelliini, Hy-
drellia and Cavatorella, were selected as
outgroups. As the exemplar of Hydrellia,
we chose the widespread type species of the
genus, H. griseola (Fallén), and for Cava-
torella, which is monotypic, we studied C.
spirodelae Deonier, which occurs in China
and Japan (Deonier 1995). Hydrellia is rep-
resented by over 200 species worldwide
(Mathis and Zatwarnicki 1995).

CHARACTERS USED IN THE PHYLOGENETIC
ANALYSIS

Head:

1. Vestiture of mesofrons: moderately mi-

VOLUME 102, NUMBER 3

Figs. 27.

Hydrellia griseola

Cavatorella spirodelae

11 13 16 17 21 22 23 24

crotomentose, appearing mostly dull
(0); sparsely microtomentose, subshiny
(1); densely microtomentose, appearing
velvety black or whitish (2); meso-
frons, except ocellar triangle, bare,
shiny (3). This multistate character is
treated as nonadditive.

. Ocellar setae: present (laterad of an-

terior ocellus, smaller than pseudopos-
tocellar setae) (0); lacking or greatly re-
duced (1).

. Arrangement of ocelli: ocelli forming

an isosceles triangle with distance be-

Table 1. Matrix of characters and taxa used in the

cladistic analysis of Lemnaphila (numbers for charac-
ters correspond with those used in the text).

OOOO0O0O00O 11111111112 222 2

123456789 01234567890 123 4
Hydrellia OOOO00000 OOODODOOOO000 OOO O
Cavatorella 000100000 10000000000 O00 O
L. longicera 111311001 02112011210 111 ?
L. grossoae 210100110 11010121001 111 1
L. scotlandae 210200100 02010111102 111 1
L. wirthi 210100120 11011101000 111 1
L. neotropica 210100110 11010101001 111 1
L. lilloana 310100110 11010101001 111 1

SIA 5. (6) 9) Zea 4 ato

10 11 16

673

Lemnaphila longicera

Lemnaphila wirthi

16
Lemnaphila grossoae

Lemnaphila neotropica

Lemnaphila lilloana

Cladogram depicting hypothetical cladistic relationships among species of Lemnaphila.

tween posterior ocelli about twice that
between either posterior ocellus and
anterior ocellus (0); ocelli forming an
equilateral triangle or with distance be-
tween posterior ocelli only slightly
more than between either posterior
ocellus and anterior ocellus (1).

. Fronto-orbital setae: anterior fronto-

orbital seta proclinate, smaller than
posterior seta, which is laterobliquely
reclinate (0); posterior seta elongate,
oriented laterally (1); both setae re-
duced but evident (2); both setae great-
ly reduced or lacking (3). This multi-
state character is treated as nonadditive.

. Color of antenna: generally unicolo-

rous, most brownish black but some-
times yellowish (0); scape, pedicel, and
basal half of Ist flagellomere yellow
(1). The derived state of this character
is an autapomorphy for L. longicera
and is therefore treated as an inactive
character in the analysis.

6. Length of Ist flagellomere: about equal

to or slightly greater than height (0);
twice height (1). The derived state of
this character is an autapomorphy for

674

Lh

. Shape of face in lateral view:

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

L. longicera and is therefore treated as
an inactive character in the analysis.
face
straight (0); shallowly concave, usually
with a shallow, transverse groove (1).

. Facial vestiture: face mostly uniform-

ly microtomentose, color varying, de-
pending on angle of view, except for
narrow, vertical stripe (0); ventral third
of face densely microtomentose, sil-
very white, distinctly contrasted with
blackish brown dorsal portion (1); ven-
tral third of face densely microtomen-
tose, dark brown (2). This multistate
character is treated as nonadditive.
Length of outer vertical seta: about *4—
¥% length of inner vertical seta (0); less
than half inner vertical seta (1). The de-
rived state of this character is an auta-
pomorphy for L. longicera and is there-
fore treated as an inactive character in
the analysis.

Thorax:

10.

UY.

|p

PS:

14.

Position of anterior dorsocentral seta:
inserted posterior of transverse suture
(QO); inserted at level of transverse su-
ture (1).

Acrostichal setulae: numerous setulae
and 1 well-developed pair of prescutel-
lar setae (0); sparse setulae anteriorly,
in 2 rows, lacking posteriorly, includ-
ing prescutellar pair (1); lacking (2).
This multistate character is treated as
nonadditive.

Comparative length of basal scutellar
seta: long, from % to being subequal
to length of apical seta (0); short, about
¥% length of apical seta (1).

Position of posterior notopleural seta:
inserted at about same level as anterior
seta (O); inserted at elevated position
relative to anterior seta (1).

Scutellar vestiture: moderately densely
microtomentose, not appearing velvety
(O); scutellum densely microtomentose,
somewhat appearing velvety (1); scu-
tellum densely microtomentose, ap-
pearing velvety, dark brown, similar to

LS!

16.

ie

18.

Nee

20.

notopleural patch (2). This multistate
character is treated as nonadditive.
Length of costal sections IIT and III:
costal sections II and III about equal in
length, vein R,,, relatively long (0);
costal section III considerably longer
than II, vein R,,, relatively short (1).
Shape of anal angle of wing: deeply
rounded, greatest depth equal to or
greater than distance between apices of
vein R,,; and M (0); very shallow
rounded, greatest depth conspicuously
less than distance between apices of
vein R,,; and M (1).

Number of setae and setulae along pos-
terior margin of anepisternum: | large
seta and 1-2 setulae (0); 1 seta, setulae
greatly reduced or lacking (1).
Presence or absence of katepisternal
seta: seta well developed, length sub-
equal to posterior anepisternal seta (0);
seta greatly reduced (1); or lacking (2).
This multistate character is treated as
nonadditive.

Color of katepisternum: brown (0);
whitish yellow, concolorous with fore-
coxa (1).

Color of legs: legs entirely yellowish,
with coxae whitish yellow and apical
half of tibiae slightly brownish yellow
(0); femora brown, tibiae and most tar-
someres yellow (1); legs unicolorous,
brown (2). This multistate character is
treated as nonadditive.

Abdomen:

Ze

Pio

25:

Development of epandrium at dorsum:
dorsum of epandrium well developed
to weakly developed (0); epandrium in-
complete dorsally with membranous
gap between lateral phalanges (1).
Condition of cercus relative to epan-
drium: cerci free from epandrium (0);
cerci fused laterally to medial margin
of epandrial phalange (1).
Development of surstyli: generally well
developed structures at ventral margin
of epandrium (0); reduced to simple

VOLUME 102, NUMBER 3

675
Table 2. Analysis of characters based on the cladogram (Fig. 27).
Character 1 2 8 4 5 6 7 8 9 10 11 12
Steps 3 1 i ) 1 1 1 2 1 1 2 1
Con. Index 100 100 100 100 100 100 100 100 100 100 100 100
Ret. Index 100 100 100 100 100 100 100 100 100 100 100 100
Character 13 14 15 16 17 18 19 20 21 22 23, 24
Steps 1 2 1 4 1 2 1 2 1 1 1 1
Con. Index 100 100 100 50 100: 100 100 100 100 100 100 100
Ret. Index 100 100 100 0) 100 100 100 100 100 100 100 100

small process at ventral margin of the
fused epandrial/cercal complex (1).

Behavior:

24. Feeding preference: miner in various,
mostly aquatic plants (0); miner in the
thalli of Lemnaceae (1). Lacking rear-
ing records for L. lonicera, we coded
this character as a ? for this species.
Like all other members of Lemnaphila,
however, we anticipate and predict that
this species will also be found to be a
miner in aquatic plants of Lemnaceae.

ANALYSIS

Using the implicit enumeration (ie*) op-
tion of Hennig86, four most and equally
parsimonious trees were generated, each
with a length of 34 steps and consistency
and retention indices of 0.91 and 0.86 re-
spectively. The basal nodes in each of these
cladograms are identical, with variation
only in the more derived four species (the
apical 2—3 nodes).

The matrix was then subjected iteratively
to successive weighing (xs w, ie*, cc) to
determine a character’s contribution or
weight and to find cladograms supported by
the most consistent characters (Carpenter
1988, Dietrich and McKamey 1995). Suc-

cessive weighing produced a single clado-
gram (Fig. 27) that is identical to one of the
first four and is our cladogram of choice.
The analysis of the characters for this clad-
ogram is given in Table 2 and the weights
of the various characters is given in Table 3.

Hydrellia was consistently the basal out-
group, with Cavatorella as the more im-
mediate outgroup to Lemnaphila. Although
the placement of Cavatorella as the sister
group to Lemnaphila is currently the best
indication of cladistic relationship, we do
not attribute great weight to this relation-
ship largely because only two synapomor-
phies (characters 2 and 4) were found to
support this sister-group relationship.

The two basal lineages within Lemna-
Phila are first L. longicera, which is the sis-
ter lineage to the remaining species in the
genus, followed by L. scotlandae. The next
four species, which are all Neotropical in
distribution (often at the same habitat), are
obviously closely related and are very sim-
ilar externally, and in the four equally par-
simonious trees, these four species varied
in position with respect to each other. In
two of these trees, L. wirthi was basal to
the other three, and in the other two trees,
L. grossoae was the basal lineage. The sin-
gle tree from successive weighing, howev-

Table 3. Status (i.e., nonadditive —; inactive ]) and weights of characters after successive weighing.

Character No. 1 2 3 4 5
Weight, status IO= | WOE IE TOs | IG |[ IKORS |
Character No. 13 14 NS) 16 iL7/
Weight, status Oar | WO= |[ MORE I Oke |p Ose I

6 7 8 9 10 11 12
OSS) TOS = 7 eee) See SP aoe
18 19 20 21 oy) 23 24
10 op Oe | Os 10s pO Ge |

676

er, has L. wirthi as the basal lineage with
the other three species forming an unre-
solved trichotomy (Fig. 27).

ACKNOWLEDGMENTS

We gratefully acknowledge the assistance
and cooperation of many organizations and
individuals who contributed to the field
work and production of this paper. To Dr.
David A. Grimaldi (AMNH), Mr. Donald E
Azuma (ANSP), Mr. John E. Chainey
(BMNH), Dr. James K. Liebherr (CU), Dr.
Steve Marshall (GUE), Dr. Mercedes Li-
zatralde de Grosso (IML), Dr. Norman
Johnson (CHSU), Dr. Gustavo Spinelli
(UNLP), Dr. Richard S. Zack (WSU) and
their institutions, who loaned specimens,
we express our sincere thanks.

The illustrations were carefully inked by
Mr. Young T. Sohn. Dr. Alan K. Graham,
Kent State University, assisted with the
preparation of the scanning electron micro-
graphs (SEMs), and Mr. George L. Venable
expertly produced the plate of SEMs (Figs.
11-15). Dr. Stephen D. Gaimari prepared
the final version of the cladogram (Fig. 27).
For reviewing a draft of this paper we thank
Drs. Stephen D. Gaimari, Benjamin A.
Foote, and Richard S. Zack.

Field work on St. Vincent, St. Lucia, and
Dominica was supported by a grant from
the Research Opportunity Fund, adminis-
tered by David Pawson and Stanwyn G.
Shetler, former Deputy Directors (USNM).
In 1995, 1996, and 1998, field work on the
West Indies was funded in large measure by
grants from the Biodiversity Program (Bi-
ological Surveys and Inventories, BSI), Na-
tional Museum of Natural History, Smith-
sonian Institution (Dr. Lynne R. Parenti,
chair). Field work on the West Indies was
greatly expedited through the able and
pleasant assistance of my wife, N. Dianne
Mathis, Ms. Hollis B. Williams, Mr. Kelvin
Guerrero, Dr. Daniel E. Pérez-Gelabert, and
Dr. Oliver S. Flint, Jr.

LITERATURE CITED

Buckingham, G. R. 1989. Lemnaphila scotlandae
(Diptera, Ephydridae) and three of its parasites

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

discovered in Florida. Florida Entomologist 72(1):
219-221.

Carpenter, J. M. 1988. Choosing among multiple
equally parsimonious cladograms. Cladistics 4(3):
291-296.

Clausen, P. J. and E. EK Cook. 1971. A revision of the
Nearctic species of the tribe Parydrini (Diptera:
Ephydridae). Memoirs of the American Entomo-
logical Society 27: 1—150.

Courtney, G. W., R. W. Merritt, H. J. Teskey, and B.
A. Foote. 1996. Chapter 22. Aquatic Diptera. Part
One. Larvae of Aquatic Diptera. Pp. 484-514. In
R. W. Merritt and K. W. Cummins, eds., An in-
troduction to the Aquatic Insects of North Amer-
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17/3)

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 678-687

ON THE VALIDITY OF HELICONIUS TRISTERO BROWER AND
HELICONIUS MELPOMENE MOCOA BROWER, WITH NOTES ON SPECIES
CONCEPTS IN HELICONIUS KLUK (LEPIDOPTERA: NYMPHALIDAE)

ANDREW V. Z. BROWER

Department of Entomology, Oregon State University, Corvallis, OR 97331-2907,
U.S.A. (e-mail: browera@bcc.orst.edu)

Abstract.—Lamas’ (1998) criticisms of Brower’s (1996a) taxonomy are shown to be
based on misinterpretations of the evidence and of the rules of nomenclature. The names
H. amaryllis amaryllis f. bellula Stichel and H. melpomene bellula Turner are unavailable.
The name H. bellula Brower is available but invalid. The name H. melpomene mocoa
Brower is the valid name for the subspecies of H. melpomene from the Putumayo region
of southeastern Colombia. The Genotypic Cluster Species Concept is contrasted unfavor-
ably to the Phylogenetic Species Concept with respect to the aims of systematics in
general, and the resolution of relationships among geographically differentiated Heliconius

taxa in particular.

Key Words:
scription, subspecies

Recently, Lamas (1998) published a crit-
ical discussion of two new names applied
to Heliconius butterflies from the upper Rio
Putumayo basin in southeastern Colombia
(Brower 1996a). The taxa in question are
Miillerian mimics: one (H. tristero) was de-
scribed as a species in the H. cydno Dou-
bleday clade, while the other was named as
a subspecies, H. melpomene mocoa. Most
members of the cydno clade exhibit blue-
and-white or blue-and-yellow wing patterns
and mimic species in the sara-sapho group
(cf. Brown 1981, Brower 1994, and Brower
and Egan 1997 for reviews of phylogenetic
hypotheses among Heliconius species), so
it was a surprise to discover a new species
mimicking sympatric races of H. melpo-
mene (L.) and H. erato (L.).

That these formerly conflated taxa rep-
resent two distinct entities is supported by

Heliconius melpomene bellula, Heliconius tristero, nomenclature, circum-

robust evidence stemming from three dif-
ferent sources (mitochondrial DNA se-
quences, genitalic morphology and wing
patterns), and that at least one of them re-
quired a new name is not at issue. Lamas’
criticisms of my paper focussed on the fol-
lowing problems: (1) the availability of H.
bellula Stichel (1923); (2) the supposed
synonymy of H. melpomene mocoa Brower
with bellula auctt.; (3) my interpretation of
the International Code of Zoological No-
menclature (1985) with regard to hybrids;
and (4) the logic of my species concept.
First, I will address these problems in nar-
row relation to Lamas’ critique of my paper
(1996a). My rebuttal of Lamas’ criticisms
is followed by a brief discussion of the spe-
cies problem in Heliconius. An English
translation of the relevant section of Lamas’
paper is included as Appendix 1.

VOLUME 102, NUMBER 3

REPLY TO LAMAS’ CRITIQUE

Is Heliconius bellula Stichel (1923) an
available name?

As Lamas (1998) pointed out, Stichel
(1923) applied the name bellula to a form
of H. amaryllis amaryllis C. & R. Felder
(H. amaryllis is now considered to be a sub-
species of H. melpomene; cf. Ackery and
Smiles, 1976). Lamas argued that Stichel’s
employment of bellula was infrasubspecific
and therefore formally unavailable. How-
ever, Neustetter (1929) cited Stichel’s name
as a trinominal form of H. amaryllis, thus
implying that the name was available
(ICZN Article 16). The subsequent employ-
ment of the name for a geographical race
by more recent authors (e.g., Turner 1971,
Brown 1979, Sheppard et al. 1985, Mallet
1993, Brower 1996b) further suggests that
the name has been treated as available (Art.
45gii), and it was under that premise that I
dealt with it in my paper (Brower 1996a)'.

However, given that Stichel’s holotype is
not a representative of the melpomene clade
(see Brower 1996a and below), Lamas’
opinion that Stichel’s name was quadrinom-
inal provides a convenient excuse to sink it
and avoid the complications described
above. If its original designation is deemed
infrasubspecific, a name remains unavail-
able until a description is provided (Art.
10c). Lamas claimed that Turner (1971)
employed the name bellula to refer to a
subspecific entity, thereby becoming its au-
thor (Arts. 23j and 50c). However, Turner’s
tentative use of the name in a figure legend
(followed by a question mark) was not ac-
companied by a description. Likewise,
Brown (1979), Sheppard et al. (1985), Mal-

'T examined and discussed Stichel’s holotypes of
bellula, permira and degener. Lamas suggested that I
was unaware of four additional forms described by Sti-
chel from the Mocoa region. Rather, these were delib-
erately omitted from discussion, because it was clear
from the original descriptions that they represent var-
ious hybrid forms with recombinant wing patterns that
are even less similar to the modern “‘bellula” concept
than the two other forms I discussed.

679

let (1993) and Brower (1996b) used the
name, but none of them provided a descrip-
tion of the taxon, nor did any of them cite
Stichel’s description (which would not per
se change its availability, in any case: Art.
11dii). All of those uses of bellula employ
it as a nomen nudum and none of them sat-
isfies the criteria of availability (Art. 13a).

Holzinger and Holzinger (1994) provided
a description and illustration of H. melpo-
mene bellula and cited Stichel (1923), but
their book is not consistently binominal (it
uncritically employs numerous infrasubspe-
cific names), and its contents are thus un-
available (Art.11c). Ironically, it appears
that my 1996 paper is the first to provide a
description for H. bellula that would render
it available under all the criteria of Art. 11
(via illustrations, description of differenti-
ating features, identification of a holotype,
and citation of Stichel 1923). But because
I circumscribed the concept narrowly, to re-
fer to a hybrid form typified by Stichel’s
original type specimen, the name is invalid.
Even so, my definition prevents subsequent
usage of bellula for any other Heliconius
taxon (Art. 23h; see hybrid section, below).
Thus, the correct authorship of Heliconius
bellula is Brower (1996a), but the name
does not refer to any valid taxon, regardless
of its repeated misapplication in recent
works (including my own).

Subjective synonymy of bellula and
mocoa?

Lamas (1998) argued that H. melpomene
mocoa Brower is a junior subjective syno-
nym of H. melpomene bellula auctt. Since
bellula is not a valid name, this is a moot
point, but it raises another important issue:
Lamas’ claim shows that has been deceived
by Miillerian mimicry! It is the holotype of
bellula and the holotype of H. tristero that
are closely-related, while the holotype of H.
melpomene mocoa represents a different
clade. Had Lamas examined the relevant
characters, he would have seen that bellula
and mocoa are not the same under any rea-
sonable circumscription H. bellula (what-

680

ever its status) is not an example of the H.
melpomene species group; instead, it is
close to H. tristero in the H. cydno species
group. To include bellula and mocoa in the
same species, one would have to view H.
cydno Doubleday as a subjective junior
synonym of H. melpomene (L.)! I described
this problem explicitly in Brower (1996a),
and it was precisely the desire to avoid such
confusion that prompted me to coin new
names for both the cydno relative (H. tris-
tero) and the melpomene relative (H. mel-
pomene mocoa) in the first place.

Even if bellula and mocoa were closely
related, it is clear from Stichel’s (1923) de-
scriptions (Appendix 2) that he felt that bel-
lula was different enough from the local
“nominate form’ to warrant a separate
name. Stichel believed (in error) that his
three “‘typical’’ specimens were H. amaryl-
lis amaryllis, today recognized as a distinct
H. melpomene subspecies with an allopatric
distribution in the Huallaga valley of Peru
(Sheppard et al., 1985). Although ideas
about what entities within Heliconius de-
serve names have changed as knowledge of
phenotypic variation and geographical dis-
tribution has grown, efforts to redefine an
old name of dubious availability in specific
contradiction to its original author’s intent
seem procrustean. A more reasonable view
is that the “‘nominate form” of H. melpo-
mene from Mocoa and environs was anon-
ymous until I (Brower 1996a) named it.

Hybrids and the ICZN

Lamas (1998) also criticized as erroneous
my invocation of Articles 1b and 23h to
curtail the current usage of bellula because
its holotype is a hybrid (Brower 1996a). He
pointed out that because the definition of
““‘hybrid”’ in the ICZN glossary states that
offspring of crosses between conspecific
subspecies are not hybrids, bellula cannot
be considered a hybrid taxon. Re-exami-
nation of Brower (1996a) shows Lamas’
point to be technically incorrect: I hypoth-
esized that the bellula holotype specimen
was the result of hybridization between H.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

heurippa and H. tristero, both of which I
explicitly claimed were species in that pa-
per. Under such circumstances ICZN article
23h specifically applies’.

Whether or not the bellula holotype ac-
tually represents a hybrid or not (and which
parental species might have spawned it) is
difficult to determine in retrospect and
without extensive data from experimental
genetic crosses. That additional specimens
displaying the same pattern have not been
collected suggests that the holotype exhibits
a rare recombinant wing pattern (typical of
the phenotypic diversity found in other Hel-
iconius hybrid zones). By contrast, there are
multiple specimens from multiple localities
that appear very similar to the holotype of
H. melpomene mocoa, and illustrations of
‘*H. melpomene bellula’’ in recent works
(e.g., Turner 1971, Brown 1979, Holzinger
and Holzinger 1994) all lack the yellow
spots that are present in the bellula holo-
type. My interpretation that the specimen
does not represent a “pure’’ geographical
race is complementary to Stichel’s (1923)
opinion that the bellula holotype is transi-
tional, as shown by his description of it as
distinct and separate from his series of
specimens of the local “‘nominate’’ form.

Logical Consistency

I made several statements that Lamas
chose to overlook in his conclusion that my
taxonomic argumentation (Brower 1996a)
“‘no tiene sustento l6gico al ser refutable.”
First, I did not argue that the current clas-
sification of Heliconius is not illogical. In-
deed, I stated that I chose the ranks I em-
ployed to preserve nomenclatorial stability,

*Lamas objected my use of the term “forbid”
(translated as “‘prohibe”’ in his paper) with reference
to the ICZN rule on names applied to interspecific hy-
brids. Article 23h says, ““A species-group name estab-
lished for an animal later found to be a hybrid . . . must
not be used as a valid name for either of the parental
species, even if it has priority over all other available
names for them, but it may enter into homonymy.” I
leave it to the reader to contemplate whether “must
not be used”’ and “forbids the use of”? mean the same
thing, or not.

VOLUME 102, NUMBER 3

even though I knew that the species delim-
itations among geographically polymorphic
Heliconius butterflies were effectively ar-
bitrary. The final points of my paper were
that,

“The mitochondrial DNA data suggest
that the degrees of relationship among the
H. melpomene races and among the H.
cydno races are similar, and that diver-
gence times within each group are also
approximately the same, implying that
the taxonomic rank of each group should
also be the same. The entire species-level
classification of the genus will probably
require revision as additional data be-
come available.”

Ignoring these qualifying remarks, Lamas
offered (without providing new data or ex-
amining the characters discussed in Brower
[1996a]) a hypothesis of circumscription
that he considered ‘“‘more valid’? than my
view: that H. timareta Hewitson, H. heurip-
pa Hewitson and H. tristero are conspecific.
He based this notion on these entities’ geo-
graphical proximity on the northeastern
slope of the Andes, evidence of hybridiza-
tion among them (which he denied on the
following page), and their hypothesized
close relationship (based on studies of
mtDNA: Brower 1996a, 1996b). However,
both the mtDNA studies and data from a
more comprehensive recent analysis (Brow-
er and Egan 1997) imply that H. timareta,
H. heurippa and H. tristero are no more
closely related to one another than any of
them is to H. cydno or H. pachinus Salvin.
Therefore, if one wanted to lump diagnos-
ably different taxa based on the symple-
siomorphy of potential interbreeding, the
logically consistent choice would be to col-
lapse all of these taxa under the oldest spe-
cies-group name (H. cydno). I said as much
in 1996a.

If Lamas (1998) were as concerned with
the consistent assignment of taxa to the ap-
propriate rank as his criticism of Brower
(1996a) implies, then according to his
‘““more valid’? hypothesis, he should have

681

named his new Heliconius subspecies (de-
scribed in the same paper) not “H. timareta
timoratus, > but ““H. heurippa timoratus”
(if timareta and heurippa are conspecific,
then the former is either a junior synonym
or a subspecies of the latter). Under the in-
terpretation Lamas claimed that he prefers,
timoratus would seem to be an infrasubspe-
cific form of the subspecies H. heurippa ti-
mareta! The complexity of sorting out
names, ranks and relationships among geo-
graphic races in Heliconius obviously pre-
sents a challenge that neither Lamas nor I
have yet resolved in a fully satisfactory
manner.

Synonymies

Heliconius melpomene mocoa Brower,
1996a.

Heliconius melpomene bellula auctt. (Turn-
er 1971, Brown 1979, Sheppard et al.
1985, Mallet 1993, Brower 1994, Holzin-
ger & Holzinger 1994) [misidentifica-
tions; unavailable name].

Heliconius melpomene mocoa Brower
1996a: 328. Holotype: Colombia, Dpto.
Putumayo, 1—3 km N. Mocoa on rd. to
Pitalito, 25 March 1992 leg. AVZ Brow-
er. Deposited in Cornell University Insect
Collection (examined).

Heliconius heurippa X Heliconius tristero,
natural hybrid

Heliconius amaryllis amaryllis f. bellula
Stichel, 1923: 262. Original type speci-
men: Colombia, Rio Putumayo, Rio Gua-
yuyaco, 7 July 1921 leg. W Hopp (see
Brower 1996a for transcript of original
labels).

Heliconius amaryllis f. bellula Stichel;
Neustetter 1929: 58.

Heliconius amaryllis f. bellula Stichel;
Brower 1996a [identified as hybrid H.
heurippa X H. tristero].

NOTES ON SPECIES CONCEPTS IN HELICONIUS

I suspect that the root of Lamas’ criti-
cisms of my concepts of tristero and mocoa
lies in his concept of species in Heliconius,

682

which differs from mine. The species prob-
lem in Heliconius has been controversial
for many years (e.g., Eltringham 1916 vs.
Kaye 1916); indeed, it is the very complex-
ity of geographical diversification within
the genus that has led it to become a model
system for the study of the evolutionary ge-
netics of mimicry (Sheppard 1960, Emsley
1964, Turner 1971, Brown et al. 1974, Mal-
let 1993). Many of the Heliconius papers
from the post-typological period applied the
biological species concept (BSC; Mayr
1940)), which unites allopatric taxa based
on their potential to interbreed. In recent
years, the BSC has been criticized on the-
oretical and practical grounds, and numer-
ous alternative species definitions have
been proposed that offer more operational
criteria for species delimitation (Eldredge
and Cracraft 1980, Mishler and Donoghue
1982, de Queiroz and Donoghue 1988, Nix-
on and Wheeler 1990, Baum and Shaw
1995, Mallet 1995). Lamas’ critique im-
plied that he favors the “genotypic cluster”
species concept (GCSC; Mallet 1995),
while I (Brower 1996a) applied the phylo-
genetic species concept (PSC) as elaborated
by Nixon and Wheeler (1990).

Evidence and Criteria of Specific
Distinctness

Mallet (1995) argued that the PSC fails
to provide “‘clear guidelines” for dividing
species because it recognizes groups based
on apomorphy: ‘‘With detailed morphology
and modern molecular techniques,”’ he as-
serted, “‘one can find apomophies for al-
most every individual,”’ which he suggested
would result in rampant splitting and pro-
liferation of species. This simplistic cari-
cature of the cladistic method misrepresents
the procedure of species delimitation,
which has been explored in depth by Davis
and Nixon (1992), Doyle (1995) and Brow-
er (1999). Cladists identify populations of
organisms that they hypothesize to be dis-
tinct, and seek discrete differences between
them. If they discover such differences, cla-
dists consider the populations to be separate

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

species; otherwise, they are collapsed into
a single species. That the cladistic approach
may yield finer resolution of the hierarchi-
cal pattern of diversity than alternative
methods is considered by many to be an
asset. In short, the guidelines of the PSC
are clear: it is Mallet’s understanding of
them that seems cloudy.

Mallet’s (1995) alternative, the GCSC,
views species as “‘identifiable genotypic
clusters” recognizable by ‘a deficit of in-
termediates”” at single and multiple loci,
and views speciation as “‘the production of
divergent populations that can coexist in
sympatry.”’ There are a number of problems
with this approach. First, there are no ex-
plicit criteria for identification of genotypic
clusters. That Mallet employed “deficit”
instead of “‘absence’’ implies that he be-
lieves that the clusters need not be fixed for
alternate alleles, but merely differ in allele
frequency by some “‘significant’’ amount.
Further ambiguities include how many
markers should be sampled (Mallet [1996]
suggested multiple loci), and what should
be done if one locus suggests continuity,
while another suggests distinction. The
across-locus averages that Mallet (1995)
employed in the hybrid indices he presented
would disguise heterogeneity among loci. A
more sophisticated and logically consistent,
but equally labor-intensive approach to
multilocus species discrimination was pre-
sented by Doyle (1995). In practice, how-
ever, methods like these that rely on ex-
haustive characterization of gene pools are
rarely employed in the study of biological
diversity, because they entail intensive,
quantitative sampling that is not feasible in
most circumstances.

Another difficulty with the GCSC is non-
dimensionality.—Maallet’s concept is only
useful for contemporaneous taxa in sym-
patry or parapatry. To cover everything
else, Mallet (1995) suggested that “‘closely
related allopatric forms should mostly be
considered conspecific.’’ But how are we to
determine that allopatric forms are “‘closely
related?’’ If allopatric populations are

VOLUME 102, NUMBER 3

“identifiable,” then they logically satisfy
Mallet’s GCSC criteria and are distinct spe-
cies by definition; if they are not identifi-
able, then why would we hypothesize that
they were different in the first place? When
the hypothetical taxa under investigation
occur in allopatry, researchers are forced to
turn to the empirical comparison of features
of organisms to draw inferences about their
taxonomic relationships. The PSC recogniz-
es separate species only when fixed differ-
ences are discovered between hypothesized
groups, which will always result in the rec-
ognition of a minimal number of taxa in
comparison to methods based on frequency
differences. The claim that the PSC over-
splits taxa relative to the GCSC is simply
false.

In summary, Mallet’s GCSC is a meth-
odologically explicit version of the nondi-
mensional BSC (Mayr 1963). It is labor-
intensive, depending upon sampling of mul-
tiple individuals at multiple loci to provide
empirical evidence of the absence of inter-
breeding. It bases the decision of specific
distinctness on an arbitrary and unstated
level of phenetic bimodality in a histogram
of average genetic scores. Although Mallet
and colleagues have made a rather convinc-
ing case for the distinctness of Heliconius
erato cyrbia Godart and H. himera Hewit-
son on the basis of this method (Jiggins et
al. 1996), that work represents a laborious
multi-year, multi-authored effort to corrob-
orate a conclusion about a single pair of
taxa that had already been hypothesized by
systematists years before (Descimon and
Mast-de Maeght 1984). How Mallet’s con-
cept could be useful in the best of circum-
stances to a museum taxonomist working
with qualitative samples of preserved, dead
specimens on pins is not clear. What is
quite evident is that Lamas has never em-
ployed the GCSC in any of his published
taxonomic work, including the descriptions
of new subspecies in Lamas (1998). Given
this lack of consistency and rigor, it is es-
pecially ironic that my (1996a) names
should be subject to such scrutiny, when

683

they are perhaps the most thoroughly-di-
agnosed Heliconius taxa that have been
published, the differentiating characters
having been drawn from the results of cla-
distic analyses of mtDNA, and corroborated
with diagnostic characters from external
and internal morphology (Brower 1996a)
and data from a nuclear gene (Brower and
Egan 1997).

Recognition and Circumscription of
Subspecies

Although the genus Heliconius contains
numerous diagnosably different populations
that bear valid species-group names, most
of these are considered to refer to intraspe-
cific variations. Since the rejection of ram-
pant typological splitting in the early 20%
Century (e.g., Riffarth 1902), Heliconius
species have been circumscribed primarily
by the BSC criterion of interbreeding. Un-
der that criterion, otherwise uniform para-
patric populations that hybridize where they
abut have been considered conspecific.
Likewise, distinct populations that hybrid-
ize with each of two otherwise allopatric
neighbors provide a transitive link that has
allowed lumping of chains of populations
into single, geographically extensive “‘bio-
logical species.’ These species’ component
*““geographical races’”’ are diagnosably dif-
ferent (i.e., they display heritable characters
that allow their unambiguous determina-
tion), and have been considered by many
researchers (e.g., Brown et al. 1974, Shep-
pard et al. 1985, Brower 1996a, 1996b) to
represent historically distinct entities. That
their names are in common use in the lit-
erature and in museum classification
schemes is a de facto acknowledgement of
their recognition as taxa, even by those bi-
ologists who would emphatically deny their
specific status. According to the PSC, diag-
nosably distinct groups that it is useful to
name are considered separate species, and
every recognized “geographical race’? in
Heliconius should be a phylogenetic spe-
cies.

The ICZN (Art. 45a) considers names at

684

both specific and subspecific levels to be
labels for taxa of a single category, the Spe-
cies Group. The concept of subspecies is
simply a convenient label for taxa at one of
the potentially many hierarchical levels
nested within the genus, as revealed by cla-
distic studies. The Code is sensibly silent
on the problems of definition and boundary
determination of concepts associated with
name-bearing type specimens; such deci-
sions are considered subjective, and left to
the judgement of the describer and subse-
quent employers of the name. In most in-
stances, the older Heliconius names were
described from one or a few dead speci-
mens in European collections, and were not
accompanied by any discussion of the cir-
cumscription of the associated concept, be-
yond designation of an intraspecific level
(e.g., subspecies, form, aberration, etc.). A
good example of such a perfunctory de-
scription is Stichel’s original diagnosis
(1923) of H. amaryllis amaryllis f. bellula
(Appendix 2).

Minimal original descriptions leave a
great deal of latitude for subsequent inter-
pretation. Such interpretations should strive
to maintain nomenclatorial stability, but
only when the names preserved are precise-
ly and accurately associated with empiri-
cally supported concepts. If a concept di-
verges from the description due to the ac-
quisition of new specimens and data to the
point that the description is no longer ade-
quate, some action is called for, ranging
from redescription of the holotype to the
separate description of differentiated con-
cepts as distinct taxa. The currently-recog-
nized “‘geographical races”’ (= phylogenet-
ic species) of Heliconius are taxa corrobo-
rated by a century of empirical research in
laboratories, museums, and the field. They
bear names originally applied to the single
holotype or short type series, and to which
the current concepts correspond. The focus
of modern systematic effort should be on
the empirical diagnosis of such taxa and the
inference of hierarchical relationships
among them, not on empty disputes over

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

the arbitrarily determined ranks that partic-
ular taxa do or do not represent. H. mel-
pomene mocoa and H. tristero are both
diagnosably distinct taxa of the Species
Group.

ACKNOWLEDGMENTS

Thanks to D. D. Judd and E C. Thomp-
son for advice on the ICZN, A. Warren for
checking my translation of Lamas’ Spanish,
and A. Kretzer for help with Stichel’s Ger-
man. Thanks to R. I. Vane-Wright for his
thorough and candid reviews, and to Ger-
ardo Lamas for making me play by the
rules.

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Publications, Oxford and Edinburgh.

APPENDIX |

Translation of Lamas’ (1998: 119-120) critique.
Translation is as precise as possible, but some idioms
were translated freely to improve comprehensibility.
For complete literature citations, see Lamas (1998)

Status of Heliconius tristero Brower and H. melpo-
mene mocoa Brower

Recently, Brower (1996) has described two new
taxa of Heliconius from Putumayo, southeastern Co-
lombia, based on preliminary data from analysis of

686

mitochondrial DNA sequences. Both names are dis-
cussed separately here.

Heliconius tristero was diagnosed by Brower (1996)
as a taxon of the species group, belonging to the H.
cydno Doubleday clade, and narrowly separated from
H. heurippa Hewitson and H. timareta Hewitson. Al-
though he did not mention it explicitly, Brower seems
to imply that heurippa, tristero and timareta could
constitute a subclade of the cydno group. The three
taxa exhibit an allopatric distribution, heurippa occur-
ring in the central eastern region of Colombia, tristero
in southeastern Colombia and fimareta in eastern Ec-
uador (and northern Peru, vide infra); while heurippa
and tristero (so far as is known) are phenotypically
monomorphic, timareta is polymorphic. Brower (1996:
330) concluded that “‘... tristero is considered a spe-
cies only because its geographically adjacent close rel-
atives, H. heurippa and H. timareta, have been tradi-
tionally considered species as well.’’ This conclusion
is not so logical as to be irrefutable. A more valid (and
refutable) hypothesis would be to consider the three
taxa conspecific, based on Brower’s own molecular
and morphological analyses (which would show their
narrow evolutionary relationship), their allopatric dis-
tributions (and geographical proximity), and the sup-
posed existence of transitional forms (“‘hybrids”’) be-
tween heurippa and tristero (represented by the
“type” of bellula Stichel, but vide infra), which might
suggest that, as Mallet (1995) put it nicely in his def-
inition of species as “‘genotypic aggregations:” “
closely related allopatric forms should mostly be con-
sidered conspecific.”’ Naturally, even though it is well
known that in the absence of evidence of specific dis-
tinctness in sympatry the conspecificity of allopatric
taxa is arbitrary, such arbitrariness is certainly less than
to assign a taxon to a particular taxonomic level fol-
lowing a “tradition.”

In synthesis, there are two opposing taxonomic hy-
potheses: 1) tristero is a taxon at the species level,
evolutionarily independent of heurippa and timareta,
with its own historical destiny; or 2) heurippa, tristero
and timareta constitute a polytypic species with three
geographical races (subspecies) which have the same
historical destiny. With the scarce taxonomic and ge-
netic information available at the moment, it is impos-
sible to decide which of these is closer to the truth.

The case of Heliconius melpomene mocoa seems
much simpler, and here I offer the hypothesis that mo-
coa is no more than a new synonym of H. m. bellula
Turner, 1971, as I will demonstrate below. The name
bellula was introduced for the first time in the literature
by Stichel (1923), who proposed it as a form of Hel-
iconius amaryllis amaryllis C. & R. Felder, thus con-
stituting an infrasubspecific name (excluded by the
ICZN). The name bellula recently became available
when Turner (1971) elevated it to the subspecific level
(as a subspecies of H. melpomene (L.), making the
name attributable to Turner (ICZN Arts. 10c, 23j, 50c).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Brower examined the male specimen upon which Sti-
chel based his infrasubspecific name bellula (and
which constitutes the holotype of bellula Turner) and
decided that it represented an individual belonging to
the cydno clade, and not to melpomene. Brower also
examined the “‘holotypes” of two other infrasubspe-
cific names proposed by Stichel (1923), permira and
degener (which have never been elevated to the sub-
specific level and thus are nomenclaturally unavail-
able), suggesting that they might represent hybrids be-
tween H. heurippa and H. tristero. What is more, he
assumed that the type of bellula appeared to be a re-
combinant backcross between these hybrids and tris-
tero. Arguing that the ICZN “‘forbids”’ the use of spe-
cies-level names that are based on hybrids, he decided
to set aside the name bellula and establish new names
for the member of the cydno clade (tristero) and the
subspecies of melpomene present in the upper Rio Pu-
tumayo (mocoa).

Brower does not seem to have realized that Stichel
(1923), in addition to bellula, permira and degener,
described four other “‘forms” of amaryillis from the
region of the upper Putumayo (anacreontica, perrara,
rufata and aglaspis), and that other authors introduced
six additional names for specimens from the same zone
(parva Neustetter, tenuifasciata Neustetter, aurofascia-
ta Neustetter, paula Neustetter, paulina Niepelt and
carminata Niepelt). In my opinion—and contrary to
Brower’s view—these 13 names pertain to examples
of melpomene, and represent transitional forms (“‘hy-
brids’’) between the subspecies bellula Turner and
malleti Lamas. All those specimens were captured
around Mocoa (01°09'N, 76°37’W) by collectors who
worked for Werner Hopp, and were very likely ob-
tained in the hybrid zone near Villa Garzon (= Villa
Amazonica; 01°05'N, 76°35’W, 420 m), a site studied
by Mallet (1993), a few km. East of Mocoa. The ho-
lotype of bellula is labeled as having been obtained on
the Rio Guayuyaco (written ““Guagzayaco”’ on the la-
bel), possibly very near the village of Guayuyaco (=
Napoles; 01°04'N, 76°26’W). The other specimens
bear the localities Rio Mulato (01°08'N, 76°36'’W),
500 m; Mocoa, 530 m; or simple ““Mocoa”’ (for a gen-
eral description of the area, see Salazar 1995). Mallet
(1993) referred to the hybrids found near Villa Garz6n
as the product of the transition between bellula and
aglaope C. & R. Felder; the correct name of the latter
subspecies is malleti Lamas, since the subspecies
aglaope is limited to the lower Rio Maranon, lower
Rio Huallaga and the Rio Ucayali basin in Peru (La-
mas 1988).

Of the 14 names applied to bellula X malleti hy-
brids, the “‘purest’’ example corresponds to the holo-
type of bellula, which only shows tiny yellow spots at
the costal inner edge of the discal red forewing band,
appearing almost identical to the holotype of mocoa
(which lacks the yellow spots). Brower based his
judgement on the presence of these yellow spots, and

VOLUME 102, NUMBER 3

the morphology of the genitalia of the bellula holo-
type, to assert that that specimen was a heurippa X
tristero “hybrid,” (the same as permira and degener).
But, in the first place, heurippa (one of the supposed
parents) is not known from the Mocoa region, only
from much further north, from the Meta and Guaya-
bero basins, while surely tristero should occur in the
basin of the Rio Orteguaza, since bellula has been re-
ported from there. (It would be very interesting to dis-
cover if heurippa, or tristero, or both, occur in the Rio
Caguan basin, between the Guayabero and the Orte-
guaza). Further, even though Brower admitted that
“male genitalia of the taxa examined are similar, and
display substantial intra-racial variability in form,’’ he
concluded that the genital morphology of the bellula
holotype corresponded to the cydno clade, based on
the examination of that one specimen.

Finally, Brower argued that the name bellula should
be set aside because it is applied to a hybrid (which,
as indicated by the discussion above, I consider com-
pletely erroneous) and because the ICZN “‘forbids”’ the
use of names applied to hybrids. In the glossary of the
Code (1985: 256) it is clearly indicated that “‘The
progeny of two individuals belonging to different sub-
species of same species are not hybrids.’’ The Code
does not “forbid” the use of names given to interspe-
cific hybrids, it simply excludes them from nomencla-
ture, except for the principle of homonymy. Therefore,
if the name bellula does not correspond to an inter-
specific hybrid, its use to designate a subspecific taxon
is perfectly valid, and in consequence, Heliconius mel-
pomene mocoa Brower, 1996 is a subjective junior
synonym of Heliconius melpomene bellula Turner,
1971.

APPENDIX 2

Translation of Stichel’s (1923) description of H. am-
aryllis amaryllis and H. amaryllis amaryllis f. bellu-
la.from the upper Putumayo of Colombia.

687

H. amaryllis amaryllis Feld.
Forma typ.

Two males.—Forewing with a somewhat variable
broad red discal patch, somewhat like the figure of H.
amaryllis euryades Riff. in Gen. Ins. v. 112, plate 3,
fig. 10, but with a more blurred distal margin, and
which is somewhat indented below the forewing me-
dian vein. The yellow transverse band of the hindwing
fragmented near the base by thin black lines, and the
veins cutting across the band are more or less black.
Mocoa (Put[umayo]), September, October.

Among other things, a character of this species is
said to be the absence of red basal streaks on the costal
margin of the forewing underside. This feature is pre-
sent in all of the observed nominate and closely allied
forms, but based on the other specific characters, par-
ticularly the position of the hindwing band, they can
only be considered forms of amaryllis. This band is
somewhat variable, but it is always positioned such
that the posterior border lies outside the posterior edge
of the discal cell. In a third male specimen (Mocoa,
May), the forewing band is somewhat smaller, so that
it scarcely touches the end of the cell, the red is faded
and greasy, which is often seen as a pathological fea-
ture of red-banded Heliconius.

Forma bellula f. nov.

One male.—Most similar to the nominate form, the
crimson forewing patch more ragged on the margin,
partly dusted with black scales, the cell almost com-
pletely free of red, an additional character is a sulfur-
yellow spot proximal to the subcostal vein. The yellow
transverse band on the hindwing is broad, extending
nearly 7 mm. from the apex to the trailing edge.
Hindwing beneath very similar to H. amaryllis rosina
Bsd. With red basal streaks on the leading edge of the
forewing and three red basal spots in the hindwing.
Forewing length 41 mm. Rio Guaqzayaco
(Put[umayo]).

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 688-740

APHID PARASITOIDS (HYMENOPTERA: BRACONIDAE: APHIDITNAE)
OF NORTHWEST USA

K. S. PIKE, P. STARY, T. MILLER, G. GRAF, D. ALLISON, L. BOYDSTON, AND R. MILLER

(KSP) Entomologist and (DA, LB, GG) Research Technicians, Washington State Uni-
versity, Irrigated Agriculture Research & Extension Center, 24106 N Bunn Road, Prosser,
WA 99350, U.S.A.; (PS) Entomologist, Institute of Entomology, Academy of Sciences of
the Czech Republic, BraniSovska 31, 370 05 Ceské Budéjovicé, Czech Republic; (RM)
Entomologist, University of Guam, CALS, AES, 303 University Drive, UOG Station,
Mangilao, GU 96923, U.S.A.; (TM) Manager, Northwest Biological Control Insectary &
Quarantine, Washington State University, Pullman, WA 99164, U.S.A.

Abstract.—A comprehensive assessment is provided of the Northwest USA aphid-par-
asitoid species (Hymenoptera, Braconidae, Aphidiinae). Eighty-one species of parasitoids
in 19 genera are recognized; 35 species are new to the region; 7 species are newly
described: Aphidius segmentatus Pike and Stary [hosts: Hyperomyzus on Hieracium, Lac-
tuca, Senecio, and other Compositae, and on Uroleucon on Achillea]; Betuloxys alnicolus
Pike & Stary [host: uncertain, possibly Boernerina variablilis Richards on Alnus]; Mon-
octonus campbellianus Pike & Stary [hosts: Brachycaudus, Hyalopterus, and Phorodon
on Prunus]; Monoctonus pacificus Pike & Stary [host: Macrosiphum tuberculaceps (Es-
sig) on Achlys triphylla|; Praon coniforme Pike & Stary [host: Aphthargelia symphori-
carpi (Thomas) on Symphoricarpos];,; Praon fulvum Pike & Stary [hosts: Macrosiphum
spp. on Achlys, Crepis, Lupinus, Spiraea]; Trioxys setaceus Pike & Stary [host: Acyrtho-
siphon macrosiphum (Wilson) on Amelanchier]. Seventy genera of aphids (200+ spp.)
were attacked by aphidiine parasitoids. Some 400+ parasitoid-aphid associations, and
hundreds more tritrophic (parasitoid-aphid-plant) combinations are reported for the first
time in North America.

Key Words: aphid, parasitoid, systematics, Northwest USA

The aphidiine parasitoids of Northwest
USA were previously understood to include
about 54 species in 13 genera; these were
linked with about 100 species of aphids
(Pike et al. 1996, 1997, 1999). Based on
new environmental studies, the parasitoid
guilds are recognized now to be much larg-
er. Presented here is a comprehensive re-
view of Northwest species, including an
abridgment of previously published records.

Eighty-one species of aphidiine parasit-
oids in 19 genera are now recognized in

Northwest USA; 35 species are new to the
region; 7 species are newly described. Col-
lectively, these parasitoids attacked more
than 200 species in 71 genera of aphids.
Some 400+ parasitoid-aphid associations,
and hundreds more tritrophic (parasitoid-
aphid-plant) combinations are reported for
the first time in North America.

The work is a foundation study of the
Northwest aphidiine fauna. It provides
background documentation for possible fu-
ture studies on a variety of fronts, such as

VOLUME 102, NUMBER 3

Table 1. Legend for listings.

689

PARASITOID

Praon americanum (Ashmead)

[Example]

Aphis coweni Palmer: WA, KITTITAS [*] — Stampede Pass, 8-VIII-96 on Veratrum viride (96K1477f).

APHID HOST STATE COUNTY SPECIFIC DATE APHID PLANT WSU CODE
COLLECTION COLLECTED Host
SITE

[*] indicates previous published record, see Pike et al. 1996, 1997, 1999.

[+] indicates collection contained more than one species of aphid.

[+] indicates parasitoid-aphid association shown is not certain.

Abbreviations: ALE, Arid Lands Ecology Reserve; CA, California; CG, Campground; Cr, Creek; det., deter-
mined; E, east; FR, forest road; G&P, Gillette & Palmer; HRL, Hille Ris Lambers; HMU, Habitat Management
Unit; ID, Idaho; Lk, Lake; mi, mile; MT, Montana; Mt, Mount; Mtn, Mountain; N, north; nr, near; OR, Oregon;
RNP, Mount Rainier National Park; Pk, Park; Rd, Road; S, south; sp., species; SP, State Park; W, west; WA,
Washington; WLA, Wildlife Area or Refuge; WSU, Washington State University; YIR, Yakama Indian Reser-

vation.

biosystematics, including new species de-
scriptions; parasitoid guilds; population ge-
netics; host adaptation, switching, and cross
habitat movement; geographic spread of ex-
otic species; interspecies population devel-
opment; ecosystem relationships and effects
on parasitoid and predator complexes, and
target pests; application and advantages of
aphid-plant biodiversity for maintaining,
stabilizing, and/or increasing parasitoids of
merit; and uses and possible export of
Northwest parasitoids to other global areas.

MATERIALS AND METHODS

Aphidiine parasitoids were obtained from
aphid rearings, with the aphids collected
from a wide range of host plants from
Northwestern USA, principally Washing-
ton. In total, more than 5,300 aphid collec-
tions were taken, of which 2,940 were par-
asitized. The aphid populations sampled
varied in size. Where possible 50 to >200
aphids were taken per sample; a subsample
of 5 to 30 aphids were preserved in 70%
ethanol for later identification. For the par-
asitoid rearings, aphids were held in-labo-

ratory at 20 + 3°C for 25-30 days in
screen-covered semi-transparent plastic
containers (two sizes used: 300 ml, 10 cm
dia X 4 cm ht; and 3500 ml, 19 cm dia X
13 cm ht) or occasionally in paper lunch
bags. After emergence, the parasitoids were
placed in 70% ethanol for subsequent de-
termination. Voucher specimens of both
aphids and parasitoids are in Washington
State University-Prosser collections, Pros-
ser, WA.

Descriptions of new species were based
on whole dry and dissected slide-mounted
specimens examined under 40—400X mag-
nification. Body lengths were measured and
recorded in millimeters. Holotypes were
dry-mounted on paper tabs and pinned. De-
scriptive terminology is after Huber and
Sharkey (1993). Holotypes are deposited in
the National Museum of Natural History,
Smithsonian Institution, Washington, DC
(USNM).

For cross reference and other information
on aphid parasitoids of North America, see
Gahan 1911; Smith 1944; Liu 1977; Marsh
1979; Johnson 1987; Pike et al. 1996, 1997,

690

1999; for world information, see Mackauer
and Stary 1967.

Aphid nomenclature follows Remaudiére
and Remaudiére 1997.

RESULTS

Aphidiine parasitoid genera and _ spe-
cies.—Parasitoid genera and species are
listed alphabetically, as are aphid hosts un-
der each species. Locations are indicated by

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

state (large caps), county (small caps), and
nearest city or identifiable site. Numbers in
parentheses represent authors’ codes of
specimens (aphid-parasitoid vouchers) in
Washington State University—Prosser col-
lections (Table 1). Collections and deter-
minations were by authors unless otherwise
specified. Aphids collected by A. Jensen
were determined by A. Jensen. Authorship
of new species is attributed to K. S. Pike
and P. Stary.

APHIDIINE PARASITOIDS OF NORTHWEST USA

Genus ACANTHOCAUDUS Smith

Acanthocaudus caudacanthus (Smith)
Uroleucon sp.: WA, PEND OREILLE—Le Clerc WLA, 3-VIII-95 on Aster sp. (95T400); SPOKANE—nr Peone,
24-VIII-95 on Centaurea pratensis (95T473*).

Acanthocaudus sp.?
Undetermined sp.: WA, GARFIELD—Pataha Cr, nr FR160 & FR40, 21-[X-95 on Chimaphila umbellata
(95T597).

Genus ADIALYTUS Foerster

Adialytus ambiguus (Haliday)
Sipha elegans del Guercio: MT, SANDERS [on Agropyron intermedium*].
Sipha sp.: WA, ASOTIN [on unknown Gramineae* ].

Adialytus fuscicornis (Ashmead)

Aphis armoraciae Cowen: OR, UMATILLA—Hermiston, 17-VI-90 on unknown plant (90J002); WA, YAKIMA—
Wenas Lk, 30-V-96 on Erysimum arenicola (96G120), & 11-VI-97 on Lomatium sp. (97G051*).

Aphis helianthi Monell: WA, YAKIMA—Wenas Lk, 11-VI-97 on Lomatium sp. (97GO0517¢).

Forda marginata Koch: WA, GARFIELD—Dye Seed Airfield, 5-[X-95 on Bromus sp. (95G542); Kirby, 1-[X-
95 on Bromus inermis (95K160).

Undetermined sp.: WA, AsoTIN [on Populus trichocarpa*}.

Adialytus salicaphis (Fitch)

Chaitophorus populicola Thomas [on Populus trichocarpa unless otherwise indicated]: ID, BOUNDARY [*];
WA, AsotTin—Asotin WLA, 31-VII-96 & 15-VIII-96 (96T355, 96T454); BENTON—Grandview, 10-X-95
(95G694); WHITMAN—Pullman, 9-X-96 on Populus tremuloides (96T582); YAKIMA—YIR, Lucy Canyon,
25-IV-96 & 16-V-96 (96G021, 96G061) & nr Satus, 1-VI-96 (96K062).

Chaitophorus populifolii (Essig) [on Populus trichocarpa]: WA, KitrirAs—Manastash Cr, 16-VIII-96
(96G485); Ellensburg, 13-VI-97 (97G083); YAKIMA—Naches, 27-IX-96 (96G600).

Chaitophorus utahensis (Knowlton) [on Salix sp.]: WA, AsoTiN—Chief Timothy HMU, 16-VIII-95 (95T442);
BENTON—nr Grandview, 24-VIII-95 (95G514); YAKIMA [*]—Grandview, 22-V-95 & 15-V-96 (95G078,
96G047).

Chaitophorus viminalis Monell [or nr viminalis] [on Salix sp.]: WA, OKANOGAN—Alta Lk, 10-L[X-97 (97K026,
97K028); YAKIMA—Wenas Cr, 13-VIII-96, (96G459).

Chaitophorus sp. {nr nigrae Oestlund & pallipes Richards]: WA, AsotiN—Heller’s Bar, 11-VII-95 on Salix
sp. (95T2107).

Chaitophorus sp. {on Salix sp. unless otherwise indicated]: WA, AsoTIN [*]; BENTON [*]—Prosser, 10-X-95
on Populus trichocarpa (95G693); GRANT—Potholes Reservoir, 13-VI-97 (97G101).

Periphyllus nr brevispinosus G&P: WA, PEND OREILLE—Sullivan Lk Rd, Harvey Cr, 3-VIII-95 on Acer sp.
(951395); SPOKANE—Mt Spokane SP, 24-VIII-95 on Acer sp. (95T487).

Periphyllus sp.: WA, ASOTIN [on Acer sp. *].

VOLUME 102, NUMBER 3 691

Genus APHIDIUS Nees

Aphidius avenaphis (Fitch)

Acrthosiphon lactucae (Passerini): WA, WHITMAN—Pullman, 13-VI-96 on Lactuca serriola (96T0327#).

Diuraphis noxia (Kurdjumov): WA, AsoTin—Anatone, 24-VII-96 & 22-VII-97 on Hordeum vulgare (961596,
97JOO5—97J023); KLICKITAT—Bickleton, 29-VI-97 & 6-VIII-97 on Triticum aestivum (97G300, 97G354).

Rhopalosiphum padi (L.): WA, GRANT—Ephrata, 8-VII-97 on Zea mays, K. Volker collector (97K058).

Sitobion avenae (FE) [on Triticum aestivum unless otherwise indicated]: ID, LArAH—Moscow Mtn, 3-VIII-96
on unknown Gramineae (9613967); SHOSHONE—Thompson Pass, 29-VII-95 on Agropyron caninum
(95G478), Arabis sp. (95G475+), & Poa nervosa (95G470); OR, UMATILLA—Harris Pk, 11-VI-96, on
Dactylis glomerata (96G230+); WA, ADAMsS—Bruce, I-VII-96 (96B004, 96B007); Othello, 26-VI-95, 1-
VII-96 (95B009, 96B009); AsoTin—Anatone, 24-VII-96 on Hordeum vulgare (961337); BENTON—Prosser,
2, 28 & 30-VI-95, 17-VII-95, 24-VI-96, 1 & 8-VII-96, 3-VII-97, (95G103, 95G276, 95G279, 95G285,
95G352, 95G354, 95G357+, 95G358, 96G271, 96G312, 96G328, 96G330, 97F201): Richland, 21-VI-95
(95G2217+); West Richland, 28-VI-95 (95G279); WSU-Prosser, 18-VII-97 on Setaria lutescens (97G243);
DouGLAs—Waterville, 24-VII-96 (96B025); FRANKLIN—Kahlotus, 6-VII-95 (95G290, 95G3087); Star
School, 6-VII-95 (96G289); GRANT—WSU-Royal Slope, 26-VI-95 (95B007); Kittiras—S. Fork
Manastash Cr, 24-VII-95 on Agropyron repens (95G375), Agropyron caninum (95G383), & T. aestivum
(95G372); 11-IX-95 on Bromus sp. (95G384); Manastash Rd, 11-IX-95, on D. glomerata (95G565);
Stampede Pass, 8-VIII-96 on Phleum sp. (96K142); KLickiTaT [on H. vulgare & Triticosecale rimpani* |
—Bickleton, 9 & 17-VII-95 (95G317+—95G3197, 95G362), 7-VIH-97 (97G224); Trout Lk, 29-VI-97 on D.
glomerata (97G160); LINcoLN—Harrington, 19-VII-96 (96B023); Davenport, 16 & i9 -VII-96 (96B011,
96B020, 96B023); Reardan, 16-VII-96 (96B014); PlERCcE—Graham, 25-V-96 on unknown grass (96T001);
WHITMAN—Colton, 31-VII-96 on H. vulgare (96T358A); Hay, 6-VII-95 (95G307); Pullman, 5-VII-94, 19-
VII-96 (94T022, 96T320A); Ridpath, 6-VII-95 (95G299+, 95G3007); Rosalia, 16-VII-96 (961294);
YAKIMA—Sunnyside, 28-VI-95 & 5-VII-95 (95G272, 95G286); Zillah, 28-VI-95 (95G270); 25-VI-95 on
D. glomerata (95G250); YIR, nr Signal Peak, 25-VI-95, on Agropyron spicatum (95G416); Chinook Pass,
24-VII-97, on Poa gracillima (97G260); 24-VII-97, on Poa nervosa (97G268).

Undetermined spp.: WA, GARFIELD—Ruckert Rd & Fitzgerald Rd, 15-VII-96 on Hordeum sp., W. Turner &
J. Rumph collectors (96J069A); KinGc—Stampede Pass, 8-VIII-96 on Caltha biflora (96G447); PEND
OREILLE—Le Clerc WLA, 3-VIII-95 on unknown grass (95T401).

Aphidius colemani Viereck
Aphis nerii Boyer de Fonscolombe: WA, YAKIMA—Union Gap, 23-X-95 on Asclepias speciosa (95G725).
Brachycaudus helichrysi (Kaltenbach): WA, WHITMAN—Pullman, 28-VI-96 on Anthemis arvensis (96T 162).
Hyadaphis foeniculi (Passerini): WA, ISLAND—Clinton, 4-VI-96 on Lonicera sp. (96G156).
Lipaphis erysimi (Kaltenbach): WA, WHITMAN—Pullman, 3-VI-96 on unknown plant (96T 1637).
Macrosiphum euphorbiae (Thomas): WA, WHITMAN—Pullman, 3-VII-96 on unknown plant (96T1637).
Myzus persicae (Sulzer): WA, SNOHOMISH—Monroe, 24-VII-96 on Pisum sativum (9615947).
Myzus sp.: WA, SPOKANE—Spokane, 11-VII-96 on Antirrhinum majus (96T258); WHITMAN—Pullman, 14-IX-

93, on Forsythia (93T019).

Undetermined sp.: WA, KING [on Hibiscus sp.*].

Aphidius ervi Haliday

Acyrthosiphon kondoi Shinji: 1D, SHOSHONE—Thompson Pass, 29-VII-95 on Melilotus alba (95G4667); WA,
KLICKITAT, Hwy 14 nr Alderdale, 14-V-96 on Melilotus sp. (96G0337).

Acyrthosiphon lactucae (Passerini) [on Lactuca_ serriola]: ID, BENEWAH—McCroskey SP, 31-VIII-95
(95T514+); WA, AsoTiIn—Anatone, 28-VI-96 (96T088); Asotin, 28-VI-96 (96T068); Asotin Cr WLA, 5-
VII-96 & 8-VIII-96 (96T 144, 96T419); Couse Cr, 10-VIII-95 (95T422); Snake River Rd, 18-VII-95 & 28-
VI-96 (95T249+, 96T072); South Fork Asotin Cr, 22-VII-95 & 11-VII-96 (95T468, 96T230); BENTON—
Prosser, 17-VII-95 (95G359+); 24-VI-96 & 8-VIII-96 (96G270, 96G434); ALE, 19-V-95 (95G071);
CHELAN—Chelan SP, 11-[X-97 (97K038); SPOKANE—Plaza, 16-IV-96 (96T279); WHITMAN—Pullman, 16,
30-VII-96 & 1, 8-VIII-96 (961304, 961362, 96T382, 961436); Ridpath, 6-VII-95 (95G296); YAKIMA—
Mabton, 28-VI-95 (95G265); Sunnyside, 22-V-95 (95G076); Wenas Lk, 11-VI-97 (97G055).

Acyrthosiphon pisum (Harris) [on Medicago sativa unless otherwise indicated]: ID, SHOSHONE—Thompson
Pass, 29-VII-95 on Melilotus alba (95G466+); MT, SANDERS [on Lathyrus odoratus*|; WA, ADAMS—
Macmaranon Rd, 2-VIII-96 (96B026); WSU-Othello, 26-VI-95 (95B008); AsoTtiIn—Couse Cr & Snake
River Rds, 18-VII-95 on Helianthus annuus (95T252); BENTON [on M. sativa & Trifolium pratense*| —
28-VI-95 (5G277+); Prosser, 27-VI-96 on Vicia angustifolia (96A009); 23-VII-9, on T. pratense (96G391);
WSU-Prosser, 26, 28-IV-97 & 14-V-97 (97G014, 97G015, 97G030); Whitstran, 1-V-97 (97G019);

692 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Co_LuMBIA—Hwy 124 & Tucannon Rd, 5-X-95 (95G678+); GRANT—WSU-Royal Slope, 26-VI-95
(95B003); IsLAND—South Whidbey, 4-VI-96 on Trifolium sp. (96G161); KLICKITAT—Hwy 14 nr Alderdale,
14-V-96 on Melilotus sp. (96G0337+); Roosevelt, 24-IV-96, on Melilotus sp. (96GO147+); LINCOLN—Lk
Roosevelt, 31-VII-97 on M. lupulina (97G304); SNoHOMISsH—Monroe, 24-VII-96 on Pisum sativum
(96T594+); SPOKANE—Four Lk, 25-VII-97 on Allium sp. (97KO005); WHITMAN—Pullman, 16, 30-VII-96 on
P. sativum (961303, 96T359); 14-IV-96 on Trifolium sp. (96T272); 26-VI-96 on Vicia sp. (96T063) &
unknown legume (96T064), & 5-VII-96 on Lathyrus oderata (96T178); Ridpath, 6-VII-95 on Lactuca
serriola (95G296), Medicago sativa (95G297), & Vicia sp. (95G302); YAKIMA—Outlook, 1-VI-96
(96K059); Rimrock Lk, 23-VU-96 on Melilotus sp. (96G383); Wenas Valley, 30-V-96 (96G111); YIR,
Signal Peak, 25-VII-95 on Vicia sp. (95G408) & 19-IX-95 on Melilotus sp. (95G625*).

Aphis craccivora Koch: WA, WHITMAN—Pullman, 25-VII-95 on Capsella bursa-pastoris (951331); 14-1V-
96, on Trifolium sp. (96T271).

Aphis sp.: ID, LATAH [on Solanum Lycopersicon *|; WA, ASOTIN—De Spain Springs, 1 1-VI-96 on Umbelliferae
sp. (96T254).

Aspidophorodon longicauda (Richards): WA, SKAMANIA—South Prairie, 11-VU-95 on Spiraea sp. (95G343).

Brachycaudus helichrysi (Kaltenbach) [on Amsinckia sp.]: WA, AsoTiIN—Anatone, 28-VI-96 (96T089);
FRANKLIN—Eureka, 14-V-96 (96G041); SkAGIT—Campbell Lk, 4-VI-96 on Prunus sp. (96G1657).

Brevicoryne brassicae (L.): WA, BENTON [on Brassica napus*).

Chaetosiphon nr hottesi Stroyan: WA, CLARK—Vancouver, 14-I[X-95 on Fragaria sp. (95VO002).

Diuraphis noxia (Kurdjumoy) [on Triticum aestivum unless otherwise indicated]: WA, AsoTIN [on Avena
fatua*| AsoTin—Anatone, 22-VII-97 on Hordeum vulgare (97JO007—97J026); BENTON [on H. vulgare & T.
aestivum* |—Wsvu-Prosser, 28-V-96 (on H. vulgare, 96G087, 96G088), 24-VII-97 (97G274); KLICKITAT—
Rock Cr, 30-V1-95 (95G283); Bickleton, 7-VII-97 (97G223, 97G238); Cleveland, 7-VII-97 (97G232,
97G234); WALLA WALLA—Hadley, 11-VI-96 on H. vulgare (96G240); WHITMAN—Pullman, 9-VII-96 on
H. vulgare, W. Turner & J. Rumph collectors (96J053—96J055); YAKIMA [*].

Hayhurstia atriplicis (L.) [on Chenopodium alba\: WA, BENTON—-WSU-Prosser, 18 & 25-VII-97 (97F238,
97F262); WHITMAN—Pullman, 16-VII-96 (96T302).

Hyalopteroides humilis (Walker) [on Dactylis glomerata|: WA, SKAGIT—Bow, 5-V-96 (96K026); Edison, 5-
V-96 (96K027).

Hyalopterus pruni (Geoffroy): WA, SkKAGIT—Campbell Lk, 4-VI-96 on Prunus sp. (96G1657#).

Hyperomyzus lactucae (L.): WA, YAKIMA—Yakima Arboretum, 26-IX-95 on Sonchus oleraceus (95G6587).

Hyperomyzus nigricornis (Knowlton): ID, BENEWAH—McCroskey SP, 31-VII-95 on Lactuca_ serriola
(95T514+); WA, GARFIELD—Stentz Springs, 22-VII-95 on Senecio sp. (95T456); YAKIMA—YIR, Cedar Cr,
19-IX-95 on Agoseris sp. (95G622).

Illinoia davidsoni (Mason): WA, KLickirAat—Trout Lk, 11-VH-95 on Rubus parviflorus (95K069).

Illinoia spp.: WA, Kirtrras—Quartz Mtn, 25-VII-96 on Rhododendron albiflorum (96G405); PIERCE—RNP,
Tipsoo Lk, 16-VIII-95 on Spiraea densiflora (95A037); SkAcit—La Conner, 5-VI-96 on Rhododendron
sp. (96G193); SNoHOMISH—Monroe, 5-VI-96 on Chaenomeles sp. (96G201).

Macrosiphum albifrons Essig: WA, PIERCE—RNP, Tipsoo Lk, 11-[X-96 on Lupinus sp. (96G547).

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow, 7-X-96 on Solanum sp. (961577); WA, ASOTIN—
De Spain Springs 11-VI-96 on Potentilla sp. (96T250), 1-VII-95 on Senecio sp. (95T381), 5 & 11-VII-
96 on Compositae (96T159, 961T249+); BENTON—Prosser, 15-V-96 on Tulipa sp. (96G045); 24-X-96 on
Medicago sativa (96G651); WSU-Prosser, 27-X-95 on Lamium amplexicaule (95K2137), 18-XI-95 on
Brassica napus (95G751+), & 1-X-97 on Malva neglecta (97G471+); GARFIELD—nr FR40 & FR44, 22-
VIII-95 on Osmorhiza chilensis (95T464+); SKAMANIA—Twin Falls CG, 4-VIII-95 on Potentilla sp.
(95K098+); YAKIMA—Bird Cr Meadows, 26-VI-95, on Castilleja (95A017+); YIR, Old Maid Canyon, 25-
VI-95 on Cynoglossum grande (95G440).

Macrosiphum gaurae (Williams): WA, AsoTin—Anatone, 28-VI-96 on Oenothera sp. (96T073).

Macrosiphum nr pallidum (Oestlund): WA, SKAMANIA—Twin Falls CG, 4-VIII-95 on Potentilla sp.
(95KO098+74).

Macrosiphum rosae (L.): WA, SNOHOMISH—Snohomish, 5-V1—96 on cultivated Rosa sp. (96G198).

Metopolophium dirhodum (Walker): OR, UMATILLA—Harris Pk, 11-VI-96 on Elymus glaucus (96G221); WA,
KLIckITAT—John Day Dam, 14-V-96 on Phalaris arundinacea (96G036+); LiINCOLN—Lk Roosevelt,
Goldsmith CG, 31-VII-97 on P. arundinacea (97G305).

Myzus nr ornatus Laing: WA, SkAGir—La Conner, 5-VI-96 on Trifolium pratense (96G1957+£).

Myzus persicae (Sulzer): WA, SkAGir—La Conner, 5-VI-96 on Trifolium pratense (96G1957+); BENTON—
WSU-Prosser, 1-X-97 on Solanum tuberosum (97G477+); WHItMAN—Pullman, 16-VII-95 on Raphanus
sativum (951248); 8-VIII-95, on Solanum nigrum (95T412); 31-VII-96 on Beta vulgaris (96T366).

VOLUME 102, NUMBER 3 693

Myzus sp.: WA, WHITMAN—Pullman, 14-IX-93 on Forsythia sp. (93TO19).

Ovatus crataegarius (Walker): WA, GRANT—Othello, 12-VII-95 on Mentha piperita (95BO10).

Phorodon humuli (Schrank): WA, SkAGIt—Campbell Lk, 4-VI-96 on Prunus sp. (96G1657+).

Rhopalosiphum maidis (Fitch): WA, BENTON—-WSU-Prosser, 18-XI-95 on Capsella bursa-pastoris (95G7507).

Rhopalosiphum padi (L.): WA, AsoTiN—De Spain Springs, 25-VII-96 on Aquilegia sp. (9613477); BENTON
[on Triticum aestivum*]|; DouGLAS—Waterville, 12-VIII-95 on 7. aestivum (95B016); FRANKLIN—nr
Kahlotus, 26-X-95 on T. aestivum (95K2077+); GRANT—Ephrata, 8-VII-97 on Zea mays (97K058); WALLA
WALLA—Prescott, 30-X-97 on Phalaris arundinacea (97G516).

Sitobion avenae (FE) [on Triticum aestivum unless otherwise indicated]: ID, BOUNDARY—Moyie Springs, 29-
VII-95, on Avena sativa (95G445); SHOSHONE—Thompson Pass, 29-VII-95 on Poa nervosa (95G470) &
Arabis sp. (95G475+); WA, ADAMs—Othello, 1-VII-96 (96B004, 96B007), WSU-Othello, 26-VI-95, 1-VI-
96 (95B009, 96B009); Asotin [*]—Clarkston, 18-VII-95 on Gramineae (95T255); Field Springs SP, 5-
VIU-93 on Gramineae (93T013); Couse Cr, 18-VII-95 on Setaria sp. (95T259); Anatone, 12-VII-94
(94T023); BENTON [on Hordeum vulgare & T. aestivum* |—Prosser, 25, 28 & 30-VI-95, (95G253, 95G274,
95G276, 95G285); 17-VII-95 (95G352, 95G354, 95G355, 95G358); 24-VI-96 (96G271); 1 & 8-VII-96
(96G312 & 96G328); WSU-Prosser, 20-I-96 on Capsella bursa-pastoris (96G003+); 18-VII-97 on Setaria
lutescens (97G243); Richland, 28-VI-95 (95G279); CoLuMBIA—Dayton, 16-VI-94 (94T020); GRANT—
WSU-Royal Slope, 26-VI-95 (95B007); FRANKLIN—Pasco, Star School, 6-VII-95 (95G289); KLICKITAT—
Bickleton, 9-VII-95 (95G318): Cemetery Rd nr Bickleton, 7-VII-97 on Hordeum vulgare (97G2267);
KitrirAs—S. Fork Manastash Cr, 24-VII-9, on Agropyron caninum (95G383); Ellensburg, 9-VII-96
(96G331); Buck Meadows, 9-VII-96 on Dactylis glomerata (96G346); Stampede Pass, 8-VII-96 on Phleum
sp. (96K142); LINcoLN—Almira, 26-VI-95 (95B002); Davenport, 16 & 19-VII-96 (96B011, 96B020);
Harrington, 19-VII-96 (96B023); Reardan, 16-VIH-96 (96B014); Wilbur, 19-VII-96 (96B019); PrERCE—
Graham, 25-V-96 on Gramineae (96TO01); RNP, Tipsoo Lk, 11-IX-96 on Poa sp. (96G543); WHITMAN
[*]—Hay, 6-VII-95 (95G307); Pullman, 5-VII-94 (94T022); Colton, 31-VII-96 on H. vulgare (96T358A);
Rosalia, 16-VII-96 (96T294); YAkIMA—Grandview, 5-VuI-95 (95G286); 23-V-96 on D. glomerata
(96G074); Zillah, 28-VI-95 (95G270, 95G272); 28-VI-95 on Avena sativa (95G271).

Uroleucon (Lambersius) sp.: WA, KirrirAs—Lost Lk Trailhead, 6-VIUI-97 on Achillea millefolium (97G318).

Undetermined spp.: WA, BENTON—Prosser, 25-X-95 on Berberis aquifolium (95G732); 29-X-95 on Senecio
vulgaris (95G735); CHELAN—Stevens Pass, 16-IX-95 on Gramineae (95K137); KLICKITAT—Cleveland, 7-
VII-97 on Lotus pinnatus (97G233); WHITMAN—Pullman, 1-VII-96 on Penstemon sp. (96T125), 5 & 9-
VII-96 on Geranium viscosissinum (961171, 961228); 9-VII-96 on Vicia sp. (96T229).

Aphidius nr hortensis Marshall
Ericaphis gentneri (Mason): WA, CHELAN—Stevens Pass, 16-[X-95 on Spiraea densiflora (95K138).
Liosomaphis berberidis (Kaltenbach): OR MULTNOMAH & WA, YAKIMA [on Berberis sp.*].

Aphidius kakimiaphidis Smith

Nasonovia alpina (G&P): WA, YAKIMA [on Mimulus lewisii*].

Nasonovia aquilegiae (Essig) [on Aquilegia formosa or sp.]: WA, BENTON—Prosser, 24-VI-96 (96G278);
Kittitas [+*]—Buck Meadows, 16-VIII-96 (96G501); YAKIMA—Yakima Arboretum, 3-VI-96 (96G249);
YIR, Old Mead Canyon, 25-VII-95 (95GG441).

Nasonovia cynosbati (Oestlund): WA, Kitriras—Lost Lk, 7-VI-94 on Ribes sp. (94K026).

Nasonovia nr cynosbati (Oestlund): WA, YAKIMA—YIR, nr Mt. Adams Lk, 4-VIII-95 on Ribes viscosissimum
(95K089).

Nasonovia nr houghtonensis (Palmer): WA, YAKIMA—Nile, 28-VII-97 on Collomia linearis (97G282).

Nasonovia wahinkae (Hottes): WA, Kittiras—Buck Meadows, 11-IX-95 on Aconitum columbianum
(95GS557).

Nasonovia sp.: MT, SANDERS [on Castilleja sp.*]; WA, Lewis—RNP, Stevens Canyon, 23-VII-96 on
Pedicularis sp. (96G3727).

Aphidius lupini Liu

Macrosiphum albifrons Essig [on Lupinus sp.]: WA, BENTON—Prosser 22-VI-95 & 24-VI-96 (95G249,
96G276); Kittrras—S. Fork Manastash Cr, 24-VII-95 (95G390); Quartz Mtn 25-VII-96 (96G408);
PIERCE—RNP, Tipsoo Lk, 11-[X-96, 5-IX-97 (96G547, 97G362); WHITMAN—Pullman, 1-VII-96 (96T129);
YAKIMA [*]—YIR, nr Howard Lk, 19-[X-95 (95G638) & nr Mt Adams Lk, 25-VII-95 (95G419): Little
Naches River, nr Timothy Meadows, 9-IX-97 (97G389).

Aphidius matricariae Haliday

Aphis fabae Scopoli [on Cirsium arvense]: WA, AsoTin—Sangster Rd, 22-VII-97 (97J003++, Turner & Rumph
collectors); KitrrrAs—Buck Meadows, 11-IX-95 (95G580).

694 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Aphis helianthi Monell: WA, YAKIMA—Chinook Pass Hwy & Bald Mt Rd, 23-X-97 on Cirsium arvense
(97G499).

Aphthargelia symphoricarpi (Thomas): WA, Kittiras—Lost Lk, 6-VIII-97 on Silene douglasii (97G345).

Brachycaudus cardui (L.): WA, ASoTIN—Sangster Rd, 22-VII-97 on Cirsium arvense, Turner & Rumph
collectors (97J0037).

Brachycaudus helichrysi (Kaltenbach): WA, BENTON—WSU-Prosser, 1-X-97 on Solanum tuberosum
(97G478); KinG [on Senecio vulgarais*]; KLIcKirAt—Bickleton, 30-V1—95 on Chrysanthemum sp.
(95G282); WHITMAN—Pullman, 26-VI-96, on Achillea millefolium (96T065).

Brevicoryne brassicae (L): WA, BENTON—-WSU-Prosser, 6-VII-95 on Brassica napus (95A024).

Capitophorus elaeagni (del Guercio) [on Cirsium arvense or sp.]: WA, Kittiras—Buck Meadows, 11-IX-97
(97G437+); YAKIMA—Mabton, Morgan Lk, 28-VI-95 (95G266); Little Naches River (FR 789 & FR 1914),
9-IX-97 (97G409); 9-IX-97 (97G408); Little Naches River, nr Timothy Meadows, 9-IX-97 (97G394).

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, BENTON—Rattlesnake Hills, 17-VII-95 (95G3577);
WSU-Prosser, 30-IX-97 (97F457+, 97463+); YAKIMA—Mabton Grade, 31-X-97 (97G5177).

Dysaphis plantaginea (Passerini): WA, CHELAN—-WSU-Wenatchee, 30-V-96 on Pyrus malus (96RO117#).

Hyalopterus pruni (Geoffroy): WA, SkKAGIT—Campbell Lk, 4-VI-96 on Prunus sp. (96G1657+).

Illinoia corylina (Davidson) [det. by A. Jensen]: WA, WHITMAN—Pullman, 14-VIII-96 on Aquilegia sp.
(96T442).

Macrosiphum euphorbiae (Thomas): WA, BENTON—-WSU-Prosser, 1-X-97 on Malva neglecta (97G4717).

Myzus persicae (Sulzer): ID, BONNER—Sandpoint, 30-IV-97 on Viola tricolor (97T006); OR, UMATILLA—
Hermiston, 23-VII-93 on unknown plant, A. Jensen collector (93J005); WA, BENTON—WSU-Prosser, 27-
X-95 on Malva neglecta (95K212+), 1-X-97 on Chenopodium album (97G470*), Solanum sarrachoides
(97G4721), & S. tuberosum (97G476); KLickiIrat—John Day Dam, 28-IV-95 on Veronica anagallis-
aquatica (95G009%+); SPOKANE—Spokane, Manito Pk, 19-IX-96 on Syngonium sp. (96T538); WHITMAN—
Pullman, 22-VII-95 on Solanum lycopersicon (9512997).

Myzus sp.: WA, WHITMAN—Pullman, 14-IX-93 on Forsythia sp. (93T019).

Ovatus crataegarius (Walker): ID, BONNER—Sandpoint, 10-IV-97 on Origanum sp. (97T003); WA, ADAMS—
Othello, 19-VI-96 & 16-VII-96 on Mentha piperita (96B006, 96B010).

Phorodon humuli (Schrank) [on Humulus lupulus|: WA, BENTON [*]—WSU-Prosser, 9-[X-95 (95K154).

Rhopalosiphum insertum (Walker): WA, CHELAN—-WSU-Wenatchee, 30-V-96 on Pyrus malus (96RO117).

Rhopalosiphum padi (L.) [on Triticum aestivum]: WA, BENTON—Rattlesnake Hills, 17-VII-95 (95G3577);
WSU-Prosser, 30-IX-97 (97F457+, 97F463+); YAKIMA—Mabton Grade, 31-X-97 (97G517+).

Sitobion avenae (F.): WA, BENTON—-WSU-Prosser, 11 & 18-VII-97 on Triticum aestivum (97F225, 97F243).

Undetermined sp.: WA, BENTON—-WSU-Prosser, 29-X-95 on Senecio vulgaris (95G735).

Aphidius nigripes Ashmead
Macrosiphum euphorbiae (Thomas): WA, Kitrrras—Manastash Rd & FR112, 9-VU-96 on Agoseris elata
(96G341) & Arnica cordifolia (96G332),; Lost Lk, 2-IX-95 on Aster sp. (95G528) & Castilleja sp.
(95G527), & 11-IX-95 on Arabis sp. (95G568) & Monardella odoratissima (95G5697).
Macrosiphum clydesmithi Robinson: WA, YAKIMA—Clear Lk, 15-VI-96 on Holodiscus discolor (96K081).

Aphidius ohioensis Smith

Macrosiphoniella ludovicianae (Oestlund) [on Artemisia ludoviciana]: CA. SHASTA [*]; WA, AsoTIN—Asotin
Cr WLA, 5-V11—96, (96T138); YAKIMA—Wenas Lk, 17-VI-97 (97G090).

Macrosiphoniella tanacetaria (Kaltenbach): WA, PIERCE—RNP, Tipsoo Lk, 5-IX-97 on Tanacetum vulgare
(97G369).

Macrosiphum creelii Davis: WA, Kittiras—Manastash Cr Rd, mi marker 8, 16-VIII-96 on Vicia sp. (96G492).

Macrosiphum sp.?: ID, LATAH—Moscow Mtn, 6-VII-96 on Holodiscus discolor (961182).

Macrosiphum valerianae (Clarke) [on Epilobium angustifolium|: WA, Kirrrras—Lost Lk, 24-VI-95
(95G399++); S. Fork Manastash Cr, 24-VII-95 (95G381); YAKIMA—YIR, Signal Peak, 11-VII-95
(95G322¢).

Microsiphoniella artemisiae (Gillette): WA, AsoTiIn—Asotin Cr HMU, 5-VII-96 on Artemisia ludoviciana
(96T141).

Uroleucon adenocaulonae (Essig): WA, KLick1rAt—Trout Lk, 29-VI-97 on Adenocaulon bicolor (97G163).

Uroleucon ur arnesense Robinson: MT, SANDERS—18 mi E of Thompson Pass, 29-VII-95 on Solidago, sp.
(95G457).

Uroleucon nr epilobii (Pergande): WA, SPOKANE—Mt Spokane SP, 27-VII-95 on Epilobium angustifolium
(95T358).

Uroleucon erigeronense (Thomas) [on Conyza canadensis unless otherwise indicated]: WA, CHELAN—
Leavenworth, 16-IX-95 (95K141); Kitrrras—Lost Lk, 6-VIII-97 on Haplopappus hirtus (97G3407), &

VOLUME 102, NUMBER 3 695

11-IX-95 on Achillea millefolium (95G574); Manastash Rd & Rd 112, 11-IX-95 on Cirsium arvense
(95G5647+); S. Fork Manastash Cr, 11-IX-95 on Achillea millefolium (95G563), & 24-VII-95 on Crepis sp.
(95G3867); LINCOLN—Lk Roosevelt, Goldsmith CG, 31-VII-97 (97G302); PIERCE—RNP, 8-IX-95 on Aster
sp. (95K106) & on Senecio sp. (95K1207), Tipsoo Lk, 5-IX-97 on Aster sp. (97G364+, 97G366), & 24-
VII-97 on Compositae (97G252); YAKIMA—Grandview, 9-VII-95 (95G314); Pleasant Valley CG, 24-VII-
97 on Crepis rucinata (97G258); Sunnyside, 23-X-95 & 13-X-96 (95G707, 96G645); YIR, nr Mt Adams
Lk, 4-VHI-95 on Eriophyllum lanatum (95K092), Signal Peak, 11-VII-95 on Aster sp. (95K049+), Cedar
Cr, 19-[X-95 on Achillea millefolium (95G620+), Klickitat Canyon (95G6287+), & White Deer Cr, 11-VII-
95 on Crepis modocensis (95K0487*).

Uroleucon nr erigeronense (Thomas): WA, LINCOLN—Lk Roosevelt, Goldsmith CG, 31-VII-97, on Aster sp.
(97G307); SPOKANE—Peone, 20-VII-95 on Solidago sp. (95T278); YAKIMA—Mabton Grade, 23-V-96, on
Balsamorhiza careyana (96G069); YIR, nr Mt Adams Lk, 4-VIII-95 on Hieracium sp. (95K0837).

Uroleucon escalantii (Knowlton): WA, BENTON—ALE, 8-VI-95 on Chrysothamnus viscidiflorus (95G148).

Uroleucon nr escalantii (Knowlton): WA, KLICKITAT—Box Springs, nr Bickleton, 29-VI-97 on Chrysothamnus
sp. (97G178). ;

Uroleucon gigantiphagum Moran: WA, YAKIMA—YIR, Signal Peak, 11-VII-95 on Solidago sp. (95G329).

Uroleucon ivae Robinson [on Iva xanthifolia|: WA, BENTON [*]—ALE, Lower Snively Spring, 12-VIII-95
(95K104); YAKIMA [*]—Grandview, 7,9 & 22-VII-95, & 30-VII-96, (95G309, 95G313, 95G366 &
96G431); Harrah, 29-IX-97 (97G465): Selah, 10-VIII-95 (95G500); Wenas-Ellensburg Rd, Ellensburg Pass,
7-VIU-96 (96G317).

Uroleucon katonkae (Hottes) [on Aster sp.]: WA, YAKIMA—Wenas Lk, 17-VI-97 (97G099); YIR, Lost Springs,
11-VII-95 (95K0497).

Uroleucon olivei Moran: WA, BENTON—WSU-Prosser, 21-VII-94 on Iva xanthifolia (94R012).

Uroleucon russellae (HRL) [on Anaphalis margaritacea unless otherwise indicated]: WA, LEwis—RNP., 8-
IX-95 on Senecio sp. (95K120+); SKAMANIA—June Lk, 4-VIII-95 (95K101); SPOKANE—Mt Spokane SP,
20-VH-95 & 24-VII-95 (95T294, 95T494); YAKIMA—YIR, Lost Springs (95G412) & Soda Springs
(95G423), 25-VII-95.

Uroleucon sonchi (L.): WA, YAKIMA—Grandview, 17-VII-95, 8-VII-96 on Sonchus oleraceus (95G348,
96G326); Sunnyside, 23-X-95 on Sonchus sp. (95G703); Satus, 1-VI-96 on Sonchus sp. (96K064).

Uroleucon suzannae ROBINSON: WA, Kittiras—Lost Lk, 6-VII-97 on Haplopappus hirtus (97G340*#).

Uroleucon spp.: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Adenocaulon bicolor (951527, 951536): WA,
CHELAN—Chiwawa Canyon, 3-VIII-96 on Adenocaulon bicolor (96R044); Kirrrras—Lost Lk, 11-IX-95
& 6-VII-97 on Aster sp. (95G579); Lost Lk Trailhead, 6-VIII-97 on Agoseris retrorsa (97G319); S. Fork
Manastash Cr, 24-VII-95 on Hieracium sp. (95G389); LINCOLN—Lk Roosevelt, Goldsmith CG, 31-VI-97
on Tragopogon dubius (97G303); PIERCE—RNP, Tipsoo Lk, 5-I[X-97 on Anaphalis margaritacea (97G367);
SPOKANE—Mt Spokane SP, 24-VIII-95 on Centaurea pratensis (95T477); YAKIMA [on Iva xanthifolia *|—
Hwy 410, Fife Peak Viewpoint, 24-VII-97 on Senecio serra (97G270+); Raven’s Roost, 28VII-97 on
Hieracium gracile (97G2987); Sawmill Flats CG, 26-VII-97 on Aster sp. (97G287); Union Gap, 23-X-95
on Centaurea maculosa (95G720); YIR, Big Spring, 4-VIII-95 on Agoseris elata (95KO090+), Lost Spring,
25-VII-95 on Cichorium intybus (95G413), Signal Peak, 11-VH-95 on Eriophyllum lanatum (95KO055), 25-
VII-95 on Achillea millefolium (95G410, 95G417); 19-[X-95 on Grindelia sp. (95G618), Signal Peak Guard
Station, 19-IX-95 on Antennaria sp. (95G616), & Windy Point, 19-IX-95 on Aster sp. (95G629).

Aphidius nr pisivorus Smith
Acyrthosiphon pisum (Harris): WA, BENTON—-WSU-Prosser, 11-VII-97 on Melilotus officinalis (97F2187*).

Aphidius polygonaphis (Fitch)

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola|: WA, WHITMAN—Kamiak Butte, 12-IX-95 (95T550);
YAKIMA—Sunnyside, 28-VI-95 (95G257).

Amphorophora parviflori Hill: WA, YAKIMA—N. Fork Tieton River & Rd 820, 27-VI-96 on Rubus parviflorus
(96G304).

Amphorophora sp.?: WA, LEwis—RNP, Stevens Canyon, 23-VII-96 on Arnica sp. (96G3597).

Aphis ceanothi Clarke: WA, YAKIMA—Windy Point CG, 3-VI-95 on Ceanothus integerrimus (95G107).

Aphis helianthi Monell: WA, YAKIMA—YIR, nr. Mt. Adams Lk., 25-VII-95 on Epilobium angustifolium
(95G402+¢).

Aphis nr oenotherae Oestlund: WA, GARFIELD—Stentz Springs, 14-IX-95 on Epilobium sp. (95T565).

Brachycaudus cardui (L.): MT, SANDERS—Thompson Pass, 17-VI-95 on Senecio sp. (95G1917#).

Capitophorus elaeagni (del Guercio): WA, KitrirAs—Frost Meadows, 11-[X-95 on Cirsium arvense
(95G5837).

Ericaphis wakibae (Hottes): WA, YAKIMA—Clear Lk, 15-VI-96 on native Rosa sp. (96K084, 96K0857).

696 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Hyperomyzus nigricornis (Knowlton): WA, YAKIMA—Bird Cr Meadows, 26-VI-95 on Hieracium sp.
(95A0167).

Hyperomyzus sp. [nr nigricornis (Knowlton) & sandilandicus (Robinson)| WA, KitrrirAs—S. Fork Manastash
Cr, 24-VII-95 on Hieracium albiflorum (95G388).

Iillinoia nr azaleae (Mason): WA, Y AKIMA—Raven’s Roost, 28-VI-97 on Rhododendron albiflorum (97G297).

Illinoia davidsoni (Mason): WA, LEwis—RNP, Stevens Canyon, 23-VII-96 on Arnica sp. (96G3597);
YAKIMA—Bird Cr Meadows, 26-VI-95 on Rubus parviflorus (95A019).

Illinoia sp. [on Vaccinium sp.]: ID, SHOSHONE—Thompson Pass, 29-VII-95 (95G472); WA, SKAMANIA—
Keenes Horse Camp, 11-VII-95 (95G337).

Macrosiphoniella ludovicianae (Oestlund) [on Artemisia ludoviciana|: WA, YAKIMA [on Artemisia
ludoviciana* |—YIR, Bedground Springs, 16-V-96 (96G0537+).

Macrosiphum clydesmithi (Robinson): WA, YAKIMA—YIR, Piscoe Meadows, 5-VIH-95 on Pteridium
aquilinum (95G4337).

Macrosiphum clydesmithi (Robinson): WA, SPOKANE—Mt Spokane SP, 20-VII-95 on unknown Polypodiaceae
(95T298); YAKIMA—Clear Lk, 15-VI-96 on Holodiscus discolor (96K081).

Macrosiphum euphorbiae (Thomas): MT, SANDERS—Thompson Pass, 17-VI-95 on Senecio sp. (95G1917);
WA, AsoTiN—De Spain Spring, 11-VII-96 on Potentilla sp. (961250); 11-VII-96, on unknown plant
(96T252); KittiraAs [on Rosa sp.*]|—Manastash Canyon, 2-1X-95 on Lactuca serriola (95G544); Lost Lk
Trail, 6-VIII-97 on Luina nardosmia (97G327); WHITMAN—Pullman, 22-VII-95 on L. serriola (95T300)
& 18-VI-96 on cultivated Rosa sp.(96T037); YAKIMA—Clear Lk, 15-VI—96 on native Rosa sp. (96K0857);
Bird Cr Meadows, 26-VI-95 on Hieracium sp. (95SA016*).

Macrosiphum nr euphorbiae (Thomas): WA, YAKIMA—YIR, Klickitat Canyon, 25-VII-95 on Cynoglossum
grande (95G418); Mill Cr Guard Station, 6-VI-95 on Lomatium nudicaule (95G115).

Macrosiphum pteridis (Wilson): OR, BENTON—McDonald Forest, 14-IV-94 on Holodiscus discolor, A. Jensen
collector (94JOO1).

Macrosiphum rosae (L.): OR, BENTON—Corvallis, 18-V-93 on unknown plant, A. Jensen collector (93J003).

Macrosiphum valerianae (Clarke): WA, YAKIMA—YIR, nr. Mt. Adams Lk., 25-VIH-95 on Epilobium
angustifolium (95G4027*).

Macrosiphum sp.: ID, BENEWAH—McCroskey SP, 7-VI-96 on native Rosa sp. 96TO05); WA, GARFIELD—
FR160 & FR40, 21-IX-95, on Streptopus sp. (9ST601); KittrrAs—Manastash Cr, mi marker 12, 6-VIII-
97 on Holodiscus discolor (97G314); YAKIMA—YIR, Piscoe Meadows, 25-VII-95 on Pteridium aquilinum
(95G4337+); Howard Lk, 19-IX-95 on P. aquilinum (95G640).

Nasonovia crenicorna (Smith & Knowlton): WA, WHITMAN—Pullman, 27-V-97 on Geranium viscosissimum
(97T035)

Nasonovia houghtonensis (G&P): WA, WHATCOM—Birch Bay, 29-V-93 on Oemleria cerasiformis, A. Jensen
collector (93J004).

Sitobion sp.. CANADA, B.C.—Vancouver, Univ. of British Columbia, 29-V-93 on Dicentra formosa, A.
Jensen collector (93J003).

Uroleucon nr arnesense Robinson: WA, CHELAN—Leavenworth, Icicle Cr, 1-VIII-95, on Solidago canadensis
(95G490).

Uroleucon nr erigeronense (Thomas): WA, KittirAs—Manastash Rd & Rd112, 9-VII-96 on Hieracium
albiflorum (96G340).

Uroleucon ivae Robinson: WA, YAKIMA—Sunnyside, 13-VIH-94 & 11-VII-97, on Iva xanthifolia (94GO009,
97G242).

Wahlgreniella nervata (Gillette): ID, SHOSHONE—Thompson Pass, 29-VH-95 on Pachistima myrsinites
(95G471).

Aphidius nr polygonaphis (Fitch)
Macrosiphum walkeri Robinson: OR, BENToN—Corvallis, 17-III-90 on Polypodium sp. (90J001).

Aphidius pulcher Baker
Acyrthosiphon pisum (Harris): WA, BENTON—-WSU-Prosser, 1-VII-96 on Melilotus officinalis (96G3117).

Aphidius rhopalosiphi DeStefani
Metopolophium dirhodum (Walker): WA, BENTON—-WSU-Prosser, 6-VIII-97 on Triticum aestivum (97F2267).
Sitobion avenae (FE): WA, BENTON—-WSU-Prosser, 11, 18 & 25-VII-97 on Triticum aestivum (assorted
collections between 97F204—97F277).

Aphidius nr rosae Haliday
Macrosiphum pallidum (Oestlund): WA, SPOKANE—nr Cheney, 23-V-95 on native Rosa sp. (95KO006).

VOLUME 102, NUMBER 3 697

Aphidius salicis Haliday

Cavariella aegopodii (Scopoli): WA, BENTON—-WSU-Prosser, 21-VII-97 on Anethum graveolens (97G247}+);
Kitritas—Reecer Cr, nr Ellensburg, 20-VI-95 on Angelica canbyi (95G203); Manastash Cr Rd, mi marker
12, 25-VI-97, on Osmorhiza occidentalis (97G132); Lost Lk, 6-VIII-97 on Osmorhiza chilensis (97G331);
Quilomene WLA, Crossover Rd & Colockum Ridge, 2-VH-97 on Osmorhiza purpurea (97G207); PEND
OREILLE—Tiger Meadows, 9-VII-96 on unknown Umbelliferae (96T219+); YAkIMA—Hwy 410, Fife Peak
Viewpoint, 24-VH-97 on Osmorhiza chilensis (97G271+); YIR, Signal Peak, 19-Ix-95 on Salix sp.
(95G623).

Cavariella pastinacae (L.) [on Heracleum lanatum unless otherwise indicated]: WA, AsoTIN—De Spain
Springs, 5-VII-96 & 25-VII-96 (96T 156, 96T350); ISLAND—N. Whidby Island, 5-VI-96 (96G171); KING—
Tinkham exit, 4-VI-96 (96G137); KirrirAs—Reecer Cr, nr Ellensburg, & S. Fork Manastash Cr, &
Manastash Rd, 20-VI-95, 24-VII-95, & 25-VII-96 (95G217, 95G393, 96G417); Taneum Lk, 11-IX-97 on
Angelica sp. (97G413); PIERCE—RNP, Ipsut Pass, 25-[X-97 (97G447); Mowich Lk, 25-IX-97 (97G450);
YAKIMA—Yakima, 28-VI-95 on Heracleum lanatum (95G269+); YIR, Klickitat Canyon, 23-VI-95
(95G235), & Signal Peak, 11-VII-95 (95G335).

Aphidius segmentatus Pike and Stary, new species
(Figs. 1-9)

Diagnosis.—The new species is distinguished by several characters, but especially a
short antenna (14—15 segments, noticeably widening to apex), and short distal abscissa
of Rl (= metacarpus). In Smith (1944), the new species keys to Aphidius ribis Haliday,
but the latter’s antenna is longer, not remarkably thickened, and the metacarpus is long.
Similar differences also distinguish the new species from a European species, A. sonchi
Marshall that attacks Hyperomyzus aphids (Stary 1963, 1966).

Etymology.—The name of the new species is derived from the somewhat unusual
segmentation of the flagellum.

Description.—Female. Head: Eye length 3X malar space. Tentorio-ocular line %
intertentorial line. Antenna 14—15 (prevalently 15) segmented, short, as long as head,
mesosoma and metasomal tergum | together, thickened to apex, segmentation of flagellum
rather distinct (Fig. 7). Flagellomeres with semierect setae (Figs. 4—6). Flagellomere 1
[F,] (Fig. 4) length 2 width, with O—1 longitudinal placodes, setae about 74 segment
diameter. F, similar to F,, with 2—3 placodes. Middle F (Fig. 5) length about 1.3 width,
distinctly wider than F,. Preapical F (Fig. 6) length about 2 width, wider than middle
E Mesosoma: Mesonotum glabrous, with rows of sparse setae indicating the effaced
notauli on the disc. Propodeum (Fig. 1) with rather small areola, carinae sometimes
indistinct to slightly indicated in upper portion. Forewing: (Fig. 8): Stigma length 3X
width; distal abscissa of R1 (metacarpus) equal or subequal to stigma width. Vein r slightly
longer than stigma width, and twice RS length; r-m equal to RS in length. Marginal setae
3X length of surface setae. Metasoma: Metasomal tergum 1 (Fig. 9) length 2.5 width
at spiracles; surface slightly rugose in anterior portion; anterolateral area costulate.
Genitalia (Fig. 2): Ovipositor sheath (Fig. 3) apex blunt. Coloration: Head brown; face,
gena, and mouthparts yellow, except brown apices of mandibles. Antenna brown; a ring
between pedicel and flagellum sometimes yellow. Mesosoma brown, propleuron yellow;
mesopleuron sometimes with yellow markings in lower portion. Tegula yellow. Wings
hyaline, venation brown. Legs light brown; coxa, trochanter and base of tibia yellow; a
dark tinge increasing from fore to hind legs, the latter the darkest. Metasoma: Tergum 1
and apical half of metasoma yellow, terga 2 and 3 and some times 4, with brown banding.
Ovipositor sheath brown. Length of body: about 2.2—2.7 mm.

Male.—Antenna 17—18 segmented. Coloration: head brown; mouthparts mainly (apices
of mandibles brown, palpi light brown). Mesosoma brown. Legs brown, base of tibia
yellow to light brown. Metasoma brown, tergum 1 yellow to light brown.

698 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-9. Aphidius segmentatus, 2 {illustrations not to equal scale]. 1, Propodeum. 2, Genitalia. 3, Ovi-
positor sheath. 4, Flagellomere 1 [F,]. 5, Middle flagellomere. 6, Preapical flagellomere. 7, Antenna [flagellum].
8, Fore wing, in part. 9, Metasomal tergum 1. Abbreviations: R1, distal abscissa of postmarginal vein [=
metacarpus]; RS, r, and r-m, wing veins.

VOLUME 102, NUMBER 3 699

Holotype.—?, USA, WA, YAKIMA Co., Yakama Indian Reservation, 25-VII-1995, G.
Graf collector, (collection code, 95G406). Host aphid: Hyperomyzus nigricornis
(Knowlton) on Hieracium cynoglossoides. Deposited in USNM.

Paratypes.—16 specimens, collection data same as holotype, deposited in part in
collections of WSU-Prosser (10 2, 4 d) and P. Stary, Ceské Budejovice, Czech Republic
(1 2,1); 11 specimens, USA, WA, Kittitas Co., L.T. Murray WLA, S. Fork Manastash
Creek, 24-VII-1996, on Hyperomyzus sp. (nr nigricornus or sandilandicus) on Hieracium
albiforum (96G388), deposited in part in collections of WSU-Prosser (6 ¢) and P. Stary,
Ceské Budejovice, Czech Republic (3 2, 2 ¢).

Other material reared from.—

Hyperomyzus lactucae (L.): WA, WHItMAN—Pullman, 12-IX-95, on Lactuca serriola
G5m559).

Hyperomyzus nigricornis (Knowlton) [on Hieracium sp. unless otherwise indicated]:
ID, LATAH—Moscow Mtn, 3-VII-96 on Compositae (96T399); WA, GARFIELD—FR40 nr
Teal Springs, 1-VIII-95 on Senecio sp. (95T378+); Pataha Cr Rd, 25-VII-95 (9513234);
Wickiup CG, 29-VIII-95 (95T508); KittiraAs—Lost Lk, 6-VIII-97 (97G317+, 97G344);
KLICKITAT—Cleveland, 7-VII-97 (97G229); PIERCE—RNP, Paradise, 8-[X-95 on Senecio
sp. (95K119), & Tipsoo Lk, 5-IX-97 (97G359); SPOKANE—Mt Spokane SP, 24-VIII-95
on Compositae (95T486); YAKIMA—Little Naches River, 5 mi W of Timothy Meadows,
9-IX-97 (97G388, 1 specimen); Hwy 410, Ravens’s Roost, 28-VII-97 (97G2987+); Pleasant
Valley CG, 24-VII-97 (97G2597); Sawmill Flats, 28-VII-97 (97G2887); Wenas Lk, 17-
VI-97 on Compositae (97G087); R, Signal Peak, 11-VII-95 (95K056).

Hyperomyzus sp. [nigricornis (Knowlton) or sandilandicus (Robinson)]: WA,
KitTTITAs—Manastash Rd & FR112, 11-IX-95 on Hieracium sp. (95G5597); L. T. Murray
Wildlife Reserve, South Fork Manastash Cr, 24-VII-95 on Hieracium albiflorum
(95G388).

Uroleucon erigeronense (Thomas): WA, KittirAs—Manastash Ridge, nr Lost Lk, 2-
IX-95 on Achillea millefolium (95G533¢£).

Aphidius smithi Sharma & Subba Rao
Acyrthosiphon pisum (Harris): WA, BENTON [on Medicago sativay;*].

Aphidius spp.

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola}: 1D, BENEWAH—McCroskey SP, 31-VIII-95
(95T534); WA, Asotin—Asotin, 10-VII-95 (95T419); CHELAN—Chelan SP, 11-IX-97 (97K038);
SPOKANE—nr Peone, 24-VIII-95 (95T470).

Acyrthosiphon macrosiphum (Wilson) [on Amelanchier alnifolia]: MT, LINCOLN—Libby, 6-VIH-96 (96G322);
WA, PEND OREILLE—Le Clere WLA, Panhandle CG, 3-VIII-95 (95T402).

Acyrthosiphon pisum (Harris): WA, BENTON [on Medicago sativa*]; Kittiras [on Rhododendron albiflorum*}).

Amphorophora geranii G&P: WA, SPOKANE—Turnbull National WLA, 25-V-95 on Geranium sp. (95T0307).

Amphorophora nr rubitox’ 1 Knowlton: WA, Is_AND—South Whidbey Island, 4-VI-96 on Rubus sp. (96G163).

Aphis armoraciae Cowen: WA, Kittitas [on Mertensia paniculata*}.

Aphis craccivora Koch: WA, WHITMAN—Pullman, 5-VII-97 on Trifolium sp. (96T170).

Aphis fabae Scopoli: WA, AsoTiN—Field Springs SP, 18-VII-95 on Cirsium arvense (951272).

Aphis helianthi Mo ell: WA, BENTON—-WSU-Prosser, 21-VU-97 on Anethum graveolens (97G247*).

Aphis holodisci Robinson: WA, AsoTIN—nr FR40 & FR44, 26-VI-96 on Holodiscus discolor (96T052).

Aphis nigratibialis Robinson: WA, YAKIMA—Nile, 28-VII-97 on Cornus stolonifera (97G280).

Aphis varians Patch: WA, PIERCE [on Epilobium angustifolium* }.

Aphis (Zyxaphis) sp.: WA, KLick1rat—Frost Meadows, 11-[X-97 on Artemisia tridentata (97G435).

Aspidophorodon longicauda (Richards): WA, SKAMANIA—South Prairie, 8-IX-94 on Spiraea douglasii
(94R192).

Brachycaudus helichrysi (Kaltenbach): WA, SPOKANE—Plaza, 16-VII-96 on unknown plant (96T278).

Brachycaudus tragopogonis (Kaltenbach): WA, BENTON—WSU-Prosser, 25-VH-97 on Tragopogon dubius
(97F 266).

700 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Capitophorus elaeagni (del Guercio): WA, YAKIMA—Rimrock Lk, 14-[X-95 on Cirsium vulgare (95G589).

Carolinaia howardii (Wilson): WA, WHITMAN—Pullman, 1-VII-96 on unknown plant (96T123).

Chaetosiphon fragaefolii (Cockerell): WA, PEND OREILLE—Le Clerc WLA, 3-VIII-95 on native Rosa sp.
(951398); STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa sp. (96T2097).

Chaetosiphon nr hottesi Stroyan: WA, SPOKANE—Mt Spokane SP, 24-VIII-95 on native Rosa sp. (9ST4917£).

Chaetosiphon tetrarhodum (Walker): WA, SPOKANE—Mt Spokane SP, 24-VIII-95 on native Rosa sp.
(95T49174).

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, AsoTiIN—Field Springs SP, 20-VI-95 (95T158);
BENTON [*].

Ericaphis gentneri (Mason) [on Crataegus sp.]: WA, CHELAN—Stevens Pass, 16-I[X-95 on Spiraea densiflora
(95K 138); Kirriras—Coleman Canyon, 2-VII-97 (97G214); SPOKANE—Chapman Lk, 16-VII-96 (96T282).

Ericaphis nr gentneri (Mason): WA, WHITMAN—Pullman, 6-VIII-96 on Crataegus (96T409).

Ericaphis scammelli (Mason): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa
sp. (96T2097).

Ericaphis wakibae (Hottes): WA, YAKIMA—N Fork Tieton River, 15-VI-96 on native Rosa sp. (96K090*).

Euceraphis gillettei Davidson: WA, KitrirAs—L.T. Murray WLA, S Fork Manastash Cr, 24-VI-95 on Alnus
sp. (95G3947+).

Hyperomyzus nigricornis (Knowlton): WA, Kittiras—Lost Lk, 6-VIII-97 on Hieracium sp. (97G344).

Hyperomyzus sp.: WA, BENTON [on Sonchus oleraceus*].

Illinoia crystleae bartholomewi (Essig) [det. by A. Jensen]: WA, Kittiras—Lost Lk, on Lonicera involucrata
(97G336).

Illinoia nr crystleae (Smith & Knowlton): WA, Kitriras—Lost Lk, 7-VII-94 on Lonicera involucrata
(94K028).

Illinoia davidsoni (Mason): WA, GARFIELD—Alder Thicket, 22-VIII-95 on Arnica latifolia (95T450).

Illinoia rhododendri (Wilson): WA, Kittitas [on Rhododendron albiflorum* |.

Illinoia spp.: WA, GARFIELD—Stentz Springs, 1-VIII-95 on Aquilegia formosa (9513627); SkAGiIt—La Conner,
6-VI-96 on Rhododendron sp. (96G193); SKAMANIA [on Vaccinium sp.*].

Liosomaphis berberidis (Kaltenbach): WA, SkAGiIr—La Conner, 5-VI-96 on Berberis sp. (96G192).

Macrosiphoniella ludovicianae (Oestlund): WA, Asotin—Anatone, 18-VIH-95 on Artemisia ludoviciana
(Osik257)):

Macrosiphum californicum (Clarke) [on Salix sp.]: WA, Kittitas [+*]; YAKIMA—YIR, Lost Springs, 11-VII-
95 (95KOS52).

Macrosiphum clydesmithi (Robinson): WA, CHELAN—Chiwawa Canyon, 5-IX-95 on Pteridium aquilinum
(95RO0267).

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow Mtn, 6-VII-96 on unknown plant (961187); 6-VII-
96 on Compositae (96T185); MT, SANDERS—Thompson Pass, 17-VI-95 on Hieracium sp. (95G185), &
Arabis sp. (95G190+); WA, GARFIELD—Stentz Springs, 1-VII-95 on Arnica latifolia (95T370); Spruce
Springs, 22-VIII-9, on Ranunculus sp. (95T459); Kittiras—Manastash Rd 25-VII-96 on Ranunculus sp.
(96G416); Taneum Lk, 11-IX-97 on Silene sp. (97G415); Lost Lk, 6-VIII-97 on Angelica canbyi
(97G330+); KLickirat—Cleveland Pk, 7-VII-97 on Lupinus sp. (97G230); SPOKANE—Mt Spokane SP, 29-
VI-95 on Ranunculus sp. (95T190); WHITMAN—Pullman, 5-VII-96 on cultivated Rosa sp. (9611727);
YAKIMA—N. Fork Tieton River, 27-VI-96 on Hieracium scouleri (96G297); Buckhorn Meadows, 14-IX-
95 on Trautvetteria caroliniensis (95G597).

Macrosiphum pallidum (Oestlund): MT, SANDERS—Thompson Pass, 17-VI-95 on Arabis sp. (95G1907#).

Macrosiphum rhamni (Clarke) [det. by A. Jensen]: WA, Kinc—Stampede Pass, 8-VIII-96, on Pteridium
aquilinum (96G441); KLickiIrat—Trout Lk, 29-VI-97 on Pteridium aquilinum (97G169).

Macrosiphum rosae (L.): WHITMAN—Pullman, 5-VII-96 on cultivated Rosa sp. (9611727).

Macrosiphum tuberculaceps (Essig): WA, YAKIMA—Hell Crossing CG, 11-[X-96 on Achlys triphylla
(96G517); N. Fork Tieton River, 15-VI-96 on A. triphylla (96K096).

Macrosiphum sp.: WA, Ktinc—Kirtiras—Waptus Lk, 27-VII-95 on Spiraea douglasii menziesii (95KO75),;
Manastash Cr, mi marker 12, 25-VI-97 on Holodiscus discolor (97G128); SPOKANE—Turnbull National
WLA, 25-V-95 on Geranium sp. (95T0307); YAKIMA—Nile, 28-VII-97 on Clematis ligusticifolia (97G278).

Metopolophium dirhodum (Walker): WA, Kittitas [on Rhododendron albiflorum*]|; SNoHOMISsH—Monroe, 5-
VI-96 on Phalaris arundinaceae (96G202+); YAKIMA—N Fork Tieton River, 15-VI-96 on native Rosa sp.
(96K0907).

Myzus cerasi (FE): WA, SKAGIT—Campbell Lk, 5-VI-96 on Prunus emarginata (96G170).

Nasonovia aquilegiae (Essig): WA, GARFIELD—Stentz Springs, 1-VIII-95 on Aquilegia formosa (9513627).

Nasonovia nr aquilegiae (Essig): WA, SPOKANE—Manito Pk, 11-VII-96 on Aquilegia sp. (9612627).

VOLUME 102, NUMBER 3 701

12

Figs. 10-12. Betuloxys alnicolus, 2 [illustrations not to equal scale]. 10, Metasomal tergum 1. 11—12, Prong.

Nasonovia polemonii (G&P): WA, Kittiras—Buck Meadows, 9-VII-96 on Polemonium pulcherrimum
(96G347).

Nasonovia sp.: MT SANDERS [on Castilleja sp.*].

Oestlundiella flava (Davidson): WA, KittrrAs—L.T. Murray WLA, S Fork Manastash Cr, 24-VII-95 on Alnus
sp. (95G39474).

Ovatus crataegarius (Walker): ID, BONNER—Sandpoint, 10-IV-97 on Origanum sp. (97T003); WA, ADAMS—
Othello, 1-VII-96 on Mentha piperita (96B003).

Prociphilus sp.: WA, YAKIMA—Clear Lk, 15-VI-96 on Lonicera sp. (96K083).

Rhopalosiphum padi (L.): WA, SNOHOMISsH—Monroe, 5-VI-96 on Phalaris arundinaceae (96G2021).

Sitobion avenae (F.): WA, BENTON—-WSU-Prosser, 18-VII-97 on Setaria lutescens (97G243).

Sitobion fragariae (Walker): WA, Is_LAND—Whidbey Island, 5-VI-96 on Dactylis glomerata (96G177).

Uroleucon erigeronense (Thomas): WA, YAKIMA—Buckhorn Meadows, 14-IX-95 on Achillea millefolium
(95G598).

Uroleucon ivae Robinson [on Iva xanthifolia}: WA, BENTON [*]; YAKIMA [*]—Grandview, 21-VIII-95
(95G503).

Uroleucon russellae (HRL): WA, SKAMANIA—Council Lk, 4-VIII-95 on Anaphalis margaritacea (95K095).

Uroleucon sonchi (L.): WA, BENTON [on Sonchus oleraceus**}.

Uroleucon sp.: WA, PEND OREILLE—Sullivan Lk, 3-VIII-95 on Adenocaulon bicolor (957391); Tiger
Meadows, 9-VII-96 on Senecio sp. (96T220).

Utamphorophora humboldti (Essig): OR. MULTNOMAH [on Physocarpus sp.*].

Undetermined spp.: WA, AsoTIN [on Chrysothamnus nauseosus*]; DOUGLAS—Waterville, 13-VII-95 on
Triticum aestivum (95B013, 95B014); KittrrrAs—Quartz Mtn 25-VII-96 on Rubus lasiococcus (96G407):
Taneum Lk, 11-IX-97 on Pedicularis bracteosa (97G425); LEwIs—RNP., Louise Lk, 1-VII-95 on Luetkea
pectinata (95K042),; SPOKANE—Mt Spokane SP, 22-VII-96 on Vaccinium sp. (96T492); YAKIMA—Clear
Lk, 15-VI-96 on Holodiscus discolor (96K080); YIR, Mabton Rd, 16-V-96 on Grindelia sp. (96G051).

Genus BETULOXYS Mackauer
Betuloxys alnicolus Pike & Stary, new species
(Figs. 10—12)

Diagnosis.—The new species is easily distinguished from B. compressicornis (Ruthe)
by its distinctly smaller size, generally brown coloration, and characters on prongs (distal
portion less differentiated and bearing three lanceolate setae). The new species was col-
lected near colonizing aphids [Boernerina variabilis (Richards)] on Alnus in Washington.
By comparison, B. compressicornis is generally yellow (exception, flagellum mostly
brown, and brown areas on mesonotum, scutellum, apices of legs and prongs), distal
portion of prong with five lanceolate setae, known as a parasitoid of Euceraphis puncti-
pennis (Zett.) on Betula, from E. Canada and Europe.

Etymology.—The name of the new species is derived from its association with Alnus.

Description.—Female. Head: Eyes medium sized; antenna 12-segmented, filiform,
reaching to half of metasoma. Mesosoma: Mesonotum smooth, with sparse setae, notauli
indicated anteriorly. Propodeum feebly granulate rugose, with sparse setae. Forewing:

702 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Stigma slightly more than 3X as long as wide (length/width [L/W] = 3.2); distal abscissa
of R1 (= metacarpus) 2 width of stigma, about % shorter than stigma length; RS vein
long, 1.2 stigma length. Metasoma: Metasomal tergum 1 (Fig. 10) smooth, length 1.6
width, spiracular tubercles distinct, prominent laterally, situated at end of first third of
tergite. Genitalia (Figs. 11-12): Prongs arcuate, apical portion somewhat dilated with
three lanceolate, perpendicular setae on dorsal side. Coloration: Head and mesosoma dark
brown. Mouthparts light brown. Antenna largely brown, except scape, pedicel, and F,
yellow, F, yellow brown. Wings subhyaline, venation light brown. Legs brown. Metasoma
brown, tergum 1 somewhat darker at base. Ovipositor sheaths and prongs brown, con-
colorous with metasoma. Length of body: about 1.8 mm.

Male.—Unknown.

Holotype.—?, USA, WA, YAKIMA Co., N. Fork of Tieton River, 15-VI-1996, K. S.
Pike collector (collection code, 96K089). Host aphid uncertain, collected near colonizing
aphids, Boernerina variabilis Richards on Alnus. Deposited in USNM.

Genus BINODOXYS Mackauer

Binodoxys carolinensis (Smith)

Aphis ceanothi Clarke: WA, KitrirAs—S. Fork Manastash Cr, 25-IV-97 on Ceanothus velutinus (97G137).

Aphis helianthi Monell: WA, CHELAN—Johnny Cr, 1-VIII-95 on Epilobium angustifolium (95G4827); PEND
OREILLE—Le Clerc WLA, 3-VIII-95 on Cornus stolonifera (95T399*+), YAKIMA—YIR, Signal Peak, 11-
VII-95 on Lomatium sp. (95G3347).

Aphis nigratibialis Robinson [on Cornus stolonifera]: WA, KLickirat—Trout Lk, 28-VI-97 (97G156);
YAKIMA—Nile, 28-VII-97 (97G280); YIR, Miller Point, 11-VII-95 (95K062).

Aphis nr oenotherae Oestlund: WA, YAKIMA—Pleasant Valley CG, 24-VH-97 on Epilobium minutum
(97G2677).

Aphis salicariae Koch: WA, PEND OREILLE—Le Clerc WLA, 3VIII—95 on Cornus stolonifera (95T399¥%).

Aphis varians Patch [on Epilobium angustifolium]: 1D, SHOSHONE—Thompson Pass, 29-VII-95 (95G4687);
MT, SANDERS [*].

Aphis nr varians Patch: WA, LEwis—RNP, Stevens Canyon, 23-VII-96 on Epilobium angustifolium (96G380).

Aphis sp. n.: WA, YAKIMA—YIR, Miller Point, 11-VII-95 on Ceanothus sp. (95K061).

Aphis sp.: WA, YAKIMA—Pleasant Valley CG, 24-VII-97 on Epilobium minutum (97G2677).

Cavariella nr aegopodii (Scopoli): WA, KirtrrAs—nr Ellensburg, Reecer Cr, 20-VI-95 on Salix sp. (95G207#).

Binodoxys clydesmithi Pike & Stary
Aphis canae Williams: WA, FRANKLIN—Kahlotus Hwy, 11-VI-96 on Artemisia tridentata (96G242*%).
Artemisaphis artemisicola (Williams) [on Artemisia tridentata]: WA, FRANKLIN—Kahlotus Hwy, 28-V-96 &
11-VI-96 (96G096, 96G2427+#).
Obtusicauda coweni (Hunter): WA, KITTITAS [on Artemisia tridentata* |.

Binodoxys conei Pike & Stary
Phorodon humuli (Schrank): WA, BENTON [on Humulus lupulus*].

Binodoxys coruscanigrans (Gahan)
Obtusicauda coweni (Hunter): WA, YAKIMA [on Artemisia tridentata*|—Wenas Lk, 30-V-96, (96G115).

Binodoxys grafi Pike & Stary

Aphis fabae Scopoli: WA, Yakima—American River, 5-IX-97 on Epilobium angustifolium (97G377}).

Aphis helianthi Monell [on Epilobium angustifolium]: WA, CHELAN—Stevens Pass, 16-IX-95 (95K1347);
Kittitas [*]—Lost Lk, 24-VII-95 (95G399+); SPOKANE—Mt Spokane SP, 27-VII-95 on unknown
Umbelliferae (95T3357).

Aphis holodisci Robinson: WA, KitrrrAS—Manastash Cr, mi marker 12, 25-VI-97 on Holodiscus discolor
(97G133).

Aphis salicariae Koch: ID, KooTENAI—Rathdrum, 29-VII-95 on Epilobium angustifolium (95G443*).

Aphis nr salicariae Koch: WA, CHELAN—Swauk Pass, 1-VIII-95 on Epilobium angustifolium (95G4927).

Aphis nr triglochinis Theobald: WA, SPOKANE—Mt Spokane SP, 29-VI-95 on native Ribes sp. (95T189).

Aphis varians Patch [on Epilobium angustifolium]: ID, KOOTENAI—Rathdrum, 29-VIH-95 (95G4437); WA,

VOLUME 102, NUMBER 3 703

CHELAN—Stevens Pass, 16-IX-95 (95K134+); Kittiras—Lost Lk, 24-VII-95 (95G399+); Stampede Pass,
8-VIII-96 (96K 139).
Aphis nr varians Patch: WA, Kittiras—Lost Lk, 24-VII-95 on Epilobium angustifolium (95G399+).

Binodoxys rhagii (Ashmead)
Uroleucon russellae (HRL) [on Anaphalis margaritacea |: WA, CHELAN—Chatter Cr, 1-VIII-95 (95G484);
YAKIMA—YIR, Delaney Springs (95G412) & Panther Butte (95G423), 25-VII-95.

Binodoxys spp.
Aphis salicariae Koch: WA, YAKIMA [on Cornus stolonifera* ].
Chaetosiphon fragaefolii (Cockerell): WA, PEND OrEILLE—Le Clerc WLA, 3-VIII-95 on native Rosa sp.
(95T398).
Illinoia rhododendri (Wilson): WA, Kittitas [on Rhododendri albiflorum* |].

Genus DIAERETELLUS Stary

Diaeretellus palustris Stary
Rhopalosiphum nymphaeae (L.): WA, YAKIMA—Outlook, 23-X-95 on Typha latifolia (95G709).

Genus DIAERETIELLA Stary

Diaeretiella rapae (M Intosh)

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola]: WA, AsoTin—Asotin, 10-VIII-95 (95T419); S. Fork
Asotin Cr, 22-VIII-95 (95T468); BENTON—WSU-Prosser, 17-VII-95 (95G353): Whitstran, 18-IX-97
(97G439); WHITMAN—Pullman, 16 & 30-VII-96, 1,6 & 19-VIII-96, 9 & 12-IX-96 (961304, 96T362,
96T382, 961407, 96T408, 96T468, 96T528, 961532).

Acyrthosiphon pisum (Harris): MT, RAvALLI—Stevensville, 25-V-96 on Melilotus sp. (96G083); WA,
BENTON—WSU-Prosser, 18 & 25-VII-97, on Melilotus officinalis (97F254+, 97F274+).

Aphis craccivora Koch: WA, WHITMAN—Pullman, 25-VII-95 on Capsella bursa-pastoris (951331).

Aphis fabae Scopoli: ID, LaTAH—Moscow Mtn, Paradise Cr, 25-IX-96 on Cirsium arvense (96T542).

Aphis nr fabae Scopoli: WA, PIERCE—RNP, Paradise, 8-I[X-95 on Spiranthes romanzoffiana (95K118).

Aphis helianthi Monell: WA, GARFIELD—Stentz Springs, 1-VIII-95 on Ranunculus sp. (9513637).

Brachycaudus cardui (L.): WA, AsoTin—De Spain Springs, 5-[X-96 on Onopordum acanthium (96T518).

Brachycaudus helichrysi (Kaltenbach): ID, BENEWAH—McCCroskey SP, 31-VIII-95 on Conyza canadensis
(95T516+); WA, AsoTiIn—Anatone, 28-VI-96 on Amsinckia sp. (96T089); Wickiup CG, 29-VIII-95 on
Sonchus sp. (95T505); BENTON—-WSU-Prosser, 1-X-97 on S. tuberosum (97G478); WHITMAN—Pullman,
28-VI-96 on Anthemis arvensis (96T162); 16-VII-96 on Anthemis cotula (96T298).

Brachycaudus tragopogonis (Kaltenbach) [on Tragopogon dubius]: WA, BENTON—WSU-Prosser, 3, 18, 25-
VII-97 (97F194, 97F242, 97F266); KLickirat—Bickleton, 8-X-96 (96G628): YAKIMA—Grandview, 24-V-
96, 25-VI-96 (96G080, 96G273).

Brachycorynella asparagi (Mordvilko) [on Asparagus officinalis]: WA, BENTON [*]|—WSU-Prosser, 6 & 25-
IX-95 (95K144, 95G548+, 95G649); YAkimA—Harrah, 29-IX-97 (97G462, 97G468, 97G469).

Braggia sp.[det. by G. Remaudiére]: WA, YAKIma [on Eriogonum elatum* |.

Brevicoryne brassicae (L.): WA, BENTON [on Brassica napus, B. oleracea & Descurainia sophia* |—WSU-
Prosser, 6-VII-95, 24-VIII-95, 5-VIII-96, 30-IX-96, 17-I-97, 23-V-97, & 1-X-97 on B. napus (95A024,
95G508, 96G438, 96G622, 97G004, 97G039, 97G475): 9-IX-95 on B. oleracea capitata (95K155); 11-
VII-97 on Sisymbrium altissimum (97G241); 1-X-97, on Malva neglecta (97G471+): West Richland 12-
VII-95 on B. oleracea acephala (95K074); CHELAN—Chelan SP. 11-[X-97 on Erysimum sp. (97K045);
CLARK—WSU-Vancouver, 21-IX-95 on B. oleracea capitata (95K179); SkAGIr—Mt Vernon, 16-X-95 on
B. sp. (95K190); PieRcE—Graham, 8-IX-96 on Cruciferae (961525); WALLA WALLA—Hwy 124, 10 mi E
of Burbank, 28-V-96 on B. napus (96G099); 28-V-96 on Sisymbrium altissimum (96G098); WHITMAN [on
B. napus*|—Pullman, 2-VIII-96 on B. napus (967390); YAKIMA— nr Bickleton 7-VII-97 on B. napus
(97G235); on Sisymbrium altissimum (97G237).

Diuraphis noxia (Kurdjumov) [on Hordeum vulgare unless otherwise indicated]: OR, UMATILLA—Hermiston,
17-VI-90 on unknown plant (90J003); WA, Asotin—Columbia Center, Pataha Cr Rd, 10-VIII-95 on
Agropyron cristatum (957431); Anatone, 24-VII-96, 1,8 & 15-VIII-96 (96T596 to 96T600): Anatone, 22
& 31-VII-97 (97J0O11 to 97J025); BENTON [*]—WSU-Prosser, 28-V-96 & 24-X-96 (96G087 to 96G090,
96G654); [KLICKITAT, on Aegilops cylindrica* |—Cemetery Rd nr Bickleton, 7-VII-97 (97G226+): WALLA
WALLA—Valley Grove, 11-VI-96 (96G240); Waitsburg, 11-VII-96 (96J056, W. Turner & J. Rumph
collectors); WHITMAN—Pullman, 9-VII-96 (96J053 to 96J055, W. Turner & J. Rumph collectors); YAKIMA

704 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

[*]; [Following on Triticum aestivum unless otherwise indicated ] ADAMs [*]; ASOTIN [*|—Couse Cr Rd,
10-VIII-95 (95T421); Field Springs SP, 10-VIII-95 (95T427); Asotin WLA, 15 & 29-VIII-96, 24-IX-96,
3-X-96 (961455, 96T457, 96T503, 961555, 961569); De Spain Spring, 5-IX-96, 3-X-96 (96T516,
96T571); BENTON [*]—WSU-Prosser, 15-V-95, 24-V-96, 3-VI-96, 24, 28-VII-97 (95G065, 96G081,
96G132, 97G274, 97G301); 29-IV-97 thru 24-IX-97, (97FO01—97F004, 97F042—97F455); 1-IV-97, 6-VI-
97 (97G005, 97G040, 97G041); 3 mi S of Prosser, 6-VI-97 (97G045); Sellards & Travis Rds, 25-VI-95
(95G252); Rotha Rd 17-VII-97 (97G245); DouGLAas—Waterville, 12-VIII-95 (95B019); FRANKLIN [*]— Star
School, 6-VII-95 (95G287); Kahlotus, 6-VII-95 (95G291); GARFIELD—Columbia Center, 10-VIII-95, 7-IX-
95 (957435, 951538, 951539); Spruce Springs, 14-[X-95 (951574); KuickiTaT [*]—Bickleton, 31-X-95,
7-VII-97 (95G743, 97G223): nr Bickleton, 18-VI-97 & 7 & 29-VII-97, 6-VIII-97 (97G119, 97G238,
97G300, 97G354); Brannon Rd nr Cleveland, 7-VII-97 (97G234); Cleveland, 18-VI-97, 27-VI-97, 7-VII-
97 (97G114 to 97G117, 97G145, 97G232); SPOKANE—Mt Spokane SP, 22-VII-95 (95T495); WHITMAN
[*]|—Hole in the Ground, 8-VI-95 (95T085); Dusty, 9-VII-96 on Triticum sp. (96J065, W. Turner & J.
Rumph collectors); YAKIMA [*]—nr Bickleton, 23-V-96, 17-VI-96 (96G073, 96G250, 96G251, 96G252);
Sunnyside, 24-VH-97 (97G273).

Dysaphis plantaginea (Passerini): WA, BENTON—Prosser, 24-VI-96 on Pyrus malus (96G279*).

Hayhurstia atriplicis (L.) [on Chenopodium album]: ID, LataH—Moscow, 28-IX-96 (96T561); WA, ASOTIN—
Anatone 28-VI-96 (96T091); De Spain Springs, 31-VII-96 (961358); BENTON—WSU-Prosser, 20-VIII-96,
& 3,18,25-VII-97 (96G509, 97F190, 97F238, 97F262); KLickirat—Bickleton 18-VI-97, 7-VII-97
(97G111, 97G118, 97G239); nr Cleveland, Dot & Brannon Rds, 27-VI-97, 7-VII-97 (97G143, 97G231);
WHITMAN—Pullman, 16 & 30-VII-96, 7-VIII-96 (961302, 961360, 96T361+, 96T417); YAKIMA [*]—
Grandview, 22-VII-95, 17-V-96, 3-VI-96, 8 & 30-VII-96, 29-VI-97 (95G367, 96G067, 96G131, 96G327,
96G430. 97G181): nr Bickleton, 29-VIII-95 (95G524); Glade Cr, 7-VII-97 (97G218); Bickleton Hwy &
County Line Rd, 7-VI-97 (97G236); Wenas Cr & Maloy Rd, 11-VI-97 (97G065); Wenas Lk, 17-VI-97
(97G091); YIR, Signal Peak Guard Station, 19-I[X-96 (96G578).

Illinoia corylina (Davidson): WA, WHITMAN—Pullman, 14-VIII-96 on Aquilegia sp. (96T442).

Lipaphis erysimi (Kaltenbach): WA, BENTON—WSU-Prosser, 30-IX-96, on Brassica napus (96G6217);
WHITMAN—Pullman, 3-VII-96 on Capsella bursa-pastoris (9611667).

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow, 7-X-96, on Solanum sp. (961577); WA, PIERCE
[on Epilobium angustifolium* }].

Macrosiphum nr euphorbiae: WA, YAKIMA—YIR, Mill Cr Guard Station, 6-VI-95 on Lomatium nudicaule
(95G115).

Macrosiphum sp.: WA, ASOTIN [on Triticum aestivum*).

Myzus persicae (Sulzer): WA, BENTON [on Raphanus sativum*|—23-VI-96 on Chrysanthemum sp.
(96G388+): WSU-Prosser, 29-VII-96 on Carthamus tinctorius (96G428); 31-VIII-95, 1-X-97 on
Chenopodium album (97G476); 27-X-95 on Malva neglecta (95K212+); 1-[X-95 on Solanum tuberosum
(95G525, 95G526): 6-VII-95 on Sonchus oleraceus (95A026+); Prosser, 23-VIH-96 on Chrysanthemum sp.
(96G388); SKAGIT [on Brassica oleracea*], SNOHOMISH—Monroe, 24-VII-96 on Pisum sativum (9615947);
WHITMAN—Pullman, 16-VII-95, 3-VII-95 on Raphanus sativum (951248, 95T407), 22-VII-95 on Solanum
lycopersicon (95T299+), & 3-VIII-95 on Spinacia oleracea (95T408); Ridpath, 6 & 22-VII-95 on
Amsinckia retrorsa (95G295+)..

Myzus sp.: WA, WHITMAN [on Forsythia sp.*].

Phorodon humuli (Schrank) [on Humulus lupulus unless otherwise indicated |]: WA, BENTON [*]—WSU-
Prosser, 9-IX-95 (95K154), & 27-X-95 on Lamium amplexicaule (95K213+); YAKIMA—Grandview, 8-VI-
95 (95G145).

Rhopalosiphum insertum (Walker): WA, DouGLas—Orondo, 28-V-96 on Pyrus malus (96RO003).

Rhopalosiphum maidis (Fitch): WA, BENTON [on Panicum milliaceum* |—WSU-Prosser, 18-XI-95 & 20-I-96
on Capsella bursa-pastoris (95G750+, 96GO03+), & 2-IX-96 on Hordeum vulgare (96G655).

Rhopalosiphum padi (L.) [on Triticum aestivum unless otherwise indicated]: WA, BENTON [on Hordeum
vulgare & Triticum aestivum*|—Prosser, 20-VIII-96 on Zea mays (96G510); WSU-Prosser, 1-X-95, 24-
VI-96 (95G668, 96G272): Cemetery Rd & Hwy 241, 12-X-95 (95G690); DouGLAs—Waterville, 12-VIII-
95 (95B016, 95B018); YAKIMA [*]—Grandview, 22-X-97 (97G490).

Sitobion avenae (FE) [on Triticum aestivum unless otherwise indicated]: WA, BENTON [*]—Sellards & Travis
Rds, 25-VI-95 (95G253); Prosser, 30-VI-95, 17-VII-95, 24-VI-96, 1 & 8-VII-96, (95G285, 95G352,
95G354, 95G356, 95G358, 96G271, 96G312, 96G330); WSU-Prosser, 22,30-VI-97, 3,11,18,25-VII-97
(assorted collections between 97F183 & 97F277), & 18-VII-97 on Setaria lutescens (97G243); nr
Whitstran, 17-VII-95, 8-VII-96 (95G355, 96G328); FRANKLIN—Star School, 6-VII-95 (95G289); Kahlotus,

VOLUME 102, NUMBER 3 705

6-VII-95 (95G290); KLickiTaT [*]—Bickleton, 9-VII-95 (95G318); WHITMAN [*]—Rosalia, 16-VII-96
(96T294); YAKIMA—Grandview, 5-VII-95 (95G286).

Uroleucon ivae (Robinson): WA, YAKIMA—Grandview, 9-VII-95, on Iva xanthifolia (95G313).

Uroleucon sp. YAKIMA—Grandview, 18-I-96, on Brassica oleracea (96GO001).

Genus EPHEDRUS Haliday

Ephedrus californicus Baker

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola]: WA, Asotin [*]; CHELAN—Chelan SP, 11-[X-97
(97K038); YAKIMA—YIR, Signal Peak Guard Station, 19-[X-95 (95G621).

Aphis fabae Scopoli: WA, AsoTiIn—Field Springs SP, 18-VII-95 on Cirsium arvense (951272).

Aphis helianthi Monell: MT, SANDERS—nr Thompson Pass, 29-VII-95 on Ligusticum verticillatum (95G460);
WA, CHELAN—Stevens Pass, 16-IX-95 on Epilobium angustifolium (95K1347); GARFIELD—Stentz Springs,
22-VIII-95 (95T452); Kitriras—Lost Lk, 24-VII-95 on Epilobium angustifolium (95G399*7), & 11-IX-95
on Heracleum lanatum (95G572); 2-[X-95 on Lomatium sp. (95G5357+); LEwis—RNP, Bench Lk, 8-[X-95
on Ligusticum sp. (95K113); PIlERCE—RNP, Tipsoo Lk, 11-[X-96 on Osmorhiza sp. (96G535+); YAKIMA—
RNP, Dewey Lk Trail, 11-[X-96 on Ligusticum sp. (96G5527+); YIR, 25-V-96 on Lomatium dissectum
(96K052), Mt Adams Lk, 25-VII-95 on Osmorhiza occidentalis (95G428), & Windy Point, 19-[X-95 on
Xerophyllum tenax (95G635).

Aphis holodisci Robinson [on Holodiscus discolor unless otherwise indicated]: WA, SPOKANE—Mt Spokane
SP, 27-VII-95 (951338); Manito Pk, 11-VII-96 (96T260); YAKIMA—Clear Lk, 15-VI-96 (96K082); Bird
Cr Meadows Rd & FR822, 26-VI-95 on Physocarpus malvaceus (95A020, 95A021).

Aphis nr oenotherae Oestlund: WA, GARFIELD—Pataha Cr Rd, 7-I[X-95 on Epilobium sp. (951541).

Aphis varians Patch [on Epilobium angustifolium]: ID, SHOSsHONE—Thompson Pass, 29-VII-95 (95G4687);
WA, CHELAN—Stevens Pass, 16-IX-95 (95K1347); Kitriras—Lost Lk, 24-VII-95 (95G3997); PIERCE—
Naches Pass, 25-IX-97 (97G460); RNP, Bench Lk, 8-IX-95 (95K114).

Aphis nr varians Patch: WA, Kittrras—Lost Lk, 24-VII-95 on Epilobium angustifolium (95G399*).

Aphthargelia symphoricarpi (Thomas) [on Symphoricarpos sp.\: WA, Kittitas [*]; SPOKANE—nr Peone,
Cooper Rd & Hwy 206, 29-VI-95 (95T178); YAkIMA—nr Rimrock Lk, 9-VI-95 (95A004).

Brachycaudus helichrysi (Kaltenbach): WA, ASoTIN—nr FR 40 & FR44, 1-VIII-95 on Achillea millefolium
(O5T3 827):

Braggia eriogoni (Cowen): WA, KittirAs—Taneum Lk, 11-IX-97 on Epilobium umbellatum (97G428).

Brevicoryne brassicae (L.): WA, FRANKLIN—Eureka, 14-V-96 on Brassica napus (96G040).

Capitophorus elaeagni (del Guercio): WA, YAKIMA—FR 1010, 8 mi S of Tieton River, 14-IX-95 on Cirsium
vulgare (95G596).

Cavariella aegopodii (Scopoli) [on Ligusticum grayi]: WA, PIERCE—RNP, Tipsoo Lk, 5-IX-97 (97G3637);
YAKIMA, YIR, Windy Point, 19-IX-95 (95G6427+).

Cavariella pastinacae (L.): WA, AsoTin—Asotin Cr Rd, 11-VII-95 on Oenothera sp. (951196).

Chaetosiphon fragaefolii (Cockerell) [Rosa sp.]: ID, BOUNDARY [{*]; WA, PEND OrREILLE—Le Clerc WLA,
3-VIII-95 (95T398).

Diuraphis noxia (Kurdjumov): WA, Asotin—Anatone, 8-VIII-96 on Hordeum vulgare (961598); Asotin Cr
WLA, 3-X-96 on Triticum aestivum (96T565).

Ericaphis wakibae (Hottes): WA, YAKIMA—Clear Lk, 15-VI-96 on native Rosa sp. (96KO0857%).

Hyperomyzus lactucae (L.): WA, KLickiIrAt—Rock Cr & Columbia River, 28-IV-95 on Sonchus oleraceus
(95GO0117). ,

Hyperomyzus nigricornis (Knowlton) [on Hieracium sp. unless otherwise indicated]: ID, LATAH—Moscow
Mtn 13-VII-96 on unknown plant (96T264); 3,15-VIII-96, on Compositae (961399, 96T446); MT,
SANDERS—nr Thompson Pass, 29-VII-95 (95G462+); WA, GarFiELD—Alder Thicket, 21-IX-95 (95T587);
KittirAs—Manastash Rd, 25-VII-96, 16-VII-96 (96G415, 96G490); Manastash Cr & FR125, 16-VIII-96
(96G493); Ellensburg, 23-VI-96 on Ribes sp. (96G267); Lost Lk, 6-VIII-97 (97G344); Klickitat—
Cleveland, 16-VI-96 on Madia minima (96K109); 7-VII-97 (97G229); PIERCE—RNP, Tipsoo Lk, 11-IX-
96 (96G541), & 5-IX-97 on Agoseris aurantica (97G3847+); SPOKANE—Mt Spokane SP, 24-VIII-95 on
Compositae (95T486); Mt Spokane Hwy, mi marker 18.5, 22-VIII-96, on Compositae (96T487, 96T488);
WHITMAN—Kamiak Butte, 7-VII-96, on unknown plant (96T205); Pullman, 10-VIII-95 on Lapsana
communis (95T436+); YAKIMA—Ravens Roost, 28-VII-97 on Hieracium albertinum (97G296); Wenas Lk,
30-V-96 on Ribes aureum (96G123); YIR, Signal Peak, 19-I[X-95 (95G622) & Fish Lk (95G637) on
Agoseris aurantica, Delaney Springs, 25-VII-95 on Hieracium cynoglossoides (95G406), & Goat Butt, 19-
IX-96 (96G585);

Hyperomyzus sp.: WA, Kittrras—S. Fork Manastash Cr, 24-VII-95 on Hieracium albiflorum (95G388).

706 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Illinoia sp.: WA, GARFIELD—FR 160 & FR 40, 21-[X-95 on Vaccinium sp. (95T596).

Macrosiphoniella ludovicianae (Oestlund) [on Artemisia ludoviciana|: WA, YAKIMA [*]; YIR, Randell
Springs, 16-V-96 (96G049¥*).

Macrosiphoniella tanacetaria (Kaltenbach): WA, CHELAN—Stevens Pass, 16-IX-95 on Tanacetum vulgare
(95K133).

Macrosiphum albifrons Essig [on Lupinus sp.): WA, KittirAs—Buck Meadows, 9-VII-96 (96G338);
KLickITAt—Trout Lk, 11-VII-95 (95K068); LEwis—RNP, Stevens Canyon, 23-VII-96 (96G364); PIERCE—
RNP, Tipsoo Lk, 11-IX-96 (96G547) & 5-IX-97 (97G362); SPOKANE—Manito Pk, 11-VI-96 (96T033); nr
Peone, 2-VII-96 (96T098); YAKIMA—American River, 5-I[X-97 (97G376).

Macrosiphum californicum (Clarke): WA, Kittrras—Murray WLA, S Fork Manastash River, 11-[X-95 on
Salix sp. (95G3917).

Macrosiphum creelii Davis: WA, Kittiras—Manastash Cr Rd, mi marker 8, 16-VIII-96 on Vicia sp. (96G492).

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow Mtn, 6-VII-96 on unknown plant (96T181); 6-VII-
96 on Compositae (96T185, 961192); MT, SANDERS—nr Thompson Pass, 29-VII-95 on Hieracium sp.
(95G462+); WA, AsoTIN—De Spain Springs, 11-VU-96 on unknown plant (96T252); nr FR 40 & FR44,
1-VIII-95 on Achillea millefolium (95T382+); BENTON — WSU-Prosser, 27-X-95 on cultivated Rosa sp.
(95K220+); GARFIELD—Stentz Springs, 1-VII-95 on Arnica latifolia (95T370); KING — Issaquah, 4-VI-96
on native Rosa sp. (96G1457+); Kittiras [on Rosa sp.*]|—Buck Meadows, 9-VII-96 on native Rosa sp.
(96G335) & 16-VIII-96 on Silene sp. (96G497); Lost Lk Trail, 6-VIII-97 on Luina nardosmia (97G327);
Stampede Pass, 8-VIII-96 on Castilleja sp. (96K1407+); KLIckIrat—Cleveland, 7-VII-97 on Lupinus sp.
(97G230); LEwis—RNP, Stevens Canyon, 23-VII-96 on Crepis sp. (96G357); PEND OREILLE—Cusick, 9-
VII-96 on native Rosa sp. (961223); SKAMANIA—Twin Falls CG, 4-VIII-95 on Potentilla sp. (95K0987);
WHITMAN—Kamiak Butte, 7-VII-96 on Compositae (96T200); Pullman, 10-VIII-95 on Lapsana communis
(95T4367) & 18-VI-96 on cultivated Rosa sp.(96T037); YAKIMA [on Ligusticum grayi+* |—Clear Lk, 15-
VI-96 on native Rosa sp. (96K0857); N. Fork Tieton River, 15-VI-96, on Hieracium albertinum (96KO094);
Buckhorn Meadows, 14-IX-95 on Trautvetteria caroliniensis (95G597); YIR, Bird Cr Meadows, 26-VI-
95, on Osmorhiza chilensis (95A014).

Macrosiphum nr euphorbiae (Thomas): WA, YAKIMA—N. Fork Wenas Cr, 30-V-96 on Arabis sp. (96G129).

Macrosiphum gaurae (Williams): WA, WHITMAN—Colton, 19-IX-95 on Gaura parviflora (95T585).

Macrosiphum pallidum (Oestlund): WA, SKAMANIA—Twin Falls CG, 4-VIII-95 on Potentilla sp. (QSKO987#).

Macrosiphum rosae (L.) [on Rosa sp. unless otherwise indicated]: OR, BENToN—Corvallis, 18-V-93 on
unknown plant, A. Jensen collector (93J002); WA, BENTON—-WSU-Prosser, 27-X-95 (95K2207); KING—
Issaquah, 4-VI-96 (96G145+); SPOKANE—Manito Pk, 19-IX-96 (96T540); YAKIMA [*]—Grandview, 13 &
21-V-97 (97G021+, 97G037).

Macrosiphum valerianae (Clarke): WA, Kirtiras—Lost Lk, 24-VII-95 on Epilobium angustifolium
(95G3997x).

Metopolophium dirhodum (Walker): WA, BENTON—-WSU-Prosser, 27-X-95 on Conyza canadensis
(95K2167+#).

Myzus cerasi (FE): WA, BENTON—Prosser, 28-V-96 on Prunus sp. (96G091); WHITMAN—Pullman, 5-VII-96 on
P. avium (9671173).

Myzus persicae (Sulzer): WA, YAKIMA—Cliffdell, 23-X-97 on Linaria dalmatica (97G496).

Nasonovia ribisnigri (Mosley): WA, WHITMAN—Pullman, 5-VII-96 on unknown plant (96T177).

Nasonovia (Kakimia) sp.: WA, WHITMAN—Pullman, 10-VIII-95 on Lapsana communis (9574367).

Obtusicauda artemisiphila (Knowlton & Allen): WA, KLICKITAT [on Artemisia tridentata*].

Obtusicauda coweni (Hunter): WA, YAKIMA [on Artemisia tridentata* |.

Obtusicauda sp.: WA, KLICKITAT [on Artemisia tridentata*}].

Pleotrichophorus nr amsinckii Richards: WA, GARFIELD—Alder Thicket CG, 14-IX-95 on Amsinckia sp.
(95T562).

Pleotrichophorus pseudoglandulosus (Palmer): WA, YAKIMA—YIR, Randell Springs, 16-V-96 on Artemisia
ludoviciana (96G049¥+¢).

Rhopalosiphum insertum (Walker): WA, Kittiras—Ellensburg, 17-V-96 on Crataegus columbiana (96G066).

Rhopalosiphum maidis (Fitch): WA, BENTON—WSU-Prosser, 18-XI-95 on Capsella bursa-pastoris
(95G750F#).

Rhopalosiphum padi (L.): WA, BENTON—-WSU-Prosser, 18-XI-95 on Capsella bursa-pastoris (95G7507#).

Sitobion avenae (F.): WA, Kinc—Stampede Pass on Dactylis glomerata (96G442); Kirtiras—Manastash Rd
& FR 112, 9-VII-96 on D. glomerata (96G346); PIERCE—Graham, 25-V-96, on Gramineae (96T001); RNP,
Tipsoo Lk, 5-IX-97 on Agoseris aurantica (97G384+);YAKIMA—Blowout Mtn, 9-IX-97 on D. glomerata
(97G401).

VOLUME 102, NUMBER 3 707

Uroleucon nr arnesense Robinson: MT, SANDERS—18 mi E of Thompson Pass, 29-VII-95 on Solidago sp.
(95G457).

Uroleucon erigeronense (Thomas): WA, BENTON—-WSU-Prosser, 27-X-95 on Conyza canadensis (95K2167);
CHELAN—Leavenworth, 16-IX-95 on Conyza canadensis (95K141); KitriraAs—Lost Lk, 2-IX-95, on
Anaphalis margaritacea (95G532) & 6-VIII-97 on Aster sp. (97G346); Manastash Rd & FR 112, 11-IX-
95 on Achillea millefolium (95G563), on Anaphalis margaritacea (95G5617) & Cirsium arvense
(95G564+); PIERCE—RNP, Tipsoo Lk, 5-IX-95 (95K106) & 8-IX-97 (97G3647+) on Aster sp.; WHITMAN—
Pullman, 10-IX-96 on unknown plant (96T530); YAKiImMA—Hells Crossing CG, 11-IX-96 on Aster sp.
(96G524+); YIR, White Deer Cr, 11-VII-95 on Anaphalis margaritacea (95G326), & Signal Peak, 19-IX-
95 on Achillea millefolium (95G6207).

Uroleucon nr erigeronense (Thomas): WA, OKANOGAN—Bridgeport SP, 9-IX-97 on Conyza canadensis
(97K023).

Uroleucon ivae Robinson: WA, YAKIMA—Grandview, on Iva xanthifolia (96G431).

Uroleucon katonkae (Hottes): WA, YAKIMA—Hells Crossing CG, 11-[X-96 on Aster sp. (96G5247).

Uroleucon russellae (HRL) [on Anaphalis margeritacea|: MT, SANDERS [*]; WA, CHELAN—Chatter Cr, 1-
VIII-95 (95G484); Kitriras—Manastash Rd & FR112, 11-[X-95, on Anaphalis margaritacea (95G5617*);
SKAMANIA—June Lk, 4-VIII-95 (95K101); SPOKANE—Mt Spokane SP, 24-VIII-95 (95T494).

Uroleucon sonchi (L.) [on Sonchus oleraceus unless otherwise indicated]: WA, CLARK—-WSU-Vancouver, 21-
IX-95 (95K172); KLickITAT—Rock Cr & Columbia River, 28-IV-95 (95G0O117); PlERCE—Graham, 8-IX-
96, on Lactuca sp. (961527); WHITMAN—Pullman, 9-IX-96 on Lactuca sp. (96T529); YAKIMA—Yakima
Arboretum, 26-IX-95 (95G6587); YIR, Satus, 1-VI-96, on Sonchus sp. (96K064).

Uroleucon spp.: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Adenocaulon bicolor (95T518, 951527,
95T536); WA, AsoTIN [on Dipsacus sylvestris*]; CHELAN—Chiwawa Canyon, 3-VIII-96 on A. bicolor
(96R044); Kitriras—Lost Lk, 6-VIII-97 on Achillea millefolium (97G318); S. Fork Manastash Cr, 24-VII-
95 on Hieracium sp. (95G389); Kiickirat—nr Bickleton, Box Springs, 27-VI-97 on Balsamorhiza
sagittata (97G154); PEND OREILLE—Sullivan Lk, 3-VIII-95 (95T391); Tiger Meadows, 9-VII-96 on Senecio
sp. (961220); PIERCE—RNP, Tipsoo Lk, 5-IX-97 on Anaphalis margaritacea (97G367); SPOKANE—Mt
Spokane SP, 24-VIII-95 on Centaurea pratensis (95T477); WHITMAN [on Lactuca serriola*|; YAKIMA [on
Aster sp.*|—Union Gap, 23-X-95 on Centaurea maculosa (95G720); YIR, Signal Peak, 19-IX-95 on
Grindelia sp. (95G618).

Wahlgreniella nervata (Gillette): WA, YAKIMA—Grandview, 13-V-97 on cultivated Rosa sp. (97G0217¢).

Undetermined spp.: WA, Spokane—Mt Spokane SP, 24-VIII-95 on Epilobium sp. (95T490); WHttmMAN—
Pullman, 25-VI-95 on Lapsana communis (951332); YAKIMA—YIR, Mabton Rd, 16-V-96 on Crepis
atrabarba (96G050).

Ephedrus chaitophori Gardenfors
Chaitophorus populifolii (Essig) [on Populus trichocarpa]: WA, Kitt1ras—Manastash Cr Rd, 5 mi marker,
16-VIII-96 (96G485); YAKIMA—Wenas Lk, 11-VI-97 (97G062).

Ephedrus clavicornis Pike & Stary

Aphis helianthi Monell: WA, PEND OREILLE [on unknown Umbelliferae+*]; WHITMAN [on Helianthus annuus*].

Aphis nr oenotherae: WA, GARFIELD, Kittitas, & Lewis [on E. angustifolium*]; YAKIMA [on E. spp. & E.
minutum* |—Wenas Cr, 13 VIII-96 (96G467) on Epilobium sp.

Aphis varians Patch [on Epilobium angustifolium]: MT, SANDERS, W. Fork Thompson River, 26-VII-94
(94R029); WA, Lewis [*] & YAKIMA [*].

Aphthargelia symphoricarpi (Thomas): WA, YAKIMA [on Symphoricarpos sp.*}.

Dysaphis plantaginea (Passerini): WA, DouGLAS [on Pyrus malus*].

Illinoia maxima (Mason): WA, SPOKANE [on Rubus parviflorus*].

Prociphilus sp.: WA, YAKIMA [on Amelanchier alnifolia*].

Sitobion avenae (EF): WA, LEwis [on Agropyron caninum*].

Undetermined spp. [on Epilobium sp. unless otherwise indicated]: CANADA, BritIsH CoLuMBIA [from
sweepings*]; WA, GARFIELD [*], Kittitas [*], & YAKIMA [*].

Ephedrus incompletus (Provancher)
Ovatus crataegarius (Walker): ID, BONNER—Sandpoint, 10-IV-97 on Origanum sp. (97T003).

Ephedrus lacertosus (Haliday)
Aspidophorodon nr longicauda (Richards): WA, SKAMANIA—nr Keenes Horse Camp, 11-VII-95 on Lupinus
sp. (95K065+#).
Ericaphis gentneri (Mason): WA, CHELAN—Stevens Pass, 16-IX-95 on Spiraea densiflora (95K138).
Macrosiphum albifrons Essig: WA, SKAMANIA—nr Keenes Horse Camp, 11-VII-95 on Lupinus sp. (95K066).

708 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Macrosiphum euphorbiae (Thomas): WA, YAKIMA—YIR, Old Maid Canyon, 25-VII-95 on Cynoglossum
grande (95G440).

Macrosiphum sp.: WA, CHELAN—Chatter Cr, 1-VII-95 on Oplopanax horridum (95G4887).

Macrosiphini aphids [immatures]: WA, YAKIMA—Morse Cr, 24-VII-97 on Achlys triphylla (97G255).

Metopolophium sp.: WA, SKAMANIA—nr Keenes Horse Camp, 11-VII-95 on Lupinus sp. (9SKO065**+).

Sitobion sp.: OR, BENTON—Mary’s Peak, 14-[X-93 on Dicentra formosa, A. Jensen collector (93J006).

Ephedrus persicae Froggatt
Brachycaudus cardui (L.): WA, SkAGiIr—La Conner, 5-VI-96 on Cirsium vulgare (96G189).
Brachycaudus helichrysi (Kaltenbach): WA, SKAGITr—La Conner, 5-VI-96 on Prunus salicina (96G196).
Undetermined sp.: WA, WHITMAN—Pullman, 3-VIII-96 on Pyrus malus (96T042).

Ephedrus spp.

Aphis coweni Palmer: WA, KittiTAs [on Veratrum viride*].

Aphis varians Patch: MT, SANDERS, Thompson Pass, 27-VII-94 on Epilobium angustifolium (94R040).

Chaetosiphon fragaefolii (Cockerell): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native
Rosa sp. (96T2097#).

Diuraphis noxia (Kurdjumoyv): WA, BENTON—WSU-Prosser, 15-V-97 on Triticum aestivum (97F045).

Ericaphis scammelli (Mason): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa
sp. (96T209+£).

Hyperomyzus nigricornis (Knowlton): ID, LATAH—Moscow Mtn, 3-VIII-96 on Compositae (96T402).

Macrosiphum nr euphorbiae (Thomas): WA, YAKIMA—N. Fork Tieton River, 27-VI-96 on Lilium
columbianum (96G299).

Macrosiphum rhamni (Clarke): ID, SHOSHONE—Mutrray, 17-VI-95 on Rhamnus purshiana (95G192).

Prociphilus sp.: WA, WHITMAN—Pullman, 13-VI-96 on unknown plant (96T029).

Undetermined sp.: WA, GARFIELD—Spruce Springs, 14-I[X-95 on Cirsium vulgare (951571).

Genus EUAPHIDIUS Mackauer

Euaphidius cingulatus (Ruthe)

Pterocomma beulahense (Cockerell): WA, KinG—Stampede Pass, 8-VIII-96 on Populus trichocarpa
(96G445+); WHITMAN—Pullman, 9-X-96 on Populus sp. (96T584).

Pterocomma bicolor (Oestlund) [on Salix sp. unless otherwise indicated]: WA, KINGc—Stampede Pass, 8-VIII-
96 on Populus trichocarpa (96G445+); KLICKIrAT—nr Bickleton, Pine Cr, 18-VI-97, on P. trichocarpa
(97G113); PreRcE [*]; SKAMANIA—Council Lk, 4-VIII-95, (95K096); WHITMAN—Pullman, 12-IX-95
(95T558+); YAKIMA—Wenas Lk, 17-VI-97 (97G093).

Pterocomma sanguiceps Richards: WA, KirrirAs—Ellensburg, 10-IV-95 on Salix (95G0047).

Pterocomma smithiae (Monell) [on Salix sp.]: WA, KittrrAs—Ellensburg, 10-IV-95 (95G004+); WHITMAN—
Pullman, 12-IX-95 (95T5587).

Pterocomma sp.: OR, MULTNOMAH [on Salix sp.*]; WA, YAKIMA—YIR, Signal Peak, 19-IX-95 on Salix sp.
(95G619).

Undetermined sp.: WA, AsoTIN—De Spain Springs, 26-VI-96 on Salix sp. (96T057).

Euaphidius setiger Mackauer
Periphyllus californiensis (Shinji): OR, BENTON—Corvallis, McDonald State Forest, 1-V-93 on unknown plant,
A. Jensen collector (93J001).
Periphyllus lyropictus (Kessler): WA, BENTON [on Acer platanoides *|; GRANT—Sun Lakes SP, 8-IX-97 on
A. sp. (97K010); YAKIMA—Grandview, 13-VI-96 on A. sp. (96G243).
Periphyllus sp.: WA, SPOKANE—Spokane, Manito Pk, 19-I[X-96 on Acer sp. (96T539).

Genus HARKERIA Cameron

Harkeria rufa Cameron
Nasonovia houghtonensis (Troop): OR, BENTON—Corvallis, McDonald State Forest, 15-IV-94 on Oemleria
cerasiformis, A. Jensen collector (94J002).

Genus LYSAPHIDUS Smith

Lysaphidus adelocarinus (Smith)
Aphis canae Williams: WA, AsoTiIn—Asotin Cr Rd, 13-VI-95 on Artemisia tridentata (95T089).

VOLUME 102, NUMBER 3 709

Aphis (Zyxaphis) sp.: WA, KirrirAs—Frost Meadows, Manastash Rd, 11-IX-97 on Artemisia tridentata
(97G435).

Artemisaphis artemisicola (Williams): WA, FRANKLIN—Kahlotus Hwy, 28-V-96 on Artemisia tridentata
(96G096).

Epameibaphis atricornis G&P [on Artemisaphis tridentata]: WA, BENTON [+*]—ALE, 19-V-95 (95G0727);
YAKIMA [{*]—nr Mabton, 9-V-95 (95G0317); Selah Cr & Hwy 821, 15-V-95 (95G0677).

Flabellomicrosiphum knowltoni Smith [on Artemisaphis tridentata]: WA, BENTON [t*]; YAKIMA—YIR, Ft.
Simcoe, 30-V-96 (95G0917).

Flabellomicrosiphum tridentatae (Wilson): WA, KLICKITAaT—Alderdale Cr & Columbia River, 28-IV-95 on
Artemisaphis tridentata (95GO137*¢#).

Flabellomicrosiphum sp.: WA, KLICKITAT [on Artemisaphis tridentatay*].

Microsiphoniella artemisiae (Gillette): WA, AsotTiIn—Asotin Cr HMU, 5-VII-96 on Artemisia ludoviciana
(96T141).

Obtusicauda coweni (Hunter): WA, AsoTiIn—Asotin Cr HMU, 7-V-96 on Artemisia tridentata (96T 1427).

Obtusicauda sp. [on Artemisia tridentata): WA, KLICKITAT [*] & YAKIMA [*].

Pleotrichophorus nr amsinckii Richards: WA, BENTON [*]—ALE, 8-VI-95 on Cryptantha sp. (95G1477).

Pleotrichophorus palmerae (Knowlton): WA, KLickirat—nr Alderdale, 9-V-95 on Chrysothamnus sp.
(95G0267).

Pleotrichophorus sp.: WA, GARFIELD—Alder Thicket, 21-IX-96 on Amsinckia sp. (95T5927); KittiraAs [on
Artemisia tridentata}*]; KLICKITAT—nr Alderdale, 9-V-95 on Chrysothamnus nauseosus (95G0267).

Pseudoepameibaphis essigi Knowlton & Smith: WA, KLICKITAT [on Artemisia tridentata}*].

Pseudoepameibaphis glauca G&P [on Artemisia tridentata): WA, Kittiras [*+*]—Taneum Rd, 11-IX-95
(95G5867); YAKIMA—YIR, Goat Butte, 9-X-96 (96K169).

Pseudoepameibaphis tridentatae (Wilson) [on Artemisia tridentata]: WA, BENTON [*]—ALE, 19-V-95
(95G072+); KittiTAs [*]; KLickiTat [+*]—Alderdale Cr & Columbia River, 28-IV-95 (95G0137); YAKIMA
[+*]—Naught Rd, 18-VI-97 (97G120); nr Mabton, 9-V-95 (95G0317+); Selah Cr & Hwy 821, 15-V-95
(95G067+); YIR, Ft. Simcoe, 30-V-96 (95G0917).

Lysaphidus ramithyrus (Smith)
Pleotrichophorus nr wasatchii (Knowlton): WA, KittirAs—Yakima Canyon, 11-IX-95 on Chrysothamnus sp.
(95G5497).
Pleotrichophorus elongatus (Knowlton): WA, LINCOLN—Lake Roosevelt, Goldsmith CG, 31-VII-97 on
Chrysothamnus sp. (97G306).

Lysaphidus rosaphidis (Smith)
Aphis (Zyxaphis) sp.: WA, KirrirAs—Taneum Rd, 11-IX-95 on Artemisia tridentata (95G5857¢£).
Chaetosiphon fragaefolii (Cockerell) [on cultivated Fragaria sp.]; WA, CLARK—Vancouver, 30-VIII-95
(95K124); WSU-Vancouver, 21-I[X-95 (95K166, 95K170).
Chaetosiphon nr thomasi HRL: WA, Clark—WSU-Vancouver, Greenhouse, 15-V-97 on Fragaria sp.
(97X020).
Pleotrichophorus sp.: KittrrAs—Taneum Rd, 11-IX-95 on Artemisia tridentata (95G5857+).

Lysaphidus spp.

Aphthargelia symphoricarpi (Thomas): WA, SPOKANE—Mt Spokane SP, 14-VI-95 on Symphoricarpos sp.
(95T116).

Chaetosiphon fragaefolii (Cockerell): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native
Rosa sp. (96T2097+#).

Ericaphis gentneri (Mason): WA, CHELAN—Stevens Pass, 16-[X-95 on Spiraea densiflora (95K138).

Ericaphis scammelli (Mason): WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa
sp. (96T2097#).

Macrosiphum euphorbiae (Thomas): WA, GARFIELD—FR40 nr Teal Springs, 1-VIH-95 on Penstemon sp.
(9513797).

Nasonovia (Kakimia) sp.: WA, SPOKANE—Mt Spokane, Scribner Rd, 29-VI-95 on Phlox sp.? (95T176).

Genus LYSIPHLEBUS Foerster

Lysiphlebus flavidus Gahan
Cedoaphis incognita Hottes & Frison: WA, YAKIMA [on Symphoricarpos sp.*}.

Lysiphlebus testaceipes (Cresson)
Acyrthosiphon lactucae (Passerini) [on Lactuca serriola|: ID, LATAH—Moscow Mtn, 15-VIII-96 (961447);

710 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

WA, ASOTIN [*]; BENTON—-WSU-Prosser, 5-VIII-96 (96G434); CHELAN—Chelan SP, 11-IX-97 (97K038);
Kittitas [*]; WHITMAN—Pullman, 17-VIII-95 (95T446); YAKIMA—YIR, Signal Peak, 19-IX-95 (95G621).

Acyrthosiphon macrosiphum (Wilson): WA, SPOKANE—nr Peone, 13-VII-95 on Ribes sp.? (95T227).

Acyrthosiphon pisum (Harris): WA, BENTON—-WSU-Prosser, 26-IV-97 on Medicago sativa (97G014).

Anoecia corni (F.): WA, YAKIMA—Ahtanum Mission, 26-IX-95 on Cornus stolonifera (95G6627¢£).

Aphis armoraciae Cowen: MT, LINCOLN—Libby, 16-VI-95 on Centaurea maculosa (95G172); WA, ASOTIN—
Anatone, 24-VII-96 on unknown plant (96T330); KittiTAs [on Mertensia paniculata *|—Reecer Cr., 20-
VI-95 on Centaurea diffusa (95G2057); KLIicKkIrat—Bickleton, 7-VII-97 on Tragopogon dubius (97G225);
WALLA WALLA—Prescott, 30-X-97 on Taraxacum officinale (97G509); YAKIMA—nr Hause CG, 15-VI-96
on Achillea millefolium (96K1047); Wenas Lk, 11-VI-97 on A. millefolium (97G053).

Aphis nr armoraciae Cowen: WA, YAKIMA—nr Parker, 28-[X-95 on Centaurea sp. (95G664).

Aphis canae Williams: WA, AsoTtin—Asotin Cr Rd, 30-V-95 on Artemisia tridentata (95T045*$).

Aphis ceanothi Clarke [on Ceanothus velutinus unless otherwise indicated]: MT SANDERS—13 mi E of
Thompson Pass, 29-VII-95 (95G459); WA, Kittiras—Manastash Cr Rd, mi marker 7, 16-VIII-96 on C.
sp. (96G487); Manastash Cr, 25-VI-9 (97G137); Quilomene WLA, 2-VIU-97 (97G213); Lost Lk, 6-VIII-
97 (97G332); KLICKITAT—Trout Lk, 29-VI-97 on C. sanguineus (97G161); YAKIMA—YIR, nr. Mt. Adams
Lk., 25-VII-95, 4-VIII-95 on C. sanguineus (95G409, 95K0787), Miller Point, 19-IX-95 (95G6277) on]),
Howard Lk Rd 19-IX-95 (95G641).

Aphis nr ceanothi Clarke: WA, YAKIMA—YIR, Signal Peak, 23-VI-95 on Ceanothus sp. (95G2317), & nr.
Mt. Adams Lk., 19-IX-95 on C. velutinus (95G607).

Aphis nr chrysothamni Wilson: WA, AsoTiIN—Asotin Cr Rd, 13-VI-95 on Chrysothamnus sp. (95T088).

Aphis coweni Palmer [on Veratrum viride unless otherwise indicated]: WA, AsoTIN—De Spain Springs, 5-
VII-96 (96T155); CHELAN—Chatter Cr, 1-VIII-85 on Pyrola asarifolia (95G4857); Kittrras—Wells
Meadow, 25-VII-96 (96G413).

Aphis craccivora Koch: WA, BENTON—-WSU-Prosser, 3-X-95 & 8-XI-95 on Asparagus officinalis (95G672,
95G747); Crow Butte, 24-VI-96, on Astragalus sp. (96GO013); Kittrras—Indian John Hill, 11-X-96 on
Kochia scoparia (96G639); KLickirat—Alderdale Rd & Hwy 14, 9-V-95 on Astragalus sp. (95G025);
Hwy 14, 4 mi W of Alderdale Rd, 14-V-96 on Medicago sativa (96G030); WHITMAN—Pullman, 25-VII-
95 on Capsella bursa-pastoris (951331); 2-VIII-96 on Melilotus alba (96T391); 3, 7,16 & 31-VII-96,
2,7,16,19 & 29-VIII-96, 9-X-96 on Robinia pseudo-acacia (961133, 96T199, 96T301, 961364, 96T393,
96T415, 961462, 96T469, 96T511 & 96T580); YAkIMA—Grandview, 30-IX-96 & 3-VIII-96 on Robinia
pseudo-acacia (95G666, 96G433); nr Union Gap, 23-X-95 on Medicago sativa (95G721+); YIR, Yakima
Chief & Pioneer Rds, 16-V-96 on Astragalus sp.(96GO059).

Aphis nr epilobiaria Theobald: WA, OKANOGAN—Bridgeport SP, 9-IX-97 on Epilobium minutum (97K017*).

Aphis fabae Scopoli [on Cirsium arvense unless otherwise indicated]: ID, LATAH—Moscow Mtn, 15-VIII-96
(961448); 13-VII-96 on Holodiscus discolor (96T267); Moscow Mtn, nr start of Paradise Cr, 21-IX-96
(96T542); WA, AsotTin—Field Springs SP, 18-VII-95 (95T272); Wickiup CG, 29-VIII-95 (95T504);
Anatone, 28-VI-96 (96T083); GARFIELD—Pataha Cr Rd, 5.6 mi S of Columbia Center, 7-[X-95 (95T544);
Columbia Center, on Philadelphus lewisii (95T430); GRANT—Sun Lakes SP, 8-IX-97 on C. sp. (97K008);
KitrirAs—Buck Meadows, 11-IX-95 (95G590), & on C. vulgare (95G580); Frost Meadows, 11-IX-95
(95G5837); Lost Lk, 11-IX-95 (95G556+, 95G564+, 95G571+) & on C. vulgare (95G577); 6-VIII-97, on
Senecio sp. (97G335); Manastash Rd, 11-IX-95, 25-VII-96, & 16-VIII-96 (95G5567, 96G424, 96G478,
[96G481 on C. vulgare]); Lost Lk Trailhead, 6-VII-97 (97G320); LEwis—RNP, Bench Lk, 23-VII-96 on
native C. sp. (96G375); KLICKIrAT—nr Bickleton, Cemetery Rd 1 mi N of Hwy, 18-VI-97, 7-VII-97
(97G107, 97G227); Klickitat River Hwy 142, 28-VI-97 on C. vulgare (97G155); OKANOGAN—Winthrop,
10-IX-97 on Arctium minus (97K031); SPOKANE—Mt Spokane SP, 20 & 27-VII-95, 24-VIII-95, (95T293,
95T340, 95T349, 951474): 13-VII-95, on Umbelliferae (95T241); 24-VIII-95 on C. ochrocentrum
(951476, 95T481); WHITMAN—Pullman, 1-VIII-95, & 7, 18-VII-96 (95T387, 96T198, 96T323); YAKIMA—
Buckhorn Meadows, 14-IX-95 on C. vulgare (95G600); American River, 5-IX-97 (97G373) & on
Epilobium angustifolium (97G377+); Blowout Mtn, end of Little River Rd, 9-[X-97 (97G390); YIR, Piscoe
Cr 25-VII-95, on Rumex crispus (95G437); Signal Peak, 19-IX-96 on C. vulgare (96G563).

Aphis nr fabae Scopoli: ID, LATAH—Moscow Mtn, 15-VIII-96 on Rumex crispus (96T450); WA, PEND
OREILLE—Sullivan Lk Rd & Harvey Cr, on Chrysanthemum leucanthemum (951394); PIERCE—RNP,
Tipsoo Lk, 11-IX-96 on Osmorhiza sp. (96G531).

Aphis filifoliae (G&P): WA, Asotin—Asotin Cr Rd, 11-VII-95 on Artemisia tridentata (95T202).

Aphis nr frangulae Kaltenbach [on Nepeta cataria]: WA, ASOoTIN—Couse Cr Rd, 10-VIII-95 (95T424); nr
FR40 & FR44, 5 & 19-VII-96 (96T161, 961242, 961243); De Spain Spring, 11-VII-96 (961242, 96T243).

Aphis gossypii Glover: WA, AsoTiIN—Heller’s Bar, 30V—95 on unknown plant (95T039).

VOLUME 102, NUMBER 3 Till

Aphis hederae Kaltenbach: OR, BENTON—Corvallis, 21-V-91 on unknown plant (91J001).

Aphis helianthi Monell: ID, KooTENAI—Fernan Lk, on Rumex sp. (95K030); SHOSHONE—Marble Cr 9-VI-96
on Philadelphus lewisii (96T012); MT, SANDERS—Thompson Pass, 17-VI-95 on Heracleum lanatum
(95G179+); 13 mi E of Thompson Pass, 29-VII-95 on Ligusticum verticillatum (95G460); WA, AsoTIN—
Asotin Cr HMU, 29-VIII-95 on Helianthus annuus (957496); Bracken Point, 7-[X-95 on H. annuus
(95T548); Anatone, 11-VII-95 on Lomatium sp. (95T217); De Spain Springs, 11-VII-96 on Lomatium sp.
(961245); CHELAN—Chatter Cr, 1-VIII-95 on Oplopanax horridum (95G4887+); Stevens Pass, 16-[X-95
(95K 1347); GARFIELD—Stentz Springs, 22-VII-96 on Osmorhiza chilensis (957452); Kittitas [on
Heracleum lanatum*|—S Fork Manastash Cr, 24-VII-95 on Angelica sp. (95G392); Buck Meadows, 24-
VII-95 on native Cirsium sp. (95G396); Manastash Rd, 24-VII-95 & 11-IX-95 on Heracleum lanatum
(95G378, 95G5607); 24-VII-95 on Penstemon sp. (95G379); Lost Lk, 11-[X-95 on Heracleum lanatum
(95G572), 2-[X-95 on Epilobium angustifolium (95G530+), & 6-VIII-97 on Angelica arguta (97G3337+);
Manastash Cr, mi marker 5, 16-VIII-96 on Heracleum lanatum (96G483); Reecer Cr, 20-VI-95 on
Ligusticum sp. (95G201) & 20-VI-95 on Lomatium nudicaule (95G209); Quartz Mtn, 25-VII-96 on
Penstemon procerus (96G399) & Ligusticum grayi (96G3987+); Colockum Ridge, 2-VII-97 on Camassia
quamash (97G196); Colockum Ridge, 1 & 2-VII-97 on Lomatium nudicaule (97G189) & Lomatium
triternatum (97G1917+); Quilomene WLA, 2-VII-97, on Lomatium sp. (97G212); LEwis—RNP, Stevens
Canyon, 23-VII-96 on native Cirsium sp. (96G355) & 8-IX-95 on Ligusticum sp. (95K113); OKANOGAN—
Bridgeport SP, 9-IX-97 on Helianthus annuus (97K016); PIERCE [on Cirsium sp. & Heracleum lanatum*|—
RNP, nr Paradise, 8-I[X-95 on native Cirsium sp. (95K122) & Xerophyllum tenax (95K121); Tipsoo Lk, 5-
IX-97 on Ligusticum grayi (97G3637+); SKAMANIA [on Ligusticum apiifolium*]; SPOKANE—nr Peone, 13-
VII-95 on Cornus stolonifera (95T236); Turnbull National WLA on Cornus stolonifera (95T009); Mt
Spokane SP, 29-VI-95 on Cornus stolonifera (95T186); nr Mt Spokane, 29-VI-95 on Heracleum lanatum
(95T171); nr Peone, 13-VII-95 on Heracleum lanatum (95T229); WHITMAN—nr Colton, Steptoe Canyon
Rd, 19-IX-95 on Helianthus annuus (957586); Pullman, I-VIII-95 on Coriandrum sativum (9513867);
YAKIMA [on Cirsium sp.*|—Mabton & Alderdale Rds, 23-V-96 on Lomatium sp. (96G071); Wenas Lk,
30-V-96 & 11-VI-97 on Lomatium sp. (96G118, 97G063+) & 17-VI-97 on unknown Umbelliferae
(97G088); American River, 5-[X-97 on Cirsium sp. (97G379), 5-[X-97 on Heracleum lanatum (97G382);
Blowout Mtn, end of Little Naches River Rd, 9-IX-97 on Cirsium sp. (97G399) & on Xerophyllum tenax
(97G395, 97G398); Sawmill Flats CG, 28-VII-97 on Cornus stolonifera (97G286); YIR, Lower Mill Cr,
25-V-96 on Lomatium dissectum (96K052) & Lomatium tritematum (96K053); Piscoe Cr, 25-VII-95 on
Ligusticum grayi (95G432); Potato Hill Guard Station, 25-VII-95 on Xerophyllum tenax (95G429):

Aphis nr helianthi Monell: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Lomatium sp. (95T517++); WA,
KitTirAs—Manastash Cr Rd, mi marker 4, 16-VIII-96 on Cornus stolonifera (96G474); YAKIMA—Chinook
Pass, 27-I[X-96 on Ligusticum sp. (96G604).

Aphis hermistonii Wilson: WA, GRANT—Sun Lakes SP, 8-IX-97 on Chrysothamnus nauseosus (97KO009).

Aphis holodisci Robinson [on Holodiscus discolor unless otherwise indicated]: MT, SANDERS—Bull River
Valley, 16-VI-95 on Physocarpus malvaceus (95G176); WA, AsoTiIN—De Spain Springs, 26-VI-96, 5-VII-
96 (96T059, 96T 150); Kittrras—Manastash Cr, mi marker 12, 25-VI-97 (97G133); KLickiIrAt—Trout Lk,
11-VH-95 (95G340+); SPOKANE—Mt Spokane SP, 27-VII-95 (951338); Spokane, Manito Pk, 11-VII-96
(961260); YAKIMA [*]—Clear Lk, 15-VI-96 (96K079, 96K082); Bird Cr Meadows Rd, FR 82 & FR 822,
26-V95 on Physocarpus malvaceus (95A020, 95A021); YIR, White Deer Cr, 11-VII-95 (95G324) & Signal
Peak (95G333), & nr Mill Cr Guard Station, 25-V-96 (96K047).

Aphis lugentis Williams: WA, YAKIMA [on Senecio triangularis & Arnica sp.*].

Aphis lupini G&P [on Lupinus sp.]: WA, GARFIELD—Pataha Cr, 7-IX-95 (95T546); STEVENS—Little Pend
Oreille Nat. WLA, 9-VII-96 (96T212).

Aphis neogillettei Palmer [on Cornus stolonifera]: WA, KLICKITAT [*]; YAKIMA—Sawmill Flats CG, 28-VII-
97 (97G285).

Aphis nerii Boyer de Fonscolombe [on Asclepias speciosa]: WA, BENTON [*]; CHELAN—-WSU-Wenatchee,
25-VIII-95 (95RO025); YAKIMA [*]—Sunnyside, 24-VIII-95, 1-X-95, 23-X-95, 13-VIII-96, 29-IX-96, 4, 22-
X-97 (95G517, 95G670, 95G704, 96G452, 96G618, 97G486, 97G489); Union Gap, 23-X-95 (95G725);
Mabton, 30-VIII-96 (96G513).

Aphis nigratibialis Robinson [on Cornus stolonifera |}: WA, Kitt1rAs—Manastash Cr, mi marker 12, 6-VIII-
97 (97G312); KLickiIrat—Trout Lk, 28-VI-97 (97G156); SPOKANE—Mt Spokane SP, 13-VII-95 (951239);
YAKIMA—Chinook Pass, 28-VII-97 (97G280).

Aphis oenotherae Oestlund: WA, OKANOGAN—Bridgeport SP, 9-IX-97 on Epilobium minutum (97K017*);
PIERCE—RNP, nr Chinook & Cayuse Passes, 16-VIII-95 on Ribes viscosissimum (95A039).

Aphis nr oenotherae Oestlund [on Epilobium sp. unless otherwise indicated]: WA, BENTON—Kennewick, 28-

712 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

[X-96 (96G597); GARFIELD—Pataha Cr Rd, 7-IX-95 (95T541, 95T545); Spruce Springs, 22-VIII-95 on
Physostegia parviflora (95T462+); OKANOGAN—Bridgeport SP, 9-I[X-97 on Oenothera strigosa (97K020);
YAKIMA—American River, 5-IX-97 on Epilobium minutum (97G375*); Fifes Peak Viewpoint, 27-[X-96
(96G603); Nile, 5-[X-97 (97G358); nr Rimrock, 14-[X-95 (95G594).

Aphis pomi de Geer: WA, DouGLAs—Bridgeport, 17-VI-96 on Pyrus malus (96X007*¥£).

Aphis rumicis L. [on Rumex crispus]: WA, BENTON [*]; YAKIMA—YIR, Satus, 1-VI-95 (96K065).

Aphis salicariae Koch [on Epilobium angustifolium]: ID, KOOTENAI—Rathdrum, 29-VII-95 (95G443+); WA,
AsoTIN—Anatone, 24-VII-96 (961336); CHELAN—1-VIII-95 (95G481+); Kirriras—Lost Lk, 2-IX-95
(95G5307); OKANOGAN—Washington Pass, 10-I[X-97 (97K033); SPOKANE—nr Cheney, 23-V-95 on Cornus
stolonifera (95K007); YAKIMA—YIR, nr. Mt. Adams Lk., 19-I[X-95 (95G6107).

Aphis nr salicariae Koch: WA, CHELAN—Swauk Pass, 1-VIII-95 on Epilobium angustifolium (95G492*).

Aphis sambuci L: WA, PIERCE, [on Sambucus racemosa*}.

Aphis spiraecola Patch: WA, BENTON [on Lactuca serriola*]; CHELAN—-WSU-Wenatchee, 7-VII-95 on Pyrus
malus (95RO18+); KitrirAs—Lost Lk, 6-VIII-97 on Angelica arguta (97G333+); YAKIMA—Sunnyside, 3-
VI-95, on cultivated Spiraea sp. (95G109); nr Sunnyside, 28-VI-95 (95G2597).

Aphis nr triglochinis Theobald: WA, SPOKANE—Mt Spokane SP, 29-VI-95 on native Ribes sp. (95T189).

Aphis nr valerianae Cowen: WA, PIERCE—RNP, Mowich Lk, 25-IX-97 on Valeriana sitchensis (97G446).

Aphis varians Patch [on Epilobium angustifolium unless otherwise indicated]: MT, SANDERS [*]: ID,
KooTenaI—Rathdrum, 29-VII-95 (95G443+); WA, CHELAN—Stevens Pass, 16-IX-95 (95K1347); PIERCE
[*]—RNP, nr Paradise, 8-IX-95 (95K114, 95K117), Tipsoo Lk, 5-[X-97 (97G368), & Mowich Lk, 25-IX-
97 (97G442): Naches Pass, 25-IX-97 (97G460); SPOKANE—Mt Spokane SP, 24-VIII-95 (95T483); YAKIMA
[*]—Green Lk, 31-VIII-94 on E. latifolium (94R140); Blowout Mtn, end of Little Naches River Rd, 9-
IX-97 (97G404); YIR, Hussen Meadows, 19-IX-95 (95G643) & Howard Lk,19-IX-95 on Lupinus sp.
(95G639);

Aphis nr varians Patch [on Epilobium angustifolium]: WA, CHELAN—Dryden, 1-VIII-95 (95G4817+); LEwis—
RNP, Bench Lk, 23-VII-96 (96G376); Stevens Cr, 23-VII-96 (96G380).

Aphis (Zyxaphis) sp.: WA, AsoTIN—Asotin Cr Rd, 22-VIII-95 on Chrysothamnus sp. (95T469).

Aphis sp. n.. WA, YAKIMA—YIR, Klickitat River nr Miller Point, 23-VI-95 & 11-VII-95 on Ceanothus sp.
(95K061, 95G230) & 19-IX-95 & 19-IX-96 on C. velutinus (95G627+, 96G580).

Aphis sp.: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Cirsium vulgare (95T531); LataH—Moscow, 15-
IX-93 on Solanum lycopersicon (93T020); WA, AsoTiIn—Anatone, 28-VI-96, on Lamium sp. (96T086);
Schumaker Rd mi marker 1, 10-VIII-95 on Cirsium arvense (957428); 28-VI-96, on Holodiscus discolor
(96T085); De Spain Spring, 5-VII-96, on Nepeta cataria (96T 161); 11,24-VH-96 on unknown Umbelliferae
(961254, 961341); CHELAN—Chelan SP, 11-IX-97, on Salix sp. (97K0427); GARFIELD—Alder Thicket (FR
40), 22-VII-95 on native Cirsium sp. (951448); Kittiras [on Pedicularis bracteosat & Senecio
triangularis*|—Lost Lk, 6-VIII-97 on Symphoricarpos sp. (97G341+); Taneum Lk, 11-IX-97 on
Pedicularis racemosa (97G411+); [KLICKITAT, on Rheum rhabarbarum*|—Trout Lk, 29-VI-97 on
Amsinckia sp. (97G172); PEND OREILLE—Cusick, 9-VII-96 on Ceanothus sanguineus (961225); PIERCE—
RNP, Tipsoo Lk, 5-IX-97 on Valeriana sitchensis (97G360); SPOKANE—Mt Spokane SP, 14-VI-95 on
Philadelphus lewisii (957117); 24-VIII-95 on Salix sp. (95T484), 29-VI-95 on unknown Umbelliferae
(95T183); Hwy 206 mi marker 16.3, 8-VI-95 on unknown Umbelliferae (95T070); WHITMAN—Pullman,
1-VII-95 on Coriandrum sativum (95T386); YAKIMA [on Umbelliferae*]—nr Chinook Pass, 11-IX-96 on
Cryptantha sp. (96G519) & on Epilobium angustifolium (96G555); Little Naches River Rd, 28-VII-97 on
Cryptantha thompsonii (97G291); Wenas Cr & Maloy Rd, 11-VI-97 on Heracleum lanatum (97G070);
YIR, Delaney Springs, 25-VII-95 on Holodiscus discolor (95G404), Windy Point, 19-I[X-95 on Cimicifuga
laciniata (95G631), & Signal Peak Guard Station, 19-[X-95 on Cornus stolonifera (95G617).

Aphthargelia symphoricarpi (Thomas) [on Symphoricarpos sp. unless otherwise indicated]: WA, KITTITAS
[*]—Colockum Rd, mi marker 4, 1-VII-97 (97G183); Lost Lk, 6-VHI-97 (97G334); KiickiTaT [on S.
albus*]; SPOKANE—Mt Spokane SP, 14-VI-95, 13-VII-95 (95T116, 95T242); Turnbull Nat. WLA, 16-VII-
96 on unknown plant (961287); Fish Lk, 14-VI-97 on S. occidentalis (97K004); YAKIMA—Wenas Cr &
Maloy Rd, 11-VI-97 (97G066); YIR, Signal Peak, 11-VII-95 (95K057).

Boernerina variabilis Richards: WA, KirrirAs—Buck Meadows, 9-VII-96 on Alnus sp. (96G336).

Brachycaudus cardui (L.) [on Cirsium vulgare unless indicated otherwise]: ID, LAtAH—Moscow Mtn, 3 &
15-VIII-96 (96T401, 961449); Nez Perce—Hwy 95 & Hwy 195, 22-VII-97 on C. arvense (97J004); WA,
AsotTin—Heller’s Bar, 11-VII-95 on C. arvense (95T212); Anatone, 24-VII-96 (961333) & on Onopordum
acanthium (961329); Asotin, 28-VI-96 on O. acanthium (961066); Weisenfels Ridge Rd nr Snake River
Rd) 28-VI-96 on O. acanthium (96T070); BENTON [*]—Prosser, Rotha Rd, 24-VIII-95 (95G512);
GaRFIELD—Spruce Springs, 22-VIII-95 on C. arvense (95T457); Pataha Cr Rd, 7-IX-95 (95T542);

VOLUME 102, NUMBER 3 TAS

KLIckITAT—Bickleton, 18-VI-96 & 7-VII-97 on C. sp. (96K112) & C. vulgare (97G228); Brooks Memorial
SP, 21-IX-95 on C. sp. (95K162+): SPOKANE—Mt Spokane SP 24-VIII-95 (951475, 95T482. 95T488):
Whitman—Pullman, 1,13-VIII-96, 3-[X-96 (96T369, 96T383, 961439, 961514); YAKIMA—nr Naches Pass,
FR 789 & FR 1914, 9-IX-97 (97G410): YIR, nr. Mt. Adams Lk., 25-VII-95 (95G405), Signal Peak, 19-
IX-95 (95G624), Satus, 1-VI-96 on native C. sp. (96K066), & Snow Cr, 19-IX-95 (95G608).

Brachycaudus helichrysi (Kaltenbach): WA, FRANKLIN—Ice Harbor Dam, 28-V-97 on Amsinckia sp. (96G097):
Kittitas [on Rudbeckia_hirta*); WHITMAN—Pullman, 19-VII-96 on Tragopogon dubius (961320):
YAKIMA—YIR, nr. Mt. Adams Lk., 4-VIII-95 on native Cirsium sp. (95KO077).

Brachycaudus tragopogonis (Kaltenbach) [on Tragopogon dubius]: ID, SHOSHONE—Kingston, 29-VII-95
(95G479); OR, UMaTILLA—Milton-Freewater, 11-VI-96 (96G239); WA, BENTON—-WSU-Prosser, 27-VI-
96, 15-VII-96, 30-V-96, 14-V-97 (96A003, 96A014, 96G100, 97G028); Rotha Rd, 18-VI-96 (96G256);
Badger Springs, 28-IX-96 (96G595): West Richland, 27-V-96 (96K056):; CHELAN—Chelan SP, 11-IX-97
(97K040); Kittiras—Ellensburg Pass, Wenas-Ellensburg Rd, 9-VI-96 (96G209); Indian John Hill, 11-X-
96 (96G638); KLicKITAT [*]—Bickleton, 17,18-VI-96, 27-VIII-96, 8-X-96 (96G254, 96K113, 96G512,
96G628); Box Springs, 27-VI-97 (97G147): WHITMAN—Pullman, 1,2,7,12-VII-96, 2,9-VIII-96, 3-X-96
(96T126, 96T132, 961197, 96T325, 961387, 961427, 961574); Yakima—Grandview, 24-V-96, 2 & 26-
VI-96, 13-X-96, 14-V-97 (96G080, 96G130, 96G273, 96G283, 96G646, 97G022): Halfway Flat CG, 27-
IX-96 (96G616); Hause CG, 15-VI-96 (96K 106); nr Bickleton, 18-VI-97 & 7-VII-97 (97G102, 97G222):
Sunnyside, 1-VI-96 (96K058): Wenas Lk, 30-V-96 (96G124); YIR, Signal Peak, 19-IX-96 (96G560) &
Goat Butte, 19-IX-96, 9-X-96 (96G583, 96G631).

Brachycorynella asparagi (Mordvilko) [on Asparagus officinalis]; WA, BENTON [*]—WSU-Prosser, 6,25-IX-
95 (95K 144, 95G649).

Braggia eriogoni (Cowen): MT. GALLATIN—Big Sky, Gallatin Rd, 26-VI-96 on Eriogonum sp. (96K119):
WA, AsoTiIN—De Spain Springs, 11,19-VII-96 on E. sp. (961246, 961317); Round Prairie Springs, 18-
VII-96 on E. sp. (96T311): KING—Stampede Pass, 8-VIII-96 on E. compositum (96G446); Kirtiras—
Colockum Ridge Rd, 2-VII-97 on E. compositum (97G190, 97G193); Taneum Lk Trail, 11-IX-97 on E.
umbellatum (97G428):; KLICKITAT [on E. heracleoides*|—Bickleton (Pine Cr Rd), 3-V-95 on E. heracleoides
(95G099); Cemetery Rd 1 mi from Hwy, 18-VI-97 on E. sp. (97G106): Cleveland, 18-VI-96 on E.
heracleoides (96K111).

Braggia nr eriogoni (Cowen) [on Eriogonum compositum unless otherwise indicated]: OR, UMATILLA—Harris
Pk, 11-VI-96 (96G234); WA, Kittiras—Lost Lk, 6-VIII-97 (97G350); KLICKITAT—Roosevelt, 14-V-96
(96G035); Bickleton, 18-VI-97 (97G110); 18-VI-97, on E. sp. (97G109): YAKIMA—Little Naches River
Rd, 28-VII-97, on E. strictum (97G290).

Braggia sp.: WA, KLickiIrat—nr Bickleton, Pine Cr Rd, 31-V-95 on Eriogonum umbellatum (95G100); Box
Springs, 29-VI-97 on E. compositum (97G176): YAKIMA [on E. compositum & E. elatum (aphid det. by
G. Remaudiére)*]—Hause CG, 15-VI-96 on E. elatum (96K105); Wenas Lk, 11-VI-97 on E. sp. (97G060);
YIR, Klickitat River, 7-VI-95 on E. compositum (95G126).

Braggia sp. n. [on Eriogonum elatum unless otherwise indicated]: WA, DouGLas—Orondo, 28-V-96
(96RO002); Kirriras—Lost Lk, 2-IX-95 on E. compositum (95G529); Reecer Cr, 20-VI-95 (95G213);
YAKIMA—Bear Cr nr Rimrock Lk, 9-VI-95 (95A006): Naught Rd, 18-VI-97 (97G104) & 7-VII-97 on E.
compositum (97G220); Naches, Hwy 410 & Hwy 12, 28-VII-97 (97G276); Naches River, Horseshoe Bend,
5-IX-97 (97G357); nr Rimrock Lk, 14-IX-95 on E. sp. (95G595); Wenas Lk, 30-V-96,11-VI-97 (96G114,
97G061); Windy Point CG, 3-VI-95, 1-VII-95, 26-VI-96 (95G105, 95K040, 96G305), & 16-VIII-95 on E.
compositum (95A031); YIR, Fort Simcoe, 6-VI-95 (95G117), Mill Cr, 19-IX-95 (95G605), Signal Peak,
19-IX-95 (95G626), & 20 mi S of Satus Pass, 7-VI-95 on E. sp. (95G142).

Brevicoryne brassicae (L.): WA, BENTON—WSU-Prosser, 6-VII-95 on Brassica napus (95A024).

Capitophorus elaeagni (del Guercio): WA, YAKIMA—Buckhorn Meadows, 14-IX-95 on Cirsium vulgare
(95G596); nr Naches Pass, FR 789 & FR 1914, 9-IX-97 on Cirsium sp. (97G408).

Cavariella aegopodii (Scopoli): WA, Kittiras—nr Lost Lk, 6-VIII-97 on Osmorhiza chilensis (97G331) &
2-IX-95 on Lomatium sp. (95G535+); YAKIMA—nr Wenas Lk, 17-VI-97 on Lomatium sp. (97G095+).
Cavariella pastinacae (L.): MT. SANDERS—Thompson Pass, 17-VI-95 on Heracleum lanatum (95G179+#);
WA, AsoTiIn—Asotin Cr Rd, 11-VII-95 on Oenothera sp. (95T196): YAKIMA—YIR, Klickitat Canyon, 23-

VI-95 on Heracleum lanatum (95G235).

Ceruraphis viburnicola (Gillette): WA, YAKIMA—Yakima Arboretum, 26-IV-95 on Viburnum sp. (9SGO08).

Chaetosiphon fragaefolii (Cockerell); WA, PEND OREILLE—Le Clerc WLA, 3-VIII-95 on native Rosa sp.
(95T398).

Chaitophorus populicola Thomas: WA, Asotin—Asotin Cr Rd, 23-V-95 on Populus trichocarpa (95T013).

714 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Chaitophorus viminalis Monell: WA, YAKIMA—Naches, 27-[X-96 on Salix sp. (96G601).

Cinara chinookiana Hottes: WA, YAKIMA—Darland Mtn, 31-VIII-94 on Abies lasiocarpa (94R138).

Cinara sp.: WA, ASOTIN—FR44 8mi S of WLA, 18-VII-96 on Pinus ponderosa (961308).

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, AsotiIN—Anatone, 10-VIII-95 (95T427); Pataha
Cr Rd, 5.6 mi S of Columbia Center, 10-VIII-95 (951435); BENTON [*]|—WSU-Prosser, 1-IV-97, 24-VII-
97 (97G005, 97G274); KLickirat—nr Cleveland, 7-VII-97 (97G232, 97G234); Bickleton, 29-VII-97
(97G300); SPOKANE—Mt Spokane SP, 22-VII-95 (95T495); YAKIMA [*]—Ridge Rd, 17-VI-96 (96G250,
96G251); Sunnyside, 24-VII-97 (97G273).

Dysaphis plantaginea (Passerini) [on Pyrus malus unless otherwise indicated]: WA, BENTON—Prosser, 22-VI-
97 (97G123); CHELAN—-WSU-Wenatchee, 7-VII-95 & 30-V-96 (95RO187+, 96ROO09, 96RO11+); WHITMAN—
Pullman, 30-VI-96, on unknown plant (96T119); YAKIMA—Grandview 26,28-VI-96 (96G282, 96G308);
nr Sunnyside, 28-VI-95 (95G2597).

Ericaphis nr gentneri (Mason): WA, SPOKANE—nr Peone, 13-VII-95 on Crataegus sp. (95T223).

Eriosoma americanum (Riley): WA, CHELAN—Swakane WLA, 3-VI-96 on Ulmus americanus (96R018%).

Eriosoma lanigerum (Hausmann): WA, CHELAN—-WSU-Wenatchee, 7-VI-95 on Pyrus malus (95RO19¢%).

Hyadaphis foeniculi (Passerini): ID, BENEWwAH—McCroskey SP, 31-VIHI-95 on Lomatium sp. (95T5177%).

Hyperomyzus nigricornis (Knowlton): WA, YAKIMA—Wenas Lk, 30-V-96, on Ribes aureum (96G123); YIR,
19-IX-95 on Agoseris sp. (95G622).

Illinoia spiraeae (MacGillivray): WA, GARFIELD—Teal Springs, 1-VIII-95 on Holodiscus discolor (951376).

Macrosiphum euphorbiae (Thomas): ID, KooTeNAI—Rathdrum, 29-VH-95 on Epilobium angustifolium
(95G4437+); LataH—Moscow, 29-VIII-96 on Solanum sp. (96T512+); WA, AsoTIN—De Spain Springs 5-
VII-96 on Compositae (96T159); BENTON—Prosser, 24-X-96 on Medicago sativa (96G651); KLICKITAT—
Trout Lk, 11-VII-95 on Holodiscus discolor (95G340*); PIERCE [on Epilobium angustifolium*].

Macrosiphum gaurae (Williams): WA, AsoTiIN—nr Weissenfels Ridge & Snake River Rds, 28-VI-96 on
Oenothera sp. (96T073).

Macrosiphum nr pallidum (Oestlund): WA, YAKIMA—American River, 5-IX-97 on Epilobium minutum
(97G375+#).

Macrosiphum sp. [nr fuscicornis MacDougall & pallidum (Oestlund)]: WA, CHELAN—Swauk Pass, 1-VIII-95
on Epilobium angustifolium (95G4927).

Myzus ascalonicus Doncaster: WA, YAKIMA [on moss*].

Myzus cerasi (F) [on Prunus sp. unless otherwise indicated]: WA, BENTON—-WSU-Prosser, 28-V-96 (96G091);
CHELAN—Wood Reservoir, 6-VI-96 (96R019); Chelan SP, 11-I[X-97 (97K041); WHITmMAN—Pullman, 5-VII-
96, on P. avium (96T173).

Myzus nr cerasi (FE): WA, WHITMAN—Pullman, 6-VII-96 on unknown plant (961411).

Myzus persicae (Sulzer): ID, LatrAH—Moscow, 29-VIII-96 on Solanum sp. (9615127).

Nasonovia (Kakimia) sp.: WA, SPOKANE—nr Peone, 13-VII-95 on native Ribes sp. (951235).

Nearctaphis bakeri (Cowen): WA, BENTON—WSU-Prosser, 5-V-95 on Crataegus sp. (95G023); KLICKITAT—
Trout Lk, 29-VI-97 on Trifolium pratense (97G170).

Nearctaphis californica HRL: WA, KLicktrat—Brooks Memorial SP, 8VI-94 on Crataegus douglasii (94—
100).

Nearctaphis kachena (Hottes): WA, Kirtrras—Taneum Lk, 11-IX-97 on Pedicularis racemosa (97G4117#).

Nearctaphis yohoensis Bradley: WA, YAKIMA—YIR, White Deer Cr, 11-VII-95 on Sorbus sp. (95KO51).

Nearctaphis sp. 1D, LataH—Moscow, 13-VI-97 on unknown plant (97T050).

Phorodon humuli (Schrank); WA, BENTON [on Humulus lupulus & Prunus salicina*|—WSU-Prosser, 19,25-
IV-95, 23-V-96, 15-IV-97, on P. salicina (95G005, 95G006, 95G007, 96G075, 97G006).

Pseudoepameibaphis tridentatae (Wilson): WA, AsoTiIN—Asotin Cr Rd, 30-V-95 on Artemisia tridentata
(95T0457#).

Pterocomma sp.: WA, YAKIMA—YIR, Signal Peak, 19-IX-95, on Salix sp. (95G619).

Rhopalosiphum cerasifoliae (Fitch) [on Prunus virginiana unless otherwise indicated]: MT, LINCOLN—Libby,
29-VII-95 (95G451); SANDERS — 20mi E of Thompson Pass, 29-VII-95 (95G454); WA, KirtrrAs—Reecer
Cr, 20-VI-95 (95G202); KLICKITAT—E end of Colockum Pass Rd, 2-VII-97, 97G215); WHITMAN—Pullman,
19-VI1-96, 16-VII-96 on unknown plant (96T046, 96T300); YAkIMA—Wenas Lk, Wenas Cr & Maloy Rd,
11-VI-97 (97G069).

Rhopalosiphum enigmae Hottes & Frison: WA, YAKIMA—Buena, 23-X-95 on Typha latifolia (95G715).

Rhopalosiphum insertum (Walker) [on Pyrus malus unless otherwise indicated]: WA, CHELAN—Chelan, 11-
VI-96 (96X001, 96X002); WSU-Wenatchee, 30-V-96 (96RO011+).; DouGLAs—Bridgeport, 17-VI-96

VOLUME 102, NUMBER 3 WAS

(96X0077+); Orondo, 25-V-95 (95RO10); 11-VI-96 (96X003, 96X004, 96X005); Bridgeport, 12,17-VI-96
(96X006, 96X008); Kitriras—Ellensburg, 17-V-96 on Crataegus columbiana (96G066).

Rhopalosiphum maidis (Fitch): WA, BENTON [on Echinochloa crus-galli+, P. milliaceum, & Triticum
aestivum*|—WSU-Prosser 23-XI-94 on Panicum milliaceum (94G113), 2-XI-96 on Hordeum vulgare
(96G655), 18-XI-95 & 20-I-96 on Capsella bursa-pastoris (96GO003+, 95G7507*).

Rhopalosiphum nymphaeae (L.): WA, YAKIMA—Outlook, 23-X-95 on Typha latifolia (95G709).

Rhopalosiphum padi (L.) [on Triticum aestivum unless otherwise indicated]: WA, BENTON [on Echinochloa
crus-gallit, T. aestivum, & ZEA MAYs*|—-WSU-Prosser, 1-X-95 (95G668); nr Paterson, 12-X-95 (95G688):
Cemetery Rd (3 mi east of Hwy 221), 12-X-95 (95G690); DouGLas—Waterville, 12-VIII-95 (95B016,
95B017, 95B018); YAKIMA [*]. [Following collections on Zea mays]: AsoTIN—Chief Timothy HMU, 16-
VIII-95 (95T439); BENTON [on Echinochloa crus-galli & Zea mays *|; BENTON—-WSU-Prosser, 24-VII-
95, 18-IX-95, 6-X-95, 28-IX-96, 3-XI-97 (95G511, 95G601, 95G683, 95G684, 96G591, 97G518); Prosser,
21-IX-96 (96G626); Grandview, 24-VIII-95 (95G513); GRANT—Ephrata, 8-VII-97 (97K058); WALLA
WALLA—Hwy 124 (nr Ice Harbor Dam), 5-IX-95, 5,18-X-95 (95G543, 95G675, 95G699): Eureka, 5-X-
95 (95G681, 95G698); Prescott, 30-X-97 on Phalaris arundinacea (97G516); YAKIMA—Grandview, 25-
VIII-95, 18-IX-95, 24-X-95, (95G520, 95G602, 95G702); Harrah, 19-[X-95 (95G604); nr Toppenish, 19-
IX-95, 16-X-95, 5-XI-95 (95G603, 95G695, 95G745).

Schizaphis graminum (Rondani): WA, BENTON—15-VI-82, (82K001).

Sitobion avenae (F) [on Triticum aestivum unless otherwise indicated]: WA, Asotin [*]; BENTON—-WSU-
Prosser, 3,11,18 & 25-VII-97 (97F192, 97F193, 97F199, 97F200, 97F209, 97F215, 97F223,
97F231,97F249, 97F252, 97F260, 97F263, 97F264, 97F265, 97F273), & 18-VII-97 on Setaria lutescens
(97G243); KLickIrAtT—Bickleton Hwy & County Line Rd, 7-VII-97 (97G224).

Sitobion sp.: WA, KLickirAt—Trout Lk, 11-VII-95 on Holodiscus discolor (95G340*#).

Tuberculatus columbiae Richards: WA, YAKIMA—YIR, Mill Cr Guard Station, 6-VI-95 on Quercus garryana
(95G112, 95G113), & on Salix sp. (95G114).

Uroleucon cirsii (L.): WA, KLICKITAT—Brooks Memorial SP, 21-[X-95 on Cirsium sp. (95K162+); PIERCE [on
Cirsium arvense*).

Uroleucon sp.: ID, KOOTENAI—Rathdrum, 29-VII-95 (95G443}).

Undetermined spp.: ID, LATAH—Moscow, 29-VIII-96 on Coriandrum sativum (96T513); BENTON—Prosser,
1-VIII-95 on Portulaca oleracea (95A028); Kittiras—Stampede Pass, 8-VIII-96 on Epilobium
angustifolium (96K146); LEwis—RNP, 8-IX-95 on Luetkea pectinata (95K115); WHITMAN [on Lupinus
sp.*]—Pullman, 19-VI-96, 3-VII-96 on P. malus (96T042); YAKIMA—Grandview, 12-X-96 on Sonchus sp.
(96G647).

Lysiphlebus utahensis (Smith)

Aphis canae Williams: WA, FRANKLIN—Kahlotus Hwy, 11-VI-96 on Artemisia tridentata (96G2427+).

Artemisaphis artemisicola (Williams) [on Artemisia tridentata]: WA, FRANKLIN—Kahlotus Hwy, 28-V-96 &
11-VI-96 on Artemisia tridentata (96G096, 96G242++); OKANOGAN—Bridgeport SP, 9-IX-97 (97K018);
YAKIMA—Naught Rd, 18-VI-97 (97G1037#).

Epameibaphis atricornis G&P: WA, BENTON—Byron Rd, 14-V-97 on Artemisia tridentata (97G024+#).

Flabellomicrosiphum tridentatae (Wilson): WA, KLICKITAT—nr Roosevelt, 16-IV-96 on Artemisia tridentata
(96G0127+#).

Obtusicauda artemisiphila (Knowlton & Allen): WA, KLIicKkirAt—nr Roosevelt, 16-IV-96 on Artemisia
tridentata (96GO012+#).

Obtusicauda coweni (Hunter) [on Artemisia tridentata}: WA, ASoTIN—Asotin Cr HMU, 7-V-96 (96T 1427);
YAKIMA [*].

Obtusicauda filifoliae (G&P) [on Artemisia tridentata): WA, BENTON—Byron Rd, 14-V-97 (97G0247#);
KLICKITAT [*]—nr Roosevelt, 16-IV-96 (97G0127#).

Obtusicauda sp.: WA, KLICKITAT [on Artemisia tridentata*}.

Pseudoepameibaphis tridentatae (Wilson) [on Artemisia tridentata}: WA, BENTON—Byron Rd, 14-V-97
(97G024++); KiLickiITAt—nr Roosevelt, 16-IV-96 (96G012+£); YAKImMA—Naught Rd, 18-VI-97
(97G1037+#).

Lysiphlebus sp.
Aphis helianthi Monell: WA, PIERCE [on Heracleum lanatum*).
Aphis sp.: WA, PIERCE [on Heracleum lanatum*]; YAKIMA [on unknown Umbelliferae*].
Brachycaudus tragopogonis (Kaltenbach): WA, KLICKITAT [on Tragopogon dubius*].

716 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Genus MONOCTONUS Haliday
Monoctonus campbellianus Pike & Stary, new species
(Figs. 13-24)

Diagnosis.—The new species does not key to any of the North American species
because of its reduced wing venation. According to the classification by van Achterberg
(1989), it is part of the M. cerasi (Marshall) group. [Note, a part of the specimens of
some species classified by Stary & Smith (1976) as European species, but distributed in
N. America, are probably this new species].

Etymology.—The name of the new species is derived from the type locality, Campbell
Lake, Skagit Co., Washington.

Description.—Female. Head: Eyes with sparse setae in lower half. Tentorio-ocular line
subequal to % of intertentorial line; malar space somewhat longer than /; of eye length,
equal to half of intertentorial line. Antenna 13-segmented, short, as long as head,
mesosoma and metasomal tergum | together, thickened to apex. Flagellomere 1 [F,] (Fig.
15) length 3.5 width, without placodes, setae equal to segment diameter; F, (Fig. 16)
slightly shorter than F,, about % wider than F,, with 3 placodes; preapical F-segments
(Fig. 17) (dength/width = 1.7) % wider than F,. Mesosoma: Antescutal depression evident
(Fig. 20). Mesonotum with notauli distinct anteriorly, smooth except for sparse setae
tracing effaced notauli on the disc (Fig. 13). Propodeum (Fig. 14) with narrow central
pentagonal areola. Forewing (Fig. 18): Stigma length about 3.5 width; distal abscissa
of Rl (= metacarpus) short, about /% of stigma length; r + 3RS vein (radial sector)
distinctly sclerotized (insertion point for completely effaced 2RS vein defines separation
between r and 3RS veins). Stigma width 2 length of r-vein; 3RS vein 2—3X length of
r-vein. Cubital cell narrow, distinctly complete or closed (Fig. 19). Legs: Femora (Fig.
22). Metasoma: Metasomal tergum | (Fig. 21) length nearly 2 width at spiracles, disc
coarsely rugose; spiracular tubercles positioned 1/3 from anterior margin; distance
between spiracles and apex greater than width at spiracles. Genitalia (Fig. 23): Ovipositor
sheath (Fig. 24) distinctly ploughshare-shaped. Coloration: Head dark brown; lower part
of gena and mandibles yellow brown, palpi yellow. Antennae brown, except scape, pedicel
and F, yellow. Mesosoma brown, sometimes propleurae, and part of mesopleurae and
propodeum yellow brown. Wings subhyaline, venation light brown. Legs yellow, apices
of tarsi darkened. Metasomal tergum | and central part of tergum 2 yellow to yellow
brown, otherwise metasoma brown, the apex yellow brown to light brown. Ovipositor
sheaths concolorous with the apex of metasoma, upper portion darker. Length of body:
about 2.0—2.3 mm.

Male.—Antenna 16-segmented. Head brown. Antennae brown, articulation between
pedicel and F, lighter. Mandibles yellow brown, palpi yellow. Mesosoma brown. Legs
yellow brown, coxae darkened. Metasoma brown, tergum 1 yellow brown.

Holotype.—?, USA, WA, SKAGIT Co., Campbell Lake, 04-VI-1996, G. Graf & P. Stary
collectors (collection code, 96G165). Host aphid (reared from a mixed collection):
Brachycaudus helichrysi (Kaltenbach), Hyalopterus pruni (Geoffroy), and Phorodon
humuli (Schrank) on Prunus sp. Deposited in USNM.

Paratypes.—31 specimens, collection data same as holotype; deposited in part in
collections of WSU-Prosser (16 2, 6 3) and P. Stary, Ceské Budejovice, Czech Republic
COR 22s):

VOLUME 102, NUMBER 3 Uy

i
insertion point
of effaced 2RS

Figs. 13-24. Monoctonus campbellianus, 2 [illustrations not to equal scale]. 13, Mesonotum. 14, Propo-
deum. 15—17, Flagellomeres 1, 2, and preapical. 18, Fore wing, in part. 19, Cubital cell. 20, Mesosoma—
pronotum with antescutal depression. 21, Metasomal tergum 1. 22, Femora. 23, Genitalia. 24, Ovipositor sheath.
Abbreviations: 2RS, 3RS, and r, wing veins.

718 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Monoctonus pacificus Pike and Stary, new species
(Figs. 25—32)

Diagnosis.—The new species is similar to M. nervosus (Haliday) (see van Achterberg
1989), but differs from the latter in coloration, number of antennal segments, absence of
placodes on flagellar segments 1—4, and arcuate r-vein in the forewing.

Etymology.—The name of the new species is derived from the general type locality,
the Pacific Northwest, USA.

Description.—Female. Head: Eyes with sparse setae in lower half. Tentorio-ocular line
subequal to 4 of intertentorial line; malar space 2 length of intertentorial line or /; of
eye length. Antenna 17—18-segmented, filiform, reaching to half of metasoma.
Flagellomere 1 [F,] (Fig. 27) length 4.0 width, the setae as long to longer than segment
diameter. F, (Fig. 28) subequal to F,. F,_, without placodes; F,, 1 placode; F, 3 placodes.
Preapical F-segments (Fig. 29) about /, wider than F,. Mesosoma: Antescutal depression
evident (Fig. 32). Mesonotum with large bare areas in central and lateral lobes. Propodeum
(Fig. 25) with narrow central areola. FOREWING (Fig. 31): Stigma length 6.0 width; distal
abscissa of RI (metacarpus) short, equal to ¥% of stigma length; r + 3RS vein (radial
sector) distinctly sclerotized; r-vein arcuate, slightly shorter than stigma width; 3RS equal
to r-vein. 2RS vein colorless but distinct. RS + M vein distinct. Metasoma: Metasomal
tergum | (Fig. 26) length 2X width at spiracles, with spiracles positioned % from anterior
margin; disc coarsely rugose. Genitalia (Fig. 30): Ovipositor sheath ploughshare-shaped.
Coloration: Largely yellow. Head, including mandibles brown; palpi yellow; antennae
light brown, except scape, pedicel, F,_, yellow (sometimes F; and F, infuscated). Length
of body: about 2.3 mm.

Male. Antenna 19—20-segmented. Coloration similar to female, metasoma somewhat
darkened.

Holotype.—?, USA, WA, YAKIMA Co., nr Clear Lake, 15-VI-1996, K. S. Pike collector,
(collection code, 96K099). Host aphid: Macrosiphum tuberculaceps (Essig) on Achlys
triphylla. Deposited in USNM.

Paratypes.—5 specimens, USA, WA, YAKIMA Co., N Fork of Tieton River, 27-VI-1996.
Host aphid: Macrosiphum tuberculaceps (Essig) on Achlys triphylla (96G298), deposited
in part in collections of WSU-Prosser (3 2, 1 6) and P. Stary, Ceské Budejovice, Czech
Republic (1 @).

Other material reared from:

Macrosiphum tuberculaceps (Essig): WA, YAKIMA, Morse Cr, 24-VII-97 on Achlys
triphylla (97G255), N Fork of Tieton River, 27-VI-1996 on Achlys triphylla (96G298);
nr Clear Lake, 15-VI-1996, on Achlys triphylla (96K099).

Monoctonus washingtonensis Pike & Stary
Aphis nr oenotherae Oestlund: WA, GARFIELD—Spruce Springs, 22-VIII-95 on Epilobium sp. (95T4627).
Diuraphis noxia (Kurdjumov): WA, BENTON [on Triticum aestivum*].
Rhopalosiphum padi (L.): WA, BENTON [on Triticum aestivum*].

Monoctonus spp.
Acyrthosiphon pisum (Harris): WA, KittiTAs [on Rhododendron albiflorum*].
Illinoia rhododendri (Wilson): WA, KittiTas [on Rhododendron albiflorum* ].
Illinoia sp.: WA, YAKIMA [on Rhododendron albiflorum*].
Macrosiphum euphorbiae (Thomas): WA, SKAMANIA—Twin Falls CG, 4-VIII-95 on Pteridium aquilinum
(95K099).
Macrosiphum walkeri Robinson: OR, BENTON—McDonald State Forest, 21-V-91 on Polypodium sp. (91JO03).
Metopolophium dirhodum (Walker): WA, Kirtiras [on Rhododendron albiflorum*}.

VOLUME 102, NUMBER 3 TS

BY)

Figs. 25-32. Monoctonus pacificus, 2 [illustrations not to equal scale]. 25, Propodeum. 26, Metasomal
tergum 1. 27-29, Flagellomeres 1, 2, and preapical. 30, Genitalia. 31, Fore wing, in part. 32, Mesosoma—
pronotum with antescutal depression. Abbreviations: R1, distal abscissa of postmarginal vein [= metacarpus];
2RS, 3RS, RS+M, and r, wing veins.

720 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Genus PAUESIA Quilis

Pauesia ahtanumensis Pike & Stary
Cinara ponderosae (Williams) [on Pinus ponderosa unless otherwise indicated]: ID, LATAH—Moscow Mtn,
6-VII-96 on Pinus sp. (961195); MT, LincoLN—Libby, 16-VI-95 (95G171); WA, AsotiIn—De Spain
Springs, 5-VII-96 (96T149); Kiickirat—Bickleton, 18-VI-97 (97G112); OKANOGAN—Carlton, 11-IX-97
(97K034); SKAMANIA & YAKIMA [*].
Cinara sp.: WA, AsoTiIN—Copper Canyon, 18-VII-96 on Pinus ponderosa (961308).

Pauesia bicolor (Ashmead)
Cinara sp.: ID, BouNDARY—Bonners Ferry, 29-VII-95 on Pinus contorta (95G444).

Pauesia juniperaphidis (Gahan)
Cinara pilicornis (Hartig): WA, YAKIMA [on Picea pungens*}.

Pauesia pahtonis Pike & Stary
Cinara ponderosae (Williams): WA, SKAMANIA [on Pinus ponderosa*}.

Pauesia ponderosaecola Pike & Stary
Cinara ponderosae (Williams) [on Pinus ponderosa unless otherwise indicated]: MT, LINCOLN—Libby, 16-
VI-95 (95G171); WA, AsoTin—De Spain Spring, 5-VII-96 (96T149, 96T151); Krirtiras—S Fork
Manastash Cr, 25-VI-97 (97G136); KLICKITAT [*]—Bickleton, 18-VI-97 (97G112); WHITMAN—Kamiak
Butte, 7-VII-96 on Pinus sp. (961203); YAKIMA [*]—YIR, White Deer Cr, 11-VII-95 (95K050).

Pauesia pseudotsugae Pike & Stary
Cinara pseudotaxifoliae Palmer: WA, Kittitas [on Pseudotsuga menziesii*].

Pauesia spp.
Cinara brevispinosa (G&P): WA, SPOKANE—Chapman Lk, 1-VI-95 on Pinus ponderosa (95T046B); YAKIMA
[on Pseudotsuga menziesii*|—YIR, Potato Hill Guard Station, 11-VII-95 on Pinus contorta (95K063).
Cinara ponderosae (Williams): WA, SPOKANE—Deer Pk, 23-V-95 on Pinus ponderosa (95KO15).
Cinara pseudotsugae (Wilson): WA, YAKIMA [on Pseudotsugae menziesii*).
Cinara sp.: WA, YAKIMA—N Fork Tieton River, 27-VI-96 on Abies amabilis (96G293).
Undetermined sp.: WA, YAKIMA—Clear Lk, 15-VI-96 on Pinus ponderosa (96K087).

Genus PRAON Haliday

Praon americanum (Ashmead)
Aphis coweni Palmer: WA, Kittrras—Stampede Pass, 8-VIII-96 on Veratrum viride (96K147).
Aphis salicariae Koch: WA, YAKIMA—YIR, nr. Mt. Adams Lk., 19-IX-95 on Epilobium angustifolium
(95G6107).

Praon artemisaphis Smith
Obtusicauda coweni (Hunter): WA, YAKIMA [on Artemisia tridentata*).
Obtusicauda sp.: WA, KLICKITATt & YAKIMA [on Artemisia tridentata *].

Praon artemisicola Pike & Stary
Artemisaphis artemisicola (Williams): WA, BENTON—Six Prong Rd, 24-IV-97 on Artemisia tridentata
(97G0087+).
Epameibaphis atricornis G&P [on Artemisia tridentata]: WA, KLIckitaT [*+], & YAKIMA [esta
Flabellomicrosiphum sp. [on Artemisia tridentata]: WA, KLICKITAT [**+], & YAKIMA [ale
Pleotrichophorus quadritrichus (Knowlton & Smith): WA, YAKIMA—Mabton, 24-IV-97 on Artemisia
tridentata (97G0077).

Pleotrichophorus sp.: WA, KLickirat—Alderdale Rd, 9-V-95 on Artemisia tridentata (95GO287¢).
Pseudoepameibaphis tridentata (Wilson) [on Artemisia tridentata): WA, BENTON [*], KLICKITAT [*+]J—
Alderdale Rd, 9-V-95 (95G0287); YAKIMA [*].

Praon coniforme Pike and Stary, new species
(Figs. 33-41)

Diagnosis.—The new species belongs to a species group characterized by the absence
of an m-cu vein in the forewing. Other distinguishing characters are antenna 16-17
segmented (in female), flagellomere 1 (F,) yellow, and ovipositor sheath coniform in

VOLUME 102, NUMBER 3 721

Ni &

SSN Nerval
\

to NaS

7

/
7
-

36

37

Figs. 33-41. Praon coniforme, @ [illustrations not to equal scale]. 33, Mesonotum. 34, Propodeum. 35,
Metasomal tergum 1. 36—38, Flagellomeres 1, 2, and preapical. 39, Fore wing, in part. 40, Genitalia. 41, Ovi-
positor sheath. Abbreviations: R1, distal abscissa of postmarginal vein [= metacarpus]; r+2RS, wing vein.

shape. The new species is similar to Praon yakimanum (see Pike & Stary 1995), but
differs from the latter in antennal segment number (P. yakimanum, 15-segmented [rarely
16]), shape of the ovipositor sheath, and aphid host. At present, the new species is only
known from snowberry aphid, Aphthargelia symphoricarpi (Thomas).

Etymology.—The name of the new species is derived from the shape of the ovipositor
sheath that resembles a cone.

Description.—Female. Head: Eyes medium sized, with sparse setae in lower half.
Tentorio-ocular line subequal to 1/4 of intertentorial line; malar space 1.5x< length of
tentorio-ocular line or slightly longer than ¥, of eye length. Antenna 16—17 segmented,
filiform, slightly thickened in apical third, reaching to half of metasoma. Flagellomere 1
[F,] (Fig. 36) slightly more than 4.0 width; 1.6 as long as F,; setae slightly longer
than segment diameter; without placodes. F, (Fig. 37) width equal to F,, with 1 placode.
Preapical F-segments (Fig. 38) 1.3 width of F,. Mesosoma: Mesonotum (Fig. 33) with
bare areas on the lateral lobes. Propodeum (Fig. 34) pubescent. FOREWING (Fig. 39):
Stigma length 3.5 width; distal abscissa of R1 (metacarpus) longer than half stigma
length; r + 3RS vein (radial sector) slightly longer than distal abscissa of R1; m-cu vein
absent. Metasoma: Metasomal tergum 1 (Fig. 35) somewhat longer than width at spiracles
(7:6); distance between spiracles and anterior margin %4 shorter than width at spiracles.
Genitalia (Fig. 40): Ovipositor sheath (Fig. 41) cone-like in shape. Coloration: Head
dark brown, mouthparts bright yellow. Antenna brown; apex of pedicel and F, yellow.
Mesosoma dark brown. Wings subhyaline, venation light brown. Legs yellow, apices of

22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tarsi infuscate. Metasomal tergum | yellow brown, darker in basal portion. Metasoma
brown. Ovipositor sheaths dark brown. Length of body: about 2.4 mm.

Male.—Antenna 19—20-segmented, filiform, longer than the body. Antenna brown, F,
with a narrow yellow base. Legs light brown.

Holotype.-—?, USA, WA, YAKIMA Co., Maloy Rd and Wenas Creek, 30-V-1996, G.
Graf collector, (collection code, 96G127). Host aphid: Aphthargelia symphoricarpi
(Thomas) on Symphoricarpos sp. Deposited in USNM.

Paratypes.—3 specimens, collection data same as holotype, deposited in part in
collections of WSU-Prosser (2 6) and P. Stary, Ceské Budejovice, Czech Republic (1 2).

Other Material reared from:

Aphthargelia symphoricarpi (Thomas) [on Symphoricarpos sp. unless otherwise
indicated]: WA, KittirAs—Lost Lake, 29-VI-1994 (94G005) & 6-VIII-1997 (97G334);
KLICKITAT—Trout Lake, 28-VI-1997 (97G159); SPOKANE—Cooper Rd, 29-VI-95
(95T178); Fish Lake, 14-VI-1997 on S. occidentalis (97KO04); YAKIMA—Maloy Rd &
Wenas Cr., 11-VI-1997 (97G066).

Praon exsoletum (Nees)

Therioaphis riehmi (Borner) [on Melilotus sp. unless otherwise indicated]: WA, BENTON—-WSU-Prosser, | 1-
VII-97 on M. officinalis (97F218+); YAKIMA—Grandview, 22-VI-96. (96G261); nr Parker, 29-I[X-95
(95G665+); YIR, Signal Peak, 19-IX-95 (95G6257).

Therioaphis trifolii (Monell) [on Medicago sativa): WA, BENTON [*]; YAKIMA, nr Union Gap, 23-X-95
(95G7217).

Therioaphis sp.: WA, BENTON—-WSU-Prosser, 28-VI-95 on Medicago sativa (95G2777).

Praon fulvum Pike and Stary, new species
(Figs. 42—50)

Diagnosis.—Yellow coloration of F, (flagellomere 1) and F, keys the new species to
P. gallicum Stary (see key by Johnson 1987), but it differs from the latter in the number
of antennal segments [P. fulvum with 18—19 (rarely 20) vs. P. gallicum with 15—16 (rarely
17)], the presence of the m-cu vein in the forewing (absent with P. gallicum), and the
mesonotum pubescence on the lateral lobes of the mesonotum (large bare areas with P.
gallicum).

Etymology.—The name of the new species is derived from its prevailing body
coloration.

Description.—Female. Head: Eyes medium sized, with sparse setae in lower half.
Tentorio-ocular line % of intertentorial line or 4% of malar space. Antenna 18-19 (rarely
20) segmented, filiform, slightly thickened towards apex, subequal to body length.
Flagellomere 1 [F,] (Fig. 45) long, length nearly 5X width, setae slightly longer than
segment diameter, without placodes. F, (Fig. 46) % shorter than F,, without placodes.
Preapical F-segments (Fig. 47) 1/8 wider than F,. Mesosoma: Mesonotum (Fig. 42) with
very small to no areas on the lateral lobes without setae. Propodeum (Fig. 49) pubescent,
somewhat bare in center. FOREWING (Fig. 48): Stigma length 3.5 width; distal abscissa
of R1 (metacarpus) slightly longer than half stigma length; RS + M vein distinct,
subcolored in basal 4—%, remaining portion colorless; m-cu vein complete, subcolored.
Metasoma: Metasomal tergum 1 (Fig. 50) longitudinal, length %4 longer than width at
spiracles; width at spiracles Y; longer than the distance from spiracles to anterior margin.
Genitalia (Fig. 43): Ovipositor sheath (Fig. 44) long, subarcuate. Coloration: Body
almost completely yellow. Head yellow, frons more or less brown. Apices of mandibles
brown. Antenna brown; scape, pedicel, F, and F, yellow. F, sometimes infuscated at apex.

VOLUME 102, NUMBER 3 123

Figs. 42-50. Praon fulvum, @ [illustrations not to equal scale]. 42, Mesonotum. 43, Genitalia. 44, Ovipositor
sheath. 45—47, Flagellomeres 1, 2, and preapical. 48, Fore wing, in part. 49, Propodeum. 50, Metasomal tergum
1. Abbreviations: R1, distal abscissa of postmarginal vein [= metacarpus]; RS+M and M-cu, wing vein.

Mesosoma yellow. Wings subhyaline, venation light brown. Legs yellow. Metasoma
yellow. Ovipositor sheaths light brown. Length of body: about 2.0—2.6 mm.
Male.—Unknown.
Holotype.— ?, USA, WA, Lewis Co., Mt. Rainier Nat. Park, Stevens Canyon, 23-VII-

724 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1996, G. Graf collector, (collection code, 96G357). Host aphid: Macrosiphum euphorbiae
(Thomas) on Crepis sp. Deposited in USNM.

Paratypes.—6 specimens, collection data same as holotype, deposited in part in
collections of WSU-Prosser (3 ¢) and P. Stary, Ceské Budejovice, Czech Republic (3 2).

Other material reared from:

Macrosiphum euphorbiae (Thomas): WA, KLICckIrAt—Cleveland, 7-VII-1997, on
Lupinus sp. (97G230).

Macrosiphum tuberculaceps (Essig): WA, YAKIMA—Wenas Creek & Maloy Rd, 11-VI-
1997 on Achlys triphylla, (96G298).

Macrosiphum sp.: WA, YAKIMA—Wenas Creek & Maloy Rd, 11-VI-1997 on Spiraea
pyramidata (97G073).

Praon gallicum Stary

Diuraphis noxia (Kurdjumoy) [on Triticum aestivum]: WA, BENTON—WSU-Prosser, 22-VII-97 (97F295);
KLickIrAt—Crider Valley Rd, 22-VII-97 (97G250++); YAkIMA—nr Sunnyside, 17-VII-97 (97G2467).

Metopolophium dirhodum (Walker) [on Triticum aestivum]: WA, YAKIMA—Glade Rd, 7-VII-97 (97G2177);
nr Sunnyside, 17-VII-97 (97G2467).

Rhopalosiphum padi (L.): WA, GRANT—Ephrata, 8-VII-97 on Zea mays (97K058); KLICKITAT—Crider Valley
Rd, 22-VII-97 on Triticum aestivum (97G2507¢).

Sitobion avenae (FE) [on Triticum aestivum]: WA, KLIcKiIrat—Crider Valley Rd, 22-VII-97 (97G2507+);
Y AKIMA—Glade Rd, 7-VII-97 (97G2177).

Praon humulaphidis Ashmead
Acyrthosiphon macrosiphum (Wilson): MT, LINCOLN—Libby, 6-VII-96 on Amelanchier alnifolia (96G322).
Macrosiphum euphorbiae (Thomas): WA, WHITMAN—Pullman, 18-VI-96 on cultivated Rosa sp. (961037) &
5-VIH-96 on unknown plant (96T176).
Macrosiphum sp. 1D, BENEWAH—McCroskey SP, 7-VI-96 on native Rosa sp. (96TO005).
Nasonovia ribisnigri (Mosley): WA, WHITMAN—Pullman, 5-VIU-96 on unknown plant (96T177).
Illinoia sp.: 1D, SHOSHONE—Thomas Pass, 29-VII-95 on Vaccinium sp. (95G472).
Illinoia sp.?: WA, CHELAN—Chatter Cr CG, 1-VIII-95 on Aruncus sylvester (95G487).

Praon occidentale Baker

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola}: ID, BENEWAH—McCroskey SP, 31-VIII-95
(95T514+); WA, BENTON—WSU-Prosser, 11,22-V-95 (95G034, 95G081); Prosser, 22-VI-97 (97G121);
WHITMAN—Chambers, 19-IX-95 (95T581); Nisqually John Landing, 19-IX-95 (95T584); Pullman, 2-VIII-
96 (961386); YAKIMA—Grandview, 23-VI-96 (96G263); Wenas Lk, 11-VI-97 (97G055).

Acyrthosiphon pisum (Harris): WA, BENTON—28-VI-95 on Medicago sativa (95G277+); Prosser, 23-VII-96
on Trifolium repens (96G393); GRANT—WSU-Othello, 26-VI-95 on Medicago sativa (95B008);
SKAMANIA—Underwood, 11-VII-95 on Lathyrus sp. (95K072).

Amphorophora geranii G&P: WA, WHITMAN—Long Rd nr Hole in the Ground, 18-V-95 on Geranium sp.
(95T0127).

Aphis craccivora Koch: ID, LAraH—Moscow Mtn, 3-VIII-96 on Trifolium sp. (96T4007+).

Aphis helianthi Monell: WA, GARFIELD—Stentz Springs, 22-VIII-95 on Osmorhiza chilensis (95T452).

Aphis lugentis Williams: WA, YAKIMA—Bird Cr Meadows, 8-[X-94 on Arnica sp. (94R172).

Brachycaudus helichrysi (Kaltenbach): WA, GARFIELD—FR160 & FR40, 21-I[X-95 on Anaphalis margaritacea
(95T5957+£); PIERCE—RNP, Tipsoo Lk, 5-IX-97 on Aster sp. (97G364++£); YAKIMA—Mabton-Bickleton Rd,
Glade Cr, 7-VII-97 on Artemisia ludoviciana (97G219).

Capitophorus elaeagni (del Guercio): WA, YAKIMA—Grandview, 28-VI-96 on Cirsium vulgare (96G3097+).

Chaetosiphon thomasi HRL: WA, PIERCE—WSU-Puyallup, 10-VI-96 on cultivated Fragaria sp. (96T022).

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, BENTON [*]; KLICKIrAT—Bickleton, 29-VII-97
(97G300).

Dysaphis plantaginea (Passerini): WA, YAKIMA—Grandview, 10-VI-96 on Pyrus malus (96G214).

Ericaphis wakibae (Hottes): WA, YAKIMA—N Fork Tieton River, 15-VI-96 on native Rosa sp. (96KO0907+).

Hyperomyzus nigricornis (Knowlton) [on Hieracium sp. unless otherwise indicated]: ID, BENEWAH—
McCroskey SP, 31-VII-95 (95T514+); LATAH—Moscow Mtn, 15-VIII-96 on Compositae (961446); MT,
SANDERS—nr Thompson Pass, 29-VII-95 (95G462+); WA, GARFIELD—Alder Thicket, 21-[X-95 (95T587);
Kitriras—Buck Meadows, 16-VIII-96 (96G493); Manastash Rd, 25-VI-96 (96G415); KLICKITAT—

VOLUME 102, NUMBER 3 725

Cleveland, 18-VI-96 on Madia minima (96K109); SPoKANE—Mt Spokane Hwy (mi marker 18.5), 22-VIII-
96 on Compositae (96T487, 96T488); YAKIMA—Bird Cr Meadows, 26-VI-95 (95A0167).

Hyperomyzus sp.: WA, Kirriras—S Fork Manastash Cr, 24-VII-95 on Hieracium albiflorum (95G388):
SPOKANE—nr Peone, 24-VIII-95 on Centaurea pratensis (95T4737+); WHITMAN—Pullman, 10-VIII-95 on
Lapsana communis (9574364).

Illinoia davidsoni (Mason): MT. SANDERS—nr Thompson Pass, 29-VII-95 on Arnica sp. (95G463); WA,
KitTTITAs—Quartz Mtn, 25-VII-96 on Arnica cordifolia (96G406++).

Illinoia richardsi: WA, GARFIELD—FR160 & FR40, 21-IX-95 on Anaphalis margaritacea (95T595+%).

Illinoia nr thalictri (MacGillivray): WA, GARFIELD—Stentz Springs (FR 40), 22-VIII-95 on Thalictrum sp.
(95T454).

Ilinoia spp.: ID, BENEWAH—McCroskey SP 7-VI-96 on unknown plant (96T006); WA, GarFIELD—Stentz
Springs, 1-VIIH-95 on Aquilegia formosa (95T362+): KITTITAS [on Aquilegia formosat*]; PlERCE—RNP.
Tipsoo Lk, 11-IX-96 on Carex sp. (96G5427 +).

Macrosiphoniella ludovicianae (Oestlund): WA, Asotin—Asotin Cr WLA, 26-VI-96 on Artemisia ludoviciana
(96T048).

Macrosiphoniella nr ludovicianae (Oestlund):; WA, YAKIMA [on Artemisia ludoviciana*].

Macrosiphum albifrons Essig: WA, PIERCE—RNP. Tipsoo Lk, 11-[X-96 on Lupinus sp. (96G547).

Macrosiphum euphorbiae (Thomas): ID, LATAH—Moscow Mtn, 6-VII-96 on Castilleja sp. (96T188); 6-VII-
96, on native Rosa sp. (96T184); 6-VII-96 on Compositae (96T185, 96T192): MT, LINCOLN—Libby, 15-
VI-95 on native Rosa sp. (95G168); SANDERS—nr Thompson Pass, 29-VII-95 on Hieracium sp. (95G4627);
WA, AsoTIN—De Spain Springs, 1-VIII-95 on Senecio sp. (951381), 26-VI-96 on unknown plant (96T0S51),
5-VII-96 on Compositae (96T159), 11-VII-96 on Potentilla sp. (96T250) & Compositae (96T251);
BENTON—WSU-Prosser, 27-X-95 on Chenopodium album (95K2187}), & 27-X-95 on cultivated Rosa sp.
(95K220+); Kirtrras—Buck Meadows, 9-VII-96 on Agoseris elata (96G341), 9-VII-96 on native Rosa Sp.
(96G335); Manastash Cr, FR125, 16-VIII-96 on Silene sp. (96G497): Manastash Ridge, 2-IX-95 on
Gnaphalium microcephalum (95G537): Lost Lk Trail, 6-VIH-97 on Luina nardosmia (97G327); Quartz
Mtn, 25-VII-96 on Arnica cordifolia (96G406+); PEND OREILLE, Tiger Meadows, 9-VII-96 on Compositae
(96T221); SPOKANE—Mt Spokane SP, 24-VIII-95 on native Rosa sp. (95T491+); STEVENS—Little Pend
Oreille WLA, 9-VII-96 on Compositae (96T210); WHITMAN—Pullman, 10-VIII-95 on Lapsana communis
(95T436+) & 18-VI-96 on cultivated Rosa sp.(96T037); WA, YaKIMA—Bird Cr Meadows, 26-VI-95 on
Hieracium sp. (95A016+); Grandview, 28-VI-96 on Cirsium vulgare (96G309+); Hwy 410 & Sawmill Flat,
28-VII-97 on native Rosa sp. (97G2897).

Macrosiphum osmaroniae (Wilson): WA, WHATCOM—Bellingham, 3-V-96 on Oemleria cerasiformis
(96K013);

Macrosiphum rosae (L.) [on cultivated Rosa sp.]: WA, BENTON—-WSU-Prosser, 27-X-95 (95K2207); nr
Prosser, Richards Rd, 22-VI-97 (97G122); YAKIMA [+*]—Grandview, 13 & 21-V-97 (97G021++4, 97G037).

Macrosiphum sp.: ID, BENEWAH—McCroskey SP. 7-VI-96 on native Rosa sp. (96T005); WA, AsotTin—nr
Anatone, 24-VIII-93 on Triticum aestivum (93T015); YAKIMA—nr Chinook Pass, Morse Cr, 24-VII-97 on
Oplopanax horridum (97G256); Sunnyside, 24-VIII-95 on Zea mays (95G5197).

Macrosiphum sp.?: WA, Kittiras—Manastash Ridge, 2 mi E of Lost Lk, 2-IX-95 on Silene sp. (95G545).

Metopolophium dirhodum (Walker): WA, YAKIMA—N Fork Tieton River, 15-VI-96 on native Rosa sp.
(96K090+F4).

Metopolophium sp.?: WA, YAKIMA—YIR, Windy Point, 19-IX-95 on Poa sp. (95G634).

Microlophium nr sibiricum: WA, YAKIMA—Sunnyside, 28-VI-95 on Urtica dioica (95G255).

Myzus persicae (Sulzer): WA, BENTON—WSU-Prosser, 27-X-95 & 1-X-97 on Chenopodium album (95K218}+,
97G476).

Nasonovia aquilegiae (Essig): WA, GARFIELD—Stentz Springs, 1-VIII-95 on Aquilegia formosa (95T3627).

Nasonovia polemonii (G&P): WA, KitrirAs—Quartz Mtn, 25-VII-96 on Polemonium occidentale (96G409).

Nasonovia (Kakimia) sp.: WA, WHITMAN—Pullman, 10-VIII-95 on Lapsana communis (95743674).

Nearctaphis bakeri (Cowen): ID, LATAH—-Moscow Mtn, 3-VIII-96 on Trifolium sp. (96T4007 +).

Nearctaphis kachena (Hottes): WA, SPOKANE—nr Mt Spokane SP, 24-VIII-95 on Castilleja (95T485).

Phorodon humuli (Schrank): WA, BENTON—WSU-Prosser, 9-IX-95 on Humulus lupulus (95K154).

Rhopalosiphum insertum (Walker): WA, DouGLas—Orondo, 28-V-96 on Pyrus malus (96RO004).

Rhopalosiphum padi (L.): WA, BENTON [on Triticum aestivum*]; GRANT—Ephrata 8-VII-97 on Zea mays
(97K058); YAKIMA—Sunnyside, 24-VIII-95 on Zea mays (95G5197).

Sitobion avenae (FE): WA, BENTON—WSU-Prosser, 3 & 11-VII-97 on Triticum aestivum (97F192, 97F211);
Kittiras—Buck Meadows, 9-VII-96 on Dactylis glomerata (96G346); PiERCE—RNP, Tipsoo Lk, 11-IX-
96 on Carex sp. (96G5427+#).

726 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Uroleucon erigeronense (Thomas): WA, Kittiras—Lost Lk, 6-VIII-97 on Aster sp. (97G346); LEwis—
PIERCE—RNP, Tipsoo Lk, 8-IX-95 & 5-IX-97 on Aster sp. (95K106, 97G3647).

Uroleucon sonchi (L.): WA, FRANKLIN—Mesa, 8-IX-97 on Lactuca serriola (97KO06).

Uroleucon zymozionense (Knowlton): WA, CHELAN—Swauk Pass, 1-VII-95, on Aster sp. (95G491).

Uroleucon sp.: ID, BENEWAH—McCroskey SP, 31-VIII-95 on Compositae (95T528); WA, PEND OREILLE—
Tiger Meadows, 9-VII-96 on Achillea millefolium (961222); PIERCE—RNP, Tipsoo Lk, 5-IX-97 on
Anaphalis margaritacea (97G367); SKAMANIA [on Aster sp.*]; SPOKANE—nr Peone, 24-VIII-95 on
Centaurea pratensis (95T4737*).

Wahlgreniella nervata (Gillette): WA, BENToN—Grandview, 13-V-97 on cultivated Rosa sp. (97G0217++);
YAKIMA—Hwy 410 & Sawmill Flat, 28-VII-97 on native Rosa sp. (97G2897$).

Undetermined spp.: WA, BENTON—Prosser, 23-VII-96 on Sonchus oleraceus (96G389); KittiIras—S Fork
Manastash Cr, 24-VII-95 on Potentilla sp. (95G395); WHITMAN—Pullman, 5-VII-96, on Geranium
viscosissimum (96T171).

Praon nr occidentale Baker
Illinoia sp.: WA, YAKIMA—Blowout Mtn, end of Little Naches River Rd, 9-IX-97 on Rhododendron albiflorum
(97G400).
Uroleucon erigeronense (Thomas): WA, Kittiras—Lost Lk, 6-VIII-97 on Haplopappus hirtus (97G3407).
Uroleucon nr ivae Robinson: WA, YAKIMA—Harrah, 29-IX-97 on Iva xanthifolia (97G465).
Uroleucon suzannae Robinson: WA, Kirtrras—Lost Lk, 6-VIII-97 on Haplopappus hirtus (97G3407 +).

Praon pequodorum Viereck

Acyrthosiphon macrosiphum (Wilson): WA, AsoTin—nr Anatone, 18-VII-95, on Amelanchier alnifolia
(95T266).

Acyrthosiphon pisum (Harris) [on Medicago sativa unless otherwise indicated]: WA, ADAMS—WSU-Othello,
26-VI-95 (95B008); BENTON [+*]—-WSU-Prosser, 27-VI-96 on Vicia angustifolia (Q96A009), 30-VI-97, &
11-VII-97 (97F186, 97F202); GRANT—WSU-Royal City, 26-VI-95 (95B003, 95B006); Mattawa, 26-VI-
95 (95B005); YAKIMA—Wenas Valley, 5 mi N of Selah, 30-V-96 on Melilotus sp. (96G109).

Amphorophora rubi (Kaltenbach): WA, KLickirat—Trout Lk, 29-VI-97 on Rubus sp. (97G168).

Illinoia sp.: WA, SPOKANE—Mt Spokane SP, 20-VII-95 on Vaccinium sp. (95T297).

Therioaphis riehmi (Borner): WA, BENTON—-WSU-Prosser, 18 & 25-VII-97, on Melilotus officinalis
(97F2547 4).

Uroleucon ivae Robinson [on Iva xanthifolia]): WA, YAKIMA [*]—Grandview, 17,22, 30-VH-95 (95G313,
95G349, 95G366).

Uroleucon sonchi (L.): WA, YAKIMA—Grandview, 17-VII-95 on Sonchus oleraceus (95G348).

Praon simulans (Provancher)
Macrosiphum rhamni (Clarke) [det. by A. Jensen]: WA, Kinc—Stampede Pass, 8-VIII-96 on Pteridium
aquilinum (96G441).

Praon unicum Smith

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola}: WA, AsoTIN—Asotin Cr WLA, 7-IX-95 (95T547);
BENTON—Prosser, 22-V-95 (95G079); CHELAN—Chelan SP, 11-[X-97 (97K038); KitTiTas [*].

Aphis coweni Palmer [on Veratrum viride]: WA, SKAMANIA [*]; YAKIMA [*]—Pleasant Valley CG, 24-VII-97
(97G266).

Aphis craccivora Koch: WA, WHITMAN—Pullman, 7-VII-96, 2-VIII-96 on Robinia pseudo-acacia (961199,
96T393).

Aphis nr fabae Scopoli: WA, YAKIMA—YIR, Windy Point, 19-I[X-95 on Cimicifuga laciniata (95G630).

Aphis helianthi Monell: WA, CHELAN—Stevens Pass, 16-[X-95 on Epilobium angustifolium (95K1347¢);
Kittiras—Lost Lk, 24-VII-95 on E. angustifolium (95G399+4); S. Fork Manastash Cr, 24-VII-95 on
Heracleum lanatum (95G378); WHITMAN—Pullman, 1-VIII-95 on Coriandrum sativum (9513867);
Y AKIMA—Ame rican River, 5-[X-97 on Heracleum lanatum (97G382); nr Wenas Lk, 17-VI-97 on Lomatium
sp. (97G095+); YIR, Piscoe Cr, 25-VII-95 on Ligusticum grayi (95G432) & Osmorhiza occidentalis
(95G428).

Aphis nigratibialis Robinson: WA, Kirtrras—S. Fork Manastash Cr, mi marker 12, 25-VI-97 on Cornus
stolonifera (97G129).

Aphis oenotherae (Oestlund): ID, BENEWAH—McCroskey SP, 31-VII-95 on Epilobium angustifolium
(95T533).

Aphis nr oenotherae (Oestlund) [on Epilobium sp. unless otherwise indicated]: WA, BENTON—Kennewick,
28-IX-96 (96G597); GARFIELD—FR 160 & FR 40, 21-IX-95 (95T594); Spruce Springs, 22-VIII-95
(9514627); Stentz Springs, 14-[X-95 (95T565); YAKIMA—American River, 5-I[X-97 on Epilobium minutum

VOLUME 102, NUMBER 3 727

(97G375+); Pleasant Valley CG, 24-VII-97 on Epilobium minutum (97G267+); Wenas Cr, 13-VIII-96
(96G467).

Aphis pomi de Geer: WA, DouGLAs—Bridgeport, 17-VI-96 on Pyrus malus (96X007+#).

Aphis rumicis L.: WA, BENTON, on [Rumex crispus*].

Aphis salicariae Koch [on Epilobium angustifolium]: ID, KOOTENAI—Rathdrum, 29-VII-95 (95G443+); WA,
CHELAN—1-VIII-95 (95G4817).

Aphis nr salicariae Koch [on Epilobium angustifolium]: WA, CHELAN—Swauk Pass, 1-VIII-95 (95G4927+);
YAKIMA—YIR, Old Mead Canyon, 25-VII-95 (95G442).

Aphis varians Patch [on Epilobium angustifolium]: ID, KOOTENAI—Rathdrum, 29-VII-95 (95G4437+); WA,
CHELAN—Stevens Pass, 16-IX-95 (95K134+#); KittrraAs—Lost Lk, 24-VII-95 (95G399+¢#).

Aphis nr varians Patch: WA, CHELAN—1-VIII-95 on Epilobium angustifolium (95G48 17).

Aphis sp.: WA, CHELAN—Chelan SP, 11-IX-97 on Salix sp. (97K042+); GARFIELD—Bear Cr Rd, 14-IX-95 on
Lilium columbianum (951577); Kirtiras—Taneum Lk, 11-[X-97 on Pedicularis racemosa (97G4117#);
WHITMAN—Pullman, 12-VIII-96 on Helianthus annuus (96T438); YAKIMA—YIR, Windy Point, 19-[X-95
on Cimicifuga laciniata (95G631).

Brachycaudus helichrysi (Kaltenbach): WA, YAKIMA—Wenas, 11-VI-97 on Prunus domestica (97G079).

Brachycaudus tragopogonis (Kaltenbach): WA, WHITMAN—Pullman, 12-IX-95 on Tragopogon dubius
(95T561).

Cavariella aegopodii (Scopoli): WA, YAkIMA—nr Wenas Lk, 17-VI-97 on Lomatium sp. (97GO95+) &
Umbelliferae (97G097).

Cavariella pastinacae (L.): WA, YAKIMA—N. Fork Tieton River, 27-VI-96 on Heracleum lanatum (96G288).

Diuraphis noxia (Kurdjumov) [on Triticum aestivum]: WA, AsoTiIN—Asotin Cr Rd, 20-VI-95 (95T161);
BENTON [*]; KLIckIrat—Bickleton, 6-VIII-97 (97G354); SPOKANE—Mt Spokane SP, 22-VII-95 (95T495).

Dysaphis plantaginea (Passerini) [on Pyrus malus]: WA, CHELAN—-WSU-Wenatchee, 30-V-96, 13-VI-96
(96RO009, 96R028); YAKIMA—Grandview, 26,28-VI-96 (96G282, 96G308).

Ericaphis gentneri (Mason): WA, BENTON—WSU-Prosser, 24-VI-97 on Crataegus sp. (97G124).

Hyperomyzus lactucae (L.): WA, WHITMAN—Pullman, 12-IX-95 on Lactuca serriola (95T559).

Hyperomyzus nigricornis (Knowlton): WA, GARFIELD—Alder Thicket, FR 40, 22-VIII-95 on Senecio sp.
(95T447); KirrirAs—Ellensburg, 13-VI-97 on Ribes sp. (97G084).

Macrosiphum euphorbiae (Thomas): ID, KOOTENAI—Rathdrum, 29-VII-95 on Epilobium angustifolium
(95G4437+); WA, KittirAs—Lost Lk, 6-VIII-97 on Castilleja (97G342+); YAKIMA [on Solanum
lycopersicon* }.

Macrosiphum gaurae (Williams): WA, YAKIMA—Wenas Lk, 11-VI-97 on Madia sp. (97G059).

Macrosiphum pallidum (Oestlund): WA, YAKIMA—American River, 5-IX-97 on Epilobium minutum
(97G3757).

Macrosiphum valerianae (Clarke): WA, Kittiras—Lost Lk, 24-VII-95 on Epilobium angustifolium
(95G399F +).

Macrosiphum sp. [nr fuscicornis MacDougall & pallidum (Oestlund)]: WA, CHELAN—Swauk Pass, 1-VIII-95
on Epilobium angustifolium (95G4927).

Metopolophium dirhodum (Walker): WA, YAKIMA—Glade Rd, 7-VII-97 on Triticum aestivum (97G2177).

Myzodium knowltoni Smith & Robinson: WA, KLickirat—John Day Dam, 28-IV-95 on Veronica anagallis-
aquatica (95GO0097F +).

Myzus cerasi (FE): WA, CHELAN—Wenatchee Heights, 6-VI-96 on Prunus sp. (96RO19).

Myzus persicae (Sulzer): WA, DoUGLAs—Orondo, 25-V-95 on Prunus armeniaca (95RO11); KLickIrat—John
Day Dam, 28-IV-95 on Veronica anagallis-aquatica (95G009+); WHITMAN—Pullman, 13-VI-96 on
unknown plant (96T0317).

Myzus sp.: WA, WHITMAN [on Forsythia sp.*].

Nasonovia ribisnigri (Mosley): WA, PIERCE—RNP, Tipsoo Lk, 5-[X-97 on Castilleja sp. (97G3617).

Nasonovia (Kakimia) sp.: WA, GARFIELD—Spruce Spring, 22-VII-95 on Castilleja sp. (95T4637+);
KitriraAs—Lost Lk, 6-VIII-97 on Castilleja sp. (97G342+); WHITMAN—Kamiak Butte, 12-IX-95 on
Penstemon sp. (95T551).

Nearctaphis bakeri (Cowen): WA, AsoTIN—De Spain Springs, 1-VIII-95 on Penstemon sp. (95T383).

Nearctaphis kachena (Hottes) [on Castilleja sp. unless otherwise indicated]: ID, BENEWAH—McCroskey SP,
31-VIII-95 (951522); WA, GaRFIELD—Spruce Spring, 22-VII-95 (95T4637£); Teal Springs, 22-VIII-96
(96T484); Kirtiras—Lost Lk, 6-VIII-97 (97G3427+); Taneum Lk, 11-IX-97 on Pedicularis racemosa
(97G411+£); SPOKANE—Mt Spokane SP, 24-VIII-95 (95T479, 95T485); YAKIMA—YIR, Mt Adams Lk Rd,
23-VI-95 (95G242).

Phorodon humuli (Schrank) [on Prunus salacina unless otherwise indicated]: WA, BENTON—-WSU-Prosser,

728 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

19,25-IV-95, 1-V-95 (95G006, 95G007, 95GO016), 9-IX-95 on Humulus lupulus (95K154); YAKIMA—
Sunnyside, 24-V-96 on H. lupulus (96G079+); Yakima Arboretum, 30-V-96 (96G104).

Rhopalosiphum insertum (Walker): WA, CHELAN—-WSU-Wenatchee, 30-V-96 on Pyrus communis (96RO15),
17-V-95 on Pyrus malus (95RO07); DouGLAs—Orondo, 25-V-95 on Pyrus communis (95RO10), 11 & 28-
V-96 on Pyrus malus (96RO03, 96RO004, 96X005); Bridgeport, 12 & 17-VI-96 on Pyrus malus (96X006,
96X0077+); KittirAas—Ellensburg, 17-V-96 on Crataegus columbiana (96G066).

Rhopalosiphum maidis (Fitch): WA, BENTON [on Panicum milliaceum*]; YAKIMA—Sunnyside, 2-X-95 on
Hordeum vulgare (95G6717).

Rhopalosiphum padi (L.): WA, BENTON [on Triticum aestivum & Hordeum vulgare* |—WSU-Prosser, 24-IV-
92, on T. aestivum (92A010); GRANT—Ephrata, 8-VII-97, on Zea mays (97K058); YAKIMA—Sunnyside,
2-X-95 on H. vulgare (95G6717).

Sitobion avenae (FE) [on Triticum aestivum unless otherwise indicated]: WA, AsoTIN—De Spain Springs, 24-
VII-96 on Aquilegia formosa (961339); BENTON—West Richland, 28-VI-95 (95G279); WSU-Prosser, 3,
11 & 18-VII-97 (97F191, 97F192, 97F215, 97F217, 97F227, 97F243, 97F249, 97F253, 97F256);
YAKIMA—Glade Rd, 7-VII-97 (97G2177).

Uroleucon sp.: ID, BENEWAH—McCroskey SP, 31-VHI-95 on unknown plant (95T520); KOOTENAI—Rathdrum,
29-VII-95 on Epilobium angustifolium (95G4437).

Undetermined spp.: WA, WHITMAN—Pullman, 19-VI-96 on Pyrus malus (96T042).

Praon nr unicum Smith
Euceraphis gillettei Davidson: WA, Kitriras—Manastash Rd, 25-VI-97 on Alnus sp. (97G1347#).
Oestlundiella flava (Davidson): WA, KittrrAs—Manastash Rd, 25-VI-97 on Alnus sp. (97G1347#).

Praon yakimanum Pike & Stary

Cavariella pastinacae (L.): WA, ASoTIN—Asotin Cr, 11-VII-95 on Oenothera (95T196).

Diuraphis noxia (Kurdjumoyv) [on Triticum aestivum]: WA, Asotin—Pataha Cr Rd, 10-VIII-95 (95T435);
BENTON [*]|—WSU-Prosser, 23-V-97 (97F100, 97F112); KLICKIrAat—Bickleton, 9-VII-95 (95G3197).

Metopolophium dirhodum (Walker): WA, YAKIMA—Sunnyside, 28-VI-95 on Phalaris arundinacea (95G254).

Rhopalosiphum padi (L.): WA, BENTON [on Triticum aestivum & Hordeum vulgare*|; GRANT—Ephrata, 8-
VII-97 on Zea mays (97K058).

Sitobion avenae (F.): WA, KLICKIrATt—Bickleton, 9-VH-95 on Triticum aestivum (96G3197).

Undetermined sp.: WA, SPOKANE—Turnbull National WLA, 13-VI-95 on Amelanchier alnifolia (95T414).

Praon spp.

Acyrthosiphon lactucae (Passerini) [on Lactuca serriola]: WA, Kitt1rAs—S Fork Manastash Cr, 24-VII-95
(95G376); Ellensburg, 22-VI-96 (96G266); YAKIMA—Grandview, 20-X-95 (95G728).

Acyrthosiphon pisum (Harris): MT, SANDER [on Lathyrus odoratus*]; WA, YAKIMA—YIR, Delaney Springs,
25-VII-95 on Vicia sp. (95G408).

Amphorophora rubi (Kaltenbach): WA, KLIick1IrAt—Trout Lk, 29-VI-97 on Rubus sp. (97G168).

Amphorophora sensoriata Mason: WA, YAKIMA—Grandview, 22-VII-95 on Rubus sp. (95G368).

Aphis armoraciae Cowen: WA, KiTTITASs [on Mertensia paniculata*}.

Aphis ceanothi Clarke: WA, KitriraAs—S Fork Manastash Cr, 25-VI-97 on Ceanothus velutinus (97G137).

Aphis coweni Palmer: WA, KiTTiITAs [on Veratrum viride*].

Aphis fabae Scopoli: WA, Kirriras—Lost Lk, 6-VIII-97 on Senecio sp. (97G335).

Aphis pomi de Geer: WA, YAKIMA [on Pyrus malus*}].

Aphthargelia symphoricarpi (Thomas) [on Symphoricarpos sp.]: WA, Kittitas [*]; KLIckIrAat—Trout Lk, 29-
VI-97 (97G159).

Aspidophorodon longicauda (Richards): WA, SKAMANIA [on Spiraea sp. *].

Cavariella aegopodii (Scopoli): WA, GARFIELD—nr FR40 & FR44, 22-VIII-95 on Osmorhiza chilensis
(95T464+4).

Ceruraphis viburnicola (Gillette): WA, SKAMANIA [on Carex sp.*].

Chaetosiphon fragaefolii (Cockerell): ID, BOUNDARY [on Rosa sp.¢*]; WA, PEND OREILLE—Le Clerc WLA,
3-VUI-95 on native Rosa sp. (95T398); WA, STEVENS—Little Pend Oreille National WLA, 9-VII-96 on
native Rosa sp. (96T2097¢#).

Epameibaphis atricornis G&P: WA, BENTON—-WSU-Prosser, 4-V-95 on Chrysothamnus nauseosus
(95G0207 +).

Ericaphis gentneri (Mason): WA, KitrirAs—Colockum Pass, 2-VII-97 on Crataegus sp. (97G214).

Ericaphis scammelli (Mason): STEVENS—Little Pend Oreille National WLA, 9-VII-96 on native Rosa sp.
(96T209F£).

Hyperomyzus lactucae (L.): WA, BENTON [on Sonchus oleraceust*].

VOLUME 102, NUMBER 3 q29

Hyperomyzus nigricornis (Knowlton): WA, GARFIELD—Wickiup CG, 29-VIII-95 on Hieracium sp. (95T508);
KittirAs—Ellensburg, 23 VI—-96 on Ribes sp. (96G267).

Hyperomyzus sp.: WA, BENTON [on Sonchus oleraceus*].

Illinoia nr crystleae (Smith & Knowlton): WA, Kitriras—S Fork Manastash Cr Rd, mi marker 8, 16-VIII-
96 on Lonicera involucrata (96G489).

Illinoia maxima (Mason): WA, KLickirAt—Trout Lk, 29-VI-97 on Rubus parviflorus (97G164).

Illinoia rhododendri (Wilson): WA, KittiTAs [on Rhododendron albiflorum*}).

Illinoia sp.: WA, GARFIELD—Stentz Springs, 1-VIII-95 on Thalictrum sp. (95T367); PIERCE—RNP, nr Tipsoo
Lk, 16-VIII-95 on Spiraea densiflora (95A037); SPOKANE—Mt Spokane SP, 27-VII-95, on Vaccinium sp.
(951346).

Macrosiphoniella ludovicianae (Oestlund): WA, AsoTIN—Couse Cr Rd, mi marker 5, 18-VII-95 on Artemisia
ludoviciana (951257).

Macrosiphoniella tanacetaria (Kaltenbach): WA, SPOKANE—nr Mt Spokane, 29-VI-95 on Tanacetum vulgare
(95T1727%).

Macrosiphum albifrons Essig [on Lupinus sp.]: WA, KING—Stampede Pass, 8-VIII-96 (96G444); KLICKITAT—
Trout Lk, 11-VII-95 (95K068).

Macrosiphum californicum (Clarke): WA, SPOKANE—Mt Spokane SP, 27-VII-95, on Salix sp. (95T348).

Macrosiphum euphorbiae (Thomas): ID, LAaTtAH—Moscow Mtn, 3-VIII-96 on Castilleja sp. (96T398); WA,
GARFIELD—nr FR40 & FR44, 22-VIII-95 on Osmorhiza chilensis (95T464++); SPOKANE—nr Mt Spokane,
29-VI-95 on Tanacetum vulgare (96T172+); YAKIMA [on Chenopodium album+*|—N Fork Tieton River,
15-VI-96 on Hieracium albertinum (96K094).

Macrosiphum nr euphorbiae (Thomas): WA, YAKIMA—YIR, Mill Cr Guard Station, 6-VI-95 on Lomatium
nudicaule (95G115).

Macrosiphum nr gaurae (Williams) [det. by A. Jensen]: WA, YAKIMA—Wenas Lk, 11-VI-97 on Mentzelia
laevicaulis (97GO52).

Macrosiphum pallidum (Oestlund): WA, SPOKANE—Cheney, 23-V-95 on native Rosa sp. (95KO006).

Macrosiphum rosae (L.): WA, WHITMAN—Pullman, 1-VII-96 on cultivated Rosa sp. (961376); YAKIMA [on
Rosa sp.*].

Macrosiphum sp.: WA, YAKIMA—YIR, Howard Lk, 19-[X-95 on Pteridium aquilinum (95G640).

Macrosiphini [immatures]: WA, BENTON—WSU-Prosser, 27-X-95 on Malva neglecta (95K2127).

Metopolophium dirhodum (Walker): WA, YAKIMA—Sunnyside, 28-VI-95 on Phalaris arundinacea (95G254).

Microsiphoniella artemisiae (Gillette): WA, AsoTiIn—Asotin Cr HMU, 5-VII-96 on Artemisia ludoviciana
(96T141).

Myzus cerasi (EF) [on Prunus avium]: WA, BENTON [*] WHITMAN—Pullman, 5-VII-96 (96T173).

Myzus persicae (Sulzer): WA, BENTON—-WSU-Prosser, 20V—97 on Prunus persica (97G036).

Myzus sp.: WHITMAN [on Forsythia sp.*].

Nasonovia nr cynosbati (Oestlund): WA, YAKIMA—N Fork Tieton River, 15-VI-96 on native Ribes sp.
(96K091).

Nasonovia polemonii (G&P): WA, Kittiras—Buck Meadows, 9-VI-96, on Polemonium pulcherrimum
(96G347).

Nasonovia (Kakimia) sp.. WA, SPOKANE—nr Peone, 29-VI-95 on Phlox sp. (95T176); YAKIMA—YIR, nr Mt
Adams Lk, 4-VIII-95 on Lonicera cilosa (95KO091).

Nearctaphis sp.: WA, ASOTIN—De Spain Springs, 11-VH-96 on Castilleja sp. (961236).

Pleotrichophorus sp... WA, BENTON—-WSU-Prosser, 4-V-95 on Chrysothamnus nauseosus (95G020*%);
KLICKITAT—Alderdale & Mabton-Bickleton Rds, 28-IV-95 on Chrysothamnus sp. (95GO12).

Rhopalomyzus lonicerae (Siebold): WA, SPOKANE—nr Peone, 13-VII-95 on unknown Gramineae (95T228).

Rhopalosiphum maidis (Fitch): WA, BENTON [on Echinochloa crus-gallit*}.

Rhopalosiphum nymphaeae (L.): WA, YAKIMA—Morgan Lk, 28-VI-95 on Nymphaea sp. (95G262).

Rhopalosiphum padi (L.): WA, BENTON [on Echinochloa crus-galli+*].

Sitobion avenae (EF) [on Triticum aestivum]: WA, BENTON—-WSU-Prosser, 25-VII-97 (97F276); YAKIMA—
Zillah, 28-VI-95 (95G270).

Uroleucon nr erigeronense (Thomas): WA, Kittiras—Buck Meadows, 9-VII-96 on Hieracium albiflorum
(96G340).

Uroleucon ivae Robinson: WA, YAKIMA [on /va xanthifolia*].

Uroleucon russellae (HRL): WA, SKAMANIA [on Anaphalis margaritacea* |.

Uroleucon sonchi (L.): WA, BENTON—-WSU-Prosser, 6-VII-95 on Sonchus oleraceus (95A025).

Uroleucon sp.: WA, YAKIMA—Union Gap, 23-X-95 on Centaurea maculosa (95G720).

Undetermined spp.: MT, SANDERS [on Senecio sp.*]; OR, UMATILLA—Harris Pk, 11-VI-96 on Preridium

730 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

aquilinum (96G235); WA, Asotin—nr Anatone, 30-V-95 on Philadelphus lewisii (95T042); De Spain
Springs, 24-VII-96 on Compositae (96T346); KittrrAs—Quartz Mtn, 25-VII-96 on Rubus lasiococcus
(96G407): KLickirat—nr Cleveland, 7-VII-97 on Lotus pinnatus (97G233); WHITMAN—Pullman, 9-VII-
96 on Lactuca serriola (961228); 25-VII-95, on Lapsana communis (951332); YAKIMA—N Fork Tieton
River, 27-VI-96 on Achlys triphylla (96G289).

Genus PSEUDOPRAON Stary

Pseudopraon mindariphagum Stary
Mindarus abietinus Koch: WA, YAKIMA—Yakima Arboretum, 30-V-96, on Abies concolor (96G102).
Mindarus sp.: WA, YAKIMA—Grandview, 10-VI-96, on Abies grandis (96G215); N. Fork Tieton River, 15-
VI-96, on Abies sp. (96K0987).

Genus TOXARES Haliday

Toxares deltiger (Haliday)
Macrosiphum adianti (Oestlund): OR, BENTON—McDonald State Forest, on unknown plant, 21-V-91 (91JO02).
Macrosiphum walkeri Robinson [det. by A. Jensen]: OR, Hoop RIveR—Starvation Cr, 21-IX-95 on
Polypodium glycyrrhiza (95K180).

Genus TRIOXYS Haliday

Trioxys artemisiarum Pike & Stary
Aphis (Zyxaphis) canae Williams: WA, DouGLas—Badger Mtn, 13-VII-95, on Artemisia tridentata
(95B0127#).
Macrosiphoniella ludovicianae (Oestlund) [on Artemisia ludoviciana]: WA, Asotin [*]—Asotin Cr WLA, 5-
VII-96 (96T 138).
Microsiphoniella artemisiae (Gillette): WA, AsoTIN—Asotin Cr HMU, 5-VII-96, on Artemisia ludoviciana
(96T141).

Trioxys bonnevillensis Smith

Aphis canae Williams [on Artemisia tridentata]: WA, DouGLAs—Badger Mtn, 13-VII-95 (95B0127);
FRANKLIN—Kahlotus Hwy, 11-VI-96 (96G2427¢#).

Artemisaphis artemisicola (Williams) [on Artemisia tridentata]: WA, BENTON—Six Prong Rd, 24-I'V-97
(97G008+); FRANKLIN—Kahlotus Hwy, 11-VI-96 (96G2427+); YAKIMA, Glade Rd, 18-VI-97 (97G1037¢).

Epameibaphis atricornis G&P [on Artemisia tridentata]: WA, BENTON [*]—ALE, 19-V-95 (95G072+); Byron
Rd, 14-V-97 (97G0247+); KLICKITAT [*]; YAKIMA [7*].

Epameibaphis utahensis Knowlton & Smith: WA, KLICKITAT [on Artemisia tridentatat*].

Epameibaphis sp.: WA, BENTON—ALE, 8-V1-95 on Artemisia tridentata (95G1517).

Flabellomicrosiphum knowltoni Smith [on Artemisia tridentata}: WA, BENTON [+*]; YAKIMA [+*]—YIR, Fort
Simcoe, 30-V-95 (95G091).

Flabellomicrosiphum tridentatae (Wilson) [on Artemisia tridentata]: WA, FRANKLIN [*]—Juniper Sand Dunes,
23-V-95 (95GO88+, 95G089+); KLickiTaT [+*]—Alderdale Cr & Columbia River Rds, 28-IV-95 (95G013).

Flabellomicrosiphum sp. [on Artemisia tridentata]: WA, KLICKITAT [+*] & YAKIMA [**].

Microsiphoniella acophorum (Smith & Knowlton) [on Artemisia tridentata]: WA, FRANKLIN [*]—Juniper Sand
Dunes, 23-V-95 (95G088+, 95G0897); FRANKLIN [{*].

Microsiphoniella sp. (Smith & Knowlton): WA, BENTON [on Artemisia tridentata }*}.

Obtusicauda artemisiphila (Knowlton & Allen): WA, BENTON—ALE, 10-V-95 on Chrysothamnus nauseosus
(95G033+); KLICKIrTAT—nr Roosevelt [on Artemisia tridentata+*].

Obtusicauda coweni (Hunter): WA, AsoTIN—Asotin Cr HMU, 7-V-96 on Artemisia tridentata (9611427).

Obtusicauda filifoliae (G&P) [on Artemisia tridentata): WA, BENTON—Byron Rd, 14-V-97 (97G0247#); nr
Prosser, Rotha Rd, 2-V-95 (95G0O17+#); KLICKITAT [F£*]; YAKIMA [+=*].

Obtusicauda sp. {on Artemisia tridentata]: WA, BENTON [+£*]; KLIcKITaT [t£*]; YAKIMA [++*].

Pleotrichophorus quadritrichus (Knowlton & Smith)?: WA, YAKIMA—Wenas Lk, 11-VI-97 on Artemisia
tridentata (97GO58t¢£).

Pleotrichophorus sp. [on Artemisia tridentata]:. WA, BENTON [t£*]; KLickrrat—Alderdale Rd, 9-V-95
(95G0287#).

Pseudoepameibaphis essigi Knowlton & Smith: WA, KLICKITAT [on Artemisia tridentata +*].

Pseudoepameibaphis tridentata (Wilson) [on Artemisia tridentata]: WA, BENTON [+*]—ALE, 19-V-95
(95G072+); Byron Rd, 14-V-97 (97G024++); KLickiTaT [+*]—Alderdale Rd, 9-V-95 (95G0287); KitTITAs

VOLUME 102, NUMBER 3 731

[+*]; YAKIMA [+*]—Naught Rd, 18-VI-97 (97G103++, 97G120); Wenas Lk, 11-VI-97 (97G0587); YIR,
Ft. Simcoe, 30-V-96 (95G0917).
Trioxys cirsii (Curtis)
Drepanosiphum platanoidis (Schrank): WA, BENTON [on Acer pseudoplatanus*].
Trioxys complanatus Quilis
Therioaphis riehmi (Borner) [on Melilotus sp.]: WA, BENTON—-WSU-Prosser, 11 & 18-VII-97 on Melilotus
officinalis (97F218+, 97F254+); KLickIrat—Roosevelt, 24-IV-96 (96G014+); YAKIMA, YIR, Signal Peak,
19-IX-95 (95G6257).
Therioaphis sp.: WA, BENTON—-WSU-Prosser, 28-VI-95 on Medicago sativa (95G2777+).
Trioxys pallidus (Haliday) [group]
Chromaphis juglandicola (Kaltenbach) [on Juglans regia]: WA, BENTON—-WSU-Prosser, 6-[X-95 (95K 1437);
KLIcKITAaT—Bickleton, 27-VIII-96 (96G511); YAKIMA [*]—Yakima Arboretum, 26-IX-95 (95G657).
Eucallipterus tiliae (L.) [on Tilia americana]: MT, LINCOLN—Libby, 29-VII-95 (95G449); WA, BENTON—
Prosser, 1-VIII-95, 25-IX-95 (95A029, 95G647, 95G648); WHITMAN—Pullman, 9-X-96 (96T586).
Monelliopsis caryae (Monell) [on Juglans nigra unless otherwise indicated]: WA, BENTON [on J. regia*];

WSU-Prosser, 6-I[X-95 (95G6507); YAKIMA [*]—Yakima Arboretum, 7-VII-94 & 26-IX-95 (94K020,
95G656).

Myzocallis coryli (Goeze): WA, CLARK—WSU-Vancouver, 21-I[X-95 on Corylus sp. (95K176).

Myzocallis multisetis Boudreaux & Tissot: WA, BENTON—-WSU-Prosser, 9-[X-95 on Quercus borealis
(95K158).

Myzocallis sp.: WA, BENTON [on Quercus rubra*|; WHITMAN—Pullman, 9-X-96 on Quercus sp. (96T579).

Panaphis juglandis (Goeze) [on Juglans regia}: WA, BENTON [*]—Prosser 25-IX-95 (95G646); WSU-Prosser,
6-IX-95 (95K1437); YAKIMA [*]—Grandview, 25-IX-95 (95G653).

Trioxys rosae Pike & Stary
Macrosiphum pallidum (Oestlund): WA, SPOKANE—nr Cheney, 23-V-95 on native Rosa sp. (95KO006).
Macrosiphum rosae (L.): WA, YAKIMA [on native Rosa sp.*].

Trioxys setaceus Pike and Stary, new species
(Figs. 51-55)

Diagnosis.—The new species is similar to T. artemisiarum Pike and Stary, from which
it may be distinguished as follows: head prevalently dark brown; antenna 1 1-segmented;
metasomal tergum 1 parallel-sided, not dilated to apex; prong dorsal surface with 6
perpendicular setae and 2 shorter setae close to the base; last sternum with a group or
longitudinal row of setae equal in length with 2 neighboring setae at dorsal base of prong.

Etymology.—The name of the new species is derived from the presence of numerous
setae on the prong of the last metasomal sternum.

Description.—Female. Head: Eye large. Malar space equal to /% eye length. Tentorio-
ocular line ¥, intertentorial line. Antenna 11-segmented, reaching to mid-metasoma.
Flagellomeres F, to F, approximately equal in size, F, to F; as long as wide. Mesosoma:
Mesosoma smooth, with a simple row of sparse individual setae tracing the effaced notauli
on the disc. Propodeum (Fig. 52) smooth, with indications of short carinae or longitudinal
rugosites in the distal portion. Forewing (Fig. 53): Stigma triangular, length 3 width;
distal abscissa of R1 (metacarpus) short, distinctly shorter than stigma width. Metasoma:
Metasomal tergum 1 (Fig. 51) length almost 2X width at spiracles (spiracles slightly
prominent), roughly parallel sided from spiracle to posterior margin. Genitalia (Fig. 54):
Prong straight, except for slight curve apically, apex with 2 longitudinally dilated setae
(Fig. 55). Prong dorsal setae perpendicular (6 setae about 2 prong diameter, 2 near base
short equal to prong diameter and to setae on distal part of last sternum); prong ventral
setae numerous, oblique, about equal to prong diameter. Coloration: Head dark brown;
lower part, clypeus, and gena yellow, palpi light yellow. Antenna scape, pedicel, and base
of flagellomere 1 yellow, remaining antenna brown. Legs yellow brown, trochanters and
base of tibia somewhat lighter. Hind leg with dark tinge. Metasomal tergum 1 yellow,

132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 51-55. Trioxys setaceus, 2 [illustrations not to equal scale]. 51, Metasomal tergum 1. 52, Propodeum.
53, Fore wing, in part. 54, Genitalia. 55, Prong, close-up of apex. Abbreviations: R1, distal abscissa of post-
marginal vein [= metacarpus].

terga 2 and 3 with yellow center and brown lateral spots; middle terga brown; distal
tergum yellow. Ovipositor sheath yellow with brown apex; prong completely yellow to
brownish at apex. Length of body: about 1.9—2.2 mm.

Male.—Antenna 13-segmented. Coloration: head brown; clypeus and mandibles yellow,
palpi light brown. Mesosoma brown. Wings hyaline, venation brown. Legs light brown,
trochanter and base of tibia yellow. Metasoma brown, except tergum | yellow.

Holotype.—?, USA, WA, AsoTIN Co., Schumaker Rd, 18-VII-1995, T. Miller collector
(collection code 95T266). Host aphid: Acyrthosiphon macrosiphum (Wilson) on
Amelanchier alnifolia. Deposited in USNM.

Paratypes.—S specimens, collection data same as holotype, deposited in collection of
WSU-Prosser (4 2, 1 3).

Other material reared from:

Acyrthosiphon macrosiphum (Wilson) [on Amelanchier alnifolia, unless indicated
otherwise]: MT, LINCOLN—Libby, 6-VII-96 (96G322); WA, AsoTiIN—Anatone, 18-VII-95
(95T266, 13 specimens); PEND OREILLE—Le Clerc WLA, 3-VIII-95 (95T402); SPOKANE,
Cooper Rd nr Hwy 206, 29-VI-1995 (95T180, 5 specimens); nr Mt Spokane, on Ribes
sp.?, (95T174).

VOLUME 102, NUMBER 3 133)

Trioxys tenuicaudus Stary
Eucallipterus tiliae (L.): WA, BENTON—Prosser, 25-IX-95 on Tilia americana, (95G647, 95G648);
KLICKITAT—Trout Lk, 29-VI-97 on T. sp. (97G174); WHITMAN—Pullman, 14-VIII-96 on T. americana,
(96T443).

Trioxys spp.

Aphis coweni Palmer: WA, KitTiTAs [on Veratrum viride*].

Chaetosiphon fragaefolii (Cockerell): WA, SPOKANE—Turnbull National WLA, 1-VI-95, on native Rosa sp.
(95T048).

Eucallipterus tiliae (L.): WA, WHITMAN—Pullman, 1-VIII-96 on Tilia americana (961377).

Euceraphis sp.: WA, SKAMANIA—Yellow Jacket Cr, 25-IX-94 on Trautvetteria caroliniensis (94G0217#).

Macrosiphoniella ludovicianae (Oestlund): WA, YAKIMA—Naches, 28-VII-97 on Artemisia ludoviciana
(97G277).

Macrosiphum rosae (L.): WA, YAKIMA—nr Yakima, 3-VI-94 on native Rosa sp. (94/063).

Monelliopsis nr caryae (Fitch): WA, BENTON—-WSU-Prosser, 24-VI-97 on Juglans nigra (97G126).

Myzocallis nr granovskyi Boudreaux & Tissot: WA, BENTON—-WSU-Prosser, 24-VI-97 on Quercus rubra
(97G125).

Oestlundiella flava (Davidson): WA, SKAMANIA—Yellow Jacket Cr, 25-IX-94 on Trautvetteria caroliniensis
(94G0217+#).

Panaphis juglandis (Goeze): WA, BENTON—-WSU-Prosser, 24-VI-97 on Juglans regia (97G127).

Pleotrichophorus sp.: WA, YAKIMA [on Haplopappus resinosus*].

Genus XENOSTIGMUS Smith

Xenostigmus bifasciatus (Ashmead)
Cinara ponderosae (Williams) [on Pinus ponderosa]: WA, AsoTiIn—De Spain Spring, 5-VII-96 (96T149);
YAKIMA [*].
Cinara sp.: WA, AsoTiN—Cooper Canyon, 18-VII-96 on Pinus ponderosa (96T308).
Undetermined sp.: WA, YAKIMA—Clear Lk, 15-VI-96 on Pinus ponderosa (96K087).

734

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

APHID HOST—PARASITOID INDEX

+—INDICATES APHID-PARASITOID ASSOCIATION NOT CERTAIN

Acyrthosiphon kondoi
Aphidius ervi
Acyrthosiphon lactucae
Aphidius avenaphis +
Aphidius ervi
Aphidius polygonaphis
Aphidius sp.
Diaeretiella rapae
Ephedrus californicus
Lysiphlebus testaceipes
Praon occidentale
Praon unicum
Praon sp.
Acyrthosiphon macrosiphum
Aphidius sp.
Lysiphlebus testaceipes
Praon humulaphidis
Praon pequodorum
Trioxys setaceus
Acyrthosiphon pisum
Aphidius ervi
Aphidius nr pisivorus
Aphidius pulcher
Aphidius smithi
Aphidius sp.
Diaeretiella rapae
Lysiphlebus testaceipes
Monoctonus sp.
Praon occidentale
Praon pequodorum
Praon sp.
Amphorophora geranii
Aphidius sp.
Praon occidentale
Amphorophora parviflori
Aphidius polygonaphis
Amphorophora rubi
Praon pequodorum
Praon sp.
Amphorophora nr rubitoxica
Aphidius sp.
Amphorophora sensoriata
Praon sp.
Amphorophora sp.?
Aphidius polygonaphis
Anoecia corni
Lysiphlebus testaceipes
Aphis armoraciae
Adialytus fuscicornis
Aphidius sp.
Lysiphlebus testaceipes
Praon sp.

Aphis nr armoraciae
Lysiphlebus testaceipes

Aphis canae
Binodoxys clydesmithi =
Lysaphidus adelocarinus
Lysiphlebus testaceipes +
Lysiphlebus utahensis ~
Trioxys artemisiarum +
Trioxys bonnevillensis +

Aphis ceanothi
Aphidius polygonaphis
Binodoxys carolinensis
Lysiphlebus testaceipes
Praon sp.

Aphis nr ceanothi
Lysiphlebus testaceipes

Aphis nr chrysothamni
Lysiphlebus testaceipes

Aphis coweni
Ephedrus sp.
Lysiphlebus testaceipes
Praon americanum
Praon unicum
Praon sp.

Trioxys sp.

Aphis craccivora
Aphidius ervi
Aphidius sp.
Diaeretiella rapae
Lysiphlebus testaceipes
Praon occidentale +
Praon unicum

Aphis nr epilobiaria
Lysiphlebus testaceipes

Aphis fabae
Aphidius matricariae
Aphidius sp.

Binodoxys grafi
Diaeretiella rapae
Ephedrus californicus
Lysiphlebus testaceipes
Praon sp.

Aphis nr fabae
Diaeretiella rapae
Lysiphlebus testaceipes
Praon unicum

Aphis filifoliae
Lysiphlebus testaceipes

Aphis frangulae
Lysiphlebus testaceipes

Aphis nr frangulae
Lysiphlebus testaceipes

Aphis gossyptt
Lysiphlebus testaceipes
Aphis hederae
Lysiphlebus testaceipes
Aphis helianthi
Adialytus fuscicornis
Aphidius matricariae
Aphidius polygonaphis +
Aphidius sp.
Binodoxys carolinensis
Binodoxys grafi
Diaeretiella rapae
Ephedrus californicus
Ephedrus clavicornis
Lysiphlebus testaceipes
Lysiphlebus sp.
Praon occidentale
Praon unicum
Aphis nr helianthi
Lysiphlebus testaceipes
Aphis hermistonii
Lysiphlebus testaceipes
Aphis holodisci
Aphidius sp.
Binodoxys graft
Ephedrus californicus
Lysiphlebus testaceipes
Aphis lugentis
Lysiphlebus testaceipes
Praon occidentale
Aphis lupini
Lysiphlebus testaceipes
Aphis neogillettei
Lysiphlebus testaceipes
Aphis nerit
Aphidius colemani
Lysiphlebus testaceipes
Aphis nigratibialis
Aphidius sp.
Binodoxys carolinensis
Lysiphlebus testaceipes
Praon unicum
Aphis oenotherae
Lysiphlebus testaceipes
Praon unicum
Aphis nr oenotherae
Aphidius polygonaphis
Binodoxys carolinensis
Ephedrus californicus
Ephedrus clavicornis
Lysiphlebus testaceipes

VOLUME 102, NUMBER 3

Monoctonus washingtonensis

Praon unicum
Aphis pomi

Lysiphlebus testaceipes +

Praon unicum
Praon sp.

Aphis rumicis
Lysiphlebus testaceipes
Praon unicum

Aphis salicariae
Binodoxys carolinensis
Binodoxys grafi
Binodoxys sp.
Lysiphlebus testaceipes
Praon americanum
Praon unicum

Aphis nr salicariae
Binodoxys grafi
Lysiphlebus testaceipes
Praon unicum

Aphis sambuci
Lysiphlebus testaceipes

Aphis spiraecola
Lysiphlebus testaceipes

Aphis nr triglochinis
Binodoxys grafi
Lysiphlebus testaceipes

Aphis nr valerianae
Lysiphlebus testaceipes

Aphis varians
Aphidius sp.
Binodoxys carolinensis
Binodoxys grafi
Ephedrus californicus
Ephedrus clavicornis
Ephedrus sp.
Lysiphlebus testaceipes
Praon unicum

Aphis nr varians
Binodoxys carolinensis
Binodoxys grafi
Ephedrus californicus
Lysiphlebus testaceipes
Praon unicum

Aphis (Zyxaphis) sp.
Aphidius sp.
Lysaphidus adelocarinus
Lysaphidus rosaphidis +
Lysiphlebus testaceipes

Aphis sp. n.

Binodoxys carolinensis
Lysiphlebus testaceipes

Aphis spp.

Aphidius ervi
Binodoxys carolinensis
Lysiphlebus testaceipes
Lysiphlebus sp.

Praon unicum

++

Aphthargelia symphoricarpi

Aphidius matricariae
Ephedrus californicus
Ephedrus clavicornis
Lysaphidus sp.
Lysiphlebus testaceipes
Praon coniforme
Praon sp.
Artemisaphis artemisicola
Binodoxys clydesmithi

Lysaphidus adelocarinus

Lysiphlebus utahensis
Praon artemisicola
Trioxys bonnevillensis

Aspidophorodon longicauda

Aphidius ervi
Aphidius sp.
Praon sp.

Aspidophorodon nr longicauda

Ephedrus lacertosus +
Boernerina variabilis
Betuloxys alnicolus ¢
Lysiphlebus testaceipes
Brachycaudus cardui
Aphidius matricariae
Aphidius polygonaphist
Diaeretiella rapae
Ephedrus persicae
Lysiphlebus testaceipes
Brachycaudus helichrysi
Aphidius colemani
Aphidius ervi
Aphidius matricariae
Aphidius sp.
Diaeretiella rapae
Ephedrus californicus =
Ephedrus persicae
Lysiphlebus testaceipes

Monoctonus campbellianus

Praon occidentale
Praon unicum

Brachycaudus tragopogonis

Aphidius sp.
Diaeretiella rapae
Lysiphlebus testaceipes
Lysiphlebus sp.
Praon unicum
Brachycorynella asparagi
Diaeretiella rapae
Lysiphlebus testaceipes
Braggia eriogoni
Ephedrus californicus
Lysiphlebus testaceipes
Braggia nr. eriogoni.
Lysiphlebus testaceipes
Braggia sp. n.
Lysiphlebus testaceipes
Braggia sp.

Diaeretiella rapae
Lysiphlebus testaceipes
Brevicoryne brassicae
Aphidius ervi
Aphidius matricariae
Diaeretiella rapae
Ephedrus californicus
Lysiphlebus testaceipes
Capitophorus elaeagni
Aphidius matricariae
Aphidius polygonaphis
Aphidius sp.
Ephedrus californicus
Lysiphlebus testaceipes
Praon occidentale +
Carolinaia howardii
Aphidius sp.
Cavariella aegopodii
Aphidius salicis
Ephedrus californicus
Lysiphlebus testaceipes
Praon unicum
Praon sp. £
Cavariella nr aegopodii

Binodoxys carolinensis +

Cavariella pastinacae
Aphidius salicis
Ephedrus californicus
Lysiphlebus testaceipes
Praon unicum
Praon yakimanum

Cedoaphis incognita
Lysiphlebus flavidus

Ceruraphis viburnicola
Lysiphlebus testaceipes
Praon sp.

Chaetosiphon fragaefolii
Aphidius sp.

Binodoxys sp.
Ephedrus californicus
Ephedrus sp. ¢
Lysaphidus rosaphidis
Lysaphidus sp. =
Lysiphlebus testaceipes
Praon sp.

Trioxys sp.

Chaetosiphon nr hottesi
Aphidius ervi
Aphidius sp. ¢

Chaetosiphon tetrarhodum
Aphidius sp. =

Chaetosiphon thomasi
Praon occidentale

Chaetosiphon nr thomasi
Lysaphidus rosaphidis

Chaitophorus populicola
Adialytus salicaphis
Lysiphlebus testaceipes

785

736 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Chaitophorus populifolii
Adialytus salicaphis
Ephedrus chaitophori

Chaitophorus utahensis
Adialytus salicaphis

Chaitophorus viminalis
Adialytus salicaphis
Lysiphlebus testaceipes

Chaitophorus sp.
Adialytus salicaphis

Chromaphis juglandicola
Trioxys pallidus

Cinara brevispinosa
Pauesia sp.

Cinara chinookiana
Lysiphlebus testaceipes

Cinara pilicornis
Pauesia juniperaphidis

Cinara ponderosae
Pauesia ahtanumensis
Pauesia pahtonis
Pauesia ponderosaecola
Pauesia sp.
Xenostigmus bifasciatus

Cinara pseudotaxifoliae
Pauesia pseudotsugae

Cinara pseudotsugae
Pauesia sp.

Cinara spp.

Lysiphlebus testaceipes
Pauesia ahtanumensis
Pauesia bicolor
Pauesia sp.
Xenostigmus bifasciatus

Diuraphis noxia
Aphidius avenaphis
Aphidius ervi
Aphidius matricariae
Aphidius sp.
Diaeretiella rapae
Ephedrus californicus
Ephedrus sp.
Lysiphlebus testaceipes
Monoctonus washingtonensis
Praon gallicum
Praon occidentale
Praon unicum
Praon yakimanum

Drepanosiphum platanoidis
Trioxys cirsii

Dysaphis plantaginea
Aphidius matricariae +
Diaeretiella rapae
Ephedrus clavicornis
Lysiphlebus testaceipes
Praon occidentale
Praon unicum

Epameibaphis atricornis
Lysaphidus adelocarinus
Lysiphlebus utahensis +
Praon sp. +
Praon artemisicola
Trioxys bonnevillensis

Epameibaphis utahensis
Trioxys bonnevillensis

Epameibaphis sp.

Trioxys bonnevillensis

Ericaphis gentneri
Aphidius nr hortensis
Aphidius sp.

Ephedrus lacertosus
Lysaphidus sp.
Praon unicum
Praon sp.

Ericaphis nr gentneri
Aphidius sp.

Lysiphlebus testaceipes

Ericaphis scammelli
Aphidius sp.

Ephedrus sp. +
Lysaphidus sp. ¢
Praon sp. =

Ericaphis wakibae
Aphidius polygonaphis
Aphidius sp.

Ephedrus californicus +
Praon occidentale +

Eriosoma americanum
Lysiphlebus testaceipes =

Eriosoma lanigerum
Lysiphlebus testaceipes +

Eucallipterus tiliae
Trioxys pallidus
Trioxys tenuicaudus
Trioxys sp.

Euceraphis gillettei

Aphidius sp. $
Praon nr unicum ¢

Euceraphis sp.

Trioxys sp. £

Flabellomicrosiphum knowltoni
Lysaphidus adelocarinus
Trioxys bonnevillensis

Flabellomicrosiphum tridentata
Lysaphidus adelocarinus +
Lysiphlebus utahensis +
Trioxys bonnevillensis

Flabellomicrosiphum sp.
Lysaphidus adelocarinus
Praon artemisicola
Trioxys bonnevillensis

Forda marginata
Adialytus fuscicornis

Hayhurstia atriplicis

Aphidius ervi
Diaeretiella rapae

Hyadaphis foeniculi
Aphidius colemani
Lysiphlebus testaceipes +

Hyalopteroides humilis
Aphidius ervi

Hyalopterus pruni
Aphidius ervi +
Aphidius matricariae +
Monoctonus campbellianus

Hyperomyzus lactucae
Aphidius ervi
Aphidius segmentatus
Ephedrus californicus
Praon unicum
Praon sp.

Hyperomyzus nigricornis
Aphidius ervi
Aphidius polygonaphis
Aphidius segmentatus
Aphidius sp.

Ephedrus californicus
Ephedrus sp.
Lysiphlebus testaceipes
Praon occidentale
Praon unicum

Praon sp.

Hyperomyzus spp.
Aphidius polygonaphis
Aphidius segmentatus
Aphidius sp.

Ephedrus californicus
Praon occidentale
Praon sp.

Illinoia nr azaleae
Aphidius polygonaphis

Illinoia corylina
Aphidius matricariae
Diaeretiella rapae

Illinoia crystleae bartholomewi
Aphidius sp.

Illinoia nr crystleae
Aphidius sp.

Praon sp.

Illinoia davidsoni
Aphidius ervi
Aphidius polygonaphis
Aphidius sp.

Praon occidentale

Illinoia maxima
Ephedrus clavicornis
Praon sp.

Illinoia rhododendri
Aphidius sp.
Binodoxys sp.
Monoctonus sp.

Praon sp.

VOLUME 102, NUMBER 3

Illinoia richardsi
Praon occidentale +

Illinoia spiraeae
Lysiphlebus testaceipes

Ilinoia nx thalictri
Praon occidentale

Illinoia spp.

Aphidius ervi
Aphidius polygonaphis
Aphidius sp.

Ephedrus californicus
Monoctonus sp.

Praon humulaphidis
Praon occidentale
Praon nr occidentale
Praon pequodorum
Praon sp.

Illinoia sp. ?

Praon humulaphidis

Liosomaphis berberidis
Aphidius nr hortensis
Aphidius sp.

Lipaphis erysimi
Aphidius avenaphis
Aphidius colemani
Diaeretiella rapae

Macrosiphoniella ludovicianae
Aphidius ohioensis
Aphidius polygonaphis
Aphidius sp.

Ephedrus californicus
Praon occidentale
Praon sp.
Trioxys artemisiarum
Trioxys sp.
Macrosiphoniella nr ludovicianae
Praon occidentale

Macrosiphoniella tanacetaria
Aphidius ohioensis
Ephedrus californicus
Praon sp. =

Macrosiphum adianti
Toxares deltiger

Macrosiphum albifrons
Aphidius ervi
Aphidius lupini
Ephedrus californicus
Ephedrus lacertosus
Praon occidentale
Praon sp.

Macrosiphum californicum
Aphidius sp.

Ephedrus californicus
Praon sp.

Macrosiphum clydesmithi
Aphidius nigripes
Aphidius polygonaphis
Aphidius sp.

Macrosiphum creelii
Aphidius ohioensis
Ephedrus californicus

Macrosiphum euphorbiae
Aphidius colemani
Aphidius ervi
Aphidius matricariae
Aphidius nigripes
Aphidius polygonaphis
Aphidius sp.
Diaeretiella rapae
Ephedrus californicus
Ephedrus lacertosus
Lysaphidus sp.
Lysiphlebus testaceipes
Monoctonus sp.

Praon fulvum
Praon humulaphidis
Praon occidentale
Praon unicum
Praon sp.

Macrosiphum nr euphorbiae
Aphidius polygonaphis
Diaeretiella rapae
Ephedrus californicus
Ephedrus sp.

Praon sp.

Macrosiphum gaurae
Aphidius ervi
Ephedrus californicus
Lysiphlebus testaceipes
Praon unicum

Macrosiphum nr gaurae
Praon sp.

Macrosiphum osmaroniae
Praon occidentale

Macrosiphum pallidum
Aphidius nr rosae

Aphidius sp. =
Ephedrus californicus +
Praon unicum

Praon sp.

Trioxys rosae

Macrosiphum nr pallidum
Aphidius ervi +
Lysiphlebus testaceipes +

Macrosiphum pteridis

Aphidius polygonaphis

Macrosiphum rhamni

Aphidius sp.

Ephedrus sp.

Praon simulans
Macrosiphum rosae

Aphidius ervi

Aphidius polygonaphis

Aphidius sp.

Ephedrus californicus

Praon occidentale

UY

Praon sp.
Trioxys rosae
Trioxys sp.
Macrosiphum tuberculaceps
Aphidius sp.
Monoctonus pacificus
Praon fulvum
Macrosiphum valerianae
Aphidius ohioensis
Aphidius polygonaphis
Ephedrus californicus +
Praon unicum ¢
Macrosiphum walkeri
Aphidius nr polygonaphis
Monoctonus sp.
Toxares deltiger
Macrosiphum spp.
Aphidius ervi
Aphidius polygonaphis
Aphidius sp.
Diaeretiella rapae
Ephedrus lacertosus
Lysiphlebus testaceipes
Praon fulvum
Praon humulaphidis
Praon occidentale
Praon simulans
Praon unicum
Praon sp.
Toxares deltiger
Macrosiphum sp. ?
Aphidius ohioensis
Praon occidetale
Macrosiphini [immatures]
Ephedrus lacertosus
Monoctonus pacificus
Praon sp.
Metopolophium dirhodum
Aphidius ervi
Aphidius rhopalosiphi
Aphidius sp.
Ephedrus californicus =
Monoctonus sp.
Praon gallicum
Praon occidentale =
Praon unicum
Praon yakimanum
Praon sp.
Metopolophium sp.
Ephedrus lacertosus ¢
Metopolophium sp. ?
Praon occidentale
Microlophium nr sibiricum
Praon occidentale
Microsiphoniella acophorum
Trioxys bonnevillensis
Microsiphoniella artemisiae
Aphidius ohioensis

738 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Lysaphidus adelocarinus
Praon sp.
Trioxys artemisiarum
Microsiphoniella sp.
Trioxys bonnevillensis
Mindarus abietinus
Pseudopraon mindariphagum
Mindarus sp.
Pseudopraon mindariphagum
Monelliopsis caryae
Trioxys pallidus
Monelliopsis nr caryae
Trioxys sp.
Myzocallis coryli
Trioxys pallidus
Myzocallis nr granovskyi
Trioxys sp.
Myzocallis multisetis
Trioxys pallidus
Myzocallis sp.
Trioxys pallidus
Myzodium knowltoni
Praon unicum <=
Myzus ascalonicus
Lysiphlebus testaceipes
Myzus cerasi
Aphidius sp.
Ephedrus californicus
Lysiphlebus testaceipes
Praon unicum
Praon sp.
Myzus nr cerasi
Lysiphlebus testaceipes
Myzus nr ornatus
Aphidius ervi ¢
Myzus persicae
Aphidius colemani
Aphidius ervi
Aphidius matricariae
Diaeretiella rapae
Ephedrus californicus
Lysiphlebus testaceipes
Praon occidentale
Praon unicum
Praon sp.
Myzus spp.
Aphidius colemani
Aphidius ervi
Aphidius matricariae
Diaeretiella rapae
Praon unicum
Praon sp.
Nasonovia alpina
Aphidius kakimiaphidis
Nasonovia aquilegiae
Aphidius kakimiaphidis
Aphidius sp.
Praon occidentale

Nasonovia nr aquilegiae
Aphidius sp.

Nasonovia crenicorna
Aphidius polygonaphis

Nasonovia cynosbati
Aphidius kakimiaphidis

Nasonovia nr cynosbati
Aphidius kakimiaphidis
Praon sp.

Nasonovia houghtonensis
Aphidius polygonaphis
Harkeria rufa

Nasonovia nr houghtonensis
Aphidius kakimiaphidis

Nasonovia polemonii
Aphidius sp.

Praon occidentale
Praon sp.

Nasonovia ribisnigeri
Ephedrus californicus
Praon humulaphidis
Praon unicum

Nasonovia wahinkae
Aphidius kakimiaphidis

Nasonovia spp.

Aphidius kakimiaphidis
Aphidius sp.

Ephedrus californicus
Lysaphidus sp.
Lysiphlebus testaceipes
Praon occidentale +
Praon unicum

Praon sp.

Nearctaphis bakeri
Lysiphlebus testaceipes
Praon occidentale +
Praon unicum

Nearctaphis californica
Lysiphlebus testaceipes

Nearctaphis kachena
Lysiphlebus testaceipes +
Praon occidentale
Praon unicum

Nearctaphis yohoensis
Lysiphlebus testaceipes

Nearctaphis spp.
Lysiphlebus testaceipes
Praon sp.

Obtusicauda artemisiphila
Ephedrus californicus
Lysiphlebus utahensis +
Trioxys bonnevillensis

Obtusicauda coweni
Binodoxys clydesmithi
Binodoxys coruscanigrans
Ephedrus californicus
Lysaphidus adelocarinus
Lysiphlebus utahensis

Praon artemisaphis

Trioxys bonnevillensis
Obtusicauda filifoliae

Lysiphlebus utahensis *

Trioxys bonnevillensis =
Obtusicauda spp.

Ephedrus californicus

Lysaphidus adelocarinus

Lysiphlebus utahensis

Praon artemisaphis

Trioxys bonnevillensis +
Oestlundiella flava

Aphidius sp. +=

Praon nr unicum +

Trioxys sp.
Ovatus crataegarius

Aphidius ervi

Aphidius matricariae

Aphidius sp.

Ephedrus incompletus
Panaphis juglandis

Trioxys pallidus

Trioxys sp.
Periphyllus nr brevispinosus

Adialytus salicaphis
Periphyllus californiensis

Euaphidius setiger
Periphyllus lyropictus

Euaphidius setiger
Periphyllus sp.

Adialytus salicaphis

Euaphidius setiger
Phorodon humuli

Aphidius ervi ¥

Aphidius matricariae

Binodoxys conei

Diaeretiella rapae

Lysiphlebus testaceipes

Monoctonus campbellianus

Praon occidentale

Praon unicum
Pleotrichophorus nr amsinckii

Ephedrus californicus

Lysaphidus adelocarinus
Pleotrichophorus elongatus

Lysaphidus ramithyrus
Pleotrichophorus palmerae

Lysaphidus adelocarinus
Pleotrichophorus pseudoglandu-

losus

Ephedrus californicus +
Pleotrichophorus quadritichus

Praon artemiscola

Trioxys bonnevillensis *
Pleotrichophorus nr. wasatchi

Lysaphidus ramithyrus
Pleotrichophorus spp.

Lysaphidus adelocarinus

VOLUME 102, NUMBER 3

Lysaphidus rosaphidis +
Praon artemisicola +
Praon sp.
Trioxys bonnevillensis +
Trioxys sp.
Prociphilus sp.
Aphidius sp.
Ephedrus clavicornis
Ephedrus sp.
Pseudoepameibaphis essigi
Lysaphidus adelocarinus
Trioxys bonnevillensis
Pseudoepameibaphis glauca
Lysaphidus adelocarinus
Pseudoepameibaphis tridentatae
Lysaphidus adelocarinus
Lysiphlebus testaceipes +
Lysiphlebus utahensis +
Praon artemisicola
Trioxys bonnevillensis
Pterocomma beulahense
Euaphidius cingulatus
Pterocomma bicolor
Euaphidius cingulatus
Pterocomma sanguiceps
Euaphidius cingulatus
Pterocomma smithiae
Euaphidius cingulatus
Pterocomma sp.
Euaphidius cingulatus
Lysiphlebus testaceipes
Rhopalomyzus lonicerae
Praon sp.
Rhopalosiphum cerasifoliae
Lysiphlebus testaceipes
Rhopalosiphum enigmae
Lysiphlebus testaceipes
Rhopalosiphum insertum
Aphidius matricariae
Diaeretiella rapae
Ephedrus californicus
Lysiphlebus testaceipes
Praon occidentale
Praon unicum
Rhopalosiphum maidis
Aphidius ervi
Diaeretiella rapae
Ephedrus californicus +
Lysiphlebus testaceipes
Praon unicum
Praon sp.
Rhopalosiphum nymphaeae
Diaeretellus palustris
Lysiphlebus testaceipes
Praon sp.
Rhopalosiphum padi
Aphidius avenaphis
Aphidius ervi

Aphidius matricariae
Aphidius sp.
Diaeretiella rapae
Ephedrus californicus +
Lysiphlebus testaceipes
Monoctonus washingtonensis
Praon gallicum +
Praon occidentale
Praon unicum
Praon yakimanum
Praon sp.
Schizaphis graminum
Lysiphlebus testaceipes
Sipha elegans
Adialytus ambiguus
Sipha sp.
Adialytus ambiguus
Sitobion avenae
Aphidius avenaphis
Aphidius ervi
Aphidius matricariae
Aphidius rhopalosiphi
Aphidius sp.
Diaeretiella rapae
Ephedrus californicus
Ephedrus clavicornis
Lysiphlebus testaceipes
Praon gallicum
Praon occidentale
Praon unicum
Praon yakimanum
Praon sp.
Sitobion fragariae
Aphidius sp.
Sitobion spp.
Aphidius polygonaphis
Ephedrus lacertosus
Lysiphlebus testaceipes
Therioaphis riehmi
Praon exsoletum
Praon pequodorum +
Trioxys complanatus
Therioaphis trifolii
Praon exsoletum
Therioaphis sp.
Praon exsoletum
Trioxys complanatus
Tuberculatus columbiae
Lysiphlebus testaceipes
Uroleucon adenocaulonae
Aphidius ohioensis
Uroleucon nr arnesense
Aphidius ohioensis
Aphidius polygonaphis
Ephedrus californicus
Uroleucon cirsii
Lysiphlebus testaceipes

139

Uroleucon nr epilobii
Aphidius ohioensis
Uroleucon erigeronense
Aphidius ohioensis
Aphidius segmentatus +

Aphidius sp.
Ephedrus californicus
Praon occidentale
Praon nr occidentale

Uroleucon nr erigeronense
Aphidius ohioensis
Aphidius polygonaphis
Ephedrus californicus
Praon sp.

Uroleucon escalantii
Aphidius ohioensis
Uroleucon nr escalantii
Aphidius ohioensis
Uroleucon gigantiphagum
Aphidius ohioensis

Uroleucon ivae
Aphidius ohioensis
Aphidius polygonaphis
Aphidius sp.
Diaeretiella rapae
Ephedrus californicus
Praon pequodorum
Praon sp.

Uroleucon nr ivae
Praon nr occidentale

Uroleucon katonkae
Aphidius ohioensis
Ephedrus californicus

Uroleucon oliveri
Aphidius ohioensis

Uroleucon russellae
Aphidius ohioensis
Aphidius sp.
Binodoxys rhagii
Ephedrus californicus
Praon sp.

Uroleucon sonchi
Aphidius ohioensis
Aphidius sp.

Ephedrus californicus
Praon occidentale
Praon pequodorum
Praon sp.

Uroleucon suzannae
Aphidius ohioensis +
Praon nr occidentale +

Uroleucon zymozionense
Praon occidentale

Uroleucon spp.
Acanthocaudus caudacanthus
Aphidius ervi
Aphidius ohioensis
Aphidius sp.

740

Diaeretiella rapae
Ephedrus californicus
Lysiphlebus testaceipes

Praon sp.

Praon occidentale Aphidius sp.

ACKNOWLEDGMENTS

We express thanks to Washington State
University collaborators W. Turner, G.
Clevenger, J. Rumph, and T. Murray, and
Wenatchee Valley College collaborator R.
Gillespie for assistance in collecting aphids
in the field and rearing parasitoids. We ex-
press thanks to G. Ahlstrand and R. Lech-
leitner for facilitating collections at Mt.
Rainier National Park, and the Yakama In-
dian Reservation for allowing access to
their lands. This work was made possible
with the support of numerous growers on
whose land we frequently collected, and in
part by funds provided by Washington State
University, the Washington Wheat and Bar-
ley Commissions, and the U.S. Department
of Agriculture.

LITERATURE CITED

Gahan, A. B. 1911. Aphidiinae of North America.
Maryland Agricultural Experiment Station Bulle-
tin 152: 147-200.

Huber, J. T. and M. J. Sharkey. 1993. Structure, pp.
13-59. In Goulet, H. & J. T. Huber, eds., Hyme-
noptera of the World: An Identification Guide to
Families. Research Branch, Agriculture Canada,
Ottawa, Ontario, Publication 1894/E, 668 pp.

Johnson, J. W. 1987. A revision of the species of
Praon Haliday in North America north of Mexico
(Hymenoptera: Aphidiidae). Canadian Entomolo-
gist 119: 999-1025.

Liu, Ching-Shen. 1977. Genus Aphidius (Aphididae,
Hym.) of California. Taiwan China, Special Bul-
letin No. 11, 118 pp.

Mackauer, M. and P. Stary. 1967. Ichneumonoidea,
World Aphidiidae, pp. 1-195. Jn Delucchi, V and
G. Remaudiere, eds., Index of Entomophagous In-
sects. Le Francois, Paris.

Marsh, P. M. 1979. Aphidiidae, pp. 295-313. Jn Krom-
bein, K. V., BP D. Hurd, Jr., D. R. Smith, and B.
D. Burks, eds., Catalog of Hymenoptera of Amer-
ica North of Mexico, Vol. 1, Symphyta and Apo-
crita (Parasitica). Smithsonian Institution Press,
Washington, D.C.

Praon unicum

Utamphorophora humboldti

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Wahlgreniella nervata
Aphidius polygonaphis
Ephedrus californicus +
Praon occidentale +

Pike, K. S. and P. Stary. 1995. New species of parasitic
wasps attacking cereal aphids in the Pacific North-
west (Hymenoptera, Braconidae, Aphidiinae).
Journal of the Kansas Entomological Society 68:
408-414.

Pike, K. S., P. Stary, R. Miller, D. Allison, L. Boyd-
ston, G. Graf, and T. Miller. 1996. New species
and host records of aphid parasitoids (Hym.: Bra-
conidae: Aphidiinae) from the Pacific Northwest,
U.S.A. Proceedings of the Entomological Society
of Washington 98: 570-591.

Pike, K. S., P. Stary, G. Graf, D. Allison, L. Boydston,
R. Miller, and T. Miller. 1997. Parasitoids (Hym.,
Braconidae, Aphidiinae) of aphids on big sage-
brush (Artemisia tridentata) and prairie sage (Ar-
temisia ludoviciana) in Washington State. Pro-
ceedings of the Entomological Society of Wash-
ington 99: 143-155.

Pike, K. S., P. Stary, T. Miller, R. Miller, D. Allison,
G. Graf, L. Boydston, and R. Gillespie. 1999.
Ephedrus clavicornis sp. n. (Hymenoptera, Bra-
conidae, Aphidiinae), an aphid parasitoid of the
Pacific Northwest, and key to Nearctic Ephedrus.
Journal of the Kansas Entomological Society 72:
10-16.

Remaudiére, G. and M. Remaudiére 1997. Catalogue
of the World’s Aphididae, Homoptera Aphidoidea.
Institut National de la Recerche Agronomique,
Paris, 473 pp.

Smith, C. E 1944. The Aphidiinae of North America
(Braconidae: Hymenoptera). Ohio State Univer-
sity Contributions in Zoology and Entomology
No. 6, 154 pp.

Stary, P. 1963. A study on the relationship of the My-
zinae and their aphidiid parasites in (Central) Eu-
rope (Homoptera: Aphidoidea, Myzinae; Hyme-
noptera: Aphidiidae). Bolletino del Laboratorio di
Entomologia Agraria “‘Filippo Silvestri” di Portici
21:199-216.

. 1966. Aphid Parasites of Czechoslovakia. W.
Junk, The Hague, 242 pp.

Stary, P. and C. FE Smith. 1976. The Nearctic distri-
bution, host plants, and nomenclature of Monoc-
tonus (Hymenoptera, Aphidiidae). Proceedings of
the Entomological Society of Washington 78:
171-175.

van Achterberg, C. 1989. Revision of the subtribe
Monoctonina Mackauer sensu stricto (Hymenop-
tera: Braconidae: Aphidiinae). Zoologische Me-
dedelingen 63: 1—22.

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 741-746

ARCHIPS GOYERANA, N. SP. (LEPIDOPTERA: TORTRICIDAE) AN
IMPORTANT PEST OF BALDCYPRESS (TAXODIACEAE) IN LOUISIANA
AND MISSISSIPPI

JAMES J. KRUSE

Division of Insect Biology, Department of Environmental Science, Policy and Man-
agement, University of California, Berkeley, CA 94720, U.S.A. (e-mail: kruse @nature.
berkeley.edu)

Abstract.—Archips goyerana, new species, is described and illustrated from southern
Louisiana and southwestern Mississippi. It appears to be host-specific on baldcypress
(Taxodium distichum; Taxodiaceae) and has become a serious pest of that tree species
since its discovery in 1983. Notes on its biology are given, and differences between it
and its sister species, A. argyrospila, are outlined. Although morphological differences
between A. goyerana and A. argyrospila are subtle, they are corroborated by differences

in forewing pattern, larval food plant preferences, and molecular data.

Key words:

Archips (Hiibner) is a large genus of at
least 80 species widely distributed in the
Nearctic, Palaearctic and Oriental regions
(Razowski 1977, 1997). The fruittree leaf-
roller, Archips argyrospila (Walker), is
probably the most widespread of the ap-
proximately 25 species of Archips in North
America. It is a notorious, polyphagous pest
of orchard trees throughout the northern
United States and southern Canada (Chap-
man and Lienk 1971, Smirle 1993, Meeker
and Goyer 1994, Goyer and Chambers
1997). Owing to the presence of numerous
geographic forms and its considerable phe-
notypic variation, MacKay (1962) and Goy-
er et al. (1995) suggested that A. argyros-
pila may consist of a number of different
sibling species that are differentiated by
food plant preference and/or geographical
distribution. In this paper I describe A. goy-
erana, a sibling species that feeds on bald-
cypress (Taxodium distichum L. Rich., Tax-
odiaceae) in Louisiana and Mississippi.
This species has inflicted serious and wide-

Tortricinae, Archipini, Taxodium

spread defoliation in southern Louisiana
since its discovery in 1983 (Braun et al.
1990, Meeker and Goyer 1993, Goyer et al.
1995). Aerial surveys conducted annually
have found as many as 60,000 ha of bald-
cypress forest affected by this insect in
southern Louisiana (Goyer and Chambers
1999):

MATERIALS AND METHODS

Institutions are abbreviated throughout
the text as follows: Essig Museum of En-
tomology (EMEC), University of Califor-
nia, Berkeley, Ca, USA; Louisiana State
Arthropod Museum (LSAM), Baton Rouge,
LA, USA; Mississippi Entomological Mu-
seum (MEM), Mississippi State, MS, USA;
National Museum of Natural History
(NMNH), Smithsonian Institution, Wash-
ington. DC, USA; University of Minnesota,
Saint Paul (UMSP), MN, USA. Specimens
were obtained from the EMEC, MEM, and
Louisiana State University Agricultural
Center (LSUAC), Baton Rouge, LA, USA.

742

Dissection methodology follows that sum-
marized in Brown and Powell (1991) ex-
cept that preparations were transferred to
95% isopropyl alcohol (instead of xylene)
after the 95% EtOH wash, and all parts
were slide-mounted with Euparol mounting
medium (Bioquip, Gardena, CA) rather
than Canada balsam. Forewing measure-
ments were made with an ocular micro-
meter mounted in a Zeiss Stemi SV6 dis-
secting microscope. Terminology for geni-
talic structures follows Horak (1984). Col-
ors were described with the standards of
Kornerup and Wanscher (1983).

SYSTEMATICS

Archips goyerana Kruse, new species
(Figs. 1-2)

Male (Fig. 1a).—Head: Vertex and up-
per frons golden yellow to brownish or-
ange, brown or brownish orange between
antennae in most specimens. Labial palpus
with golden yellow and brownish orange
scaling. Mesonotum: Golden yellow and
brownish orange scaling, dark reddish
brown scaling also present in many speci-
mens; tegulae usually concolorous with me-
sonotum, but often with more brownish or-
ange or reddish brown scaling. Forewing:
Length 6.8-8.5 mm (mean 7.4 mm, n =
37). Costal fold extending from base to near
proximal margin of median fascia, i.e., ca.
30-35% length of FW costa. Upper side
with underscaling gray to grayish red, gray-
ish red concentrated near center, extending
basally and dorsally; overscaling pattern el-
ements include distinct but broken reddish
brown fasciae and black, broken strigulae
and irregular white, yellowish white, gray,
and golden yellow scaling throughout; bas-
al fasciae indistinct; subbasal and median
fasciae distinct and strongly contrasting
with yellowish white interfascial regions
near costa; postmedian fasciae distinct at
costa, breaking dorsally; subterminal and
terminal fasciae indistinct, represented by
reddish brown strigulae or reddish brown
patches, broken by yellowish white and

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

golden yellow scaling. Fringe gray with
reddish brown and/or brownish orange
scales replaced by gray scales at tornus.
Under side with upper side pattern near cos-
ta, becoming gray dorsally in area of
hindwing overlap. Hindwing: Upper side
uniformly gray, except some dark gray and/
or orange scaling near apex in some indi-
viduals. Under side gray, yellowish white
near costa, orange strigulae or dark gray or
orange scaling near apex. Genitalia: As in
Fig. 1b (slide JJK 220; n = 6). Uncus large,
prominent, slightly narrowed from base,
slightly broadened apically, width of apex
generally less than or equal to base (Fig.
lc). Gnathos arising from triangular projec-
tions of the tegumen, gently concavely
curved and fused together apically, extend-
ing slightly beyond coastal margin of valva;
socius vestigal; valva ovate; sacculus grad-
ually broadening terminad with produced
tip arising subapically of valva. Aedeagus
straight or gently curved, terminating in
slender tip strongly curved to left. Phallo-
base gently curved (Fig. 1d).

Female (Fig. 2a).—FW length 8.0—10.0
mm (mean 8.9 mm, n = 50). Superficially
as in male except forewing ground color
gray, with very little grayish red near center
of wing, and generally with less yellowish
white and golden yellow overscaling, but
more black strigulae throughout. Genitalia:
As in Fig. 2b (slide JJK 221; n = 7). Pa-
pillae anales setose, without projections;
sterigma short, produced as a shallow bowl;
antrum ovate; ductus bursae long; signa
long, bladelike. Genitalia are virtually in-
distinguishable from A. argyrospila except
for a more weakly produced sterigma (Fig.
2G).

Types.—Holotype ¢: LOUISIANA: As-
sumption Parish, Pierre Part, 27 April 1999
as pupa (R. A. Goyer), in LSAM. Paraty-
pes: (n = 101). LOUISIANA: Assumption
Parish, Pierre Part, 27 April 1999 as pupae,
14 3, 23 2 (R. A. Goyer); St. Charles Par-
ish, Norco, 27 April 1999 as pupae, 19 6,
42 2 (R. A. Goyer); MISSISSIPPI: Hinds
County, Clinton, 2 2, 18-19 May 1996 (M.

VOLUME 102, NUMBER 3 743

( L a :

Figs. 1-2. Adults and genitalia of Archips goyerana. la, Male of A. goyerana, holotype, Assumption Parish,
Louisiana; scale in mm. 1b, Male of A. goyerana; valvae spread, aedeagus removed, holotype, Assumption
Parish, Louisiana, slide number JJK 220.; scale bar = 1 mm. Ic, Detail of uncus of male A. goyerana (left) and
detail of uncus of male A. argyrospila (right), Harrison County, Long Beach, Mississippi, drawn from slide
number JJK 117. 1d, Detail of aedeagus of male A. goyerana (left) and detail of aedeagus of male A. argyrospila
(right). 2a, Female of A. goyerana, paratype, St. Charles Parish, Louisiana. scale in mm. 2b, Female of A.
goyerana, paratype, St. Charles Parish, Louisiana, slide number JJK 221; scale bar = 1 mm. 2c, Detail of
sterigma of female A. goyerana (left) and detail of sterigma of female A. argyrospila (right), Bossier Parish,
Barksdale Air Force Base, Louisiana, drawn from slide number JJK 214.

and E. Roshore); Harrison County, Long Diagnosis.—Archips goyerana was dis-
Beach, | 2, 13 May 1997 (R. Kergosien). covered in 1983 in Iberville Parish, Loui-
Paratypes are deposited in LSAM (36), | siana (Goyer and Lenhard 1988). It is very
MEM (10), UMSP (5), NMNH (20), and — similar in appearance to A. argyrospila but
EMEC (30). it is far less variable. In A. goyerana the

744

fasciae are usually dark reddish brown, in-
terfascial spots are strongly contrasting with
fasciae, and black striae are usually present.
Interfascial spots are generally less well de-
fined in A. argyrospila, and fasciae typical-
ly consist of various shades of brown or
brownish orange. A pale form of A. goy-
erana is brownish orange on the forewing
and pale brownish orange or light gray on
the hindwing. This form appears to domi-
nate in Mississippi and is difficult to distin-
guish from co-occurring A. argyrospila. In
male genitalia of A. argyrospila the uncus
is distinctly widened terminally, weakly
club shaped (Fig. Ic). In addition, the phal-
lobase in A. argyrospila is typically curved
at an angle greater than 45° from the plane
of the aedeagus as opposed to less than 35°
in A. goyerana (Fig. 1d). Female genitalia
of A. goyerana are nearly indistinguishable
from those of A. argyrospila, although the
sterigma tends to be slightly more robust
and squarish in A. argyrospila, in compar-
ison to the shallow mildly angled bowl of
A. goyerana (Fig. 2c). Molecular data in-
dicates a divergence ranging from 1.47 to
2.53% between A. argyrospila and A. goy-
erana in 475 base pairs of mitochondrial
DNA in the Cytochrome Oxydase I gene,
while widely separated populations of A.
argyrospila differ by 1.26% or less (Kruse
and Sperling, unpublished data).
Distribution and biology.—Archips goy-
erana occurs in southern Louisiana and
southwestern Mississippi (Fig. 3). It origi-
nally may have been endemic to forested
wetlands, but has undergone a population
explosion and expansion of its range since
1983 (Goyer et al. 1990, Zhou et al. 1993,
Goyer et al. 1995). In 1988, A. goyerana
was found for the first time on baldcypress
east of the Mississippi River (in Baton
Rouge) and by 1993 had spread eastward
to the suburbs of New Orleans (Goyer et al.
1995). Noticeable defoliation was evident
Over portions of at least eleven Louisiana
parishes (Meeker and Goyer 1993, 1994).
It was collected in southwestern Mississippi
in 1996 (Clinton) and in 1997 (Long

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Beach). Archips argyrospila and A. goyer-
ana are known to co-occur near Baton
Rouge, Louisiana and Long Beach, Missis-
sippi. Individuals captured away from for-
ested wetlands are speculated to feed on or-
namental baldcypress.

Like A. argyrospila, A. goyerana 1s uni-
voltine throughout its range, with overwin-
tering egg masses attached to the bark of
thin twigs of the host plant in obligate dia-
pause (Braun et al. 1990, Goyer et al. 1990,
Brown 1991, Meeker and Goyer 1993,
Goyer and Chambers 1997). Dormant eggs
hatch by bud break of baldcypress trees
during late February and early March (Goy-
er et al. 1990, Meeker and Goyer 1993,
Goyer and Chambers 1997). First instar lar-
vae disperse and seek out the terminal por-
tions of expanding baldcypress foliage, bur-
row within the cluster of young needles,
and begin feeding inconspicuously (Goyer
and Chambers 1997).

Larvae of A. goyerana feed on the open-
ing leaf buds and elongating leaves (Braun
et al. 1990). As the foliage expands, devel-
oping A. goyerana larvae produce silk to
roll adjacent needles and branchlets into a
tight mass, surrounding themselves individ-
ually and then feeding on the foliage inside
(Braun et al. 1990, Goyer and Chambers
1997). Larvae remain inconspicuous on
baldcypress unless the tree is severely de-
foliated, at which time they become very
active, crawling on branches and spinning
down on silk (Braun et al. 1990). Massive
mortality to mature larvae by drowning in
standing water was observed on many oc-
casions where baldcypress were completely
defoliated (Braun et al. 1990).

Larvae undergo five larval instars before
pupation (Braun et al. 1990). Adults emerge
between late-April and mid-May (Goyer
and Chambers 1997). The development
time for A. argyrospila from egg hatch to
adult emergence takes about 6 weeks
(Braun et al. 1990). For A. goyerana, de-
velopment time is considerably longer,
about 8 to 10 weeks on baldcypress in the
field (Braun et al. 1990, Goyer and Cham-

VOLUME 102, NUMBER 3

745

Fig. 3.
occur are shaded.

bers 1997). Previous studies have shown
that A. goyerana is virtually unable to sur-
vive on oaks, a major food source for near-
by populations of A. argyrospila (Goyer et
al. 1995). Differences in pheromonal re-
sponses between citrus-feeding A. argyros-
pila and baldcypress-feeding A. goyerana
populations suggest evolutionary diver-
gence in mating behavior (Goyer et al.
1995). The forested wetland habitat of A.
goyerana and the distant relationship be-
tween its food plant and the food plants of
A. argyrospila further confirm that the two
are distinct.

Archips goyerana is a wetlands pest in
Louisiana. Repeated defoliation of baldcy-

Range of Archips goyerana in Louisiana and Mississippi. Counties where A. goyerana is known to

press has been observed in several areas of
southern Louisiana, with dramatic reduc-
tions in radial growth and death of portions
of the tree canopy in all age groups, and
mortality primarily in the small, understory
trees that do not recover fully from com-
plete and repeated defoliation (Goyer and
Lenhard 1988, Goyer et al. 1990, Goyer
and Chambers 1997). In addition, baldcy-
press defoliation is closely linked with ex-
cessive flooding, affecting tree successional
patterns in forest wetlands (Goyer and
Chambers 1997). With 70% of Louisiana’s
baldcypress growing stock located within or
near areas now infested with A. goyerana,
the persistence and continued spread of de-

746

foliation poses an additional threat to this
dwindling and unique forest resource
(Meeker and Goyer 1993).

Etymology.—The species is named for
Richard A. Goyer of the Department of En-
tomology, Louisiana Agricultural Experi-
ment Station, Louisiana State University
Agricultural Center, Baton Rouge, Louisi-
ana. He collected the holotype as well as
most of the paratypes, and is responsible for
the majority of the research on the biology
and ecology of the species.

ACKNOWLEDGMENTS

I thank Richard A. Goyer (LSUAC) and
Richard L. Brown (MEM) for the contri-
bution of specimens, and the collectors cit-
ed in the text, who made this study possi-
ble. I thank the following for reviewing and
providing helpful comments on various
drafts of the manuscript: John W. Brown,
Systematic Entomology Laboratory,
USDA, Washington, DC; Jerry A. Powell,
University of California, Berkeley, CA;
Brent D. Mishler, University of California,
Berkeley, CA; and Felix A. H. Sperling,
University of Alberta, Edmonton, Alberta.
This work was supported by a National Sci-
ence Foundation Partnerships in Enhancing
Expertise in Taxonomy (PEET) grant to Fe-
lix A. H. Sperling and Jerry A. Powell.

LITERATURE CITED

Braun, D. M., R. A. Goyer, and G. J. Lenhard. 1990.
Biology and mortality agents of the fruit tree leaf-
roller (Lepidoptera: Tortricidae), on baldcypress in
Louisiana (USA). Journal of Entomological Sci-
ence 25: 176-184.

Brown, J. J. 1991. Diapause, pp. 175—185. Jn vari der
Geest, L. P. S. and H. H. Evanhuis ,eds., Tortricid
Pests: Their Biology, Natural Enemies and Con-
trol. World Crop Pests,. Vol. 5. Elsevier, Holland.
808 pp.

Brown, J. W. and J. A. Powell. 1991. Systematics of
the Chrysoxena group of genera Lepidoptera: Tor-
tricidae: Euliini). University of California Publi-
cations in Entomology 111, 87 pp.

Chapman, P. J. and S. E. Lienk. 1971. Tortricid fauna

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of apples in New York (Lepidoptera: Tortricidae).
New York State Agricultural Experiment Station
Special Publication, 122 pp.

Goyer, R. A. and J. L. Chambers. 1997. Evaluation of
insect defoliation in baldcypress and its relation-
ship to flooding. National Biological Service Bi-
ological Science Report 8, 36 pp.

Goyer, R. A. and G. J. Lenhard. 1988. A new insect
pest threatens baldcypress. Louisiana Agricultural
Experiment Station Bulletin 31: 16—21.

Goyer, R. A., G. J. Lenhard, and J. D. Smith. 1990.
Insect herbivores of a baldcypress-tupelo ecosys-
tem. Forest Ecology and Management 33-34:
517-522.

Goyer, R. A., T. D. Paine, D. P. Pashley, G. J. Lenhard,
J. R. Meeker, and C. C. Hanlon. 1995. Geographic
and host-associated differentiation in the fruittree
leafroller (Lepidoptera: Tortricidae). Annals of the
Entomological Society of America 88: 391-396.

Horak, M. 1984. Assessment of taxonomically signif-
icant structures in Tortricinae (Lep., Tortricidae).
Bulletin de la Société Entomologique Suisse 57:
3-64.

Kornerup, A. and J. H. Wanscher. 1983. Methuen
handbook of colour (3rd ed.). Methuen and Com-
pany, London.

MacKay, M. R. 1962. Larvae of the North American
Tortricinae (Lepidoptera: Tortricidae). Canadian
Entomologist Supplement 28: 1—182.

Meeker, J. R. and R. A. Goyer. 1993. Relationships
between patterns of defoliation by the fruittree
leafroller (Lepidoptera: Tortricidae) and foliage
morphology of baldcypress in forested wetlands
of Louisiana. Journal of Entomological Science
28: 317-326.

. 1994. Fruittree leafroller, Archips argyrospila
(Walker) (Lepidoptera: Tortricidae), performance
as influenced by host foliage type. Journal of En-
tomological Science 29: 1-9.

Razowski, J. 1977. Monograph of the genus Archips
Hiibner (Lepidoptera, Tortricidae). Actazoologica
Cracoviensia 22: 55—206, 223.

. 1997. Generic composition of the New World
Archipini (Lepidoptera, Tortricidae) with descrip-
tion of two new genera and two new species. Mis-
cel-lania Zoologica 20.1: 125-130.

Smirle, M. J. 1993. Larval performance of two leaf-
roller species on known and potential hosts. En-
tomologia Experimentalis et Applicata 67: 223-—
Bil.

Zhou, J., R. A. Goyer, and G. J. Lenhard. 1993. Sea-
sonal occurrence of Calleida viridipennis Say and
Plonchionus timidus Haldeman (Coleoptera: Car-
abidae) predators of the fruit tree leafroller Ar-
chips argyrospila Walker in Louisiana. Journal of
Entomological Science 28: 254—262.

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 747-754

DISCOVERY OF THE SMALL JAPANESE CEDAR LONGHORNED BEETLE,
CALLIDIELLUM RUFIPENNE (MOTSCHULSKY)
(COLEOPTERA: CERAMBYCIDAE), IN LIVE ARBORVITAE
IN CONNECTICUT

CHRIS T. MAIER AND CAROL R. LEMMON

Department of Entomology, Connecticut Agricultural Experiment Station, PO. Box
1106, New Haven, CT 06504, U.S.A. (e-mail: chris.maier@po.state.ct.us)

Abstract.—The small Japanese cedar longhorned beetle, Callidiellum rufipenne (Mot-
schulsky) (Coleoptera: Cerambycidae), was discovered principally in three cultivars of
American arborvitae, Thuja occidentalis L., in four towns in Fairfield and New Haven
Co. in southwestern Connecticut. Between September 1998 and March 1999, infestations
of C. rufipenne were found in 102 (0.5%) of 20,000 cupressaceous plants that were
inspected in garden centers, nurseries, landscaped areas, and the wild in Connecticut.
Within 2 years of planting, 94 (92.2%) of infested plants had been balled and burlaped,
which may have stressed them. Of the 102 plants, 101 (99%) were 0.9-2.1 m high ar-
borvitae that averaged 6.3 beetles/plant, and the other one was yellow cedar, Chamae-
cyparis nootkatensis (D. Don) Sprach. Arrivals of beetles in wood material at ports of
entry, increased international trade and travel, and the high abundance of potential hosts
in coastal North America probably have facilitated introduction and establishment. Adults,

life cycle, and damage of C. rufipenne are described briefly.

Key Words:

Arborvitae, Callidiellum, Cerambycidae, Chamaecyparis, Coleoptera, Con-

necticut, Cupressaceae, Juniperus, larval damage, plant pest, state record,

Thuja occidentalis

The small Japanese cedar longhorned
beetle, Callidiellum rufipenne (Motschul-
sky) (Coleoptera: Cerambycidae), is best
known as a borer of Japanese cedar, Cryp-
tomeria japonica D. Don (Taxodiaceae),
and Japanese cypress or Hinoki false cy-
press, Chamaecyparis obtusa (Siebold &
Zuccarini) Endlicher (Cupressaceae) (Ma-
kihara 1984, Shibata 1994), which are im-
portant trees in Japanese plantations (Ko-
bayashi 1985). Larvae of C. rufipenne,
however, also tunnel into the wood of other
species of Cupressaceae and rarely Pina-
ceae (Bates 1873, Gressitt 1951, Shiraki
1952, Campadelli and Sama 1988, Bahillo
and Iturrondobeitia 1995). Shibata (1994)

and others have suggested that C. rufipenne
attacks only dead or dying wood of its co-
niferous hosts. Other species of Callidiel-
lum, C. cupressi (Van Dyke) and C. vires-
cens Chemsak and Linsley in the western
United States (Chemsak and Powell 1964,
Chemsak and Linsley 1966) and C. flavo-
signatus Pu and C. villosulum (Fairmaire)
in China (Gressitt 1951, Pu 1991), also are
restricted to conifers in the Cupressaceae,
Pinaceae, and Taxodiaceae.

In May 1997, a single adult of C. rufi-
penne was collected on wild eastern red ce-
dar, Juniperus virginiana L., at Manteo,
Dare Co., North Carolina (E.R. Hoebeke, in
litt.). This represented the first record of this

748

eastern Asian beetle in the wild in the Unit-
ed States. After the initial discovery of the
lone adult, K. Ahlstrom (in litt.) reared ad-
ditional adults from larvae collected in the
wood of dead eastern red cedar at Manteo.
Before its recent introduction into Italy
(Campadelli and Sama 1988), Spain (Bah-
illo and Iturrondobeitia 1995), and this
country, C. rufipenne had a distributional
range of Russia (Sakhalin), Japan (includ-
ing the Ryukyu Islands), Korea, Taiwan,
and northeastern China (e.g., Gressitt 1951,
Duffy 1968, Makihara et al. 1989). Gressitt
(1951) suggested that C. rufipenne probably
was introduced into Taiwan. Port inspectors
have repeatedly intercepted this beetle in
dunnage and other wood products shipped
to New Zealand (Bain 1974, 1977), the
United States (Mumford 1965, 1966, 1967;
Girard) 1968; 1969; 119715,.1972a;-01972b,
1973, 1974; USDA 1979, 1980, 1981,
1982, 1984), and other countries.

Here we report the discovery of C. rufi-
penne in Connecticut, document its infes-
tation of live plants, mainly American ar-
borvitae or northern white cedar, Thuja oc-
cidentalis L., and briefly describe its life
history and the appearance of damage. We
initiated this and additional studies because
C. rufipenne may pose a threat to the nurs-
ery industry and to northern forests. Arbor-
vitae are important to the nursery trade be-
cause they are popular ornamental shrubs
grown widely and planted especially near
homes and commercial buildings. In addi-
tion, northern white cedars are an integral
part of some forests in northeastern and
north-central North America.

MATERIALS AND METHODS

Adult records from Connecticut are
based on inspecting plants for larval injury
and then removing or rearing beetles from
plants with damage. We extracted adults
from wood mainly between late September
1998 and February 1999 by finding holes
(the eventual adult exit routes that had been
plugged with chewed wood by larvae) in
branches and then by splitting the wood to

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

reveal the individual adults in their pupal
chambers. Between November 1998 and
March 1999, additional infested arborvitae
(about 80% of ‘all infested plants) were
transported from garden centers or land-
scaped areas, mostly residential yards in
Fairfield and New Haven Co., to two out-
door enclosures with double screening at
Lockwood Farm, Hamden, New Haven
Co., Connecticut. In late March, branches
that appeared to be uninfested were pruned
from these shrubs and burned in accordance
with state regulations. The remaining wood
that had evidence of larval boring was di-
vided into two equal groups. Wood of each
of the shrubs in one lot was placed in a
sealed white plastic drum (18.9 liters) in an
environmental chamber adjusted to 21 +
1°C and to a 15L:9D photoperiod. The re-
mainder of the wood was put into screened
cages of various sizes in one of the outdoor
screened houses. Adults were collected
from drums or cages every 3—7 days until
emergence ended.

Data for infestations of C. rufipenne dis-
covered by federal inspectors at ports of en-
try between 1964 and 1982 (Mumford
1965, 1966, 1967; Girard 1968, 1969, 1971,
1972a, 19726, 1973, 1974; USDASIC7=
1980, 1981, 1982, 1984) were used in Fig.
1. Yearly records were based on the fiscal
year, which changed during the mid-1970’s.
Between 1964 and 1982, C. rufipenne had
a quarantine status of an actionable pest. In
1982, the Animal and Plant Health Inspec-
tion Service (APHIS) changed its quaran-
tine status from an actionable to a non-re-
portable pest (J. Cavey, in litt.), which re-
sulted in a rapid decrease in records of its
interception. In 1998, C. rufipenne again
became an actionable pest.

Voucher specimens are deposited at the
Connecticut Agricultural Experiment Sta-
tion, New Haven; Cornell University, Itha-
ca, New York; National Museum of Natural
History, Smithsonian Institution, Washing-
ton, D.C.; and Essig Museum of Entomol-
ogy, University of California, Berkeley,
California.

VOLUME 102, NUMBER 3

Table 1.

749

Location and characteristics of live arborvitae, Thuja occidentalis, infested by the small Japanese

cedar longhorned beetle, Callidiellum rufipenne, in Connecticut during 1998.

Location of Infested Plants No. Beetles

County Town Variety of Arborvitae Height (m) No. Infested Mean Range
Fairfield Greenwich Smaragd 1.8—2.1 7 4.4 2-9
Fairfield Greenwich/Stamford! Smaragd 1.5-1.8 4 10.3 2-27
Fairfield Stamford Brandon 1.8—2.1 2 4.0 3-5
Fairfield Stamford Smaragd 1.5-1.8 26 Hitt 2-37
Fairfield Stamford/Greenwich! Smaragd 1.5-1.8 16 S55) 2-11
New Haven Milford Nigra 1.5-1.8 2 2.0
New Haven Milford Smaragd 0.9-1.5 8 IBS 1-3
New Haven Milford Smaragd 1.2-1.8 Si 4.5 1-16
New Haven North Haven Smaragd 1.5-1.8 5 3.4 1-9

'in 1998, these arborvitae initially were at the garden center in the town listed first, and then, usually in
spring of the same year, they were planted in yards in the town listed second.

RESULTS AND DISCUSSION

Infestations in Connecticut.—On 22 Sep-
tember 1998, one adult of C. rufipenne was
found in a branch of a emerald green ar-
borvitae, T. occidentalis ‘Smaragd’, planted
in spring 1998 near a small factory in Mil-
ford, New Haven Co. Subsequent inspec-
tion of the planting of 42 arborvitae at this
site revealed 7 live plants (16.7% of the to-
tal) with injury from larval boring. The 42
arborvitae planted by the factory were pur-
chased at a nearby garden center that also
had infested arborvitae that had not been
sold. This find represents the second North
American record of this beetle outside a
port of entry, and the first from a live ar-
borvitae that visually appeared to be
healthy. In 1967, an inspector at a port in
the United States did intercept C. rufipenne
in dunnage of a Thuja species on a cargo
ship (Girard 1968).

Between autumn 1998 and spring 1999,
state inspectors found 102 (0.5% of total)
infested plants among 20,000 cupressa-
ceous shrubs or trees examined in garden
centers, nursery fields, landscaped areas,
and the wild throughout Connecticut. Plants
infested with C. rufipenne were located
mainly in four garden centers and in nearby
residential areas with arborvitae that re-
cently had been purchased from the garden
centers (see Table 1 for the locations). Of

102 infested plants, 100 (98.0%) had been
imported from western North America, and
94 (92.2%) had been balled and burlaped
within 2 years of inspection for C. rufipen-
ne. Thorough examination of nursery fields
at the points of origin in British Columbia
(J. Bell and B. Gill, in litt.) and Oregon (R.
Westcott, in litt.) and inspection of ship-
ments from the western North America in
1999 did not reveal any damage or speci-
mens of C. rufipenne. The plants with C.
rufipenne apparently became infested in
Connecticut; however, the origin and time
of introduction are unknown.

Of the 102 infested plants, 101 (99.0%)
were arborvitae (Table 1), and one was a
yellow cedar, Chamaecyparis nootkatensis
(D. Don) Spach. Among the 101 infested
arborvitae of three cultivars, 97 (96.0%)
were the variety ‘Smaragd’, which current-
ly is the most widely grown and planted
arborvitae in Connecticut. Infested arbor-
vitae had between 1 and 37 beetles, and
ranged in height from 0.9 to 2.1 m. In all,
632 beetles developed in the wood of the
101 arborvitae (mean = 6.3 live beetles/
plant), and 6 in that of yellow cedar. An
unknown number of arborvitae that may
have been infested by C. rufipenne at the
four garden centers were sold and probably
planted in southwestern Connecticut.

Even though most infested plants that

750

100

No. Interceptions

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

64 66 68 70 72 74 76 78 80 82

Fiscal Year

Figaale

Frequency of interceptions of the small Japanese cedar longhorned beetle, Callidiellum rufipenne, at

ports of entry in the United States between 1964 and 1982. The change in the fiscal year in the early 1970's
apparently caused some inconsistencies in compilation of data between 1973 and 1975.

were balled and burlaped appeared healthy,
they may have been susceptible to attack by
C. rufipenne because they were stressed by
root and branch breakage and possibly in-
adequate watering before, during, and after
shipment to Connecticut. Furthermore,
these arborvitae occasionally were infested
by bark beetles (Scolytidae), such as
Phloeosinus canadensis Swaine and Poly-
graphus rufipennis (Kirby).

Factors contributing to accidental intro-
duction and establishment.—The small Jap-
anese cedar longhorned beetle has repeat-
edly been intercepted by federal and state
inspectors at ports of entry in the United
States (Mumford 1965, 1966, 1967; Girard
19685 1969, 197 13 197 2a 19726; 6 1973;
1974; “OSDA 11979, 1980) 19811982,
1984; R. Penrose, in litt.). In most cases,
the intercepted beetles were associated with

dunnage or other wooden material from
eastern Asia. In Vancouver, British Colum-
bia, in 1927, C. rufipenne was reared from
imported wood of Japanese origin (Leech
1949). Hatch (1971) recorded another spec-
imen from Seattle, Washington, but this
beetle, too, was likely associated with an
imported wood product.

Between 1964 and 1982, when federal
inspectors actively reported infestations of
C. rufipenne, they repeatedly discovered
them in imported wood material (Fig. 1).
During these years, infestations were found
in dead wood, especially dunnage, in 21
states in the United States and in Puerto
Rico (Mumford 1965, 1966, 1967; Girard
1968, 1969, 1971; 1972a,. 1972b; 799723
1974; USDA 1979, 1980; 198171982;
1984). Although C. rufipenne was not dis-
covered at ports of entry in Connecticut, it

VOLUME 102, NUMBER 3

was found in ports of the nearby states of
New Jersey, New York, and Pennsylvania.
In coastal counties in Connecticut and near-
by states, potential hosts abound in residen-
tial areas and forests. In particular, over-
mature, often dead and dying trees of J. vir-
giniana are numerous in successional areas
along major highways and in regrowth for-
ests. We suggest that the frequency of ar-
rivals of C. rufipenne in imported wood, in-
creased international trade and travel, and
the high abundance of potential hosts near
coastal areas probably have facilitated the
accidental introduction and establishment
of C. rufipenne in the United States.

Brief description of adult appearance, life
history, and damage.—Males are iridescent
deep blue with a single brownish red to red
patch at the anterolateral corner of the up-
per surface of each elytron. The females
(Fig. 2) have entirely reddish brown elytra
and a reddish abdomen. The antennae of
males are longer than the body, whereas
those of females are shorter, about % to %
the length of the body. Thus, the sexes can
be separated by both color and antennal
length. When they are reared from arbor-
vitae, adults vary from 5—13 mm in length.

The life history of C. rufipenne (Shibata
1994, in litt.; Maier 1999; Y. Soma, in litt.)
is similar to that of other species of Calli-
diellum (Chemsak and Powell 1964, Nak-
amura and Kojima 1981). Adults of C. ru-
fipenne emerge from wood in spring and
soon mate on the host. Females of C. rufi-
penne then lay eggs singly or in small
groups in cracks or crevices in stems or
bark. After hatching, larvae bore into the
cambium and phloem. They expand their
feeding tunnel until they become full-grown
(Fig. 3) in late summer. After feeding is
completed, larvae bore into the sapwood
and carve an ellipsoidal pupal cell. The pu-
pal chamber is connected to the surface of
a branch by an exit tunnel (Fig. 4) that is
oval in cross-section. Larvae plug the even-
tual exit route with fragments of chewed
wood. They pupate in late summer or early
fall, and adults eclose within 2—3 weeks,

ial

usually in the autumn. Adults remain in the
pupal cell until the following spring when
they remove the plug to the pupal cell and
emerge. In Connecticut, C. rufipenne has
one generation per year. In Japan, the life
cycle also usually requires 1 year. In some
northern regions of Japan, however, it may
require 2 years to complete the life cycle
(Y. Soma, in litt.).

In live arborvitae, injury from larval bor-
ing becomes most obvious between late
summer and the following spring when the
bark of small arborvitae often splits to re-
veal the sinuous larval tunnels (Fig. 5). The
larval gallery is packed with frass, which
apparently is also mixed with chewed wood
fragments. In some cases, these tunnels may
encircle branches and, thereby, disrupt the
flow of water and nutrients within the plant.
The material packed in the larval gallery
has both light and dark particles, giving it
a “‘salt-and-pepper”’ look.

ACKNOWLEDGMENTS

We thank Xavier Asbridge, Joan Bravo,
Dora Brown, Jeffrey Fengler, Verryn Jen-
nings, Tracy LaProvidenza, Roland Leval-
lee, Gale Ridge-O’Connor, Stephen San-
drey, Michael Thomas, and Peter Trenchard
for assistance in inspecting plants and rear-
ing beetles. We especially want to acknowl-
edge Gale Ridge-O’Connor, who recog-
nized that the beetle was not native to Con-
necticut. The screened buildings for storing
infested plants were constructed by Richard
Cecarelli, Jeffrey Fengler, Rollin Hannan,
and Edward Naughton. Adults of C. rufi-
penne were identified by John A. Chemsak,
University of California, Berkeley, E. Ri-
chard Hoebeke, Cornell University, Ithaca,
and Steven A. Lingafelter, USDA System-
atic Entomology Laboratory, Washington,
D.C. Donald E. Bright, Jr., Research
Branch, Agriculture and Agri-Food Canada,
Ottawa, Ontario, identified the scolytids.
Kenneth Ahlstrom, North Carolina Depart-
ment of Agriculture, Raleigh, Jon Bell, Ca-
nadian Food Inspection Agency, New West-
minster, British Columbia, Joseph Cavey,

752 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 2-5. Small Japanese cedar longhorned beetle, Callidiellum rufipenne. 2, Adult female, 12 mm in length.
3, Full-grown larva exposed after removal of bark of arborvitae, 13 mm in length. 4, Larva in its ellipsoidal
pupal cell; note the pupal plug of chewed wood fragments. 5, Branch injury caused by larval boring. Arrows
show frass and chewed wood exposed after the bark splits in late summer.

VOLUME 102, NUMBER 3

USDA, APHIS, PPQ, Riverdale, Maryland,
Bruce Gill, Canadian Food Inspection
Agency, Ottawa, Ontario, E. Richard Hoe-
beke, Richard Penrose, California Depart-
ment of Food and Agriculture, Sacramento,
Eiichi Shibata, Nagoya University, Japan,
Yukihiro Soma, Yokohama Plant Protection
Station, Yokohama, Japan, and Richard
Westcott, Oregon Department of Agricul-
ture, Salem, provided valuable information
about C. rufipenne or surveys for it. Robert
Tracy, USDA, APHIS, PPQ, Elizabeth,
New Jersey, kindly provided access to pub-
lications that could not easily be obtained
elsewhere. Finally, we appreciate the co-
operation of Patricia Douglass and Wayne
Petrie, USDA, APHIS, PPQ, Wallingford,
Connecticut.

LITERATURE CITED

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. 1977. Overseas wood- and bark-boring insects
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Chemsak, J. A. and J. A. Powell. 1964. Observations
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Duffy, E. A. J. 1968. A Monograph of the Immature
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753

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through June 30, 1967). United States Department
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Plant Quarantine Division, Hyattsville, Maryland.
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. 1969. List of intercepted plant pests, 1968

(pests recorded from July 1, 1967, through June

30. 1968). United States Department of Agricul-

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antine Division, Hyattsville, Maryland. ARS 82-

6-3, 86 pp.

. 1971. List of intercepted plant pests, 1969

(pests recorded from July 1, 1968, through June

30, 1969). United States Department of Agricul-

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antine Division, Hyattsville, Maryland. ARS 82-

6-4, 77 pp.

. 1972a. List of intercepted plant pests, 1970

(pests recorded from July 1, 1969, through June

30, 1970). United States Department of Agricul-

ture, Animal and Plant Health Inspection Service,

Hyattsville, Maryland. APHIS 82-1, 85 pp.

. 1972b. Addendum to list of intercepted plant

pests, 1970 (pests recorded from July 1, 1969,

through June 30, 1970). United States Department

of Agriculture, Animal and Plant Health Inspec-

tion Service, Hyattsville, Maryland. APHIS 82-1

(we suspect that this issue should have been la-

beled 82—2), 13 pp.

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(pests recorded from July 1, 1970, through June

30, 1971). United States Department of Agricul-

ture, Animal and Plant Health Inspection Service,

Hyattsville, Maryland. APHIS 82-3, 81 pp.

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1-662.

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injuring insect damage in Japanese cedar and cy-
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Maier, C. T. 1999. Beetle thought to attack only dead

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wood found in live arborvitae in Connecticut.
Frontiers of Plant Science 51(2): 4—5.

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Nakamura. 1989. A list of cerambycid beetles
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(pests recorded from July 1, 1964, through June

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PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 755-756

NOTE

Ectoparasitic Insects from Migrating Saw-Whet
Owls (Aegolius acadicus) in Central Wisconsin

A total of 833 saw-whet owls (Aegolius
acadicus Gmelin) were trapped, providing
an opportunity to survey these birds for ec-
toparasitic insects. This is the largest sur-
vey of ectoparasitic insects from this bird
ever conducted during migration and
should provide baseline data for future in-
vestigators.

Mist nets were used with a tape-recorded
conspecific call to trap migrating saw-whet
owls at Linwood Springs Research Station
(44°28'’N, 89°40’W) in central Wisconsin
during their fall and spring nocturnal mi-
grations (IX-23 to XI-08-1997, II-18 to III-
26-1998, IX-28 to XI-11-1998, and II-20 to
III-27-1999). The birds were placed in tem-
porary holding compartments for less than
two hours. Of the 833 trapped owls, 644
were checked for ectoparasitic insects by
hand during banding activities. One hun-
dred and sixty-nine of the 644 trapped owls
were randomly selected (1/5 of the sample)
and extensively examined for a 2-minute
period. Collected insects were fixed in 70%
ethanol, dehydrated through an ethanol se-
ries to xylene, and mounted in Canada Bal-
sam. Voucher specimens are deposited in
the National Museum of Natural History,
Smithsonian Institution, Washington, DC,
accession number TM2014200.

Sixty-four of 644 (9.9%) owls searched
during banding and 19/169 (11.2%) when
examined for 2 min. were positive for ec-
toparasitic insects. Hippoboscids (costa
americana Leach, Ornithoica vicina Walk-
er, Ornithomyia fringillina Curtis) were har-
bored by 1.7% of the birds, Mallophaga
(Kurodaia acadicae Price and Beer, Strigi-
philus acadicus Emerson and Price) by
5.8%, and Siphonaptera (Cediopsylla sim-

plex Baker, Orchopeas leucopus Baker) by
3.8%. Table 1 shows the prevalence of sin-
gle and multiple infestations during fall and
spring migrations of owls.

Except for hippoboscids and a single
specimen of C. simplex (new host record),
all ectoparasites were collected during both
fall and spring. The three hippoboscids
were only collected during the fall (Table
1). As part of a larger study on Wisconsin
hippoboscids during 11 autumns and 6
springs, Mueller et al. (1969. Transactions
of the Wisconsin Academy of Sciences,
Arts and Letter 57: 189—207) collected two
O. vicina and two O. fringillina from 234
saw-whet owls during migration. These au-
thors also noted that hippoboscids are un-
common or absent during the spring.

Orchopeas leucopus was collected from
31 birds. Holland (1985. Memoirs of the
Entomological Society of Canada, No. 130,
pp. 130, 631) reported O. leucopus on 4
saw-whet owls from Ontario, Canada.
Specimens of this flea most likely trans-
ferred to the owls from rodents, while our
sole specimen of C. simplex may have
come from a rabbit. Rodents and rabbits are
natural hosts for these fleas. Forty-eight
birds harbored K. acadicae and S. acadicus.
The relatively low numbers of fleas and lice
may be explained as follows: Fleas are in-
termittent parasites on mammalian or avian
hosts, and, in the case of raptors, may be
accidental. Mallophagans complete their en-
tire life cycle on their host but are extremely
small, attach themselves firmly to feathers,
and are sometimes difficult to detect on liv-
ing birds. Additionally, lice may be present
only as nits during bird migration (Dogiel.
1964. In General Parasitology. Oliver and
Boyd, Edinburgh and London 516 pp.).

756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Number of single and multiple infestations and prevalence of ectoparasitic insects from saw-whet
owls.
Fall Spring Fall Spring Parasite
Ectoparasites 1997 1998 1998 1999 Totals
Diptera
Hippoboscidae
Icosta americana 2/644 = 0.31% 0/644 = 0.00% 0/644 = 0.00% 0/644 = 0.00% 2
Ornithoica vicina 6/644 = 0.93% 0/644 = 0.00% 0/644 = 0.00% 0/644 = 0.00% 6
Ornithomyia fringillina 2/644 = 0.31% 0/644 = 0.00% 3/644 = 0.47% 0/644 = 0.00% 5
O. vicina, O. fringillina 1/644 = 0.16% 0/644 = 0.00% 0/644 = 0.00% 0/644 = 0.00% 1
Mallophaga
Menoponidae
Kurodaia acadicae 3/644 = 0.47% 2/644 = 0.31% 9/644 = 1.40% 0/644 = 0.00% 14
Philopteridae
Strigiphilus acadicus 10/644 = 1.55% 3/644 = 0.47% 7/644 = 1.09% 0/644 = 0.00% 20
K. acadicae, S. acadicus 1/644 = 0.16% 1/644 = 0.16% 1/644 = 0.16% 0/644 = 0.00% 3
Siphonaptera
Pulicidae
Cediopsylla simplex 0/644 = 0.00% 1/644 = 0.16% 0/644 = 0.00% 0/644 = 0.00% 1
Ceratophyllidae
Orchopeas leucopus 2/644 = 0.31% 8/644 = 1.24% 4/644 = 0.62% 6/644 = 0.93% 20
Multispecies Infestations
O. leucopus, K. acadicae 2/644 = 0.31% 2/644 = 0.31% 0/644 = 0.00% 0/644 = 0.00% 4
O. leucopus, S. acadicus 0/644 = 0.00% 3/644 = 0.47% 0/644 = 0.00% 3/644 = 0.47% 6
O. leucopus, K. acadicae, 1/644 = 0.16% 0/644 = 0.00% 0/644 = 0.00% 0/644 = 0.00% 1
S. acadicus
Seasonal Totals 30 20 24 9 83

Stephen J. Taft, Corenna D. Kerstner, De-
partment of Biology, University of Wiscon-
sin-Stevens Point, Stevens Point, WI 54481,
U.S.A. (e-mail: staft@uwsp.edu), and Eu-

gene A. Jacobs, Linwood Springs Research
Station, 1601 Brown Deer Lane, Stevens
Point, Wisconsin 54481, U.S.A.

VOLUME 102, NUMBER 3

dvi)

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 757-758

NOTE

First Record of the California Pine Needle Aphid, Essigella (Essigella)
californica (Essig) (Homoptera: Aphididae: Lachninae), in Southern Brazil

The genus Essigella del Guercio 1909 is
the only native Nearctic representative of
the subtribe Eulachnina (Sorensen. 1994.
The Pan-Pacific Entomologist 70: 1—102).
The genus has recently been revised and in-
cludes three subgenera, 13 species, and two
subspecies, all of which are linear-bodied
and feed on needles of Pinaceae, mainly Pi-
nus, but also Pseudotsuga and Picea (Sor-
ensen 1994; Remaudiere and Remaudiére.
1997. Catalogue of the World’s Aphididae.
Institut National de la Recherché Agrono-
mique, Paris, 473 pp.). Essigella is close to
the native Palearctic Eulachnus del Guer-
cio, 1909, and to Pseudessigella Hille Ris
Lambers, 1966 (Sorensen. 1990. Annals of
the Entomological Society of America 83:
394—408). The genus has bifid tarsal claws;
a sclerotized dorsum; head and pronotum
fused; meso- and metanotum fused dorsal-
ly; abdominal tergite I usually free from the
metanotum; and abdominal tergites HI-VII
fused (Sorensen 1994).

Essigella (Essigella) californica (Essig
1909) is a lime green, small-sized (1.5—2.0
mm) aphid found in western North America
from southern British Columbia and Alber-
ta to southern Mexico (Sorensen 1994;
Blackman and Eastop. 1994. Aphids on the
World’s Trees—An Identification and Infor-
mation Guide. CAB International and The
Natural History Museum, London, 987 pp.
+ 16 pls.). One confirmed record from Mi-
ami, Florida, suggests that it may occur in
the Caribbean and have a pan-Mexican dis-
tribution (Sorensen 1994). This species has
been recently introduced into Europe. In
France, it has been found causing damage
on several Pinus species (Turpeau and Re-
maudiére 1990, as cited by Sorensen 1994).

In Spain, it has been captured by suction
pan traps (Seco Fernandez and Mier Duarte.
1992. Boletin de la Asociacion Espanhola
de Entomologia 16: 255—256).

In Brazil, E. californica was first col-
lected by early June 1999 on slash pine Pi-
nus elliotti Engelm. in Rolandia, Parana
State (23°19'S, 51°22'W, altitude about 540
m a.s.l.) and on Mexican weeping pine Pi-
nus patula Schiede & Deppe, in the Agron-
omy Campus of the Universidade Federal
do Paranda, Curitiba, Parand (25°25’S,
49°14’W, altitude 945 m as.l.) (R. C. Z.
Carvalho, collector). Both localities repre-
sent urban areas where the plants are used
for landscaping. In October 1999, E. cali-
fornica was collected at a farm in Corbélia,
Parana (24°45’'S, 53°20'W, altitude 750 m
a.s.l.), on P. elliotti (S. M. N. Lazzari, col-
lector).

Alate and apterous viviparae and nymphs
were found on the branch tips of isolated
plants, feeding on the base of the pine nee-
dles, and moving quite rapidly when dis-
turbed. Colonies of E. californica on P. pa-
tula from Curitiba were small and associ-
ated with Cinara pinivora (Wilson 1919)
(Lachninae: Cinarini), while the popula-
tions from Rolandia and Corbélia were
more numerous, only on a few trees, and
were not associated with other aphid spe-
CIES:

According to J. Sorensen (in litt.), E. cali-
fornica is quite variable geographically, oc-
curs on various hosts, and might be a com-
plex. The one collected from Brazil is in the
same phena as those which have been taken
from Spain, France, Australia, and New Zea-
land. It seems to be the same that occurs in
Mendocino County, along the northern coast

758

of California, principally on Pinus attenuata
Lemmon and Pinus muricata D. Don. These
pines are in the subsection Oocarpae as is
the Mexican weeping pine, which is closely
related to and hybridizes well with slash
pine in the Australis subsection. In Europe
and Australia, E. californica occurs on Pinus
radiata D. Don, another Oocarpae pine. In
California, this aphid can be found on P.
radiata, but they are less robust and much
less common than other Essigella species.
Sorensen also mentions that E. californica
doesn’t do much damage, if any, on pines in
New Zealand.

Another Eulachnina that has been record-
ed from Brazil feeding on Pinus, Eulachnus
rileyi (Williams 1911) (Eastop, Costa, and
Blackman. 1993. Pesquisa Agropecuaria
Brasileira 28: 269-280) may be confused
with E. californica, but they can be distin-
guished by the following characters: E. ri-
leyi has 6-segmented antennae, claws with-
out bifid tips, color in life varying from
dark olive green to gray, with a dusting of
bluish-gray wax; E. californica has 5-seg-
mented antennae, tarsal claws incised with
double tips, and lime green color in life.

Slide-mounted specimens of E. califor-
nica are deposited in the Pe. Jesus S. Moure

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Entomological Collection, Departamento de
Zoologia, Universidade Federal do Parana
(DZUP). One sample of P. elliotti from Ro-
landia is deposited under the number 8599
and one of P. patula under the number
8598 in the Herbarium of the Forestry De-
partment (EFC) of the Universidade Federal
do Parana, Brazil.

We are thankful to Dr. John Sorensen,
Plant Pest Diagnostics Center, California
Department of Food & Agriculture, for con-
firming the aphid identification and for his
valuable comments; to Dr. David Voegtlin,
Illinois Natural History Survey, for his
valuable suggestions; to Dr. Jarbas Yukio
Shimizu, EMBRAPA/Centro Nacional de
Pesquisa de Florestas; and to Dr. Carlos
Vellozo Roderjan, Forestry School of the
Universidade Federal do Parana for the
identification of the pine species.

Regina Célia Zonta de Carvalho, Centro
de Diagnostico Marcos Enrietti—SEAB,
Rua Jaime Baldo 575, 80040-340, Curitiba,
Paranda, Brasil (e-mail: regcarva@zaz.
com.br) and Sonia Maria Noemberg Laz-
zari, Departamento de Zoologia, Universi-
dade Federal do Paranda, Caixa Postal
19020, 81531-990, Curitiba, Parana, Brasil
(e-mail: lazzari@ bio.ufpr.br).

VOLUME 102, NUMBER 3

759

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 759-760

NOTE

New Distributional and First Specific Host-Plant Records for Thionia acuta Doering
and 7. producta Van Duzee (Auchenorrhyncha: Fulgoroidea: Issidae)

The mostly Neotropical issid genus
Thionia Stal (ca. 72 spp.) includes eight
species in America north of Mexico
(Wheeler and Wilson. 1987. Journal of the
New York Entomological Society 95: 440—
451). Information on host plants of North
American Thionia species and descriptions
of their nymphal stages have been pub-
lished mainly in the last few years (Wheeler
and Wilson 1987; 1988. Journal of the New
York Entomological Society 96: 266—273;
Wheeler. 1996. Proceedings of the Ento-
mological Society of Washington 98: 374—
375). Wilson et al. (1994. pp. 7-113 In
Denno and Perfect, eds., Planthoppers:
Their Ecology and Management, Chapman
and Hall, New York) provided a list of re-
corded host plants of the Fulgoroidea. Here-
in, I provide new state records and the first
host-plant associations for the seldom-col-
lected T. acuta Doering, as well as the first
specific host records for the morphological-
ly similar T. producta Van Duzee.

Thionia acuta Doering

Doering (1939. University of Kansas Sci-
ence Bulletin 25: 447—575[1938]) described
this issid under the name 7. naso Fowler,
relying on the identification of E. D. Ball,
who compared Doering’s specimens from
Concan, Tex., with Fowler’s holotype of T.
naso from Jalapa, Mexico. Once Doering
had access to the holotype of T. naso, she
realized the Texas material represented a
new species, which she named acuta (Doer-
ing. 1941. University of Kansas Science
Bulletin 27:185—233) and described by bib-
liographic reference to her previous paper
(Doering 1939). No records of 7. acuta
have been published since Doering’s (1939)

description, and host relationships have re-
mained unknown.

In 1999, I collected nymphs and adults
of 7. acuta in Kansas and Oklahoma on ju-
niper (Juniperus spp.; Cupressaceae) in na-
tive stands and landscape plantings. Vouch-
er specimens of 7. acuta (and T. producta
from Arizona) have been deposited in the
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM), and the S.W. Wilson Collection,
Central Missouri State University, Warrens-
burg.

Collection records.—KANSAS: Riley
Co., Kansas State University, Manhattan
(39°11'39"N, 96°34'57’W; elev. 330 m), 22
June 1999, ex Juniperus virginiana trees
(96, 162, 3 5" instars). OKLAHOMA:
Cleveland Co., University of Oklahoma,
Norman (35°12’26"N, 97°26'54’W; elev.
292 m), 12 June 1999, ex Juniperus chi-
nensis, J. virginiana (11d, 122, 3 5™ in-
stars); Garvin Co., unnamed road SW of
Rts. 7 & 35, W of Davis (34°29'32’N,
97°11'46"W; elev. 252 m), 13 June 1999,
ex Juniperus virginiana (16, 62, 1 5 in-
star); Woodward Co., Rt. 34, 3.7 km N of
Vici (36°10'45"N, 99°19'55”W, elev. 690
m), 25 June 1999 (24, 72).

Thionia producta Van Duzee

Described from Colorado by Van Duzee
(1908. Proceedings of the Academy of Nat-
ural Sciences of Philadelphia 59:467-—
498[1907]), T. producta has since been re-
corded from Arizona, Texas, and Utah
(Doering 1939; Metcalf. 1958. General Cat-
alogue of the Homoptera, Fasc. IV, Part 15
Issidae, Waverly Press, Baltimore). The
host plant, based on R. H. Beamer’s col-
lecting at Leakey, Tex., on 8 July 1936, is

760

“cedar” (Doering 1939). My collections of
T. producta in Arizona’s Huachuca Moun-
tains confirm cedar (Juniperus) as a host,
specifically alligator juniper, J. deppeana
Steudel.

Collection records.—ARIZONA: Co-
chise Co., Ash Canyon Rd. nr. Twin Oaks
Rds (0:4) iki W of (REG92 NG 123: 3IN,
110°14.1'W; elev. 1540 m), 4 June 1997, ex
Juniperus deppeana (96, 7¢; 1 3", 1 4",
and 13 5" instars); Miller Canyon Rd., 1.2
km Wor Rt 925Gil2seNe tO" l'S.2' W;
elev. 1525 m), 5 June 1997, ex Juniperus
deppeana (66, 12; 13 5" instars).

Discussion.—Kansas and Oklahoma are
new state records for T. acuta, known pre-
viously only from Concan, Tex. The Kansas
collection extends the range of this plant-
hopper northeast by more than 1,100 km.
In addition, collections of nymphs and
adults from native stands of Juniperus vir-
giniana and in landscape plantings from J.
virginiana trees (10-15 m high) and a J.
chinensis hedge (ca. 10 m long) represent
the first published host associations for T.
acuta. One specimen from the series that
P.W. Oman collected at the type locality in
Texas bears a “‘juniper”’ label, but host data
were not mentioned by Doering (1939,
1941). Collections of 7. producta nymphs
from alligator juniper in the Huachuca
Mountains of Arizona represent the first
specific host records for a species known
previously only from “‘cedar.”’

In describing 7. acuta (erroneously as T.
naso), Doering (1939) noted its resem-
blance to the juniper-feeding 7. producta.
Because male genitalia of both species are
similar—she alluded to minor differences—
she did not describe those of T. acuta, but
referred to the genitalic description under T.
producta in the same publication. Although
Doering (1939) acknowledged that some
taxonomists might consider the two issids
varieties of a single species, she felt it less

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

confusing if they were recognized as sepa-
rate species.

Doering’s (1939) hesitancy in recogniz-
ing T. acuta and T. producta as distinct spe-
cies might have been influenced not only
by the morphological similarity of these
planthoppers, but also by her awareness that
both species had been collected on juniper
at Concan, Tex. (assuming she saw the sin-
gle specimen of T. acuta that bears host in-
formation). Only in the case of 7. producta,
however, did she publish a host association
(Doering 1939).

Adults of 7. acuta and T. producta, al-
though similar, differ morphologically more
than those of many cryptic or sibling spe-
cies. The two species can be distinguished
by length of the vertex (see Doering 1939:
plate 51), even accounting for the intraspe-
cific variation that she noted in T. producta,
and by total body length with wings in re-
pose (T. acuta = 6.8—8.4 mm, T. producta
= 5.5-6.8 mm). These morphologically
similar, juniper-feeding issids, nymphs of
which are both green, might prove to be
sister taxa when Thionia is analyzed cladis-
tically.

I thank Thomas J. Henry (Systematic En-
tomology Laboratory, c/o National Muse-
um of Natural History, Washington, D.C.)
for companionship in the field, for helping
collect issids, for comparing specimens of
T. acuta collected in 1999 with determined
material in the USNM, and for calling my
attention to host information on a specimen
of T. acuta; and Stephen W. Wilson (De-
partment of Biology, Central Missouri State
University, Warrensburg) for verifying (and
in some cases correcting) my preliminary
sorting of Thionia nymphs to instar, and for
verifying the identification of both Thionia
species.

A. G. Wheeler, Jr., Department of Ento-
mology, Clemson University, Clemson,
South Carolina 29634, U.S.A. (e-mail:
awhlr@clemson.edu)

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 761-767

OBITUARY
A TRIBUTE TO TED SPILMAN

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Theodore James Spilman

(1925-1996)

762

All who knew Theodore James Spilman
will remember his earnest and varied inter-
ests in many subjects, from U.S. presidents
to music, in addition to his thorough com-
mand of all things entomological. Ted was
fascinated by the lives of entomologists as
much as by entomology itself. As a devoted
member of the Entomological Society of
Washington, serving over the years in sev-
eral offices including President, he had an
unofficial role as the Society’s historian. On
the occasion of the one-thousandth meeting
of the ESW in February 1995, he told sto-
ries of the early meetings and formation of
the ESW, complete with century-old “gos-
sip’’ and detailed knowledge of the lives of
some founding members, leaving those of
us present wanting to hear much more. This
was however to be Ted’s last address to the
membership, as an ongoing battle with can-
cer ended his productive life in September
1996. But Ted’s recollections and writings
on the growth of the ESW certainly still
give its present members, and those to
come, a sense of their organization’s unique
history and purpose. As fellow ESW mem-
bers representing three generations of en-
tomologists who knew and worked with
Ted, we offer some personal remembrances
of him, followed by a list of his published
contributions to the science and history of
entomology.

REMINISCENCES OF TED SPILMAN
by John M. Kingsolver

Theodore J. Spilman died September 22,
1996 at 71 years of age. He and I worked
together for 23 years in our positions with
the U.S. Department of Agriculture in
Washington, D.C., and were the closest of
friends. My first meeting with Ted was in
March, 1960, while I was on a type-study
trip to eastern museums. He and Rose Ella
Spilman (then Warner) helped me find the
type specimens I needed to see. By 1962,
when I joined the U.S. Department of Ag-
riculture staff at the National Museum, Ted
and Rose Ella were married. The three of

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

us were part of a group of seven USDA and
two Smithsonian Coleopterists with offices
in the National Museum, although ento-
mological offices and the insect collection
were then housed temporarily in a building
on Lamont St. north of the main museum.
We were a congenial group although our
respective institutional duties were some-
what divergent. The USDA “‘crew”’ includ-
ed Donald Anderson, George Vogt, Ted,
Rose Ella and myself, whereas the SI group
consisted of Oscar Cartwright and Paul
Spangler. Later, Robert Gordon and Richard
White joined the USDA group and Terry
Erwin the SI group.

Of course, I picked brains to learn pro-
cedures, and everyone on both staffs was
helpful. Ted and I became near “‘brothers”’
after we learned that he was one day older
than I. This led to joint birthday celebra-
tions in March for as long as we were as-
sociated. Several years later, I discovered
that one of my ancestors was named Spil-
man. Although Ted tried desperately, he
could never connect our family lines.

Ted was well liked by everyone. He had
a thorough knowledge of the darkling bee-
tle family, the click beetles, and several oth-
er smaller coleopterous families. He didn’t
publish many large papers but seemed fated
to put out small emergency fires that con-
stantly cropped up in agricultural entomol-
ogy. His papers were meticulous, practical,
and useful. He was well-read in the arts and
the classics, and had a considerable library
at home.

Ted was very professional, usually wear-
ing a lab coat or jacket and tie whereas
some of the others of us dressed more ca-
sually. He was especially at home with vis-
itors to the museum, helping them to find
specimens in the collection, and making
certain that they had everything they need-
ed during their visit.

For several reasons, Ted and Rose Ella
and my wife and I seldom socialized. We
had children in school, my wife worked
evenings in a hospital, and opportunities for
visiting were scarce. One series of social

VOLUME 102, NUMBER 3

gatherings thrived for several years in the
1960’s. Several couples and singles from
both the USDA and Smithsonian staffs
formed a loosely organized group to meet
and read plays. Parts would be assigned
ahead of time, and we would meet in a
member’s home where the plays would be
read but not acted out. Refreshments would
be furnished by the hosts to round out the
evening.

Ted often came to work at the museum
after his retirement in 1985 although the
last few years of his life he was seriously
ill. I last spoke with him shortly before he
died.

R. J. Gagne (1997, American Entomol-
ogist 43 (3):191—192) wrote a fine obituary
for Ted and listed his vital statistics and
many accomplishments, especially of his
association with the Entomological Society
of Washington.

TED SPILMAN, FRIEND
by Ross H. Arnett, Jr.

I first met Ted when I returned to Cornell
for a short visit soon after I went to work
at the U.S. National Museum. V. S. L. Pate
told me that a friend of his had a former
student at the University of Louisville, now
registered at Cornell, who wanted to do
graduate work on beetle taxonomy. Pate
wanted me to meet him. When I was in
Comstock Hall that morning, Ted was in
class, so I didn’t get to meet him there. I
drove to College Town and was parking the
car to get some lunch when this young fel-
low ran up to the car and said: “‘You are
Ross Arnett?” ‘Yes.’ “I am Ted Spilman
and I want to work on Pythidae!”’

Thus began a close friendship that lasted
the rest of our lives. We were in close touch
both entomologically and personally in so
many ways. When I left the Museum to
teach, Ted took over my job, and the groups
of beetles I was working on. As I wrote the
“beetle book’’ Ted helped in so many un-
acknowledged ways. He looked up things
for me. He found specimens for me. He

763

criticized (most valuable). As everyone
knows who has tried to do a big piece of
work, without friends of the caliber of Ted,
they never would complete their project.

When I moved back to Washington to
teach I had no place to live until we sold
our house in Rochester. Ted put me up in
his apartment (this was before the days of
Rose Ella). I slept on a “‘hide-a-bed”’ in the
living room. My head was near a fish tank.
During the night the fish clicked their gills
(courtship sounds?). Ted didn’t believe me.
I don’t know if he ever found out about his
talking fish.

We sold our house up north. Ted and I
went house hunting in Bladensburg. One
day coming back from such a trip, Ted had
an accident with his Studebaker. No one
was hurt, but the front of the car was a
mess. Ted never got it fixed. He drove it for
quite awhile, and then traded it off. Ted and
I picked out a house which my wife Mary
and the children never saw until the day we
moved in.

Ted took the job in Washington before he
finished his doctoral thesis and took his
orals at Cornell. I suggested that he finish
his degree at Catholic University of Amer-
ica under my direction. He did register for
a semester as a graduate student, but he was
much too busy at the museum to bother
with the trivial academic hurdles on a cam-
pus across town, so finishing the degree
was always put aside for something more
important—his research. Therefore, he re-
mained ‘‘Mr.”’ Spilman, but in our minds
he was “Doctor.”

Then came Rose Ella Warner. Ted and
Rosa Ella were made for each other. It was
my distinct honor and pleasure to be Ted’s
best man at their wedding. Nothing between
close friends can ever be as wonderful as
having this role in the ceremony of the
Holy Matrimony. I offered the toast the best
man is supposed to do, not very elegant, I
am afraid, but I wished them a happy mar-
ried life, and it was to the very last.

No matter where I moved, Ted was al-
ways back there at the USNM (entomolog-

764

ical Mecca) willing to answer questions,
send specimens, and help in every way I
asked. He called me a few days before he
died to say good-bye. I said “‘Good-bye,
dear friend.”

TED SPILMAN’S CONTRIBUTIONS TO
ENTOMOLOGY

by Warren E. Steiner, Jr.

The wide range of entomological topics
that attracted Ted Spilman’s interest, paired
with his cordial willingness to share his
findings with others, made him a great re-
source to the research community at all lev-
els. In the late 1970s when I was at the
National Museum as a student and contract
technician, I first got to know Ted and
found him to be a living encyclopedia. Be-
ing able to tap this irreplaceable source of
information, then simply chat about topics
of mutual interest, was then a real help to
a beginning student, and is now truly
missed.

I often relied on Ted for a quick identi-
fication, or help in finding references on a
particular beetle or natural history topic.
Because so many unrelated beetle taxa were
the subjects of his research over the years,
primarily the tenebrionoid families but also
Ptinidae, Bostrichidae, Elateridae, Ceram-
bycidae, and the odd little “‘jumping shore
beetles” (intertidal Limnichidae), asking
Ted was much faster (and more entertain-
ing) than searching the library.

Fieldwork was not a major part of Ted’s
entomological career, but he did travel to
Dominica for about 3 months in 1964 to
participate in the Bredin-Archbold-Smith-
sonian biological survey of the island,
where he made some significant collections.
Most of his collecting and work on rearing
of larval beetles was centered in the eastern
U.S. where he helped show me that, when
it came to knowledge of beetles, the back
yard was frontier. He and I made at least
two local field trips in the early 1980s in
search of new finds—we picked at polypore
fungi along the Potomac, sorted through

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

sand of Delaware dunes, salvaged drowned
specimens from the beach drift line, and
had a good time trading trivia. But we
turned back at Assateague Island when, to
our dismay, we unknowingly had chosen
the day of the annual wild pony roundup
for our visit. Glamorous megafauna had
overwhelmed entomology once more, but
my adaptable travel partner was equally
happy to find a camp along the Pocomoke,
where the mosquitoes are at least smaller
than those at the beach.

Ted’s research products are a complex
quilt of subject matter. Without hesitation
he can be called a coleopterist. Fossil bee-
tles, beetles of medical importance, cave
beetles, morphology of beetle genitalia,
beetles in mammal nests, introduction of
foreign beetles, beetle nomenclature, rear-
ing the larvae and pupae of beetles, the
lives (and even grave sites) of people who
studied beetles, were among his topics ad-
dressed. His curatorial and identification re-
sponsibilities covered several of the largest
families of beetles and many unrelated
smaller groups. He liked island faunas and
was keeping and building separate survey
collections of West Indian and Hawaiian
Tenebrionidae. He seemed to enjoy solving
complex nomenclatural problems, and in
several papers, delved into the definition of
terms and usage of names. The odd and
atypical attracted his attention—he even
named a beetle species “‘extraordinaria.”’

The diversity of subjects was equally
broad when it came to the books that Ted
reviewed. These included systematic revi-
sions, catalogs, identification manuals,
treatments of biology, morphology, and ag-
ricultural importance of insects, and biog-
raphies of entomologists. His reviews are
still useful and amusing to read—he talks
frankly and informally about any shortcom-
ings and yet gives the book a good sell,
often with humorous flair. It is obvious that
he enjoyed reading and commenting on the
writings of others, including works by col-
leagues, and he sometimes published the
same review in two or more places.

VOLUME 102, NUMBER 3

Ted was a long-time member of the Co-
leopterists’ Society and served as Editor of
the Coleopterists’ Bulletin for five years,
succeeding Ross Arnett in 1962. With his
writing style and acute command of taxo-
nomic principles, terms and literature, he
was an excellent and dependable reviewer
of manuscripts. Alas, production of some
taxonomic works surely must have been set
back because of his loyal service to others
in many ways and his constant attention to
many duties on the job.

A lot of Ted’s research pursuits went un-
finished, but he leaves behind a wealth of
information for the next students of these
special topics, in the form of literature col-
lections and notes, card files, manuscripts
in progress, copies of correspondence with
colleagues, and identified specimens. His
sixty published works (not including book
reviews) listed below are only the tip of a
great iceberg of accomplishments—Ted’s
separate files on research topics, grouped by
subject and by taxon, will be most useful
and unique products of his labors. These
will perpetuate his contributions to the sci-
ence, as will the professional example he
set for all of us and the generations to fol-
low.

Publications by T. J. Spilman

1952a. The male genitalia of the Nearctic Salpingidae.
Coleopt. Bull. 6(1): 9-13.

1952b. Dragonfly oviposition. The Kentucky Natural-
ist 6(1): 15

1953a. An odd case of gynandromorphism in the ex-
ternal genitalia of Boros discicollis (Salpingidae).
Coleopt. Bull. 7(5): 41—44.

1953b. Corrections to Leng’s catalogue and Gebien’s
Coleopterorum Catalogus: Tenebrionidae and Ka-
talog der Tenebrioniden. J. New York Ent. Soc.
61: 215-216.

1954. Generic names of the Salpingidae and their type
species (Coleoptera). J. Washington Acad. Sci.
44(3): 85-94.

1958. The transfer of Anamphidora from the Tene-
brionidae to the Alleculidae (Coleoptera). Proc.
Ent. Soc. Washington 60(6): 288.

1959a. Notes on Edrotes, Leichenum, Palorus, Eup-
sophulus, Adelium, and Strongylium (Tenebrioni-
dae). Coleopt. Bull. 13: 58—64.

1959b. A study of the Thaumastodinae, with one new

765

genus and two new species (Limnichidae). Co-
leopt. Bull. 13: 111-122.

1960a. The tenebrionid Diaclina fagi as medicine in
the Orient. Coleopt. Bull. 14(1): 12.

1960b. Ptinus variegatus Rossi, new to the United
States (Coleoptera: Ptinidae). Ent.
Washington 62(2): 103-104.

1960c. Some synonymy in Oryzaephilus (Coleoptera:
Cucujidae). Proc. Ent. Soc. Washington 62(4):
Daile

1961a. Remarks on the classification and nomenclature
of the American tenebrionine genus Adelonia (Co-
leoptera: Tenebrionidae). Pan-Pac. Ent. 37(1): 49—
Syl.

1961b. On the immature stages of the Ptilodactylidae
(Coleoptera). Ent. News 72(5): 105-107.

1961c. (with W. H. Anderson). On the immature stages
of North American Pyrochroidae (Coleoptera).
Coleopt. Bull. 15(2): 38—40.

1961d. A few tenebrionids new to Cuba. Coleopt. Bull.
15(4): 127.

196le. Uloma extraordinaria, a new species from
Cuba (Tenebrionidae). Coleopt. Bull. 15(4): 113—
Hits).

1962a. A few rearrangements in the Tenebrionidae,
with a key to the genera of the Ulomini and Te-
nebrionini of America, north of Mexico (Coleop-
tera). Coleopt. Bull. 16: 57—63.

1962b. The New World genus Centronopus, with new
generic synonymy and a new species (Coleoptera:
Tenebrionidae). Trans. Amer. Ent. Soc. 88: 1—19.

1963a. The American genus Mycotrogus: A synopsis,
a new species from Cuba, and a note on a larva
(Coleoptera: Tenebrionidae). Proc. Ent. Soc.
Washington 65(1): 21—30.

1963b. On larvae, probably Tauroceras, from the Neo-
tropics (Coleoptera: Tenebrionidae). Coleopt.
Bull. 17: 58-64.

1965. Ptinus variegatus in Georgia and Kentucky (Co-
leoptera: Ptinidae). Proc. Ent. Soc. Washington
67(4): 230.

1966a. On the generic names Alphitobius, Phtora, Cla-
moris, and Cataphronetis (Coleoptera: Tenebrion-
idae). Proc. Ent. Soc. Washington 68(1): 6—10.

1966b. Larva and pupa of Amarygmus morio from Ha-
waii (Coleoptera: Tenebrionidae). Proc. Hawaiian
Ent. Soc. 19(2): 297-301.

1966c. A new species of Martinius from Cuba (Cole-
optera: Limnichidae). Coleopt. Bull. 20(4): 123-
5).

1967a. A new North American Ulomine genus and
species, Doliodesmus charlesi (Coleoptera: Tene-
brionidae). Pan-Pac. Ent. 43(2): 149-154.

1967b. Gmelin’s 13th Edition of the Systema Naturae:
A case of neglect. Ent. News 78(7): 169-172.

1967c. The heteromerous intertidal beetles (Coleop-
tera: Salpingidae: Aegialitinae). Pac. Ins. 9(1): 1—
2

Proc. Soc.

766 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1968. Two new species of Niptus from North Ameri-
can caves (Coleoptera: Ptinidae). Southw. Nat.
13(2): 193-200.

1969. Larva and pupa of Pyanisia tristis from Ala-
bama (Coleoptera: Tenebrionidae). Coleopt. Bull.
23(3): 57-61.

1970. A new species of Tribolium from North Amer-
ica. The results of past confusion. U.S. Dept. Agr.
Coop. Econ. Ins. Rpt. 20(24): 396.

1971a. Bredin-Archbold-Smithsonian Biological Sur-
vey of Dominica: Bostrichidae, Inopeplidae, La-
griidae, Lyctidae, Lymexylonidae, Melandryidae,
Monommidae, Rhipiceridae, and Rhipiphoridae
(Coleoptera). Smithsonian Contr. Zool. 70: 1—10.

1971b. The longitudinal lines on beetle elytra: A def-
inition of stria. Coleopt. Bull. 25(4): 121-122.

1972a. Fossil Stichtoptychus and Cryptorama in Mex-
ican amber (Coleoptera: Anobiidae). Univ. Calif.
Publs. Ent. 63: 87-89.

1972b. A new genus and species of jumping shore
beetle from Mexico (Coleoptera: Limnichidae).
Pan-Pac. Ent. 40(2): 108-115.

1972c. Uloma Dejean, 1821 (Insecta: Coleoptera): Pro-
posed designation of a type-species under the ple-
nary powers. Z.N.(S) 1979. Bull. Zool. Nomencl.
29(1): 32-34.

1972d. An amendment to the proposal concerning-
Uloma (Insecta: Coleoptera). Z.N.(S) 1979. Bull.
Zool. Nomencl. 29(4): 194-195.

1973a. A list of the Tenebrionidae of Michigan (Co-
leoptera). Great Lakes Ent. 6(3): 85-91.

1973b. Nomenclatural problems in six genera of Te-
nebrionidae (Coleoptera). Proc. Ent. Soc. Wash-
ington 75(1): 39—44.

1973c. (with C. A. Triplehorn). A review of Strongy-
lium of America north of Mexico, with descrip-
tions of two new species (Coleoptera: Tenebrion-
idae). Trans. Amer. Ent. Soc. 99(1): 1—27.

1974. Temnocerus and Heterolabus, a new synonymy
(Coleoptera: Curculionidae). Proc. Ent. Soc.
Washington 76(3): 246.

1976a. A new species of fossil Ptinus from fossil wood
rat nests in California and Arizona (Coleoptera:
Ptinidae), with a postscript on the definition of a
fossil. Coleopt. Bull. 30(3): 239-244.

1976b. (Witn J. D. Solomon and R. E. Doolittle). Cer-
ambycid beetles captured in sticky-traps in Mis-
sissippi. Coleopt. Bull. 30(3): 289-290.

1978a. Lagria villosa in Brazil, with new descriptions
and illustrations of the larva and pupa (Coleop-
tera: Lagriidae). Ciencia e Cultura 30(3): 342—
347.

1978b. Descriptions of the larva, pupa, and adult of
Alegoria dilatata, associated with banana plants
in the Neotropics (Coleoptera: Tenebrionidae).
U.S. Dept. Agr. Coop. Plant Pest Rpt. 3(6): 47-51.

1978c. Pests not known to occur in the United States
or of limited distribution: Lined click beetle, Agri-

otes lineatus (L.); a wireworm, Agriotes obscurus
(L.); a wireworm, Agriotes sputator (L.). U.S.
Dept. Agr. Coop. Plant Pest Rpt. 3(48—52): 731-734.

1978d. (with R. E. Woodruff and E. J. Gerberg). A false
powderpost beetle new to the United States (Co-
leoptera: Bostrichidae). Fla. Dept. Agr. Cons. Serv.
Div. Plant Ind., Entomology Circular 195, 2 pp.

1979a. (with E. J. Ford, Jr.). Biology and immature
stages of Dirrhagofarsus lewisi, a species new to
the United States (Coleoptera: Eucnemidae). Co-
leopt. Bull. 33(1): 75-83.

1979b. Larvae and pupae of Centronopus calcaratus
and Centronopus supressus (Coleoptera: Tene-
brionidae), with an essay on wing development in
pupae. Proc. Ent. Soc. Washington 81(4): 513-521.

1980. (with P. G. Crook and J. A. Novak). The lesser
mealworm, Alphitobius diaperinus, in the scrotum
of Rattus norvegicus, with notes on other verte-
brate associations (Coleoptera, Tenebrionidae; Ro-
dentia, Muridae). Coleopt. Bull. 34(4): 393-396.

1982. False powderpost beetles of the genus Dinoderus
in North America (Coleoptera: Bostrichidae). Co-
leopt. Bull. 36(2): 193-196.

1983. (with V. E Wright). An annotated bibliography
on Prostephanus truncatus (Horn), (Coleoptera:
Bostrichidae): A pest of stored grain. Trop. Stored
Prod. Inf. 46: 25—30.

1984a. Vignettes of 100 years of the Entomological
Society of Washington. Proc. Ent. Soc. Washing-
ton 86(1): 1-10.

1984b. Identification of larvae and pupae of the larger
grain borer, Prostephanus truncatus (Coleoptera,
Bostrichidae), and the larger black flour beetle,
Cynaeus angustus (Coleoptera: Tenebrionidae).
Proc. Third Intl. Working Conf. Stored-Product
Ent. 44-53.

1985. Review of Lanelater of North America (Cole-
optera: Elateridae). Insecta Mundi 1(1):1—12.
1989a. A visit to the tomb of Thomas Say in New
Harmony, Indiana. Ent. News 100(5): 224—227.

1989b. Vignettes of the Presidents of the Entomolog-
ical Society of America, 1889-1989. Bull. Ent.
Soc. Amer. 35(3): 33-65.

1991a. (with J. E Lawrence). Tenebrionidae (Tene-
brionoidea) (including Alleculidae, Cossyphodi-
dae, Lagriidae, Nilionidae, Rhysopaussidae, Ten-
tyriidae), pp. 520-528. Jn Stehr, E W., ed., Im-
mature Insects. Kendall-Hunt, Dubuque, Iowa.

1991b. (In Gorham, J. R., Insect and mite pests in
food) Part 1.6. Spider beetles (Ptinidae, Coleop-
tera). U.S. Dept. Agr. Handbook 655(1): 137-147.

1991c. (In Gorham, J. R., Insect and mite pests in
food) Part 1.11. Darkling beetles (Tenebrionidae,
Coleoptera). U.S. Dept. Agr. Handbook 655(1):
185-214.

1995. (with R. L. Aalbu and K. W. Brown). The sys-
tematic status of Amblycyphrus asperatus, Thren-
us niger, Pycnomorpha californica, Emmenastus

VOLUME 102, NUMBER 3

rugosus, and Biomorphus tuberculatus Motschul-
sky (Coleoptera: Tenebrionidae). Proc. Ent. Soc.
Washington 97(3): 481—488.

ACKNOWLEDGMENTS

For discussions and comments on Ted’s
“life history” and reviews of early drafts
of this remembrance, we thank D. M. An-
derson, R. J. Gagné, Stuart E. Neff, Paul
and Phyllis Spangler.

767

Warren E. Steiner, Jr., John M. Kingsol-
ver, and Ross H. Arnett, Jr. (WES) Depart-
ment of Entomology, NHB-187, Smithsoni-
an Institution, Washington, DC 20560,
U.S.A.; (JMK) Florida State Collection of
Arthropods, P.O. Box 147100, Gainesville,
FL 32614, U.S.A.; (RHA), 2406 NW 47th
Terrace, Gainesville, FL 32606, U.S.A. (de-
ceased).

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 768-770

Book REVIEW

The Genus Rhipicephalus (Acari, [xodi-
dae): A Guide to the Brown Ticks of the
World. Jane B. Walker, James E. Keirans
and Ivan G. Horak. Cambridge Univer-
sity Press, The Edinburgh Building,
Cambridge CB2 2RU, United Kingdom.
Kit (643 4pp..7.27 9ehies/ de tabs:, 74
maps. 2000. $100.00/cloth. ISBN 0-521-
48008-6. www.cup.cam.ac.uk.

Through nearly two centuries of study,
no tick genus has proved as taxonomically
intractable as Rhipicephalus. Particularly in
tropical Africa, exclusive home to 60 of the
world’s 74 known species, misidentifica-
tions of rhipicephalids have rendered sus-
pect numerous otherwise valuable veteri-
nary and parasitological papers. Now, how-
ever, as if in tribute to the new millennium,
acarology’s own Gordian knot lies sundered
by the peerless team of Drs. Walker (On-
derstepoort Veterinary Institute, South Af-
rica), Keirans (U.S. National Tick Collec-
tion, Georgia Southern University, States-
boro) and Horak (Faculty of Veterinary Sci-
ence, University of Pretoria, South Africa),
whose decades-long synergy has yielded
the most comprehensive—and attractive—
volume ever published on a single genus of
ticks, the sixth in a series of monographs
on the Ixodoidea begun by George Henry
Falkiner Nuttall (1862-1937) and col-
leagues in 1908 and championed from the
outset by Cambridge University Press.

Introductory sections of this sumptuous
work provide a definition of Rhipicephalus
in the context of the family Ixodidae (sec-
tion 2), augmented by illustrations of key
characters (section 3) and a glossary (sec-
tion 4) of all morphological, taxonomic and
other terms that are associated with mem-
bers of this genus (some of these, like “‘sha-
greened” (p. 19), are uniquely rhipicephalid
descriptors). There follows (section 5) a
worldwide annotated checklist of all the
Rhipicephalus species names ever pub-

lished, including junior synonyms, new
combinations and lapsus calamorum, to-
gether with full citations for original de-
scriptions, and collection data and deposi-
tories for types and type series. The book
then neatly divides itself in two: sections 6,
7 and 8 describe the rich rhipicephalid fau-
na of the Afrotropical Zoogeographic Re-
gion; sections 9, 10 and 11 address the 10
Rhipicephalus species that occur outside the
Afrotropics. As noted by the authors, this
dichotomy does not work for all taxa. In-
cluded in their Afrotropical discussion is
the circumglobal R. sanguineus, the world’s
most widespread tick species and the only
representative of its genus in the Western
Hemisphere, as well as R. fulvus, most col-
lections of which have come from Tunisia
(i.e., the Palearctic Zoogeographic Region),
and R. camicasi and R. turanicus, with vast
ranges in Africa and adjacent lands. Also,
the trans-African subspecies R. e. evertsi
apparently has been recently introduced by
humans into Yemen and the Red Sea coast
of Saudi Arabia. But for the most part, the
Rhipicephalus faunas of Africa and the rest
of the world do not intersect, and the pre-
liminary discussions of each (sections 6 and
9) comprise fascinating historical reviews
of regional tick research, exhaustive lists of
country-specific references, and the au-
thors’ long-anticipated male and female
identification keys, whose user-friendliness
is immeasurably enhanced by inclusion of
information on preferred hosts and distri-
butions within the key couplets themselves.

At the core of this effort are the individ-
ual species accounts (sections 7 and 10),
each wholly self-contained, which collec-
tively constitute over three quarters of the
text. In these accounts, thorough descriptive
diagnoses of all known active stages are ac-
companied by stunning scanning electron
photomicrographs, conventional light mi-
crographs of slide-mounted female genital

VOLUME 102, NUMBER 3

apertures a la Feldman-Muhsam (1956,
Bull. Res. Council Israel 5B: 300—306),
and, for both sexes, the heartbreakingly
beautiful pen-and-ink drawings of A. Ol-
wage, Walker’s long-time artistic collabo-
rator. Olwage’s preeminence as a tick illus-
trator gained world renown with the publi-
cation of his series of color plates depicting
the adults of Amblyomma species capable
of transmitting the agent of heartwater,
Cowdria ruminantium (1987, Onderste-
poort J. Vet. Res. 54: 353-379). In the opus
at hand, his skill is perhaps best exemplified
in drawings of the ornate R. pulchellus,
whose specific epithet means “beautiful”
(p. 366), and the densely punctate male of
R. ramachandrai (p. 562). Olwage also pro-
duced the final versions of each tick distri-
bution map, wherein data as diverse as type
locality Gf known), and confirmed and un-
confirmed records of various authors are
precisely but harmoniously plotted (excel-
lent examples are the distribution of R. guil-
honi, p. 207, and R. sulcatus, p. 431). Given
the biomedical importance of this genus, it
is surprising how many diagnoses are in-
complete for at least one life history stage.
While males and females are known for all
species, both nymphs and larvae are un-
known for the Afrotropical R. aquatilis, R.
bequaerti, R. bergeoni, R. boueti, R. deltoi-
deus, R. dux, R. interventus, R. longiceps,
R. masseyi, R. oreotragi (one of two new
species described in this book; the other is
R. warburtoni, known from all stages), R.
punctatus, R. serranoi and R. supertritus. In
addition, the nymph of R. longicoxatus is
unknown, as is the larva of R. pseudolon-
gus. Preimagines of R. complanatus and R.
ziemanni exist in collections but have not
been described. Outside Africa, the imma-
tures of R. scalpturatus are unknown, as
well as the larva of R. leporis, while the
larva of R. pusillus was not available for
study.

Each species diagnosis closes with one or
more paragraphs of critically important
“notes on identification,” in which the au-
thors offer insights for separating similar

769

species, often discerning helpful differences
in minor morphological expressions, such
as those accompanying engorgement, or in
tick ecology, distribution, and host specific-
ity. This discussion leads seamlessly to a
tabulation of all known hosts of each tick
species, these data having been gleaned
from large and small tick collections world-
wide and from the literature (hosts are en-
tirely unknown for R. deltoideus and R.
scalpturatus). Statements on tick zoogeog-
raphy, incorporating altitudinal data, rain-
fall and vegetation zones, and often ex-
plaining or clarifying the map records,
round out each account, together with a
summary of disease relationships (only 27
Rhipicephalus species (36%) have ever
been linked to any human or animal dis-
ease, several transmitting only in the labo-
ratory to splenectomized hosts) and a list of
species-specific references. At the close of
both the African and non-African species
accounts, the tick/host tables of the ac-
counts are combined and inverted, yielding
checklists of all Rhipicephalus species col-
lected from each host animal (sections 8
and 11). In this reversed arrangement, tick
names in boldface generally indicate that
the animal under which they are listed is a
preferred host of the adults. However, in the
Afrotropical R. follis, R. gertrudae and R.
simus, the hosts of immatures (usually
small rodents) are entirely unrelated to the
hosts of adults (usually ungulates), so these
three tick species appear twice at opposite
ends of the checklist. With additional field
collections of immatures, this bimodal pat-
tern of parasitism is likely to apply to many
more rhipicephalids. Dual entries also ap-
pear for the immatures of the Afrotropical
R. pravus, R. sp. near pravus and R. war-
burtoni, which are often found on lago-
morphs (like most African Rhipicephalus)
but chiefly parasitize elephant shrews.
Section 12, ““Species groups based on the
immature stages,”’ is a first attempt to help
taxonomists identify the maddeningly sim-
ilar immatures of this genus by sorting
known nymphs and larvae into 10 more or

770

less recognizable groups (8 Afrotropical, 2
non-Afrotropical) based on diagnostic ca-
pitular characters, again admirably drawn
by Olwage. Brief descriptions of these char-
acters are provided for each group and, as
in the adult key, summaries of each species’
geographic distribution assist in clinching
identifications. The authors take pains to
note that these groups may not be “natu-
ral,’ although the immatures of the Afro-
tropical R. evertsi group—R. bursa, R. ev-
ertsi and R. glabroscutatum—are both mor-
phologically similar and the only species in
their genus known to have a two-host, rath-
er than three-host, life cycle. As well, spe-
cies in the ‘“‘pointy-palp”’ groups of nymphs
and larvae (1.e., the Afrotropical R. capen-
sis, R. follis and R. simus groups), which
are exceedingly difficult to identify, seem
to share the habit of feeding on specific
small mammals that are not parasitized by
the adult stages.

To describe in any detail the variety of
diseases associated with members of the ge-
nus Rhipicephalus, and the research that has
been conducted in this sphere, would ne-
cessitate doubling the present volume. This
is, after all, a book about ticks: how to iden-
tify them, how they live, and where and
when they may be found. But just as the
authors employ host/tick checklists to sum-
marize the discussions of host relationships
in the individual species accounts, in their
final section (13) they leave disease-orient-
ed readers with two multipage, landscape-
format tables on the transmission of diseas-
es to animals and humans. Both are superb
synopses. The animal table lists 25 Rhipi-
cephalus-borne diseases and 28 causative
agents, plus two afflictions (tick paralysis
and tick toxicosis), with additional headers
for the animals affected, the tick vectors
(including, if known, the primary natural
vectors and any that have transmitted ex-
perimentally), the number of hosts utilized
by each tick species (in Rhipicephalus gen-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

erally 3, sometimes 2), the tick stages that
acquire and transmit disease, and one or
more classic references (an appended bib-
liography provides 80 additional reviews of
animal diseases vectored by rhipicephalid
ticks). The table for tickborne diseases of
humans is similar, listing 15 diseases and
causative agents, with further headers for
tick vectors, number of hosts, the mode of
infection or isolation of the causative
agents, and classic references (the appended
bibliography for this table provides 33 ad-
ditional reviews).

Even at the dawn of the 21st century,
books of this length and complexity are cer-
tain to contain at least a few minor typo-
graphical errors. Accordingly, the publish-
ers have agreed to issue an erratum slip,
which currently makes the following emen-
dations: the correct date of publication for
the new species R. oreotragi (p. 31) and R.
warburtoni (p. 38) is 2000, the date 1999
being an unfortunate carryover from pub-
lication delays; the Rocky Mountain Lab-
oratory is in Montana, a state that should
be abbreviated MT, not MA (Massachu-
setts) (p. 45); and under the “notes on iden-
tification’”’ for R. serranoi (p. 408), “XX”
and ‘“‘YY”’ should be replaced by male and
female symbols. Such minutiae aside, this
authoritative and beautiful book is the best
beginning we could wish for acarology’s
third century. No one who visits its pages
will fail to find instruction or inspiration.
Surely somewhere Cecil Warburton (1854—
1958), whose meticulous studies of Rhipi-
cephalus intraspecific variation made him
the obvious choice for this work’s dedica-
tory frontispiece, is smiling serenely at his
reward: an infinite number of ticks and an
eternity in which to examine them.

Richard G. Robbins, Armed Forces Pest
Management Board, Walter Reed Army
Medical Center, Washington, DC 20307-
5001, U.S.A. (E-mail: robbinrg @acq.osd.
mil)

PROC. ENTOMOL. SOC. WASH.
102(3), 2000, pp. 771-772

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CONTENTS

(Continued from front cover)

MATHIS, WAYNE N. and JAMES F. EDMISTON—A revision of the shore-fly genus Lemnaphila
Cresson (Diptera: Ephydridae)

PAPP, JENO and SCOTT R. SHAW—A study of the genus Falcosyntretus Tobias from the
New World with five new species and a key to known species (Hymenoptera: Braconidae:
Euphorinae)

PIKE, K. S., P. STARY, T. MILLER, G. GRAF, D. ALLISON, L. BOYDSTON, and R. MILLER—
Aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) of northwest USA

POLAVARAPU, S., JOHN A. DAVIDSON, and DOUGLASS R. MILLER—Life history of the
Putnam scale, Diaspidiotus ancylus (Putnam) (Hemiptera: Coccoidea: Diaspididae) on blue-
berries (Vaccinium corymbosum, Ericaceae) in New Jersey, with a world list of scale insects
on blueberries

SITES, ROBERT W. and MICHAEL R. WILLIG—Morphometric variation among populations of
Ambrysus mormon Montandon (Heteroptera: Naucoridae)

VASU, V., DAVID R. SMITH, and MALKIAT S. SAINI—Review of the Asian sawfly genus
Anisoarthra Cameron (Hymenoptera: Tenthredinidae)

NOTES

TAFT, STEPHEN J., CORENNA D. KERSTNER, and EUGENE A. JACOBS—Ectoparasitic
insects from migrating saw-whet owls (Aegolius acadicus) in central Wisconsin

WHEELER, A. G., JR.—New distributional and first specific host-plant records for Thionia acuta
Doering and T producta Van Duzee (Auchenorrhyncha: Fulgoroidea: Issidae)

ZONTA DE CARVALHO, REGINA CELIA, and SONIA MARIA NOEMBERG LAZZARI—

First record of the California pine needle aphid, Essigella (Essigella) californica (Essig)
(Homoptera: Aphididae: Lachninae), in southern Brazil

OBITUARY §

STEINER, WARREN E., JR., JOHN M. KINGSOLVER, and ROSS H. ARNETT, JR.—A tribute
to Ted Spilman

BOOK REVIEW

ROBBINS, RICHARD G.—The Genus Rhipicephalus (Acari, Ixodidae): A Guide to the Brown
Ticks of the World, by Jane B. Walker, James E. Keirans, and Ivan G. Horak

MISCELLANEOUS

Instructions for authors

549

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601

| VOL. 102 OCTOBER 2000 NO. 4
: QL. (ISSN 0013-8797)

HG |

© PROCEEDINGS

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ENTOMOLOGICAL SOCIETY

CONTENTS

ALUJA, MARTIN, JAIME PINERO, MAURILIO LOPEZ, CESAR RUIZ, ALBERTO ZUNIGA,
ENRIQUE PIEDRA, FRANCISCO DiAZ-FLEISCHER, and JOHN SIVINSKI—New host
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Gerstacker, Rhagoletis zoqui Bush, Rhagoletis sp., and Hexachaeta sp. (Diptera: Tephritidae) 802

BORKENT, ART—The larva and pupa of Schizonyxhelea forattinii Wirth and Grogan (Diptera:
Ceratopogonidae) with a discussion of the phylogenetic relationships of the genus .......... 862

BRANHAM, MARC A. and MIGUEL ARCHANGELSK Y—Description of the last larval instar
and pupa of Lucidota atra (G. A. Olivier 1790) (Coleoptera: Lampyridae), with a discussion
Gidodomuinal sepment homology across life stages) )) 22.2020. dices we enna voce eee edesedsevse 869

BROWN, JOHN W. and JON LEWIS—Catalogue of the type specimens of Tortricidae (Lepidop-
tera) in the collection of the National Museum of Natural History, Smithsonian Institution,

WTSS HR EEAWOPeS ID OMS Se GS a ea As eee Se eRe a DL SAGY 4 eR apa RTRSY MR ed AP Ue ete A at 1014
CONTRERAS-RAMOS, ATILANO—A new species of Chloronia Banks (Megaloptera: Corydal-

idae) from southeastern Brazil, with a key to the species of Brazil .......................4.. 919
DARSIE, RICHARD FE, JR.—Description of the pupa of Armigeres (Leicesteria) omissus (Edwards)

and a key to the larvae and pupae of the Armigeres occurring in Nepal (Diptera: Culicidae) .. 964
ECKERLIN, RALPH P. and HARRY F. PAINTER—New records of fleas (Siphonaptera) from

SSISURSIND) VIWEG NATE SCE Ne Sg ea Re yn oe aNd A ea Papa Neer WES Aha ae 969

EVANS, HOWARD E.—Three new species of Dipogon Fox (subgenus Dipogon) (Hymenoptera:
Pompilidae) from central and western North America ................ 0.02 cece eee cece eee eens 1010

GAGNE, RAYMOND J., HELGA BLANCO-METZLER, and JEAN ETIENNE—A new
Neotropical species of Clinodiplosis (Diptera: Cecidomyiidae), an important new pest of culti-
Seca pels (Capsicuin Spps:, SOlAnACEAG), (02 pahie slo ekret Nafatac se cies eae ate emclayh aie Sine 4 abscissa 831

GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota pubescens
Freidberg and Mathis (Diptera: Tephritidae) on Lessingia filaginifolia (Hooker and Arnott) M. A.

Mave i AStErACcAC) Min SOULAe Ti CallfOrnlay., oleic :.°4 2 Sake ayes tials Sisiele Beis Ret sete ek ae Ge Gisele es ce seicet 878
GUILBERT, ERIC—Revision of the genus Parada Horvath (Hemiptera: Tingidae) with cladistic
SLT STIS 92 sai hk Tah os Baad Se NRG to ANS Boa ORI AT (5 gg a ae 816

(Continued on back cover)

THE

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PROC. ENTOMOL. ;
102(4), 2000, ;

“”. WASH.
773-193

FLOWER FLIES OF THE SUBGENUS OCYPTAMUS (MIMOCALLA HUi-
(DIPTERA: SYRPHIDAE)

FE CHRISTIAN THOMPSON AND MANUEL A. ZUMBADO

(FCT) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S.
Department of Agriculture, % National Museum of Natural History Smithsonian Insti-
tution, Washington, DC 20560-0169 U.S.A. (e-mail: cthompso@sel.barc.usda.gov);
(MAZ) Entomology, Instituto Nacional de Biodiversidad, Santo Domingo, Heredia, Costa
Rica (e-mail: mzumbado @inbio.ac.cr)

Abstract.—The subgenus Ocyptamus (Mimocalla) is revised. Ocyptamus (Mimocalla)
tristani Zumbado, new species, is described and two new synonyms are proposed (Sal-
pingogaster flukei Curran 1941 and Baccha polista Hull 1943 = Baccha bonariensis
Bréthes 1905). A key to the species, along with synonymies, descriptions, distributions,

and figures for all species are given.

Key Words: Neotropical, key

The New World tropics have the richest
diversity of flower flies in the World. This
diversity holds great promise for sustain-
able agriculture. The flower fly diversity
largely resides in three genera (Toxomerus
(150 spp.), Ocyptamus (300 spp.) and Co-
pestylum (320 spp.)). The flies of the genus
Ocyptamus are predators of a diverse array
of plant pests (mites, scales, plant hoppers,
aphids, etc.). To fully utilize their potential,
the species must be named and described
so that their life-histories can be studied and
reported. Then the relative contribution of
these predators to agroecosystems can be
assessed and, perhaps, enhanced.

Consider coffee production: While most
coffee is today grown in large, open, single
crop systems which require insecticide use,
there is a growing trend to return to more
natural shade-grown systems. In such sys-
tems, the new species described herein may
be a useful biological control agent. The
species is a predator of black, soft-brown
and hemispherical scales, which are pests
of coffee and various ornamental plants.

This paper is the second in a series to
treat various components of the large di-
verse genus Ocyptamus. The first part
(Thompson, in preparation) includes a de-
scription and diagnosis of the genus and a
key to the components (subgenera and spe-
cies groups) along with the treatment of
several species groups. This paper treats the
species group named and treated as a sub-
genus by Hull. Until a cladistic analysis is
done of the genus as a whole, we, for prag-
matic reasons, retain Mimocalla as a sub-
genus. The likely sister to Mimocalla is O.
conjunctus Wiedemann and the sister to O.
conjunctus + Mimocalla is O. sargoides
Macquart. Using these species as out-
groups, we include a preliminary cladistic
hypothesis for the species of Mimocalla.
Redescriptions, complete synonymies, illus-
trations, and distributions are given for all
species. Terminology follows Thompson
(1999), the abbreviations found in the syn-
onymies follow Thompson (2000), the use
of the asterisk in the distribution statement
means verified records to be found below

774 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Habitus of Ocyptamus tristani, dorsal.

in the material examined section. Color im-
ages of these flies along with the rest of the
information in this paper may be found on
the Diptera WWW site (http://www.sel.
barc.usda.gov/Diptera/) and on the Diptera
Data Dissemination Disk.

Genus Ocyptamus
Ocyptamus Macquart 1834: 554. Type spe-
cies, fascipennis Macquart (Coquillett
1910: 577) = fuscipennis Say.
Subgenus Mimocalla Hull

Baccha subg. Mimocalla Hull 1943d: 46.
Type species, Baccha capitata Loew

T. Litwak

(orig. des.); Hull 1949: 97 (discussion,
key).

Description.—Face usually yellow, rare-
ly with weak brownish vitta or black with
narrow yellow vitta, with strong tubercle;
frons prominent; vertex short, broad, with
ocellar triangle about 1% times its length
anterior to hind margin of eye; occiput with
2 rows of cilia, with cilia normal, not scale-
like; metasternum bare; metathoracic epis-
ternum pilose; postmetacoxal bridge incom-
plete; scutellum yellow or rarely entirely
black, with 1 row of strong and long ventral

VOLUME 102, NUMBER 4

pile; plumula long to absent; calypter with
strong fringe only on ventral edge; meson-
otum usually with bright yellow pollinose
vittae, rarely entirely dark; metatrochanter
always with a tuft of long black pile, fre-
quently males with long ventral pile also on
metacoxa and femur; abdomen clavate,
constricted at base of 2nd tergum and grad-
ually widening apically, brown to black,
with yellow fasciae; male genitalia greatly
enlarged, with cercus and apicolateral cor-
ners of 9th tergum enlarged; male 5th ster-
num variously modified on apical half.

Wing: Alula normal, about 1% times as
broad as cell BM, bare or microtrichose;
crossvein r-m at basal % of cell DM, slight-
ly beyond termination of vein Sc; vein M1
sinuate, ending approximately at termina-
tion of vein R1; vein R4+5 sinuate, making
a shallow loop into celi R4+5.

Length: 12-16 mm; wing, 10—14 mm.

Discussion.—The subgenus Mimocalla is
readily distinguished from all other groups
of Ocyptamus by: 1) strongly petiolate and
clavate abdomen; 2) large size; 3) enlarged
male genitalia including a modified 4th
sternum; 5) vein R4+5 sinuate; 6) vein M1
sigmoid; and 7) metatrochanter with a
strong pile tuft. In appearance Mimocalla
can only be confused with O. (Theranta)
conjunctus, but Mimocalla is readily distin-
guished by the sinuate vein R4+5 and short
antenna.

While a full cladistic analysis has not
been done, the characters of the male gen-
italia suggest these relationships (Mimocal-
la = (giganteus + willistoni) + ((bonarien-
sis + erebus) + (nymphaea + (capitatus +
tristani)))). In most Ocyptamus, the cercus
is unmodified, small, and oval, but in Mim-
ocalla the cercus is enlarged. In one group
(giganteus + willistoni), the shape of the
cercus is triangular and the surstylus short,
blunt apically and directed ventrally. In the
other group, the cercus is elongate and the
surstylus is also elongate, tapered apically
and directed posteriorly. Within this second
group, the genitalia of capitatus and tristani
are the same, and those of nymphaea are

ads

very similar. These three species als. hare
the more or less simple 5th sternum. “he
other two species (bonariensis, erebis)
have the 5th sternum produced apicolater-
ally on the right side.

Little is known about the biology of
Ocyptamus species. However, the life his-
tory of one species of Mimocalla has been
published. We review that work in light of
our rearing of tristani, new species. Bruch
(1923) noted that bonariensis larvae com-
pletely suck dry a scale insect before mov-
ing on to the next and that the complete
larval cycle probably lasts about three
weeks. Once the larva finishes feeding, the
larva remains quiescent for about two days,
then evacuates the gut and changes position
in order to pupate, either on the same plant
or nearby, and finally resting about one
more day before pupating. The larva slowly
becomes smaller and releases a viscous
transparent liquid, which when hardens to
anchor the puparium. During the process of
transformation the posterior part of the lar-
va becomes narrower, while undulatory and
contraction movements of the pupa forming
inside inflates bit by bit the anterior portion;
this process lasts about two hours. The re-
cently formed puparium is colored the same
as the mature larva but shortly after pupa-
tion it becomes pale brown. The pupal stage
lasts about two weeks.

Observations by M. Zumbado and J. A.
Ugalde on tristani confirm Bruch’s obser-
vations. Mature tristani larvae (Figs. 20-—
22) measure between 12 and 14 mm long
and about 5 mm wide. The body is subpar-
allel along its length, rather convex dorsally
and flat ventrally; anteriorly it is narrower
and posteriorly is obliquely truncated. The
color is similar to that of bonariensis. The
puparium (Fig. 23) is smaller than the larva
and measures 8 to 9 mm long; in a dorsal
view, its shape is pyriform; the anterior por-
tion is highly convex dorsally, becoming
narrower and cone shaped posteriorly. Ven-
trally the puparium is flat posteriorly and
slightly concave anteriorly. The surface is

776

coarse, without tubercules. The posterior
breathing tube is very short and black.

We reared tristani from July Ist to Oc-
tober 13, 1993 and from June 20 to October
10, 1994, at INBio headquarters in Santo
Domingo, Heredia Province, Costa Rica, at
an elevation of 1,100 m. During the first
season we did not successfully rear a single
adult, just parasitic wasps (Hymenoptera:
Pteromalidae and Encyrtidae). Larvae and
puparia were found from June to October,
during the rainy season in Costa Rica, when
humidity favors scale insect development.
Adults were present from February to Au-
gust.

We found tristani larvae feeding on scale
insects (Homoptera: Sternorrhyncha: Coc-
coidea: Coccidae) on various plant species
(Rutaceae: Citrus sp., Fabaceae: Acacia sp.,
Gliricidia sepium (Jacq.) Kunth ex Walp,
Solanaceae: Datura sp., Rubiaceae: Coffea
arabica Linnaeus, Lamiaceae: Ocimum
basilicum Linnaeus).

The larvae use a glue-like saliva (Roth-
eray 1986) to defend themself against ag-
gressive organisms such as ants. Two Cam-
ponotus ants were seen attacking a larva
that responded by turning the anterior por-
tion of the body towards the ant and re-
leasing a drop of this liquid substance on
them. The ants were trapped for at least 2
minutes before freeing themselves.

Several species of parasitic wasps were
found to attack tristani, all of them emerg-
ing from puparia. All reared parasitic wasps
belong to the Encyrtidae, mainly Syrpho-
phagus sp. which attacks the larval stage
(Noyes 1995), and Pteromalidae, mainly
Pachyneuron sp. In most cases, several to
many (4 to 75 encyrtids, 1 to 33 pterom-
alids) emerged from a single puparium.
These natural enemies seem to be very ef-
fective against tristani larvae. Neither Pa-
chyneuron nor Syrphophagus are host spe-
cific in Ocyptamus, both are hyperparasi-
toids in aphids (Homoptera: Aphididae) or
primary parasitoids in syrphids (Hanson
and Gauld 1995). Syrphophagus also para-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

sitizes Braconidae and Aphelinidae hyme-
nopterans (Noyes 1995).

In our first attempts to rear the fly (60
puparia), we got only parasitic wasps (636
Encyrtidae and 260 Pteromalidae). So we
started taking very recently formed puparia
(17) and were then successful in getting
adult flies (12). This suggests that female
wasps are ovipositing on freshly formed pu-
paria, perhaps attracted by the chemicals re-
leased during the transformation into pupae.

KEY TO SPECIES OF OCYPTAMUS
(MIMOCALLA)

1. Legs black; mesonotum black; wing broadly
black anteriorly; large, body length 20 mm . .
A seenta co: Circe ctekn aioe here mene Sb sc giganteus
— Legs partially or entirely pale, always with me-
sotibia yellow; wing hyaline or at most narrow-
ly light brownish anteriorly
. Metafemur and tibia black; mesonotum black;
face yellow; large, body length 16 mm or more
FRR eR ee in arg coro Bnala o-oo & willistoni
— Metafemur and tibia partially pale, brownish
orange to yellow; mesonotum at least yellow
on postalar callus; if mesonotum extensively
dark, then face extensively dark; smaller, body
length less than 16mm -:--. °° 2232. 3
3. Abdomen yellow on Ist, 2nd and basal 4% of
3rd segments, elsewhere black; pleuron entire-
ly yellow; legs yellow. Alula microtrichose . .
Seer 2 ged aa Oe, AY Ree nymphaea
— Abdomen not as such, with Ist and 2nd seg-
ment partially dark, brown to black; pleuron
and legs partially dark
4. Postpronotum brownish black, only margins
pale; notopleuron extensively brownish black;
face extensively brownish black, only narrowly
yellow laterally; abdomen appearing totally
dark, abdominal pale fasciae greatly reduced
SRP ee PROTA Lae chars Co cof erebus
— Postpronotum and notopleuron extensively yel-
low; face yellow or with narrow brown vitta;
abdomen with distinct yellow fasciae
5. Alula almost entirely bare. Antenna pale, or-
ange to brownish orange; scutellum yellow or
obscurely grayish on disc (West Indies)
a age a gailcy Dea = ita ks Ae COR capitatus
— Alula more extensively microtrichose, entirely
(2) or partially bare medially (d). Antenna and
scutellum usually partially dark (not West In-
dies)
6. 2nd tergum brown to black, except broadly yel-
low basolaterally and with medial triangular
macula; scutum with lateral yellow vitta con-
tinuous behind transverse suture; abdominal

WN

VOLUME 102, NUMBER 4

fasciae on 3rd & 4th terga broad, about % as

Wi GeraSpSeCIMenterps vas seri ase « Alghct ce lees tristani
— 2nd tergum reddish brown except narrowly

yellow basolaterally; scutum with lateral yel-

low vitta broadly interrupted behind transverse

suture; abdominal fasciae on 3rd & 4th terga

narrow, about '/, as wide as segment.....

Ocyptamus (Mimocalla) bonariensis
(Bréthes)
(Figs. 2—4)

Baccha bonariensis Bréthes 1905: 340 Ar-
gentina, Buenos Aires (HT ¢6 MACN
lost); Kertész 1910: 157 (cat. cit.); Fluke
1957-5 153.(Cat:. cit.).

Ocyptamus bonariensis: Thompson et al.
19762 13 .(Cat:-cit.).

Salpingogaster flukei Curran 1941: 284 Ec-
uador, Tungurahua, Banos (HT 4
AMNH); Arnaud and Owen 1981: 125
(type data). New synonym.

Baccha flukei: Hull 1949: 98, 105 (key ref.,
note); Fluke 1956: 246 (cat. cit.).

Ocyptamus flukei: Thompson et al. 1976:
18 (cat. cit.).

Baccha (Mimocalla) phobia Hull 1943b: 51
Ecuador, Tungurahua, Banos, Chaupi
(HT ¢ AMNH); Hull 1949: 240 (fig. (ab-
domen pattern)). Syn. Hull 1949: 98.

Baccha (Mimocalla) polista Hull 1943c: 89
Brazil, Santa Catarina, Nova Teutonia
(HT 56 AMNH); Hull 1949: 98, 105, 236
(fig. (abdomen pattern), key ref.). New
synonym.

Baccha nigriventris (misidentifications):
Sack 1920: 249, 1921: 135 (puparium);
Bruch 1923: 1 (cat. cit.); Knutson 1971:
29 (notes).

Male.—Head: Face yellow, with light
brown medial vitta, sparsely white polli-
nose laterally, only densely pollinose nar-
rowly along eye margin, white pilose ex-
cept black pilose ventrad to antenna; gena
yellow, shiny, bare; lunule yellow except
brown medially; frontal triangle brown on
anterior 42, yellow posteriorly, black pilose;
vertical triangle black, black pilose; occiput
black except yellow on ventral ¥, densely
whitish gold pollinose ventrally becoming

TY

more golden dorsally, white pilose v« \trally
becoming more golden dorsally; anicnna
brownish black except reddish brown ba-
soventrally on basoflagellomere, black pi-
lose; basoflagellomere elongate, about %
longer than wide.

Thorax: Mainly brownish black with
yellow maculae; postpronotum yellow, with
rest of prothorax brownish yellow; scutum
black except with broad yellow vitta run-
ning from postpronotum to and including
anterior % of postalar callus but broadly in-
terrupted behind transverse suture and dor-
sad to wing base, with vitta about as wide
as postpronotum, with black areas black
pollinose except for broad golden pollinose
submedial vitta which tapers posteriorly
and does not reach scutellum and a very
narrow indistinct brown pollinose medial
vitta on anterior % which expands into a
broader golden pollinose vitta at level of
postalar callus, long yellow pilose anteri-
orly and laterally, short intermixed black
and yellow pile elsewhere; scutellum
brownish black except yellow base and api-
cal margin, short sparse black pilose with a
few intermixed yellow pili basally; subscu-
tellar fringe singular, white; pleuron black-
ish brown except yellow dorsad to procoxa
and broadly yellow on posterior anepister-
num, dorsal % of katepisternum, anterior %2
anepimeron and anterior % katatergum,
very sparsely white pollinose except denser
dorsally on katepisternum, yellow pilose;
plumula very short, white; calypter yellow-
ish white except dorsal margin brownish;
halter brownish yellow. Legs: Procoxa or-
ange, sparsely gray pollinose, yellow pi-
lose; mesocoxa brown, black pilose medi-
ally, yellow pilose laterally; metacoxa
black, long black pilose with a few inter-
mixed yellow pili laterally; protrochanter
orange; mesotrochanter brownish black;
metatrochanter black, long black pilose;
profemur orange, yellow pilose; mesofemur
orange except blackish basally and apico-
posteriorly, yellow pilose except for a few
black pili on dorsoposterior surface; meta-
femur orange except black ventrally, orange

778 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

| / yy i
WHat

Figs. 2-7. Male genitalia. 2, Ocyptamus bonariensis, 9th sternum, aedeagus and associated structures, lateral
view. 3. O. bonariensis, 4th sternum, ventral view. 4, O. bonariensis, 9th tergum and associated structures,
lateral view. 5, O. tristani, 9th sternum and associated structures, lateral view. 6, O. tristani, 4th sternum, ventral
view. 7, O. tristani, 9th tergum and associated structures, lateral view.

VOLUME 102, NUMBER 4

pilose except black pilose ventrally; pro and
mesotibiae and tarsi yellow, yellow pilose;
metatibia orange except yellow basal % and
apex, yellow pilose; metatarsus brownish
orange, yellow pilose. Wing: Narrowly
brown anteriorly, hyaline posteriorly, mi-
crotrichose except bare cell R1 anterior to
vein Rs, cell R, cell BM, anterobasal % of
cell CuP, cell R4+5 posterior to spurious
vein, narrowly on basoposteriorly in cell
DM, basomedial % of cell CuA1, anterior
to vein A2; alula entirely microtrichose to
bare basomedially, normal, as wide as cell
CuP; brown area including base and ex-
tending posteriorly to veins R and R1.
Abdomen: Dark brown with yellow fas-
ciae; 1st tergum brownish black except yel-
low basally, white pilose basolaterally,
black pilose elswhere; 2nd tergum brownish
black except narrowly yellow basolaterally,
yellow pilose basolaterally, black pilose
elsewhere; 3rd tergum yellow on basal \,
brownish black apically, yellow pilose on
yellow area, black pilose elsewhere; 4th ter-
gum brown except yellow basal ;, black
pilose except yellow pilose basolaterally;
5th tergum brown, black pilose except yel-
low laterally; genitalia brown and yellow,
black pilose; 1st sternum brown, white pi-
lose; 2nd sternum brown except yellow bas-
al %, yellow pilose; 3rd & 4th sterna brown
except narrowly yellow basally, black pi-
lose; 5th sternum brown, yellow pilose.
Male genitalia: See Figs. 2—4.
Female.—Similar to male except for nor-
mal sexual dimorphism and frons black an-
teriorly except broadly yellow along eye
margin, brown on medial %, black posteri-
orly, shiny on anterior %, sparsely gray pol-
linose medially, black pollinose on poste-
rior %4, short black pilose.
Length.—15 mm; wing, 12 mm.
Distribution.—Ecuador*, Peru*, Brazil
(Santa Catarina*), Uruguay*, Argentina
(Buenos Aires*, Misiones*).
Types.—Baccha bonariensis was de-
scribed from a unique male from Palermo,
Buenos Aires, Argentina, collected on 11
April 1904. A search of the Diptera collec-

7719

tion of MACN failed to find this spc -imen.
However, there is a single reared female
from Buenos Aires (9 Dec 1904) which was
identified by Bréthes.

Salpingogaster flukei was described from
a series of 4 males from Ecuador collected
by W. Clark McIntyre. The holotype is now
in the AMNH and was examined. Paratypes
are in the CNC and UWEC.

Baccha phobia was described from a se-
ries of 2 males from Ecuador collected by
W. Clark McIntyre. The holotype is now in
the AMNH and paratype is in the CNC.
Both were examined.

Baccha polista was described from a se-
ries of 6 specimens from Brazil (Santa Ca-
tarina) collected by Fritz Plaumann and in
the Fluke Collection. The holotype is now
in the AMNH and was examined. Paratypes
are in the CNC and UWEC.

Material examined (15 ¢, 6 2).—EC-
UADOR. Types of flukei and phobia.
PERU. Cuzco, Kilometer 94, Machu Picchu
RailRoad, 7 Mar 1978, PB M. Marsh (¢
USNM ENT 00037937 USNM); Lima,
Bosque Carrion, 2,400 m, 16 April 1987, P.
Hocking (6 USNM ENT 00030147
USNM). BRAZIL. Santa Catarina: types of
polista; Nova Teutonia, 300-500 m, Jan
1965, E Plaumann (26 2 USNM ENT
00037924-6 USNM); ... Feb 1965 (¢
USNM ENT 00037927 USNM); ... Feb
1972 (6 USNM ENT 00037930 USNM);
... Nov 1971 (6 USNM ENT 00037929
USNM); ... Feb 1972, E Plaumann (¢d
USNM ENT 00037977 USNM); ... Dec
1964 (6 USNM ENT 00037928 USNM).
URUGUAY. Montevideo, “Rec. 3.1.42,”
H. L. Parker (2 with puparium USNM
ENT 00037938 USNM). ARGENTINA.
Buenos Aires: Buenos Aires, 4 Dec 1904
(MACN); Moreno, Dec 1971, Fritz (6
USNM ENT 00037931 USNM); ... Dec
1972 (2 2 USNM ENT 00037932-3
USNM); ... Jan 1973 (2 USNM ENT
00037934 USNM); Villa Elisa, 15—29 Dec
1979,°G; My & O: S: FlinteIr (2 USNM
ENT 00037935 USNM); Santa Catalina,
near Buenos Aires, 10 Nov 1970, J. W.

780

Boyes (6 USNM ENT 00022541 CNC).
Misiones: Dos de Mayo, Dec 1973, M.
Fritz (6 USNM ENT 00037936 USNM).

Discussion.—Ocyptamus bonariensis 1s
very similar to erebus, but differs in having
distinct yellow pollinose vittae on the me-
sonotum and broader yellow fasciae on the
abdomen. The shape of the male cercus is
also distinctive.

The concepts of flukei Curran and polista
Hull are very close. Hull (1949: 105) sep-
arated them by two characters (shape of the
2nd tergum (2nd tergum narrow, only as
wide basally as apically (flukei) versus
broader, about twice as wide apically as ba-
sally (polista)) and color of hind femur
(hind femur entirely brownish yellow (flu-
kei) versus black ventrally (polista))).
While these differences are supported by
the material we have examined, we consid-
er them to be a function of geographic var-
iation which will be bridged once material
from intermediate localities are studied.
Hence, we have synonymized the two con-
cepts. The name bonariensis was unknown
to Hull, hence he could not have known
that it was an older synonym for his polista.
Also, we include here a single male from
Peru (Cuzco) with a bare alula, no basal
yellow fascia on 4th tergum, greatly re-
duced yellow fascia on 3rd tergum and
which is slightly larger than typical Mimo-
calla (16 mm body length, 14 mm wing
length). As the male genitalia are the same
as those of typical specimens, we consider
this to be an aberrant specimen.

Ocyptamus (Mimocalla) capitatus (Loew)

Baccha capitata Loew 1863: 14 Cuba (LT
3 MCZ here designated); Williston 1887:
124 (Cuba, redescr.); Aldrich 1905: 354
(cat. Git.): Kertész 1910:.157 (cat: \cit.);
Fluke 1956: 239 (cat. cit.); Wolcott 1923:
217, 1936: 346, 1948: 461 (Puerto Rico,
with Saissetia hemisphaerica on coffee).

Ocyptamus captitatus: Thompson et al.
1976: 14; Thompson 1981: 44 (figs. (ab-
domen pattern, male genitalia, wing),
West Indies).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Doros insularis Bigot 1883: 330 Cuba (HT
2 BMNH); Williston 1887: 299 (note).
Syn. Thompson 1981: 44.

Xanthogramma insulare: Aldrich 1905: 369
(cat. cit.); Kertész 1910: 152 (cat. cit.).

Ocyptamus insularis: Thompson et al.
1976: 20 (cat. cit.).

Baccha carlota Curran 1929: 491 Cuba,
Trinidad Mountain, Mina Carlota (HT @
AMNH); Hull 1949: 98, 105. 238, 240
(fig. (abdomen pattern), key ref.); Fluke
1956: 239 (cat. cit.); Arnaud and Owen
1981: 83 (type data). Syn. Thompson
1981: 44.

Ocyptamus carlota: Thompson et al. 1976:
14 (cat. cit.).

Male.—Head: Face yellow, with light
brown medial vitta, sparsely white polli-
nose laterally, only densely pollinose nar-
rowly along eye margin, white pilose ex-
cept black pilose ventrad to antenna; gena
yellow, shiny, bare; lunule brown except
yellow submedially; frontal triangle yellow
except brown along lunule, black pilose;
vertical triangle black, black pilose; occiput
black except yellow on ventral ’;, densely
golden pollinose becoming more brownish
on dorsal %, yellow pilose except pile more
golden dorsally; antenna orange except
more brownish orange on dorsoapical %4 of
basoflagellomere, black pilose; basoflagel-
lomere elongate, slightly less than twice as
long as wide.

Thorax: Mainly black with yellow mac-
ulae; postpronotum yellow; scutum black
except with broad yellow vitta running
from postpronotum to and including ante-
rior % of postalar callus but broadly inter-
rupted behind transverse suture and dorsad
to wing base, with vitta about as wide as
postpronotum, with black areas black pol-
linose except for broad golden pollinose
submedial vitta which tapers posteriorly
and does not reach scutellum and a very
narrow indistinct brown pollinose medial
vitta on anterior 7% which expands into a
broader golden pollinose vitta at level of
postalar callus, long yellow pilose anteri-

781

VOLUME 102, NUMBER 4

hin ‘i
Wr
a iS
‘ Ww
5
feet
og

a, Ab 12

Male genitalia. 8, Ocyptamus giganteus, 9th sternum, aedeagus and associated structures, lateral

Figs. 8-13. italia. 8, ) é
view. 9, O. giganteus, 4th sternum, ventral view. 10, O. giganteus, 9th tergum and associated structures, lateral
view. 11, O. nymphaea, 9th sternum and associated structures, lateral view. 12. O. nymphaea, 4th sternum

ventral view. 13, O. nymphaea, 9th tergum and associated structures, lateral view

782

orly and laterally, short intermixed black
and yellow pile elsewhere; scutellum yel-
low except disc blackish, short sparse black
pilose with a few intermixed yellow pili ba-
sally; subscutellar fringe singular, white;
pleuron black except yellow dorsad to pro-
coxa and broadly yellow on posterior ane-
pisternum, dorsal % of katepisternum and
anterior % anepimeron, very sparsely white
pollinose except denser dorsally on katepis-
ternum, yellow pilose; plumula very short,
yellow; calypter yellowish white except
dorsal margin brownish; halter yellow.
Legs: Procoxa brownish black except yel-
low apex, sparsely gray pollinose, yellow
pilose; mesocoxa black, black pilose me-
dially, yellow pilose laterally; metacoxa
black, long black pilose with a few inter-
mixed yellow pili laterally; protrochanter
yellowish brown; mesotrochanter brownish
black; metatrochanter black, long black pi-
lose; profemur yellow, yellow pilose; me-
sofemur yellow except black on basodorsal
¥%, yellow pilose except for a row of black
pili on dorsoposterior surface; metafemur
black except yellow apical ¥,, black pilose
except yellow pilose on yellow area; pro
and mesotibiae and tarsi yellow, yellow pi-
lose; metatibia black except yellow basal %
and apex, black pilose; metatarsus yellow,
yellow pilose. Wing: Brown anteriorly, hy-
aline posteriorly, microtrichose except bare
cell R posterior to spurious vein on apical
¥%, cell BM, anterobasal % of cell CuP, cell
R4+5 posterior to spurious vein, narrowly
on basoposteriorly in cell DM, basomedial
¥% of cell CuA1, anterior to vein A2; alula
bare, normal, as wide as cell CuP; brown
areas including base and extending poste-
riorly to vein M to level of end of vein RS,
to spurious vein in apical portion of cell R,
to vein R4+5 except narrowly hyaline
along sinuate portion.

Abdomen: Dark brown with yellow fas-
ciae and orange apex; Ist tergum yellow ex-
cept brownish black apicomedial %, yellow
pilose; 2nd tergum brownish black except
for yellow basolateral triangular macula,
with maculae broadly separated medially,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

yellow pilose basolaterally, black pilose
elsewhere; 3rd tergum yellow on basal %,
brownish black apically, yellow pilose on
yellow area, black pilose elsewhere; 4th ter-
gum brown except with yellow basal fascia
which is narrowly isolated from lateral mar-
gin and occupies basal %4, with apical por-
tion of tergum becoming more orange,
black pilose; 5th tergum orange, black pi-
lose; genitalia yellowish orange, black pi-
lose; 1st sternum yellow, yellow pilose; 2nd
sternum brown except yellow basally, yel-
low pilose; 3rd sternum brown except yel-
low basal %, black pilose except yellow pi-
lose basally; 4th sternum brown, brown pi-
lose; 5th sternum orange except yellow api-
cally, brown pilose. Male genitalia. See
Figs. 56a—c in Thompson (1981: 59); es-
sentially the same as tristani (Figs. 5—7).

Female.—Similar to male except for nor-
mal sexual dimorphism and frons yellow
except may be darker medially and along
lunule, orange pollinose except maybe
more brownish medially, short black pilose.

Length.—14 mm; wing, 11 mm.

Variation.—The brownish black color in
some examined individuals is brownish or-
ange; also the antenna may be entirely or-
ange and scutellum entirely yellow.

Distribution.—West Indies (Cuba*, Do-
minican Republic*, Puerto Rico*, Lesser
Antilles*).

Types.—Baccha capitata was described
from an unspecified number of male spec-
imens from Cuba from Gundlach. In the
MCZ, there is a male with a silver square
and labelled ‘‘capitata mihi’? in Loew’s
hand. This specimen is here designated lec-
totype so as to fix and stabilize the current
concept of the name, and has been so la-
belled.

Doros insularis was described fron a
unique female specimen from Cuba. The
holotype is now in the BMNH and has been
examined.

Baccha carlota was described from a
unique female from Cuba collected by G.
Salt and in the Curran collection. The ho-

VOLUME 102, NUMBER 4

lotype is now in the AMNH and was ex-
amined.

Material examined (8 3, 7 2).—CUBA.
Types of capitata, insularis & carlota. DO-
MINICAN REPUBLIC. Pedernales, 23.5
km N Cabo Rojo, 18-16N 71-38W, 540 m,
13 July 1990, C. Young, J. Rawlins & S.
Thompson (6 USNM ENT 00037907
USNM). PUERTO RICO. Comerio, 10
Aug 1913, J. R. Johnston (6 USNM ENT
00037900 USNM); Mayaguez, 4 Mar 1912,
C. W. Hooker (2 USNM ENT 00037901
USNM); ... 26 Jul 1915, R. H. van Zwa-
lenberg (6 USNM ENT 00037902
WSNM); .... > May’ 1939;H. K.\Plank.G
66 2 22 with puparia USNM ENT
00037904-6, ... 72-73 USNM) VIRGIN
ISLANDS. St. Croix, 6 Jul 1963, R. M.
Bond & R. Boyd (¢6 % USNM ENT
00037899, ... 903 USNM).

Discussion.—Ocyptamus capitatus is
similar to tristani and bonariensis in having
distinctive yellow pollinose vittae on me-
sonotum, but differs from these species in
having a yellow scutellum and the apex of
the abdomen orange to brownish orange.

Ocyptamus (Mimocalla) erebus (Hull)
(Figs. 14—16)

Baccha (Mimocalla) erebus Hull 1943d:
54; 1943e: 137 Brazil, Santa Catarina,
Nova Teutonia (HT ¢ AMNH); Hull
1949: 98, 105, 240 (fig. (abdomen pat-
tern), key); Fluke 1956: 245 (cat. cit.).

Ocyptamus erebus: Thompson et al. 1976:
17 (cat: Cit.).

Male.—Head: Face yellowish brown,
with dark brown medial vitta, sparsely
white pollinose laterally, only densely pol-
linose narrowly along eye margin, white pi-
lose except black pilose ventrad to antenna;
gena yellow, shiny, bare; lunule black ex-
cept brownish submedially; frontal triangle
black except narrowly yellow along eye
margin, shiny except dull pollinose on pos-
terior %, black pilose; vertical triangle
black, black pilose; occiput black except
yellow on ventral Y,, densely gray pollinose,

783

white pilose; antenna black excepi more
brownish orange basoventrally on basofia-
gellomere, black pilose; basoflagellomere
elongate, slightly less than twice as long as
wide.

Thorax: Mainly brownish black; post-
pronotum reddish brown; scutum black ex-
cept with obscure narrow reddish brown
vitta posterior to postpronotum and another
anterior to and including anterior % of pos-
talar callus, with area behind transverse su-
ture and dorsad to wing base dark, with
black areas black pollinose except for nar-
row golden pollinose submedial vitta which
tapers posteriorly and does not reach scu-
tellum and a narrow golden pollinose me-
dial vitta on posterior % at level of postalar
callus, long yellow pilose anteriorly and lat-
erally, short intermixed black and yellow
pile elsewhere; scutellum dark brown ex-
cept brownish yellow on base and apical
margin, short sparse black and yellow pi-
lose; subscutellar fringe singular, white;
pleuron black except obscurely yellow on
posterior anepisternum, dorsal % of katepis-
ternum and medially on katatergum, very
sparsely white pollinose except denser dor-
sally on katepisternum, white pilose; plu-
mula very short, yellow; calypter brownish
white except dorsal margin brown; halter
brown. Legs: Procoxa yellow, sparsely
white pollinose, yellow pilose; mesocoxa
orange, brownish-orange pilose; metacoxa
black, long black pilose with a few inter-
mixed yellow pili laterally; pro and meso-
trochanter brown; metatrochanter black,
long black pilose; pro and mesofemur red-
dish brown, yellow pilose except for a few
black pili on dorsoposterior surface; meta-
femur reddish brown except yellow apical
/;, Orange pilose except black apicoventral-
ly; pro and mesotibiae and tarsi yellow, yel-
low pilose; metatibia yellow except reddish
brown medially, orange pilose; metatarsus
dark brown, yellow pilose. Wing: Light
brown anteriorly, hyaline posteriorly, mi-
crotrichose except bare cell R1 anterior to
vein Rs, cell R, cell BM, anterobasal % of
cell CuP, cell R4+5 posterior to spurious

784 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 14-19. Male genitalia. 14, Ocyptamus erebus, 9th sternum, aedeagus and associated structures, lateral
view. 15, O. erebus, 4th sternum, ventral view. 16, O. erebus, 9th tergum and associated structures, lateral view.
17, O. willistoni, 9th sternum and associated structures, lateral view. 18, O. willistoni, 4th sternum, ventral view.
19, O. willistoni, 9th tergum and associated structures, lateral view.

VOLUME 102, NUMBER 4

vein, narrowly on basoposteriorly in cell
DM, basomedial % of cell CuA1, anterior
to vein A2; alula microtrichose, normal, as
wide as cell CuP; brown areas including
base and extending posteriorly to vein R
and R1.

Abdomen: Dark brown with greatly re-
duced yellow fasciae; Ist tergum brownish
black, white pilose; 2nd tergum brownish
black except for small yellow basolateral
triangular macula, with maculae broadly
separated medially, yellow pilose basolater-
ally, black pilose elsewhere; 3rd and 4th
terga brownish black except for very nar-
row obscure yellow fasciate maculae broad-
ly separated medially and occupying less /,,
of tergal length, yellow pilose laterally,
black pilose elsewhere; 5th tergum black,
black pilose; genitalia brownish black,
black pilose; Ist sternum brown, white pi-
lose; 2nd sternum brown except yellow ba-
sally, yellow pilose; 3rd and 4th sterna
brown except yellow apical margin, black
pilose; 4th sternum brown, brown pilose;
5th sternum brownish orange, brown pilose.
Male genitalia. See Figs. 14-16.

Female.—Similar to male except for nor-
mal sexual dimorphism and frons black ex-
cept narrow yellow vitta along eye margin,
shiny anteriorly, gray pollinose on medial
¥%, black pollinose posteriorly, short black
pilose.

Length.—13 mm; wing, 11 mm.

Distribution.—Brazil (Minas Gerais*,
Sao Paulo*, Santa Catarina*).

Type.—Baccha erebus was described
from 4 male specimens collected in south-
ern Brazil (Santa Catarina) by Fritz Plau-
mann and in the Fluke Collection. The ho-
lotype is now in the AMNH and was ex-
amined. Paratypes are in the CNC and the
University of Wisconsin Entomology Col-
lection. The name was first validated in a
key which appeared before the original de-
scription.

Material examined (10 ¢d 6 ?).—BRA-
ZIL. Minas Gerais: Arceburgo, FE Fortaleza,
Dec 1946, Barretto (2 22 USNM ENT
00037974—5 MZUSP). Sao Paulo: Cantar-

785

eira, Chapadao, Nov 1946, Barreiic (2
USNM ENT 00037976 MZUSP). Santa “a-
tarina: types of erebus; Nova Teutonia, 27-
11S 52-23W, 300-500 m, Feb 1965, FE
Plaumann (2 66 USNM ENT 00037908—
9 USNM); ... Feb 1972 (2 USNM ENT
00037916 USNM); > Mar 21957, (2
USNM ENT 00037910 USNM); ... Mar
1960 (2 USNM ENT 00037912 USNM);
... Nov 1952 (6 USNM ENT 00037911
USNM); ... Nov 1971 (3 66 USNM ENT
00037913—5 USNM).

Discussion.—Ocyptamus erebus is a dark
version of bonariensis, the yellow vittae on
the mesonotum and abdominal fasciae are
greatly reduced, obscure or absent. The
shape of the male cercus is also very dis-
tinctive.

Ocyptamus (Mimocalla) giganteus
(Schiner)
(Figs. 8-10)

Baccha gigantea Schiner 1868: 340 *‘South
America’?> = Venezuela (LT 6 NMW
here designated); Williston 1886: 314
(cat. cit.); Kertész 1910: 160 (cat. cit.);
Fluke 1956: 154 (cat. cit.).

Ocyptamus giganteus: Thompson et al.
1976: 19 (cat. cit.).

Male.—Head: Mainly black; face black,
with narrow yellow medial vitta, white pol-
linose except narrowly shiny medially, with
pollinosity denser along eye margin becom-
ing sparser medially, black pilose; gena
black, shiny, bare; frontal triangle black ex-
cept narrowly yellow medially along eye
margin, black pilose, black pollinose except
white pollinose on yellow areas; lunule
black except brownish submedially; vertical
triangle black, black pollinose and pilose;
occiput black, black pilose except mesial
row of cilia mainly white and white pilose
on ventral /;; antenna black except slightly
reddish basoventrally on basoflagellomere,
black pilose, elongate, with basoflagellom-
ere about twice as long as wide.

Thorax: Black, black pilose except white
pilose along anterior margin of scutum and

786 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

/ wes i Prt «
hah 4 ; “a f ( ae

J »

LDONZO

21

ie

[

A

¥
x

Saal st eee
whit st ae ; : ed Dia

C
Figs. 20-23.

LDONZO

Immature stages of Ocyptamus tristani. 20, Larva, a, lateral view, b, dorsal view, c, ventral

view. 21, Larva, anterior view. 22, Larval posterior spiracle, posterior view. 23, Puparium, dorsoblique view.

posterior anepisternum; scutum brownish
gray to black pollinose, with whitish to gray
submedial and medial vittae; scutellum
brownish black pollinose, subscutellar
fringe singular, white; pleuron black polli-
nose except katepisternum white pollinose
on dorsal %; calypter black; plumula long;

halter black. Legs: Black except grayish
white basal %4 of mesotibia, black pilose;
metatrochanter with distinct pile tuft; me-
tafemur without long pile. Wing: Dark an-
teriorly, hyaline posteriorly, microtrichose
except bare on anterobasal % of cell CuP;
alula dark, normal, as wide as cell CuP, mi-

VOLUME 102, NUMBER 4

crotrichose; black area covering base of
wing including base of cell CuP, all of cell
BM, most of cell R except posterior to spu-
rious vein on apical %, anterobasal corner
of cell R4+5, extending almost to level of
middle of cell R4+5; brownish orange area
between black area and gray apex; gray
area covering apex of wing, anterior to vein
R4+5, distal to end of vein R1.

Abdomen: Black except yellow basola-
teral triangular macula on 2nd tergum and
narrow yellow fascia on base of 3rd, black
pilose except white pilose on Ist, bases of
2nd and 3rd terga and on yellow areas and
male genitalia. Male genitalia. See Figs. 8—
10.

Female.—Similar to male except for nor-
mal sexual dimorphism and frons black ex-
cept very narrowly yellow on ventrolateral
73, black pollinose sublaterally becoming
gray pollinose medially, black pilose.

Length.—20 mm; wing, 14 mm.

Distribution.—Costa Rica*, Colombia™*,
Venezuela*.

Types.—Baccha giganteus was described
from an unspecified number of specimens
from an unspecified locality in South Amer-
ica. In NMW, there are a male and two fe-
males from Venezuela, Lindig collector, la-
belled as giganteus. The male is here des-
ignated lectotype so as to fix and stabilize
the current concept of the name, and has
been so labelled.

Material examined (3 6 4 ?).—VENE-
ZUELA. Types of giganteus. COSTA
RICA. Puntarenas: Las Alturas, Cerro Chai,
2,100 m, 14 Aug 1995, T. Pape (2 6 USNM
ENT 00037918 NRS, 00030146
USNM); Quijada del Diablo, 3.1 km NE de
Mellizas, 1,800 m, LS 316900_600600, 17
Aug 1997, A. Picado, Lot# 47696 (2 IN-
BIOCRIO02546773 INBIO). COLOMBIA.
Antioquia: Caldas, Oct 1973, R. Velez (2
USNM ENT 00037917 USNM).

Discussion.—Ocyptamus giganteus is
readily recognized by its overall dark col-
oration, black thorax, legs, anterior wing
margin and abdomen except for a single
yellow fascia on the base of the 3rd tergum.

787

Also, true to its name, giganteus is the \arg-
est species of the genus.

Ocyptamus (Mimocalla) nymphaea (Hull)
(Figs. 11-13)

Baccha nymphaea Hull 1943a: 40 Brazil,
Sao Paulo, Campinas (HT 6 USNM);
Hull 1949: 98, 105, 238 (fig. (abdomen
pattern), key ref.); Fluke 1956: 256 (cat.
Cit):

Ocyptamus nymphaea: Thompson et al.
1976: 23.(Cat. Cit.):

Male.—Head: Face orange, shiny except
narrowly white pollinose laterally, white pi-
lose; gena yellow, shiny, bare; frontal tri-
angle orange on posterior % along eye mar-
gins, brownish black anteromedially, black
pilose, shiny anteriorly, dull pollinose pos-
teriorly; lunule orange except black medi-
ally; vertical triangle black, black pilose;
occiput black on dorsal %, orange basally,
densely yellow pollinose, yellow pilose; an-
tenna orange except more brownish orange
on dorsoapical % of basoflagellomere,
densely black pilose; basoflagellomere
elongate, slightly less than twice as long as
wide.

Thorax: Orange except brownish black
medially on scutum; scutum broadly orange
laterally and anterior to scutellum, brown-
ish black and brown-black pollinose medi-
ally, with broad golden pollinose submedial
vitta and a very narrow indistinct brown
pollinose medial vitta on anterior % which
expands into a large triangular golden pol-
linose macula on posterior 4%, long yellow
pilose anteriorly and laterally, short inter-
mixed black and yellow pile elsewhere;
scutellum orange, short sparse black pilose;
subscutellar fringe singular, yellow; pleuron
sparsely white pollinose, yellow pilose; ca-
lypter orange except dorsal margin brown-
ish; plumula absent; halter orange. Legs:
Orange except metatarsus brownish orange,
yellow pilose except black ventral pile tufts
on metacoxa and trochanter and scattered
intermixed black pile on posterior surface
of mesofemur and medially on metatibia.

788

Wing: Hyaline except cell C brownish or-
ange and cell Sc brownish, extensively mi-
crotrichose except bare: cell R posterior to
spurious vein, cell BM, anterobasal % of
cell CuP, cell R1 anterior to spurious vein;
cell R4+5 posterior to spurious vein, nar-
rowly on base of cells DM and CuA1.

Abdomen: Orange and orange pilose on
Ist, 2nd and basal %4 of 3rd segment; black
and black pilose elsewhere. Male genitalia.
See Figs. 11-13.

Female.—Similar to male except for nor-
mal sexual dimorphism and frons black and
shiny on anterior 4%, orange and orange pol-
linose on posterior %4, short black pilose.

Length.—14 mm; wing, 11 mm.

Distribution.—Brazil (Sao Paulo*, Minas
Gerais*).

Type.—Baccha nymphaea Hull was de-
scribed from two reared males from Brazil.
Although Hull headed his description as
‘female,’ the types are males. The holo-
type is preserved in the USNM and has
been examined. The paratype is now in
CNC.

Material examined (5 6 2 ?).—BRA-
ZIL. Minas Gerais: Arceburgo, E Fortaleza,
Dec 1946, Barretto (3 ¢6 2 2 USNM ENT
00037978—82 USNM MZUSP). Sao Paulo:
Campinas, 14 Oct 1939, H. E G. Sauer
(Holotype 6 with puparium USNM ENT
00037983 USNM); Campininha, Mogi
Guacu, 23 Oct 1970, J. W. Boyes (¢
USNM ENT 00022540 CNC).

Discussion.—Ocyptamus nymphaea is
readily recognized by the orange and black
abdomen. The other species of Mimocalla
have the abdomen fasciate or mostly dark.

Ocyptamus (Mimocalla) tristani
Zumbado, new species
(Figs. 1, 5—7)

Male.—Head: Face yellow, with light
orange medial vitta, sparsely white polli-
nose laterally, only densely pollinose nar-
rowly along eye margin, white pilose ex-
cept black pilose ventrad to antenna; gena
yellow, shiny, bare; lunule yellow except
brown medially; frontal triangle yellow ex-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

cept brown along lunule, black pilose; ver-
tical triangle black, black pilose; occiput
black except yellow on ventral /;, densely
yellowish-white pollinose ventrally becom-
ing more golden dorsally, white pilose ven-
trally becoming more golden dorsally; an-
tenna brown except more brownish orange
basoventrally on basoflagellomere and yel-
low on inner side of scape and pedicel,
black pilose; basoflagellomere elongate,
slightly less than twice as long as wide.
Thorax: Mainly yellow with black mac-
ulae; prothorax yellow; scutum black ex-
cept with broad yellow vitta running from
postpronotum to and including anterior %
of postalar callus but narrowly attenuated
dorsad to wing base, with vitta about as
wide as postpronotum, with black areas
black pollinose except for broad golden
pollinose submedial vitta which tapers pos-
teriorly and does not reach scutellum and a
very narrow indistinct brown pollinose me-
dial vitta on anterior % which expands into
a broader golden pollinose vitta at level of
postalar callus, long yellow pilose anteri-
orly and laterally, short intermixed black
and yellow pile elsewhere; scutellum yel-
low except disc blackish, short sparse black
pilose with a few intermixed yellow pili ba-
sally; subscutellar fringe singular, white;
pleuron yellow except black narrowly on
anterior convex surface of posterior anepis-
ternum, ventral % of katepisternum, poste-
riorly on katatergum and anteriorly on an-
atergum, very sparsely white pollinose ex-
cept denser dorsally on katepisternum, yel-
low pilose; plumula absent short, yellow;
calypter yellowish white except dorsal mar-
gin brownish; halter yellow, with capitulum
brownish. Legs: Proleg yellow except
slightly brownish subapically on posterior
of femur, sparsely gray pollinose, yellow pi-
lose; mesoleg yellow except sligthly brown-
ish on base of and subapically on posterior
of femur, yellow pilose except for a row of
black pili on dorsoposterior surface; meta-
coxa brown, long black pilose with inter-
mixed yellow pili laterally; metatrochanter
brown, long black pilose; metafemur yel-

VOLUME 102, NUMBER 4

low except black ventrally and posteriorly
except apically, black pilose except yellow
pilose on yellow area; metatibia yellow ex-
cept brownish medial %, yellow pilose;
metatarsus yellow except brownish-orange
basotarsomere, yellow pilose. Wing: Hya-
line except stigma brownish and cell Sc yel-
lowish orange, microtrichose except bare
cell R posterior to spurious vein on apical
¥, cell BM, anterobasal % of cell CuP, cell
R4+5 posterior to spurious vein, narrowly
on basoposteriorly in cell DM, basomedial
¥% of cell CuAl, anterior to vein A2; alula
microtrichose, normal, as wide as cell CuP.
Abdomen: Dark brown with broad yel-
low fasciae; Ist tergum yellow on basal %,
black apically, yellow pilose basally, black
pilose apically; 2nd tergum brownish black
except for large yellow basolateral trian-
gular macula and medial inverted V-shaped
macula, with basolateral maculae narrowly
separated medially and occupying basal %
of tergal length, with V-shaped macula
slightly beyond middle of tergum and com-
pletely surrounded by black areas, yellow
pilose basolaterally, black pilose elsewhere;
3rd tergum yellow on basal *%, brownish
black apically, yellow pilose on yellow
area, black pilose elsewhere; 4th tergum
yellow on basal 4%, margin and apicolateral
corner, elsewhere black, yellow pilose; 5th
tergum yellow except with broad medial
black fasciate macula, black pilose; geni-
talia yellow, black pilose; Ist sternum yel-
low, yellow pilose; 2nd sternum brown ex-
cept yellow basally, yellow pilose; 3rd ster-
num brown except yellow basal %, black
pilose except yellow pilose basally; 4th
sternum brown, brown pilose; 5th sternum
orange except yellow apically, brown pi-
lose. Male genitalia. See Figs. 5—7.
Female.—Similar to male except for nor-
mal sexual dimorphism and frons black ex-
cept yellow broadly along eye margin, short
black pilose.
Length.—14 mm, wing, 12 mm.
Distribution.—Mexico*, El Salvador*,
Costa Rica*, Colombia*, Venezuela*.
Holotype.—Male, Costa Rica, Heredia,

789

Santo Domingo, Instituto Nacion:! de
Biodiversidad, 1,100 m, LN 217300
526200, 2 Jul 1994, M. A. Zumbado,
voucher# 94-MAZ-28, with puparium
(INBIOCRI001146854), deposited in IN-
Bio, Santo Domingo.

Paratypes (25 6, 25 ?).—MEXICO.
Vera Cruz: Fortin de la Flores, 9 June 1959,
H. E. Evans (6 2 USNM ENT00022536-7
CNC). EL SALVADOR. Santo Tecla, 7 Sep
L970; lipes’ (Aq i2t) USNMRENE
00037962—3 USNM). COSTA RICA. Ala-
juela: Sarchi, 27 Jul 1954, L. A. Salas, ex
larva feeding on Saisseta hemisphaerica on
coffee (2 6 2 USNM ENT 00037957-9
USNM); Upala, Dos Rios, Sect. San Ra-
mon, 620 m, LN 318100 381900, 18 Mar-
13 Apr 1995, EF A. Quesada, lot# 5274 (d
with puparium INBIOCRI002246265 IN-
BIO). Guanacaste: Volcan Cacao, Cerro
Pedregal, 1,000 m, Feb—Apr 1989, I. Gauld
(2 USNM ENT 00037970 MIUCR). He-
redia: same data as holotype (2 INBI-
OCRIO01146855 INBIO, 3 with puparium
INBIOCRIO02153879 INBIO); same local-
ity as holotype, 4 Jul 1994, M. A. Zumbado
(36 with puparium, INBIOCRIO02153880
INBIO); ... 7 July 1994, M. A. Zumbado
(6 with puparium INBIOCRI002153881
INBIO); ... Oct 1994, M. A. Zumbado (34
22 INBIOCRO002153891—5 INBIO);
11-13 Aug 1994, L. Donzo lot# 3317, (¢
INBIOCRIO02112689 INBIO); ... 2 Jul
1994, J. A. Ugalde (d with puparium, IN-
BIOCRI002153889 INBIO); ... 4 Jul
1994, J. A. Ugalde (6 with puparium, IN-
BIOCRIO02153887 INBIO); 13 Jul
1994, J. A. Ugalde (2 with puparium, IN-
BIOCRI002153888 INBIO); Heredia, 1
Apr 1960, A. Morales, “‘feeding on black
scale” (6 USNM ENT 00037960 USNM);
Santo Domingo, 7 Jun 1974, E. Solera (@
USNM ENT 00037961 USNM); Belén, San
Antonio, 950 m, LN 218800 516175, 8 Aug
1996, M. A. Zumbado, reared from scales
on Ocimum basilicum (Lamiaceae) (6 2
with puparia INBIOCRI00215877-8 IN-
BIO). San José: San José, May 1930, J. E
Tristan (2 USNM ENT 0037956 USNM);

790

San Sebastian, 16 May 1988, A. Retana,
‘“Comiendo escama”’ (¢ with puparium, 3
2 USNM ENT 00037964—7 USNM); Cui-
dad Colén, 800 m, Feb 1990, L. Fournier
(2 USNM ENT 00037968 USNM); San
Antonio de Escazt, 1,300 m, Apr 1989, P.
Hanson (6 USNM ENT 00037969
MIUCR). COLOMBIA. Antioquia: Medel-
lin Valley, Apr—Jun 1947, FE L. Gallego (32
USNM ENT 00037939—41 USNM); .
[no date], EF L. Gallego, “larva predator of
aphids’? (2 USNM ENT 00037942
USNM); Bello, 25 Dec 1970, G. Sanchezg,
“larva feeding on Saisetia coffeae on or-
namental palm’ (7 3 5 @ with puparia
USNM ENT 00037943-54 USNM). VEN-
EZUELA. Distrito Federal: Cumbre de Bo-
queron, Frente a Bajo Seco, 1,700 m, 7 Mar
1970, larva predator of Coccus hesperidum
on “‘naranjo” (6 with puparium USNM
ENT 00037955 USNM).

Etymology.—This species is named af-
ter José Fidel Tristan Fernandez (1874-—
1932), the outstanding Costa Rican natu-
ralist who published the first list of the in-
sects of Costa Rica (Tristan 1897) and col-
lected the first specimen of this species
known to us.

Discussion.—This species has been
reared a number of times, and specimens
have been submitted for identification to
the Systematic Entomology Laboratory,
USDA. The species has been reared as a
predator of various scales (Saissetia cof-
faea (Walker), S. oleae (Olivier) and Coc-
cus hesperidum Linnaeus) on a range of
hosts. The determination labels associated
with the specimens in the USNM indicate
that the species was first incorrectly iden-
tified as Baccha bromleyi by Hull and this
mistake was perpetuated by subsequent
specialists (e.g., W. W. Wirth, L. V. Knut-
son) as either bromleyi or new species near
bromleyi. Apparently none of these mis-
identifications were published.

Ocyptamus tristani is similar to capita-
tus but differs by the presence of a yellow
fascia on the 2nd tergum and broader yel-
low fasciae on the 3rd and 4th terga.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Ocyptamus (Mimocalla) willistoni
Thompson
(Figs. 17-19)

Baccha dolosa Williston 1891: 37 (preocc.
Walker 1857) AH* H* Mexico, Guer-
rero, Omiltemi, 8,000 ft.; Amula, 6,000
ft.; Xucumanatlan, 7,000 ft.; Guatemala,
Cerro Zunil, 4,000—5,000 ft. (ST 3
AMNH, BMNH); Aldrich 1905: 355
(cit.); Kertész 1910: 158 (cit)? al
1949: 98, 105, 234 (abdomen pattern,
key ref.); Fluke 1956: 244 (cit.).

Ocyptamus willistoni Thompson in
Thompson et al. 1976: 30 (new name for
dolosa Williston).

Male.—Head: Face yellow, white pol-
linose laterally, shiny medially, white pi-
lose; gena yellow, shiny, bare; frontal tri-
angle broadly yellow along eye margin,
black medially, black pilose, sparsely gray
pollinose medially; lunule yellow except
black medially; vertical triangle black,
black pilose; occiput black, densely white
pollinose, white pilose; antenna black ex-
cept reddish basally on scape and basofla-
gellomere; basoflagellomere about % lon-
ger than wide.

Thorax: Bluish gray to black except
yellow base of scutellum and pre-alar area;
scutum grayish-white pollinose except
with black pollinose submedial and sub-
lateral vittae, long white pilose along an-
terior margin and laterally, short black pi-
lose elsewhere; scutellum yellow on basal
%, black elsewhere, short black pilose;
subscutellar fringe multiple, white; pleuron
sparsely silver pollinose except dense sil-
ver pollinose on dorsal % of katepisternum
and posterior anepisternum, white pilose
except black pilose on anepimeron; calyp-
ter white except black dorsal margin; plu-
mula short; halter yellow. Legs: Coxae
and trochanters black, silver pollinose,
black pilose except white pilose apicola-
terally; pro and mesofemora black except
yellow apex, white pilose except black pi-
lose on anteroapical %; metafemur black,
white pilose except black pilose on apical

VOLUME 102, NUMBER 4

¥%; pro and mesotibiae yellowish white,
white pilose; metatibia black, black pilose;
pro and meso tarsi black except brownish
basal % of mesobasitarsomere, white pilose
on basotarsomere and basally on 2nd tar-
somere, black pilose elsewhere; metatarsus
yellowish white to orange on basal 3 tar-
someres, black apically, white pilose; pile
tufts distinct on metacoxa and trochanter.
Wing: Hyaline except black apex and cell
Sc, extensively bare; microtrichose on api-
calstaror cell Cy celluSc)apical’/.of cell
R1, apicomedial % of cell R2+3, apico-
medial %4 of cell R4+5 and DM, apico-
medial % of cell CuAl, apicoposterior “4
of cell CuP, along posterior edge from api-
cal % of cell CuP to apex; dark apex ex-
tending apically from end of vein R1 and
posterior to vein R4+5; alula hyaline,
bare, normal, as wide as cell CuP.

Abdomen: \st segement black, white pi-
lose, sparsely grayish pollinose; 2nd seg-
ment black except yellow basolaterally,
black pilose except white pilose basolater-
ally; 3rd segment yellow on basolateral %,
black apically, shiny, black pilose on dark
areas, white pilose on yellow areas; 4th
segment black, shiny, black pilose; 5th
segment red except black basomedially on
tergum, black pilose; genitalia red, yellow
pilose, see Figs. 17-19.

Female.—Similar to male except for
normal sexual dimorphism and: frons yel-
low on lateral %4, black elsewhere, sparsely
gray pollinose medially, black pilose; wing
with black area more extensive, including
all of cell C and RI.

Length.—16—17 mm; wing, 12—14 mm.

Types.—Baccha dolosa was described
from 5 specimens collected in southern
Mexico (Guerrero) by H. H. Smith, and
Guatemala by G. C. Champion. A male
specimen labelled ‘‘Omilteme, Guerrero,
8,000 ft., July, H. H. Smith,”’ ‘“‘Biol. Centr.
Amer., Dipt.—Syrphidae, E D. Godman,
O. Salvin, 1903-—51,’’ [yellow cotype la-
bel] and printed Williston determination
label, in the BMNH is here designated lec-
totype to fix and stabilize the current con-

791

cept of the name, and has been so labelled.
A paralectotype is in the AMNH.

Distribution.—Mexico (Guerrero*, Mo-
relos*, Mexico*, Chiapas*), Guatemala
(Williston), Costa Rica*.

Material examined (19 6,8 2 ).—MEX-
ICO. [no data] (2 with puparium USNM
ENT 00037920 USNM); “‘Cuernavaca, 8/
24” [handwritten, illegible], G. Lassman
(2 USNM ENT 00022539 CNC). Chiapas:
Teopisca, 2 miles W of, 4 Aug 1957, J. A.
Chemsak & B. J. Rannells (6¢6 32 USNM
ENT 00022547—-55 CNC). Distrito Feder-
al: Mexico City, Nov 1916, R. Muller (d
USNM ENT 00037919 USNM); Carretera
Pedrevall yAjusco, “Kin 7,56"seo. 1979e
Butze (6 USNM ENT 00028744 UNAM);
we ZorOct 1979. J Butze, Ie Gutierrez (2
36 USNM ENT 00028745, 7 UNAM).
Guerrero: Tuxpan, Torre microondas,
1,650 m, 16 Aug 1984, J. Butze (6 USNM
ENT 00028738 UNAM). Mexico: Teques-
quinahuac, Cerro Tlaloc, 12 Oct 1983, M.
Paz (3 6 USNM ENT 00028742 UNAM);
Tequesquinahauc, Encinar, 12 Oct 1983,
G. Sampedro (6 USNM ENT 00028746
UNAM). Morelos: Tetela del Volcan, 8 Jul
1978, G. Aratz (2 USNM ENT 00028748
UNAM); 6 mi E Cuernavaca, | Sep 1974,
G. Bohart & W. Hanson (3 6 USNM ENT
00037921—3 USNM); Cuernavaca-Cuautla
Highway, Canyon de Lobos, 26 Nov 1965,
M. W. McFadden (2 USNM ENT
00037971 USNM) COSTA RICA. Puntar-
enas: Gravel pit near Las Alturas, 8-57-9N
82-51-1W GPS, 1,600 m, 16 Aug 1995, M.
A. Metz (6 USNM ENT 00021672 USNM);
Las Alturas, Cerro Chai, 2,100 m, 14 Aug
1995, T. Pape (6 USNM ENT 00037984
NRS); Send. a c. Pittier, 1 km de la Estacion,
1,800—2,000 m, LS 331800.577400, 8—20
July 1997, M. M, Moraga Red, Lot#47388
(2 INBIOCRIO02567298 INBIO).

Discussion.—Associated with a pupari-
um in the AMHH are the notes from its
collector (Marston Bates) which declares
the adult was reared “from aphis colony on
Citrus (probably Aphis sp.),” the larva be-

792 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ing a “beautiful rose and blue, looking
much like a limacodid.”’

Ocyptamus willistoni is readily recog-
nized by the lack of yellow pollinose vittae
on the mesonotum, the red apex of the ab-
domen, and the blackish apical wing mac-
ula.

ACKNOWLEDGMENTS

We thank Drs. A. O. Bachmann and Ad-
riana Olvia, Museo Argentino de Ciencias
‘‘Bernardino Rivadavia,’’ Buenos Aires
(MACN); A. Contreras-Ramos, Instituto de
Biologia, Universidad Nacional Aut6noma
de Mexico, Mexico (UNAM); Jeffrey M.
Cumming and J. R. Vockeroth, Canadian
National Collection, Agriculture Canada,
Ottawa (CNC); David Grimaldi, American
Museum of Natural History, New York
(AMNH); Paul Hanson, Museo de Insectos,
Universidad de Costa Rica, San Pedro de
Montes de Oca, San Jose (MIUCR); Steven
Krauth, Department of Entomology, Uni-
versity of Wisconsin, Madison (UWEC);
Ruth Contreras-Lichtenberg, Naturhisto-
risches Museum Wien, Vienna (NMW);
Thomas Pape, Naturhistoriska Riksmuseet,
Stockholm (NRS); Philip Perkins, Museum
of Comparative Zoology, Cambridge
(MCZ); Richard Vane-Wright, The Natural
History Museum, London (BMNH); E C.
do Val, Museu de Zoologia, Universidade
de Sao Paulo, SAo Paulo (MZUSP); for per-
mission to study material in their care. Oth-
er collection acronyms used are USNM for
United States National Museum (= the Na-
tional Museum of Natural History, Smith-
sonian Institution), Washington, D.C. and
INBIO for Institudo Nacional de Biodiver-
sidad, Santo Domingo de Heredia.

We are indebted to our colleague, Jestis
A. Ugalde, for rearing a good portion of the
tristani material (flies and parasites).

We also thank Drs. Stephen D. Gaimari
and Wayne N. Mathis, Smithsonian Insti-
tution, Washington, D.C.; Alexander Kon-
stantinov, Allen L. Norrbom, and Michael
E. Schauff, Systematic Entomology Labo-
ratory, USDA, Washington, D.C. & Belts-

ville, Md., for their critical reviews of the
manuscript. Finally, thanks are due to Ti-
tana Litwak and Leonardo Donzo (figs. 20—
23) for the fine illustrations. Titana Litwak
as copyright holder of the color habitus
hereby makes this image available for non-
commerical and scientific use only.

This work began as a review of the Me-
soamerican species done as a class project
by the junior author under the direction of
the senior, who later expanded the manu-
script to cover the subgenus as a whole.

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PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 794-801

A NEW SPECIES OF TRIASPIS HALIDAY (HYMENOPTERA: BRACONIDAE)
PARASITIC ON THE PEPPER WEEVIL, ANTHONOMUS EUGENI CANO
(COLEOPTERA: CURCULIONIDAE)

ROBERT A. WHARTON AND VICTOR LOPEZ-MARTINEZ

(RAW) Department of Entomology,Texas A&M University, College Station, TX 77843,
U.S.A. (e-mail: rawbaw2@acs.tamu.edu); (VL-M) Colegio de Postgraduados, IFIT-En-
tomologia Km 35.5 carr. México-Texcoco Montecillo, edo. de México C.P. 56230 Mexico

Abstract.—A new species of braconid wasp, Triaspis eugenii, is described. This species
is a parasitoid of the pepper weevil, Anthonomus eugenii, in Nayarit, Mexico. It is being
investigated as part of a classical biological control program directed against this pest in
Florida. Diagnostic features are provided for separating this species from other species of
Triaspis attacking weevil pests. The overall similarity to another pepper weevil parasitoid

in the genus Urosigalphus is also noted.

Key Words:
control, Mexico

The genus Triaspis was first described by
Haliday (1835), and is currently placed in
the tribe Brachistini of the subfamily Hel-
coninae by most workers (Mason 1974,
Sharkey 1997). Though confused with Si-
galphis Latreille for nearly 80 years, the
true identity of Triaspis was fixed in the
early 1900s and has remained stable ever
since (Martin 1955). Triaspis is character-
ized by the presence of a completely fused
carapace and the absence of the r-m cross-
vein in the fore wing. In typical Triaspis,
the carapace retains the two transverse fur-
rows: all that remain of the sutures that
originally separated the three segments
comprising the carapace. Schizoprymnus
Foerster, sometimes recognized as a sepa-
rate genus and sometimes treated as a sub-
genus of Triaspis, is separated from typical
Triaspis by the absence of the two trans-
verse furrows, at least dorsally. Most other
brachistines have at least the first two me-
tasomal segments separated by a flexible
suture. The only exception in the New

Braconidae, Helconinae, Brachistini, pepper weevil, parasitoid, biological

World is the genus Urosigalphus Ashmead,
otherwise distinguished by having the outer
tarsal claw of the hind leg much larger than
the inner claw. In the New World, various
species of Triaspis have been reared from
the same weevil hosts as other brachistines
in the genera Urosigalphus, Nealiolus Ma-
son, and/or Aliolus Say.

The most recent revisions or reviews of
Triaspis include those of Martin (1955) for
North America, Papp (1984) for Australia,
Tobias (1986) for the European part of the
former USSR, Priore and Tremblay (1987)
for Italy, Chou and Hsu (1996) for Taiwan,
and Belokobylskij (1998) for Eastern Rus-
sia. The neotropical fauna has never been
revised and isolated descriptions of only
five species have been published to date
(Shenefelt 1970). A Palaearctic species at-
tacking the pea weevil, Bruchus pisorum
(L.), has also been introduced to the Neo-
tropics (Shenefelt 1970).

In general, members of the genus Trias-
pis are poorly known biologically (Shaw

VOLUME 102, NUMBER 4

and Huddleston 1991). Nevertheless, a few
species have been reared from weevils, bru-
chids, and anthribids, and those that are best
studied oviposit in the host egg and emerge
from late instar larvae. The species de-
scribed here has been reported as the most
abundant parasitoid of the pepper weevil,
Anthonomus eugenii Cano, in the Mexican
state of Nayarit, attaining 50% parasitism
in some of the samples from commercial
hot pepper crops there (Mariscal et al.
1998). The species is currently being tested
for possible use in the biological control of
the pepper weevil in Florida. Information
on the host weevil has recently been sum-
marized by Clark and Burke (1996).

MATERIAL AND METHODS

All of the material used for this study
was collected in Nayarit, Mexico, and was
reared from peppers containing the pest
weevil A. eugenii. The majority of the spec-
imens were collected by Eugenio Mariscal
and by Marco Toapanta and Phil Stansly as
part of a biological control program through
the University of Florida, in cooperation
with Mario Urias of the National Institute
of Agriculture and Forestry Research (IN-
IFAP).

Terminology for the desciption generally
follows Sharkey and Wharton (1997). Ad-
ditionally, the 3 segments of the carapace
are referred to as Tl, T2, and T3. Total
length of ovipositor was determined by dis-
section of 10 individuals, and comparing
the total length with that of the dissected
metasoma. Approximate length of oviposi-
tor is also indicated in the description, as
distance protruding beyond apex of cara-
pace. The carapace is bordered by a thin
flange called the marginal lamella (Martin
1955). Variation in quantitative traits is in-
dicated by a range and mean, based on 10
males and 10 females.

Figures were produced using a scanning
electron microscope (SEM). All SEM im-
ages were shot at 10kv on a JOEL JSM-
5600. Specimens were coated with about 20

795

nm gold, using a Hummer 6.2 sputtering
unit with argon as a source gas.

RESULTS

Triaspis eugenii Wharton and
L6épez-Martinez, new species
(Figs. 1-17)

Holotype female.—Top label = ““MEX-
ICO: Nayarit Mpio. Santiago Ixcuintla,
Puerta de Mango, 24.11.1999 Toapanta &
Stansly”’ Second label = “reared from An-
thonomus eugenii on Serrano pepper
emerged 7.iv.1999°’. Deposited in collec-
tion of Universidad Nacional Aut6noma de
Mexico, Mexico City. Paratypes: 9 females,
10 males, same data as holotype. 25 fe-
males, 26 males, ““MEXICO: Nayarit Bal-
adero 2.11.1997 E. Mariscal’’; 3 females,
“Santiago Ixc. Nay. 23 febrero 1997 E.
Mariscal Chile Serrano 11 msnm Hosp. An-
thonomus eugenii’’. Paratypes deposited in
the entomological collections of the follow-
ing institutions: Texas A&M University,
College Station; The Natural History Mu-
seum, London; Universidad Aut6énoma de
Yucatan, Mérida; Universidad Postgradua-
dos, Montecillos; Universidad Aut6noma
de Nuevo Leon, Monterrey; The Canadian
National Collection, Ottawa; El Colegio de
la Frontera Sur, Tapachula; National Mu-
seum of Natural History, Smithsonian In-
stitution, Washington, DC; and Instituto de
Ecologia, Xalapa.

Diagnosis.—This species is characterized
by the nearly bare mesoscutum and scutel-
lum (Fig. 13), the propodeum with a very
short anterior face and sharply declivous
posterior face (Figs. 10—11), the pattern of
sculpture on the metasoma (Figs. 14-17),
and a clypeus that is twice wider than high
(Figs. 1-2). Triaspis eugenii is most similar
to Triaspis azteca Martin from Mexico and
especially Triaspis vestiticida Viereck from
Peru. The flagellum is uniformly dark in 7.
azteca but the basal flagellomeres are pale
(especially ventrally) in 7. eugenii and most
specimens of T. vestiticida. The mesonotum
is almost uniformly sparsely setose in T.

796 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

se

}

3G tira 5
Figs. 1-6. Head of Triaspis eugenii. 1, Clypeus in frontal view, arrows showing median and lateral lobes.
2, Face in frontal view showing setal pattern. 3, Anterior-dorsal view of clypeus showing rugulose and pitted

surface. 4, Head in dorsal view, arrow showing depression on frons immediately posteriorad antennal base. 5,
Mandible. 6, Lateral view of head.

9)

vestiticida but largely bare in T. eugenii. Of giniensis, the species described here differs
the North American species treated by Mar- in having more extensive sculpture on the
tin (1955), 7. eugenii most closely resem- carapace, and a taller, narrower clypeus
bles Triaspis virginiensis Ashmead and (clypeus ‘“‘transverse’”’ in T. virginiensis,
Triaspis rectangulata Martin. From T. vir- about 0.75 X width of face). From T. rec-

VOLUME 102, NUMBER 4

ee
S36 bre

Figs. 7-9.

197)

Antenna of female Triaspis eugenii. 7, Composite view of flagellum (scape and pedicel absent).

8, Basal 3 flagellomeres showing relative lengths of the 3 segments and patch of short sensilla (arrow) on
flagellomere 1. 9, Flagellomeres 7—9, showing patch of irregular setae and placode sensilla; see Fig. 7 for scale.

tangulata, it differs in antennal coloration
and shape of the head.

Description.—Female. Length: 1.85—2.15
mm.

Head: Polished; somewhat trapezoidal,
slightly narrowed ventrally in frontal view
(gena thus not inflated: Fig. 2); vertex, oc-
ciput, temple, and gena sparsely setose
(Figs. 4, 6), the setae arising from indistinct
punctures; face smooth, more densely cov-
ered with ventrally and ventral-medially di-
rected setae (Fig. 2); frons (Fig. 4) sparsely
setose laterally, bare medially, with shallow
depressions posteriorad scape on either side
of a low, median ridge, depressions weakly
sculptured in part, frons otherwise polished;
eye large, about twice longer than temple
in dorsal view, with temple strongly reced-
ing behind eye; malar space 0.25—0.3 X eye
height, 0.4—0.45 xX height of face; face
1.65—1.85 (m = 1.75) X wider than high;

epistomal suture deeply impressed through-
out; clypeus narrow: width 0.57—0.65 (m =
0.62) X width of face, about twice wider
than high, surface uneven and distinctly
punctate (Fig. 3), ventral margin sinuate:
with weak, broadly rounded median lobe
and a large, very obtuse angulation near the
base of each mandible (Figs. 1, 5), weakly
convex in profile; mandible (Fig. 5) lacking
obvious sculpture; ocelli not surrounded by
an impressed groove; antenna (Fig. 7) with
18-19 segments; first two flagellomeres
equal in length (Fig. 8), flagellomeres 3—9
gradually shortening towards apex, flagel-
lomeres 9 through 14—15 bead-like (Fig. 9),
nearly as wide as long; maxillary palps
short, about equal in length to eye height,
6-segmented; labial palps with only three
apparent segments.

Mesosoma: Pronotum dorsally deeply
pitted along posterior margin, the sculpture

798

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 10-15.

11, Lateral view of thorax and propodeum. 12—13, Dorsal view of mesosoma. 14—15, Dorsal view of metasoma
showing slightly different degrees of development of sculpture.

weakening laterally, extending for a vary-
ing distance ventrally as a crenulate line;
lateral face of pronotum (Fig. 11) bare me-
dially, largely unsculptured in most speci-
mens except at extreme ventral and poste-
rior corners. Mesoscutum with anterior de-
clivity and extreme base of notauli setose
(Fig. 11), with 1—2 rows of scattered setae
along notauli and around lateral margin of
scutum (Figs. 12—13), median and lateral
lobes otherwise bare and highly polished;
notauli very finely crenulate and weakly

Triaspis eugenii. 10, Lateral view of body, showing relative lengths of antenna and ovipositor.

impressed throughout, very narrowly sepa-
rated posteriorly, the area of convergence
completely impunctate (Fig. 13); scutellar
sulcus finely rugulose and with a distinct
midridge; scutellum polished, nearly bare;
mesopleuron (Fig. 11) bare medially, setose
only along anterior and posterior margins,
dorsally along subalar depression, and ven-
trally beneath sternaulus; sternaulus shal-
low, sinuate, unsculptured or nearly so. Me-
tanotum (Figs. 11, 13) short, length along
midline, when viewed in profile (Fig. 11),

VOLUME 102, NUMBER 4

Figs. 16-17.
16, Posterior-lateral view. 17, Posterior view.

equal to or shorter than median carina of
propodeum; metapleuron densely and uni-
formly rugose beneath a mat of short, white
setae. Propodeum (Figs. 10—12) very short
anteriorly, steeply declivous posteriorly;
short-setose throughout, the sculpture read-
ily visible through the setae; with well-de-
fined pentagonal areola occupying roughly
median third of declivity, declivity (includ-
ing areola) otherwise irregularly carinate-
rugulose, the sculpture varying in intensity,
posterior-lateral ridge distinctly protruding.
Hind femur short, broad, 2.85—3.1 (m =
3.0) X longer than wide; tarsal claw with
distinct basal tooth. Fore wing 2.4—2.5 xX
longer than maximal width; basal cell
sparsely setose; 1M distinctly bowed; Rla
1.55—2.1 (m = 1.8) X longer than R1b, the
latter not extending to wing tip; m-cu 1.7—
2.3 (m = 1.9) X longer than (RS+M)b;
lcu-a distinctly postfurcal, 1.15—2.5 (m =
1.5) X longer than 1CUa; 2cu-a absent, 2-
1A usually very short, thus Ist subdiscal
cell broadly open posterior-distally.
Metasoma: Approximately equal in
length to mesosoma. Carapace short and
broadly oval, in dorsal view 1.25—1.35 (m
= 1.27) X longer than maximum width; ex-
tensively sculptured; Tl and T2 carinate
over a finely rugulose and variously pitted
background, the surface of T1+T2 usually
(80%) completely sculptured (Figs. 15—16),
more rarely with sculpture evanescent pos-

799

‘ a Se
EG nt ee ae

afl wit :

Metasoma of Triaspis eugenii showing marginal lamella and apically sinuate carapace margin.

terior-medially adjacent T2/T3 suture (Fig.
14), sculpture usually more strongly cari-
nate laterally, with at least some carinae ex-
tending across the foveolate T1/T2 suture;
T3 polished and unsculptured over medial
0.2—0.4 (Fig. 17), strigose laterally, T2/T3
suture (Figs. 14—15) complete, very broadly
u-shaped, finely crenulate; dorsal carinae of
T1 distinctly elevated above surrounding
sculpture and weakly converging over basal
half, separated by about 0.4 X carapace
width at this point, parallel-sided and grad-
ually weakening over distal half, usually
reaching suture as a pair of simple carinae
similar in appearance to adjacent carinate
sculpture; carapace somewhat truncate pos-
teriorly, with distinct concavity mid-ven-
trally (Fig. 17); marginal lamella (Figs. 10,
16—17) present as a very narrow ridge along
T1 and T2, widening posteriorly on T3, ex-
tending completely around posterior margin
of carapace as a broad, thin flange. Ovipos-
itor 1.15—1.4 (m = 1.25) X longer than me-
tasoma; at rest, extending roughly 0.7 X
carapace length beyond apex (Fig. 14).
Color: Black; tegula mostly black except
outer margin brown; antenna dark brown
with scape, pedicel, and basal 3—6 flagel-
lomeres yellowish ventrally; clypeus vari-
ously dark reddish brown; mandible yellow
except apically; palps pale yellow to whit-
ish; legs patterned as follows: hind coxa
dark brown to black basally and posteriorly,

800

yellow anteriorly and apically, fore and mid
coxae more extensively yellow; trochanters
yellow; hind femur yellow with broad, sub-
apical brown band, band somewhat more
diffuse and largely limited to dorsal and
posterior sides of mid femur, fore femur
yellow with dark brown spot dorsally over
much of basal half; hind tibia gradually
darkening from base to apex, at least basal
fourth dark brown, fore and mid tibiae yel-
low; tarsi brown to light brown dorsally,
yellowish ventrally.

Male.—As in female except face 1.75—
2.0 (m = 1.85) X wider than high; antenna
with 19-20 segments, flagellomeres grad-
ually becoming shorter towards apex, but
subapical flagellomeres longer and usually
not as distinctly bead-like as in female, first
flagellomere shorter, 0.75—0.9 (m = 0.8) X
length of second. Hind femur somewhat
more slender, 2.95—3.55 (m = 3.25) X lon-
ger than maximum width. Carapace dis-
tinctly narrower, in dorsal view 1.3—1.45 (m
= 1.4) X longer than maximum width. Sub-
apical brown band on hind femur often
more diffuse, fore and mid femora more ex-
tensively yellow.

Discussion.—Martin (1955), in his revi-
sion of the North American species, delin-
eated several species groups in Triaspis.
The species described here, however, is not
readily assignable to any of Martin’s infor-
mal groups. To accommodate the neotrop-
ical species in general, species groups will
have to be re-defined using a larger suite of
characters including the propodeal structure
and shape of the clypeus. The sculpture of
the carapace is of value in separating some
species and species groups, but examination
of the large amount of reared material avail-
able to us indicates that caution is needed
when using sculpture alone to identify iso-
lated individuals. In T. eugenii, for exam-
ple, the sculpture underlying the carinations
may vary from rugulose and distinctly
punctate to nearly smooth.

The two previously described neotropical
species that most closely resemble T. eu-
genii have both been reared. Triaspis azteca

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

attacks the bean pod weevil, Apion god-
mani Wagner (Martin 1952, Perez 1985),
and the description of 7. vestiticida was
based on material reared from Anthonomus
vestitus Boheman infesting cotton (Viereck
1912).

Members of the genus Triaspis are su-
perficially similar to members of the genus
Urosigalpus, since both have a carapace-
like abdomen and similar wing venation.
Since both can be reared from the same
weevil hosts (including pepper weevil), it is
important to exercise care when identifying
reared material. As noted above, the species
of Urosigalphus have an unusually large
outer claw on the hind leg.

Etymology.—tThe species epithet is taken
from the only known host, Anthonomus eu-
genil.

ACKNOWLEDGMENTS

We are grateful to Ed Riley and Horace
Burke (Texas A&M University) for infor-
mation on weevil hosts of 7. azteca, Terry
Junek for assistance with the literature, and
Dave Smith (USDA/ARS, Systematic En-
tomology Laboratory) for faciliting exami-
nation of the Triaspis collection at the Na-
tional Museum of Natural History. Phil
Stansly, Marco Toapanta, Eugenio Maris-
cal, José Antonio Sanchez-Garcia, and Jor-
ge Leyva-Vazquez kindly made material
available for the description, and provided
additional information on hosts and the bi-
ological control program. Special thanks to
Matt Buffington (Texas A&M University)
and Jim Ehrman for preparation of the im-
ages. All SEM work was done by Jim Ehr-
man at the Digital Microscopy Facility,
Mount Allison University, Sackville, NB,
Canada. This work was supported in part
by the Texas Agricultural Experiment Sta-
tion. The Texas Parks and Wildlife Depart-
ment is gratefully acknowledged for per-
mission to collect in state parks (permits
39-49, 41-95, and 10-98), from which we
obtained specimens of Triaspis useful for
comparison of species-level characteristics.

VOLUME 102, NUMBER 4

LITERATURE CITED

Belokobylskij, S. A. 1998. 12 Subfamily Brachistinae
(Calyptinae), pp. 440—489. Jn Ler, P. A., ed., Key
to the Insects of Russian Far East, Vol. ITV. Neu-
ropteroidea, Mecoptera, Hymenoptera. Pt. 3.
Vladivostok: Dal’nauka, 708 pp. [In Russian]

Chou, L.-y. and T.-C. Hsu. 1996. The Braconidae (Hy-
menoptera) of Taiwan. 7. Subtribe Triaspina. Jour-
nal of Agricultural Research of China 45: 436—
497.

Clark, W. E. and H. R. Burke. 1996. The species of
Anthonomus Germar (Coleoptera: Curculionidae)
associated with plants in the family Solanaceae.
Southwestern Entomologist Supplement 19: 1—
114.

Haliday, A. H. 1835. Essay on parasitic Hymenoptera
of the Ichneumones Adsciti (cont.). Entomological
Magazine 3: 121-147.

Mariscal Mejorado, E., J. L. Leyva Vazquez, and R.
Bujanos Muniz. 1998. Parasitoides del picudo del
chile, Anthonomus eugenii Cano (Coleoptera:
Curculionidae), en Nayarit, México. Vedalia 5:
39-46.

Martin, J. C. 1952. A new braconid from Mexico. The
Canadian Entomologist 84: 30-31.

. 1955. A taxonomic revision of the triaspidine
braconid wasps of Nearctic America (Hymenop-
tera). Canada Department of Agriculture, Science
Service, Entomology Division. 157 pp.

Mason, W. R. M. 1974. A generic synopsis of Brach-
istini (Hymenoptera: Braconidae) and recognition
of the name Charmon Haliday. Proceedings of the
Entomological Society of Washington 76: 235—
246.

Papp, J. 1984. First survey of the Triaspidini species
of the Indo-Australian region (Hymenoptera: Bra-
conidae, Calyptinae) 1. The genus Triaspis Hali-
day. Acta Zoologica Academiae Scientarum Hun-
garicae 30: 137-158.

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Pérez, G. 1985. Himenopteros parasitoides de Apion
spp. (Coleoptera: Curculionoidea: Apionidae) en
Tepoztlan, Morelos. Folia Entomologica Mexi-
cana 63: 39—46.

Priore, R. and E. Tremblay. 1987. Contributo alla re-
visione delle specie italiane dei generi Triaspis
Haliday e Schizoprymnus Foerster (Hymenoptera
Braconidae). Bollettino del Laboratorio di Ento-
mologia Agraria ‘‘Filippo Silvestri” di Portici 44:
47-61.

Sharkey, M. J. 1997. Subfamily Helconinae, pp. 260—
272. In Wharton, R. A., P. M. Marsh, and M. J.
Sharkey, eds., Manual of the New World Genera
of the Family Braconidae (Hymenoptera). Inter-
national Society of Hymenopterists. Special pub-
lication 1, 439 pp.

Sharkey, M. J. and R. A. Wharton. 1997. Morphology
and Terminology, pp. 19-37. In Wharton, R. A.,
P. M. Marsh, and M. J. Sharkey, eds., Manual of
the New World Genera of the Family Braconidae
(Hymenoptera). International Society of Hymen-
opterists. Special publication 1, 439 pp.

Shaw, M. R. and T. Huddleston. 1991. Classification
and biology of braconid wasps. Handbooks for the
Identification of British Insects 7(11): 1-126.

Shenefelt, R. D. 1970. Braconidae 2: Heliconinae [sic],
Calyptinae, Mimagathidinae, Triaspinae. /n Ferri-
ére, Ch. and J. Van der Vecht, eds., Hymenopter-
orum Catalogus (nov. ed.) 5: 177-306.

Tobias, V. I. 1986. 7. Subfam. Brachistinae (Calypti-
nae). Handbook to the Insects of the European
Part of the USSR, 3, Hymenoptera, Part 4: 158—
180. Akademia Nauk, Leningrad, 501 pp. [In Rus-
sian]

Viereck, H. L. 1912. Contributions to our knowledge
of bees and ichneumon-flies, including the de-
scription of twenty-one new genera and fifty-sev-
en new species of ichneumon-flies. Proceedings of
the United States National Museum 42: 613-648.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 802-815

NEW HOST PLANT AND DISTRIBUTION RECORDS IN MEXICO FOR
ANASTREPHA SPP., TOXOTRYPANA CURVICAUDA GERSTACKER,
RHAGOLETIS ZOQUI BUSH, RHAGOLETIS SP., AND HEXACHAETA SP.
(DIPTERA: TEPHRITIDAE)

MARTIN ALUJA, JAIME PINERO, MAURILIO LOPEZ, CESAR RUIZ, ALBERTO ZUNIGA,
ENRIQUE PIEDRA, FRANCISCO DIAZ-FLEISCHER, AND JOHN SIVINSKI

(MA, JP, ML, CR, AZ, EP) Instituto de Ecologia, A.C., Apartado Postal 63, C.P. 91000
Xalapa, Veracruz, México (MA, e-mail: alujam @ecologia.edu.mx); (FD-F) Departamento
de Desarrollo de Métodos, Campania Nacional Contra las Moscas de la Fruta, 2a Avenida
Sur No. 5 (Altos), C.P. 30700 Tapachula, Chiapas, México; (JS) Center for Medical,
Agricultural and Veterinary Entomology, USDA ARS, 1700 SW 23rd Drive, P.O. Box
14565, Gainesville, FL 32604, U.S.A.

Abstract.—We report the results of a nine-year study (1990-1998), aimed at determin-
ing the distribution of fruit flies (Diptera: Tephritidae) and identify their host plants in
Veracruz, Mexico. Twenty Anastrepha species, Toxotrypana curvicauda and several un-
identified Hexachaeta species were captured in McPhail traps. Among the 20 Anastrepha
species, A. limae, A. dentata, and A. canalis are new records for Veracruz. Out of a total
of 51 species of native, wild plants and exotic, cultivated plants collected in Veracruz,
Mexico (representing 3,736 kg of fruit), 34 were infested by 13 Anastrepha species, 4
plants were infested by 4 Rhagoletis species, 2 plants were infested by Toxotrypana
curvicauda, and one by an undescribed Hexachaeta species. Our observations include
new host plant records for Anastrepha hamata (larvae feed only on seeds), A. bahiensis,
A. chiclayae, A. fraterculus, A. obliqua, T. curvicauda, Rhagoletis zoqui, R. sp. and Hex-
achaeta sp. We present host plant data based on size of fruit and degree of infestation,
provide information on local names and fruiting phenology, and discuss our findings in
light of their practical implications and with respect to the zoogeography of Mexican fruit
flies.

Resumen.—Reportamos los resultados de un estudio de nueve anos (1990-1998), en-
caminado a determinar la distribucidn de moscas de la fruta (Diptera: Tephritidae) e
identificar sus plantas hospederas en Veracruz, México. Veinte especies de Anastrepha,
Toxotrypana curvicauda y varias especies no identificadas de Hexachaeta fueron captu-
radas en trampas McPhail. De estas especies A. limae, A. dentata y A. canalis representan
nuevos reportes para Veracruz. De un total de 51 especies de plantas nativas (silvestres)
y exOticas (cultivadas) colectadas en Veracruz, México (representando 3,736 kg de fruta),
34 fueron infestadas por 13 especies de Anastrepha, otras 4 por 4 especies de Rhagoletis,
2 por Toxotrypana curvicauda y una por una especie no descrita de Hexachaeta. Nuestras
observaciones incluyen nuevas plantas hospederas para Anastrepha hamata (las larvas se
alimentan sdlo de semillas), A. bahiensis, A. chiclayae, A. fraterculus, A. obliqua, T.
curvicauda, Rhagoletis zoqui, R. sp. y Hexachaeta sp. Presentamos informacion sobre
plantas hospederas basada en el tamafio de fruto y grado de infestacidén, proporcionamos
informacion acerca de nombres locales de los frutos, su fenologia de fructificacién y

VOLUME 102, NUMBER 4

803

discutimos nuestros descubrimientos en relaci6n a sus implicaciones practicas y con res-
pecto a la zoogeografia de las moscas de la fruta presentes en México.

Key Words:
zoogeography

Flies in the genera Anastrepha Schiner,
Rhagoletis Loew, Toxotrypana Gerstacker,
and Hexachaeta Loew are widely distrib-
uted in the Neotropics (Norrbom et al.
1999). In Anastrepha, there are 197 de-
scribed species distributed from the south-
ern USA to northern Argentina (Aluja
1994, Norrbom et al. 2000). The host plants
of Anastrepha were recently reviewed by
Norrbom (2000) and Norrbom et al. (2000).
These combined reports indicate that plant
species in more than 75 families are used
as hosts. For Toxotrypana curvicauda Ger-
stacker, host plants are restricted to two
families only: Caricaceae and Asclepiada-
ceae. They include species such as Carica
papaya L., Carica cauliflora Jacq., Morren-
ia odorata Lindl., and Gonolobus sorodius
Gray. (Mason 1922, Castrej6n-Ayala and
Camino-Lavin 1991, Landolt 1994). For
flies in the genus Hexachaeta, almost no
host-plant data are available. This genus in-
cludes 25 described species and many un-
described species, and the few host plant
reports available are predominantly plants
in the family Moraceae (Norrbom et al.
1999).

Here, we report the results of a long-term
study (nine years of year-round fruit collec-
tions and trapping activities) aimed at de-
termining the distribution of the fruit fly
genera Anastrepha, Rhagoletis, Toxotry-
pana, and Hexachaeta and identifying their
host plants in the State of Veracruz, Mexi-
co.

MATERIALS AND METHODS

Study sites.—During nine years (1990-—
1998) we collected all wild or cultivated
fruit (potential fruit fly hosts) we could find
in 42 sites located in the state of Veracruz,

Anastrepha, Rhagoletis, Toxotrypana, Hexachaeta, distribution, host plants,

Mexico. Two of these 42 sites are located
in northern Veracruz (Morgadal and Ala-
mo), one site is located in southern Vera-
cruz (Los Tuxtlas), and the rest (39 sites)
are located in central Veracruz. All study
sites, their exact location (latitude, longi-
tude) and altitude are described in Table 1.

Fruit sampling methods.—Ripe, and on
occasion unripe, fruits of the plants listed
in Table 2, were collected in areas with un-
perturbed and perturbed native vegetation,
in backyard gardens, and in semicommer-
cial and commercial orchards as described
by Aluja et al. (1987). Samples were
placed in plastic washbowls and transport-
ed to our laboratory in Xalapa, Veracruz,
Mexico. There, samples were individually
weighed and placed in plastic trays which
in turn were placed over a plastic wash-
bowl containing vermiculite as a pupation
medium.

Processing of larvae and pupae.—We in-
spected, on a daily basis, the pupation me-
dium in the washbowls described above and
collected all pupae and larvae. These were
transferred to labeled, 500 ml plastic con-
tainers with vermiculite. All containers (one
per sample) were kept in the laboratory at
26 + 2°C and 60-70% RH until all adult
flies had emerged from the pupae.

Trapping of adult flies —Traps to survey
adult flies were placed in Apazapan, Llano
Grande, Monte Blanco, and Martinez de la
Torre, Veracruz. With the exception of Mar-
tinez de la Torre, we worked in commercial
mango orchards surrounded by patches of
native vegetation, coffee plantations, and
other orchards. In Martinez de la Torre we
worked in a commercial grapefruit (Citrus
paradisi Macfad.) orchard. At all sites we
placed 16 McPhail traps baited with 250 ml

804

of a mixture of hydrolyzed protein, water,
and borax in each orchard. In Apazapan we
also placed 16 traps in a commercial chi-
cozapote (Manilkara zapota [L.] P. Royen)
orchard adjacent to the mango orchard. Bait
preparation, and placement and service of
traps are described in detail in Aluja et al.
(1996).

Identification of adult flies and plants.—
All flies emerging from puparia or caught
in McPhail traps were identified at the In-
stituto de Ecologia, A. C., by Vicente Her-
nandez-Ortiz. Plants were dried, pressed,
and identified by Carlos Duran, staff mem-
ber of the IXAL herbarium at the Instituto
de Ecologia, A. C., in Xalapa, Veracruz.
Voucher specimens of plants and insects not
already represented in the permanent col-
lections were kept in the IXAL herbarium
and IXAL permanent insect collection. All
plant names used here correspond to those
listed in the USDA’s GRIN online database
(USDA 2000).

RESULTS

Fruit sampling.—A total of 3,736 kg of
fruit, representing 51 species from 22 plant
families, was processed during this study
(Table 2). Of these, 42 species from 17 fam-
ilies were found to be natural hosts of 13
Anastrepha species, 4 Rhagoletis species,
Toxotrypana curvicauda, and an undescri-
bed Hexachaeta species (Table 3). The
fruiting phenology of all of these plant spe-
cies is described in Table 4.

The more polyphagous species were A.
ludens (Loew) (8 plant species attacked),
followed by A. obliqua (Macquart) and A.
fraterculus (Wiedemann) with 7 plant spe-
cies attacked. In Apazapan we found Myr-
ciaria floribunda (H. West ex Will.) O.
Berg (Myrtaceae) simultaneously infested
by three species of Anastrepha: A. frater-
culus, A. obliqua, and A. bahiensis Lima
(new host plant records for the latter two
species). We were also able to confirm that
Terminalia catappa L. (Combretaceae) is a
natural (i.e., field-infested) host plant of A.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

fraterculus (first report in the State of Ve-
racruZz).

In contrast to the polyphagous A. obli-
qua, A. ludens and A. fraterculus, A. aphel-
ocentema Stone and A. alveata Stone were
monophagous species attacking Pouteria
hypoglauca (Standley) Baehni (Sapotaceae)
and Ximenia americana L. (Olacaceae), re-
spectively. Anastrepha hamata (Loew) was
an oligophagous species attacking seeds of
both Chrysophyllum mexicanum Brandegee
ex. Stand. and Pouteria campechiana
(Kunth) Baehni fruits, belonging to the
family Sapotaceae. Rhagoletis zoqui Bush,
R. pomonella Walsh, R. turpiniae Hernan-
dez and R. sp. adults were obtained from
Juglans pyriformis Liebm., Crataegus mex-
icana DC., Turpinia insignis (H.B. & K.)
Tul., and Solanum ionidium Bitter, respec-
tively. Toxotrypana curvicauda adults were
obtained from Gonolobus niger (Cav.) R.
Br. (Asclepiadaceae) fruits collected in the
Llano Grande region. Hexachaeta sp. adults
attacked Trophis mexicana (Liebm.) Bur.
(Moraceae) fruits in the Monte Blanco re-
gion (details of all the above are in Table
5).

Infestation patterns (i.e., number of lar-
vae per kg of fruit) are described in Table
5. Notably, the smaller, wild fruits were the
most infested (Table 5, Fig. 1).

The highest infestation levels (larvae/kg
of fruit) were found in Spondias mombin
L., followed by Ximenia americana L.
(fruits in the families Anacardiaceae and
Olacaceae attacked by A. obliqua and A. al-
veata, respectively), while the lowest infes-
tation levels were found in Jnga jinicuil G.
Don (Leguminoseae), infested by A. dis-
tincta Greene, followed by Tabernaemon-
tana alba Mill. (Apocynaceae) attacked by
A. cordata Aldrich (Table 5).

Trapping.—We captured 20 Anastrepha
species in McPhail traps. Of these, 16 spe-
cies were captured in Apazapan, 12 species
in Llano Grande, 14 in Monte Blanco and
11 in Martinez de la Torre (Table 6). In the
mango orchards, A. obliqua was by far the
most abundant species, followed by A. lu-

VOLUME 102, NUMBER 4 805
Table 1. Study site locations.
Northern Western Altitude
No. Study site latitude longitude (meters)
1 El Conchal 19°04’ 96°06’ 17
D La Mancha 19°35’ 96°22’ 18
3 Cardel 193227 96°22’ 31
4 Playa Azul 19D 96°23’ 47
5 Playa Oriente LOPE 96°19’ 121
6 Paso de Ovejas 19°17’ 96°26’ 122
oT Morgadal (Poza Rica) 20°24’ 97°21’ 128
8 Alamo OSS) 97°40’ 180
9 Puente Nacional 19°19’ 96°28’ 198
10 Los Tuxtlas 18°25’ 95°06’ 200
11 Coyolar Ome 96°36’ 228
12 El Aguaje 1925! 96°36’ 250
13 El Crucero Om We 96°35’ 281
14 Actopan 19°30’ 96°36’ Sui
i) Plan del Rio 1hOF23% 96°39’ 319
16 Rinconada 1S 2 96°33’ B22,
17 Apazapan 19°19’ 96°42’ 331
18 Martinez de la Torre 19°58’ 96°47’ 400
19 Emiliano Zapata eal 96°33’ 416
20 Cerro Gordo 19"25¢ 96°41’ 559
ay Llano Grande 19°22’ 96°53’ 680
22 Corral Falso 19°27’ 96°45’ 744
23 San Pedro 19°24’ 96°52’ 904
24 Tejeria 19°22’ 96°56’ 924
25 Miradores 19°28’ 96°46’ 966
26 Limones 19°20’ 96°55’ 1,021
DF Monte Blanco 19°23’ 96°56’ 1,026
DS Bella Esperanza 19°26’ 96°52’ 1,057
29 Vaqueria 19°19" 96°57’ 1,103
30 Coatepec 19°26’ 96°57’ 1,142
Sil Xico 19°25’ 97°00’ EAS)
32 Teocelo 19°24’ 96°58’ 1,163
33 Jilotepec 19°37 96°56’ 1,292
34 Cosautlan 19°20’ 96°59’ 1,318
35 Texin 19°20’ 97°01’ 1,464
36 Xalapa LOSSY 96°54’ 1,468
37] Amatla 19°19’ 97°03’ 1,478
38 El Diez 9 2ie 97°01’ 1,498
39 La Perla 19°19’ 97°03’ 1,578
40 El Arenal 19°19’ 97°05’ 1,649
41 Acajete 19°34’ 96°59’ 1,875
42 La Joya esr 97°02’ 2,004

dens (Monte Blanco and Llano Grande) and
A. serpentina (Wiedemann) (Apazapan). At
Martinez de la Torre (citrus orchard), A. lu-
dens was the most common Anastrepha
species captured. All of these species are
considered agricultural pests of economic
importance in Mexico.

Many Anastrepha species of non-eco-

nomic importance were also captured at all
four trapping sites (e.g., A. bicolor [Stone],
and A. spatulata Stone). In contrast, A. bah-
iensis, A. dentata (Stone), A. robusta
Greene, and A. zuelaniae Stone were found
only in Apazapan, while A. canalis Stone
was captured only in Monte Blanco. Adults
of Hexachaeta spp. were commonly cap-

806

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Plant species sampled in central Veracruz, Mexico, to determine their host status to the local tephritid

fruit flies.

Plant family

Scientific name

Local common name

Locality code*

Anacardiaceae

Annonaceae
Apocynaceae
Asclepiadaceae
Bombacaceae

Caricaceae
Combretaceae
Euphorbiaceae
Guttiferae
Juglandaceae
Leguminoseae

Malpighiaceae
Moraceae

Myrsinaceae
Myrtaceae

Olacaceae

Passifloraceae

Rosaceae

Rutaceae

Mangifera indica L.

var. criollo

var. manila

var. Kent
Spondias mombin L.

Spondias purpurea L.

Spondias radlkoferi Donn. Sm.

Spondias sp.

Tapirira mexicana Marchand

Annona muricata L.

Tabernaemontana alba Mill.

Gonolobus niger (Cav.) R. Br.

Quararibea funebris (La Llave)
Vischer

Carica papaya L.

Terminalia catappa L.

Ricinus communis L.

Mammea americana L.

Juglans pyriformis Liebm.

Inga jinicuil G. Donht.

Inga spuria Humbl.

Inga vera Willd.

Byrsonima crassifolia (L.) Kunth

Trophis mexicana (Liebm.) Bur.

Brosimum alicastrum Sw.

Icacorea compressa (Kunth) Standley

Myrciaria floribunda (H. West ex
Willd.) O. Berg

Psidium guajava L.

Psidium guineense Sw.

Psidium sartorianum (O. Berg.) Nied.

Syzygium jambos (L.) Alston
Ximenia americana L.

Bunchosia biocellata Schlecht.
Passiflora ciliata Aiton

Passiflora edulis Sims.

Passiflora edulis f. flavicarpa Deg.
Cordia dodecandria Sesse & Moc.
Crataegus mexicana DC.
Eriobotrya japonica (Thunb.) Lindl.
Prunus persica (L.) Batsch
Casimiroa edulis La Llave & Lex.
Citrus maxima (Burm.) Merr.
Citrus reticulata Blanco

Citrus paradisi Macfad.

Citrus sinensis (L.) Osbek

var. valencia
var. navel

Mango corriente
Mango manila
Mango petacon
Jobo

Ciruelo tropical

Jobo Cimarron
Cundoria
Cacao silvestre
Guanabana
Huevo de gato
Vaquitas
Canela

Papaya
Almendro
Higuerilla
Zapote domingo
Nogal

Jinicuil

Chalahuite peludo
Chalahuite
Nanche

Jobo Cimarron
Ramon (Ojite)
Capulin silvestre
Guayabilla

Guayaba

Guayaba acida
Guayaba Tejon
Pomarrosa
Ciruela de monte
Granada de arbol
Granada roja
Granada amarilla
Maracuya

Copite

Tejocote

Nispero

Durazno

Zapote Blanco
Pomelo
Mandarina
Toronja

Naranja Valencia

Naranja Ombligona

1-9, 11-12, 14-15, 17,
19-21, 24, 27

1-3, 5—6, 10, 16-17,
19-21, 23, 24, 27-28

1-6, 9-11, 13-17, 19—
223

24, 27

3235.25

27, 30

5, 16-17

10

PONS PST

10

ts WS), 17

Dy Sy MS

21

2

34, 36

2, 233 24, 20) 2 ee
30, 34

21, 24, 27

21, 24, 27

=H Nnnnu

SEE Sse
—

by

S)

1-6, 9-17, 19-34, 36—

21, 24, 27, 32, 36
215 24.27 So

2. 18: 235255355
5; 6; LO sss OS iee2 ete
26, 27, 29, 32, 34, 36

VOLUME 102, NUMBER 4

Table 2. Continued.

Plant family Scientific name

807

Local common name Locality code*

Citrus aurantium L.

1, 9, 14-16, 19-21, 23-
DD), AM, Xd SO), Dil.

Naranja Cucha

33, 40
Sapotaceae Bumelia spiniflora A. DC. Pionche 17
Calocarpum mammosum (L.) Zapote Mamey D2 23524 202i
P. Royen
Chrysophyllum mexicanum Zapote Nino 10
Brandegee ex. Stand.
Chrysophyllum cainito L. Caimito 8
Manilkara zapota (L.) P. Royen Chico Zapote 14-17, 19
Pouteria hypoglauca (Standl.) Baehni Zapote calentura DT
Pouteria campechiana (Kunth) Baehni Zapote Nifio DY
Solanaceae Solanum ionidium Bitter Tomatillo 36
Staphyleaceae Turpinia insignis (H.B. & K.) Tul. Turpinia 36

* Places in which plant species were sampled. Numbers correspond to those presented in Table 1.

tured in McPhail traps in all four trapping
sites. Toxotrypana curvicauda was captured
only in Apazapan, Veracruz.

DISCUSSION

We discovered new host plants for An-
astrepha bahiensis, A. hamata (2 plant spe-
cies), A. chiclayae Greene, A. fraterculus,
and most significantly, for the economical-
ly-important species, A. obliqua. New host
plants are also reported for Toxotrypana
curvicauda, Rhagoletis zoqui, R. sp. and
Hexachaeta sp. We were further able to
document, for the first time, the presence of
A. canalis, A. dentata, and A. limae Stone
in the State of Veracruz, Mexico.

Of the 17 species of Anastrepha reported
by Hernandez-Ortiz (1992) in the state of
Veracruz, we found all in fruits, or captured
in McPhail traps. In addition, we also found
the previously unrecorded A. alveata (Pie-
dra et al. 1993), A. bicolor, A. canalis, A.
dentata, and A. limae.

Of the new records found for the genus
Anastrepha, A. hamata stands out, not only
because this is the first report of its host
plants, but also because the larvae were
found feeding only on seeds of two fruit
species. Thus, this species is added to the
list previously published by Hernandez-Or-
tiz and Aluja (1993), in which A. cordata,

A. sagittata (Stone), A. pallens Coquillett,
A. crebra Stone, A. anomala Stone, and A.
pickeli Lima are reported as seed-feeding
species. We note however, that A. cordata
and A. crebra also apparently feed in the
mesocarp of fruits (Norrbom et al. 2000).
Anastrepha bahiensis, A. fraterculus, and
A. obliqua were found simultaneously at-
tacking the previously unreported host Myr-
ciaria floribunda (local common name:
Guayabilla) (Myrtaceae), in the Apazapan
region. We wonder if these species are un-
able to recognize the host marking phero-
mone of their congeners or if the extremely
scarce supply of alternative host plants dur-
ing the time of year when this plant is in
fruit, causes flies to oviposit in previously
utilized and marked fruit (due to a large eg-
gload). For A. fraterculus and A. obliqua, it
is relevant to mention that this plant can
function as an alternative host from March
until May. This could allow A. fraterculus
to bridge the period when its previously re-
ported hosts are unavailable (Psidium spp.
prior to May, and S. jambos [L.] Alston
from May to July). In the case of Anastre-
pha obliqua, we believe that this host could
also serve as a “‘bridge’’ between the fruit-
ing periods of two wild hosts (Tapirira
mexicana Marchand, which locally bears
fruits from September to November, and

808 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 3. Native and exotic host plants that harbored fruit flies in Veracruz, Mexico, during our nine year
study (1990-1998).

Common local name

Scientific name

Fruit fly species of host plant of host plant Plant family
Anastrepha
alveata Ciruela de monte Ximenia americana Olacaceae
aphelocentema Zapote calentura Pouteria hypoglauca Sapotaceae
bahiensis Guayabilla* Myrciaria floribunda Myrtaceae
Ramon or Ojite Brosimum alicastrum Moraceae
chiclayae Granada amarilla Passiflora edulis Passifloraceae
Granada roja Passiflora ciliata Passifloraceae
cordata Huevo de gato Tabernaemontana alba Apocynaceae
crebra Canela Quararibea funebris Bombacaceae
distincta Jinicuil Inga jinicuil Leguminosae
Chalahuite peludo Inga spuria Leguminosae
Chalahuite Inga vera Leguminosae
fraterculus Almendro** Terminalia catappa Combretaceae
Guayabilla Myrciaria floribunda Myrtaceae
Guayaba Psidium guajava Myrtaceae
Guayaba acida Psidium guineense Myrtaceae
Guayaba tejon Psidium sartorianum Myrtaceae
Pomarrosa Syzygium jambos Myrtaceae
Durazno Prunus persica Rosaceae
hamata Zapote Nino* Chrysophyllum mexicanum Sapotaceae
Zapote Nino* Pouteria campechiana Sapotaceae
ludens Mango Mangifera indica Anacardiaceae
cultivar Criollo
cultivar Manila
cultivar Kent
Durazno Prunus persica Rosaceae
Zapote Blanco Casimiroa edulis Rutaceae
Naranja cucha Citrus aurantium Rutaceae
Pomelo Citrus maxima Rutaceae
Toronja Citrus paradisi Rutaceae
Mandarina Citrus reticulata Rutaceae
Naranja dulce Citrus sinensis Rutaceae
obliqua Mango Mangifera indica Anacardiaceae
var. Manila
var. Kent
var. Criollo
Ciruelo Spondias purpurea Anacardiaceae
Cundoria Spondias sp. Anacardiaceae
Jobo cimarron Spondias radlkoferi Anacardiaceae
Jobo Spondias mombin Anacardiaceae
Cacao Tapirira mexicana Anacardiaceae
Guayabilla* Myrciaria floribunda Myrtaceae
serpentina Zapote mamey Calocarpum mammosum Sapotaceae
Zapote nino Chrysophyllum mexicanum Sapotaceae
Caimito Chrysophyllum cainito Sapotaceae
Chico Zapote Manilkara zapota Sapotaceae
Zapote calentura Pouteria hypoglauca Sapotaceae
striata Guayaba Psidium guajava Myrtaceae
Guayaba acida Psidium guineense Myrtaceae

Guayaba tejon

Psidium sartorianum

Myrtaceae

VOLUME 102, NUMBER 4

Table 3. Continued.

809

SsSesas"\VTQVWVWKK.® 5

Common local name

Fruit fly species of host plant

Rhagoletis
pomonella Tejocote
turpiniae Turpinia
sp. Tomatillo
zoqui Nogal

Hexachaeta

(undescribed species) Jobo Cimarr6n*

Toxotrypana
curvicauda Papaya

Vaquitas*

Scientific name

of host plant Plant family

Rosaceae
Staphyleaceae
Solanaceae
Junglandaceae

Crataegus mexicana
Turpinia insignis
Solanum iodinium
Juglans pyriformis

Moraceae

Trophis mexicana

Caricaceae
Asclepiadaceae

Carica papaya
Gonolobus niger

* First report for Mexico.
** First report for Veracruz State.

Spondias purpurea L., available from April
to June). Interestingly, M. floribunda is
heavily preyed upon local mammals who
eat practically every available fruit (trees
bear fruit during the peak of the dry season
and thus represent one of the few available
food items for wildlife). Because of this, it
is very difficult to find fruit in the field.

The presence of A. fraterculus in fruits
of Terminalia catappa corroborates the pre-
vious report by Patino (1989) working in
Papantla and Gutiérrez Zamora, Veracruz.
However, we find it noteworthy that it was
never found infesting any of the several Cit-
rus species we sampled.

Unfortunately, we were not able to rear
A. bicolor, A. canalis, A. dentata, A. limae,
A. pallens, A. robusta, A. spatulata, and A.
zuelaniae from host fruits. These species
were only captured in McPhail traps. How-
ever, Hernandez-Ortiz (1992) pointed out
that species belonging to the dentata group
(e.g., A. dentata), daciformis group (e.g., A.
bicolor and A. pallens) and robusta group
(e.g., A. robusta) probably attack plants be-
longing to the family Sapotaceae. Future ef-
forts at discovering their host plants should
thus be directed to all wild species of this
family. According to Stone (1939) and Bak-
er et al. (1944), A. pallens attacks Bumelia
spiniflora A. DC. (Sapotaceae), while No-
trbom (1998) pointed out that it infests

fruits of Sideroxylon celastrinum (Kunth)
T.D. Pennington and S. lanuginosa Michx.
(Sapotaceae). We note that Norrbom (1998)
considers that B. spiniflora was likely mis-
spelled (the correct name should be B. spi-
nosa A.DC.). Should the latter be true, then
B. spiniflora is a synonym of S. celastrinum
(Norrbom 1998). Anastrepha leptozona
Hendel was also not reared from any of the
fruits collected. However, Aluja et al.
(1987) found it in Micropholis mexicana
Gilly ex Cronquist (Sapotaceae), in the state
of Chiapas, and Norrbom and Kim (1988)
report its occurrence in 6 plant families.
Norrbom et al. (1999) also note that A. zue-
laniae, A. limae and A. canalis are associ-
ated with plants in the families Flacourti-
aceae, Passifloraceae and Staphylaceae, re-
spectively.

In the Los Tuxtlas region, Hernandez-Or-
tiz and Pérez Alonso (1993) reported the
presence of 13 species of Anastrepha. Of
these, they hypothesized that A. crebra, A.
minuta and Anastrepha sp. (close to A. per-
dita) are apparently restricted in Mexico, to
tropical, evergreen rain forests. Our results,
indicating that these species are only pre-
sent in the tropical decidous forests of cen-
tral and northern Veracruz, lend support to
this hypothesis.

For Toxotrypana curvicauda, our report
corroborates the previous records of Mason

810 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 4. Fruiting phenology of fruit fly host plants in Veracruz, Mexico.

Month
Plant species J F M A M J J A S O N D

Anastrepha host plants

Brosimum alicastrum 4
Calocarpum mammosum

xx xX
xx x
x xX
x
*

Casimiroa edulis
Chrysophyllum mexicanum x xX
Chrysophyllum cainito xX x

Citrus aurantium

*
~

~ x

Citrus maxima
Citrus paradisi

x MK xX

Citrus reticulata

xx KK XM
*
xx KK XK
xa KKM

Citrus sinensis
Inga jinicuil Xx Xx Ȣ
Inga spuria
Inga vera
Mangifera indica
cultivar Criollo
cultivar Manila x
cultivar Kent
Manilkara zapota
Myrciaria floribunda
Passiflora edulis
Passiflora ciliata
Pouteria hypoglauca x x x
Pouteria campechiana x x x
Prunus persica x x xX xX
Psidium guajava
0-500* x Xx x x
500—1,000*
1 ,000—1,500*
Psidium guineense

~*~ xX

xx KK
xx K KK
x x x x
x x*
x mK KM

x mK x

~*~ x

~*~
*

x x
~ x
xxx

Psidium sartorianum
Quararibea funebris
Spondias purpurea x x x x

Spondias mombin xX x
Spondias radlkoferi x
Spondias sp. Xx x
Syzygium jambos x
Tabernaemontana alba

xx x
x x*K xX
xx mK mK MK
x mex MK
xx

x x

~*~ x
nK %
*

Tapirira mexicana
Terminalia catappa x xX xX x
Ximenia americana x x x

x xK x
~*~ x

Hexachaeta host plants
Trophis mexicana x xX

Toxotrypana host plants
Carica papaya xX x x x x xX xX xX
Gonolobus niger x xX x

Rhagoletis host plants

Crataegus mexicana

x x
~*~

Juglans pyriformis
Solanum ionidium

mM XM
ax KK

Turpinia insignis

* Meters above sea level.

VOLUME 102, NUMBER 4

S316)

300

2S0

200

100

MEAN NUMBER OF LARVAE/KG. OF FRUIT
ul
Oo

@) 100

300

811

400 500 700

FRUIT WEIGHT

Fig. 1.

(1922), Castrej6n-Ayala and Camino-Lavin
(1991), and Landolt (1994), who pointed
out that adults of this species feed mainly
on fruits belonging to the families Carica-
ceae and Asclepiadaceae. Here we report
this species infesting Gonolobus niger, a
plant also belonging to the family Ascle-
piadaceae.

For Hexachaeta sp., we are the first to
report its host plant as Trophis mexicana.
This confirms that various species belong-
ing to this fly genus develop in fruits of the
Moraceae.

We are able to report a new host plant
(Juglans pyriformis) for R. zoqui. The only
other known host plant record for this fly
species is J. mollis Engelm. (Bush 1966).
Further, we also report for the first time that
Solanum ionidium is a host plant of an un-
described species in the genus Rhagoletis.
This adds further evidence about the im-
portance of the family Solanaceae as hosts
of this genus.

The polyphagy observed in A. ludens (8
plant species belonging to 3 families), A.

Number of larvae per kilogram of fruit as influenced by size of fruit.

obliqua (7 plant species belonging to 2
families) and A. fraterculus (7 species be-
longing to 4 families) has previously been
discussed by Aluja et al. (1987) and Her-
nandez-Ortiz and Aluja (1993). In our opin-
ion, this polyphagy permits these four spe-
cies to be abundant during practically the
whole year. Anastrepha bahiensis was also
found to be a polyphagous species, with lar-
vae feeding on fruits belonging to two gen-
era in two different families (M. floribunda
[Myrtaceae] and Brosimum alicastrum Sw.
[Moraceae]).

In contrast, some species in our study ap-
peared to be monophagous. This is the case
for A. alveata feeding on Ximenia ameri-
cana (Olacaceae) (Piedra et al. 1993), and
A. aphelocentema on Pouteria hypoglauca
(Sapotaceae). Anastrepha aphelocentema
was found only in the northern part of Ve-
racruz (Papantla) (Patifio, 1989), in spite of
the fact that this plant is also present in lo-
calites such as La Mancha, approximately
150 km away from Papantla. In La Mancha,
fruits were attacked only by A. serpentina.

812 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 5. Infestation patterns by Anastrepha flies based on size of host fruit.

eT

Degree of
infestation
Total no. Total no. Fruit fly (larvae/kg
Host plant Kg. sampled of pupae of adults species of fruit)
Large (300—1,000 gr)
Calocarpum mammosum 143.73 3,942 1, TSI A. serpentina 27.43
Citrus paradisi 1,345.71 22,739 19,428 A. ludens 16.90
Mangifera indica
cultivar Kent 56.57 902 ES95
27 A. obliqua
716 A. ludens
Citrus maxima 79.66 853 S52 A. ludens 10.70
Carica papaya 126.50 529 477 T. curvicauda 4.18
Medium (50-300 gr)
Chrysophyllum cainito 11.00 2,210 1,926 A. serpentina 201.00
Prunus persica 10.62 338 31.83
228 A. fraterculus
Si) A. ludens
Manilkara zapota 101.83 3,178 Dab A. serpentina 31.21
Pouteria hypoglauca? 19.05 506 328 A. aphelocentema 26.56
Casimiroa edulis B95) 952 477 A. ludens 24.10
Pouteria hypoglauca' 17.28 369 Dray A. serpentina 21.35
Juglans pyriformis WL ES aV7/ 899 R. zoqui 20.51
Citrus sinensis 264.27 Spyd57/ 2,742 A. ludens 20.27
Citrus aurantium 669.29 1S F503 525 A. ludens 20.20
Mangifera indica
var. criollo 121.78 eZ 14.55
20 A. ludens
995 A. obliqua
Citrus reticulata 2.80 36 25 A. ludens 12.86
Chrysophyllum mexicanum 59.36 531 342 A. hamata 8.94
Mangifera indica
var. manila 42.97 337 7.84
58 A. ludens
222 A. obliqua
Terminalia catappa 10.09 55 10 A. fraterculus 5.45
Inga jinicuil 22.38 Vi 64 A. distincta 3.44
Small (<5O gr)
Spondias mombin 64.67 28,138 4,241 A. obliqua 435.10
Ximenia americana 16.08 4,960 2,082 A. alveata 308.46
Turpinia insignis 12225 3,416 674 R. turpiniae 278.90
Psidium guajava 97.60 16,619 170.28
4,879 A. striata
8,923 A. fraterculus
Spondias sp. 29.20 4,514 2ASS A. obliqua 154.59
Inga vera 0.56 76 76 A. distincta PS S2k
Psidium guineense 4.76 509 106.93
ie A. striata
359 A. fraterculus
Solanum ionidium 0.29 29 11 R. sp. 100.00
Psidium sartorianum 3.62 282 77.90
47 A. striata
43 A. fraterculus
Tapirira mexicana 14.93 1,060 325 A. obliqua 71.00
Spondias radlkoferi TST 384 131 A. obliqua 50.72

nn

VOLUME 102, NUMBER 4 813

Table 5. Continued

Degree of

infestation

Total no. Total no. Fruit fly (larvae/kg

Host plant Kg. sampled of pupae of adults species of fruit)

Inga spuria 10.52 495 373 A. distincta 47.05

Crataegus mexicana 22.50 986 559 R. pomonella 43.83

Spondias purpurea 182.34 6,170 ils} A. obliqua 33.84

Passiflora ciliata 0.84 24 18 A. chiclayae 28.57

Syzygium jambos DN (9) 615 389 A. fraterculus 28.01

Quararibea funebris 3.26 89 67 A. crebra 2-30

Trophis mexicana 0.21 5) 72 Hexachaeta sp. 23.81

Passiflora edulis 0.67 10 A. chiclayae 14.93

Tabernaemontana alba 16.43 62 49 A. cordata Seyi

Gonolobus niger 5.00 “3 ce T. curvicauda —

Brosimum alicastrum ca. 0.30 “A A. bahiensis —

Myrciaria floribunda ca. 0.30 2 A. obliqua —

A. fraterculus
A. bahiensis

' Fruit collected in La Mancha.
? Fruit collected in Morgadal (Poza Rica).
* Pupae and adults that emerged were not counted.

Table 6. Fruit fly species captured in McPhail traps
in Central Veracruz, Mexico.

Llano Monte M. de la
Fruit fly species Apazapan Grande Blanco Torre
A. alveata xX xX xX
A. bahiensis x
A. bicolor xX xX xX xX
A. canalis xX
A. chiclayae xX x x xX
A. cordata xX
A. dentata xX
A. distincta xX x 4 xX
A. fraterculus 4 Xx Xx XxX
A. hamata xX xX
A. leptozona x Xx
A. limae xX »4
A. ludens x xX xX xX
A. obliqua x x x x
A. pallens Xx 4
A. robusta xX
A. serpentina xX x x x
A. spatulata x xX Xx X
A. striata xX xX xX
A. zuelaniae Xx
T. curvicauda xX
Hexachaeta spp. Xx Xx x Xx

It is possible that the recent heavy habitat
alteration caused local extinction of this
species.

It is significant that in the Monte Blanco
region (central Veracruz) fruits of Pouteria
campechiana were attacked only by A. ha-
mata (on seeds), whereas in Los Tuxtlas
(southern part of the state), Chrysophyllum
mexicanum fruits were attacked by both A.
serpentina on pulp, and A. hamata on
seeds. It is noteworthy that two species of
Anastrepha can utilize the same host with-
out competition. Pouteria campechiana
seeds and pulp were also reported attacked
by both A. sagittata and A. serpentina, re-
spectively, by Baker et al. (1944). However,
Aluja et al. (1987) reported only A. serpen-
tina larvae feeding on pulp of P. campe-
chiana fruits in the State of Chiapas (Mex-
ico), but no trace of A. sagittata.

In other Anastrepha species, we found
that their host plants belonged to multiple
species of only one genus (stenophagous
species). This was true for A. distincta in
Inga spuria Humbl. and J. vera Willd. (Leg-
uminoseae), for A. striata Schiner in Psi-
dium guajava L., P. guineense Sw. and P.
sartorianum (O. Berg) Nied. (Myrtaceae)

814

and for A. chiclayae in Passiflora edulis
Sims. and P. ciliata Aiton (Passifloraceae).
We also confirm that A. serpentina is an
oligophagous species with larvae found
feeding in four fruit species belonging to
the Sapotaceae (see Hernandez-Ortiz 1992).
Interestingly, A. hamata was also found to
be an oligophagous species (larvae feed on
seeds of 2 plant species belonging to the
Sapotaceae).

Even though trapping efforts were made
in an extended region of central Veracruz,
A. dentata, A. robusta, and A. zuelaniae
were captured only in Apazapan, where the
vegetation has been characterized as low
deciduous forest. In contrast, adults of A.
canalis were captured only in Monte Blan-
co, where the vegetation was originally
comprised of montane cloud forest. Anas-
trepha limae was found in both Llano
Grande and Monte Blanco. This suggests
that the geographic distribution of some An-
astrepha species is restricted to certain
zones, possibly because their host plants are
present only in these zones.

We also found that the timing of capture
of certain monophagous species coincided
with the fruiting periods of their host plants.
For example, A. hamata and A. alveata
adults were captured in McPhail traps al-
most exclusively when their host fruits
were available (P. campechiana and C.
mexicanum, and X. americana, respective-
ly). In our opinion, this is probably due to
an extraordinary capacity of adult survival
for long periods of time. Under laboratory
conditions, A. alveata adults can live up to
431 days (Aluja and Jacome, unpublished
data). Such adult longevities probably allow
adults to lay eggs in the fruits produced the
following year. However, the locations
where the adults of such species persist dur-
ing most of the year are unknown.

Finally, our data on the relationship of
fruit size and larval infestation levels rep-
resents further evidence that small, wild
fruits are more heavily infested than larger,
commercially grown fruits that were for the
most part, recently (on an evolutionary

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

scale) introduced to the Americas. This re-
lationship was recently discussed by Aluja
et al. (2000) who indicated that native flies
have probably still not fully developed the
ability to metabolize the toxic chemicals
that these exotic plants contain. As a result,
and despite the fact that females lay many
eggs in them, few eggs hatch or most larvae
die or develop poorly.

ACKNOWLEDGMENTS

We thank Jestis Reyes Flores for having
encouraged us to do this study, Vicente
Hernandez-Ortiz for identifying all fly spe-
cies reported here, and Isabel Jacome, Anita
Sanchez, Braulio Cérdoba, Guadalupe Tru-
jillo, Gloria Lagunes, and Anabel Davila
(all Instituto de Ecologia, A.C., Xalapa, Ve-
racruz, Mexico) for technical assistance. We
also thank Leticia Lagunes, Pedro Cérdoba,
Augusto Martinez, Faustino Cabrera and
the Bigurra-Armida family for allowing us
to collect fruits and install McPhail traps in
their orchards. Thanks are also due to Car-
los Duran, and Sergio Avendano (Herbari-
um IXAL, Instituto de Ecologia, A.C.) for
identifying plant material and to Diana
Pérez-Staples for reviewing an early ver-
sion of the manuscript. Useful critical re-
views were also made by A.L. Norrbom
and an anonymous referee. Partial finan-
cial support was provided by the Consejo
Nacional de Ciencia y Tecnologia
(CONACyT) (Grant D111-903537), the
Comisi6n Nacional para el Conocimiento y
Uso de la Biodiversidad (CONABIO)
(Grant FB325/H296/96), the United States
Department of Agriculture-Agricultural Re-
search Service (Agreement 58-6615-3-
025), the Sistema Regional de Investigaci6n
del Golfo de México (SIGOLFO) (Grant
96-01-003-V) and the Campafia Nacional
Contra las Moscas de la Fruta Mexico Se-
cretaria de Agricultura, Ganaderia y Desar-
rollo Rural—Instituto Interamericano de
Cooperacion para la Agricultura (SAGAR-
IICA).

VOLUME 102, NUMBER 4

LITERATURE CITED

Aluja, M. 1994. Bionomics and Management of An-
astrepha. Annual Review of Entomology 39:
151-174.

Aluja, M., J. Pinero, I. Jacome, E Diaz-Fleischer, and
J. Sivinski. 2000. Behavior of flies in the genus
Anastrepha (Trypetinae: Toxotrypanini), pp. 375—
406. In Aluja, M. and A.L. Norrbom, eds., Fruit
Flies (Diptera: Tephritidae): Phylogeny and Evo-
lution of Behavior. Boca Raton, FL, USA. CRC
Press, 944 pp.

Aluja, M., J. Guillén, G. de la Rosa, M. Cabrera, H.
Celedonio-Hurtado, P. Liedo, and J. Hendrichs.
1987. Natural host plant survey of the economi-
cally important fruit flies (Diptera: Tephritidae) of
Chiapas, Mexico. Florida Entomologist 70: 329—
338.

Aluja, M., H. Celedonio-Hurtado, P. Liedo, M. Ca-
brera, E. Castillo, J. Guillén, and E. Rios. 1996.
Seasonal population fluctuations and ecological
implications for management of Anastrepha fruit
flies (Diptera: Tephritidae) in commercial mango
orchards in Southern Mexico. Journal of Econom-
ic Entomology 89: 654—667.

Baker, A. C., A. C. Stone, C. C. Plummer, and M.
McPhail. 1944. A review of studies on the Mex-
ican fruit fly and related Mexican species. USDA-
APHIS Miscellaneous Publication No. 531. 154
Pp.

Bush, G. L. 1966. The taxonomy, cytology, and evo-
lution of the genus Rhagoletis in North America
(Diptera: Tephritidae). Bulletin of the Museum of
Comparative Zoology (Harvard University) 134:
431-562.

Castrejon-Ayala, F and M. Camino-Lavin. 1991. New
host plant record for Toxotrypana curvicauda
(Diptera: Tephritidae). Florida Entomologist 74:
466.

Hernandez-Ortiz, V. 1992. El género Anastrepha Schi-
ner en México (Diptera: Tephritidae). Taxonomia,
distribucion y sus plantas hospederas. Instituto de
Ecologia Publ. No. 33, Xalapa, Veracruz, México,
162 pp.

Hernandez-Ortiz, V. and M. Aluja. 1993. Listado de
especies del genero neotropical Anastrepha (Dip-
tera: Tephritidae) con notas sobre su distribucion
y plantas hospederas. Folia Entomologica Mexi-
cana 88: 89-105.

Hernandez-Ortiz, V. and R. Pérez-Alonso. 1993. The

815

natural host plants of Anastrepha (Diptera: Te-
phritidae) in a tropical rain forest of Mexico. Flor-
ida Entomologist 76: 447—460.

Landolt, P. J. 1994. Mating frequency of the papaya
fruit fly (Diptera: Tephritidae) with and without
host fruit. Florida Entomologist 77: 305-312.

Mason, A. C. 1922. Biology of the papaya fruit fly
Toxotrypana curvicauda in Florida. U.S. Depart-
ment of Agriculture Bulletin No. 1081, 10 pp.

Norrbom, A. L. 1998. A revision of the Anastrepha
daciformis species group (Diptera: Tephritidae).
Proceedings of the Entomological Society of
Washington 100: 160-192.

. 2000. Host plant database for Anastrepha and
Toxotrypana (Diptera: Tephritidae: Toxotrypani-
ni), Diptera Data Dissemination Disk 2.

Norrbom, A. L. and K. C. Kim. 1988. A list of the
reported host plants of the species of Anastrepha
(Diptera: Tephritidae). USDA-APHIS Miscella-
neous Publication No. 81-52, 114 pp.

Norrbom, A., L. E. Carroll, FE C. Thompson, I. M.
White, and A. Freidberg. 1999. Systematic Data-
base of Names, pp. 65-251. Jn Thompson, E C.,
ed., Fruit Fly Expert Identification System and
Systematic Information Database. Myia 9, 524 pp.

Norrbom, A., R. A. Zucchi and V. Hernandez-Ortiz.
2000. Phylogeny of the Genera Anastrepha and
Toxotrypana (Trypetinae: Toxotrypanini) Based
on Morphology. pp. 343-362. In Aluja, M. and A.
Norrbom, eds., Fruit Flies (Diptera: Tephritidae):
Phylogeny and Evolution of Behavior. CRC Press.
Boca Raton, Florida, 944 p.

Patino, J. 1989. Determinacion de las especies de An-
astrepha Schiner (Diptera: Tephritidae) en frutales
y citricos de Papantla y Gutiérrez Zamora, Vera-
cruz, Bsc. Thesis, Universidad Veracruzana, Tux-
pan, Veracruz, México.

Piedra, E., A. Zuniga, and M. Aluja. 1993. New host
plant and parasitoid record in Mexico for Anas-
trepha alveata Stone (Diptera: Tephritidae) Pro-
ceedings of the Entomological Society of Wash-
ington 95: 127.

Stone, A. 1939. A new genus of Trypetidae near An-
astrepha (Diptera). Journal of the Washington
Academy of Science 29: 340—350.

USDA, ARS. 2000. National Genetic Resources Pro-
gram. Germoplasm Resources Information Net-
work—(GRIN). [Online Database] National Germ-
plasm Resources Laboratory, Beltsville, Mary-
land.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 816-830

REVISION OF THE GENUS PARADA HORVATH (HEMIPTERA: TINGIDAE)
WITH CLADISTIC ANALYSIS

ErIc GUILBERT

ESA 8043 du CNRS, Museum National d’Histoire Naturelle,

Laboratoire

d’Entomologie, 45 rue Buffon, F-75005 Paris, France (e-mail: guilbert@mnhn.fr)

Abstract.—Three new species of Parada from New Caledonia are described: P. minuta,
P. paitae and P. majuscula. The monotypic Alloeocysta approba Drake is considered a
junior synonym of Parada and to which its only included species is transferred. The
evolution of certain characters, and the relationships among species of Parada are pre-
sented together with a cladistic analysis. An identification key for these species is also

given.

Key Words:

Horvath (1925) erected Parada as a sub-
genus of Cysteochila Stal for the reception
of Cysteochila (Parada) taeniophora Hor-
vath (1925). Cysteochila has the paranota
widely reflected, inflated, covering the lat-
eral carinae, and the internal margins (« mar-
gine interno ») rounded, while Parada has
the paranota narrowly reflexed, superficially
adpressed onto the pronotum and not cov-
ering the lateral carinae, and the internal
margins slightly sinuate and never rounded
(Horvath 1925). Drake (1942) raised Par-
ada to generic level, and add two new spe-
cies to the genus (P. torta and P. popla).
Later, he added four more new species to
the genus (P. absona, P. darlingtoni and P.
hackeri Drake 1952; P. solla Drake and Ru-
hoff 1961).

The genus presently comprises seven
Australian species (New South Wales and
Queensland) (Drake and Ruhoff 1965).
Three new species from New Caledonia are
described here. In addition, Drake (1961)
described a new genus and a new species
from a single specimen caught in New
South Wales: Alloeocysta approba. This
species, the only included species of the ge-

Parada, Tingidae, new species, evolution, phylogeny, key

nus, shares with Parada the form of the lat-
eral carinae, the paranota and the hood, and
is considered a junior synonym. Alloeocysta
approba Drake is transferred to Parada. In
addition, Cysteochila cubens Guilbert (in
press), described from New Caledonia
shows affinities with the species of Parada
and could be an intermediate species be-
tween these two genera. Among these re-
lationships, the evolution of some charac-
ters getting narrower are discussed through
a cladistic analysis. Deposition of types is
specified: MNHN for Muséum National
d’Histoire Naturelle, Paris; BPBM for
Bernice P. Bishop Museum, Hawaii. All
measurements are in millimeters.

SYSTEMATICS
Parada Horvath 1925

Cysteochila (Parada) Horvath 1925: 3.
Type species: Cysteochila (Parada) tae-
niophora Horvath 1925: 3, by orig. desig.

Parada: Drake 1942: 4.

Alloeocysta Drake 1961: 109. New Syn-
onymy. Type species: Alloeocysta appro-
ba Drake 1961: 109, by orig. desig.

VOLUME 102, NUMBER 4

Diagnosis.—Head with at least two fron-
tal and two occipital spines. Bucculae wide,
with apices meeting in front. Antenna long,
slender, first segment short, second small-
est, third longest and fourth longer than first
two together, pubescent.

Pronotum gibbose, deeply punctuate, are-
olate at apex, pubescent near collar, tricar-
inate. Median carina long, slightly erected,
contiguous to hood. Lateral carinae elevat-
ed, ending before collar, curved inward, not
resting on pronotum, modified as to form a
long inflated areolate cyst. Hood inflated,
raised. Paranota long, wide, reflexed up-
right, not resting on pronotum, three to four
areolae broad, almost reaching lateral cari-
nae.

Hemelytra wider and longer than abdo-
men, slightly wider than pronotum width.
Costal area uni- to biseriate, outer margins
bent upward. Subcostal area bi- to triseriate,
outer margins bent downward. Discoidal
area five to seven areolae broad. Sutural
area large, with larger areolae than subcos-
tal and discoidal areas. Hypocostal laminae
uniseriate.

Parada approba (Drake), new
combination
(Figs. 7—8)

Alloeocysta approba Drake 1961: 109.

Comments.—According to Drake (1961)
Alloeocysta approba is different “from oth-
er Australian genera by having the hood
distinctly wider than long and the lateral ca-
rinae inflated, cyst-like on disc of prono-
tum’”’ (Holotype measurements, body length
2.99; body width 1.26; hood length 0.37;
hood width 0.55). All species of Parada
have lateral carinae inflated like A. approba
and some of them have a hood wider than
long. This is the case of P. majuscula and
P. paitae. Alloeocysta approba has anten-
nae as long as that of the species of Parada
mesOA5: I;.0:12: Ill, 0.85: IV, 0.3); a pron-
otum gibbose, deeply punctuate and areo-
late at apex, a hood forming a vesicule wid-
er than long, lateral carinae modified as to

817

form a subcylindrical areolate cyst that is
closed inward by the internal margins
touching the pronotum, while it is not the
case for the species of Parada. The hem-
elytra are a little longer than abdomen with
a costal area uniseriate, a subcostal area bis-
eriate and a discoidal area four to five are-
olae broad. Despite some characters which
distinguish A. approba from species of Par-
ada; approba being the unique species of
the genus Alloeocysta, the latter is consid-
ered a synonym of the genus Parada. Thus,
I propose the new combination Parada ap-
proba (Drake) n. comb.

Type.—Holotype: ¢, Bogan river, New
South Wales, Australia (Australian Muse-
um, Sydney).

Parada majuscula Guilbert, new species
(Figs. 1-3)

Description.—Head and body black, legs
and antenna yellowish to brown fuscous,
distal part of fourth antennal segment and
tarsi brown to black. Pronotum clear brown
to fuscous. Hemelytra yellowish with three
dark brown spots, one near base that may
occupy just basal third of costal area or be
expanded to cover all of the basal third of
the hemelytron, plus clear brown spots at
apical third and apex of costal area. Length
3.05; width 1.17.

Head with two frontal and two small,
slender occipital spines. Bucculae wide,
triseriate, with apices meeting in front. La-
bium reaching hind coxae and labial sulcus
sinuate. Sternal laminae widely separated
on metasternum. Antennal measurements: I,
0.192 I, 0.12: If, 1.01: IV, 045).. last an-
tennal segment pubescent.

Pronotum gibbose, deeply punctuate, are-
olate on hind process, pubescent near collar,
tricarinate. Median carina long, slightly el-
evated, contiguous to hood. Lateral carinae
elevated, ending before collar, curved in-
ward, not resting on pronotum, modified as
to form a long inflated open cyst, four are-
olae broad. Hood inflated, raised, slightly
wider than long (length 0.33; width 0.38).
Paranota long, wide, reflexed and partially

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

818

1 mm.

Parada majuscula, habitus. Scale

orale

VOLUME 102, NUMBER 4

TON
ac CaS ee

819

? SS

Figs. 2-3.

recurved above pronotum, not resting on
pronotum, four small areolae broad, almost
reaching lateral carinae.

Hemelytra wider, longer than abdomen,
slightly wider than pronotum width. Costal
area bent upward, mostly biseriate but un-
iseriate at apex, outer areolae larger than
inner. Subcostal area bent downward, tris-
eriate. Discoidal area pentaseriate. Sutural
area large, with larger areolae than subcos-
tal and discoidal areas at apex. Hypocostal
laminae uniseriate.

Types.—Holotype: 36, New Caledonia,
Riviere Bleue P7, dense evergreen rainfo-
rest, 24.X.1992, L. Bonnet de Larbogne, J.
Chazeau & E. Guilbert colls (fogging)
(MNHN). Paratypes: 1 2, same data as ho-
lotype (MNHN); 3 6 and 1 2, New Cale-
donia, Yahoue III.1978, N.L.H. Krauss
coll., Acc. #1978.114 (BPBM); 1 2°, New
Caledonia, 6 km N of Paita, 25.1.1963,

Parada majuscula. 2, Profile. 3, Ventral face. Scale = 1 mm.

C.M. Yoshimoto coll. (BPBM); 1 ¢, New
Caledonia, Mt Koghis, 400—600 m, II.1980,
N.L.H. Krauss coll., Acc. #1980.128
(BPBM).

Comments.—The body length of this
species may be sexually dimorphic (Table
1), but this characteristic cannot be assessed
here because of the small number of spec-
imens.

Parada paitae Guilbert, new species
(Figs. 4—5)

Description.—Head and body brown.
Pronotum, hemelytra, legs and antenna
clear to yellowish, apex of last antennal
segment brown fuscous. Length 2.98; width
1.17; hood length 0.31; hood width 0.31.

Same characters as P. majuscula, except
rostrum extends between median coxae but
does not reach hind coxae, costal area is
uniseriate, areolae are rather large, except

820 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Means of body, hood and antennal segments measurements for 5 males and 3 females of P. ma-
Juscula in mm.

el
Body Hood

Antennal Segments
Length Width

Length Width I II Il IV
em
Males POON 1.102 0.32 0.372 1.003 0.366 0.191 0.123
Females 3.149 L27Z 0.349 0.400 1.010 0.292 0.200 0.123
Both 3205 1.165 0.331 0.383 1.006 0.451 0.194 0.123

O83

wee

SS

SV /
4

S
<
ao

Figs. 4-6. Parada spp. 4, P. paitae, right hemelytra, paranota and lateral carina of the habitus. 5, P. paitae,
ventral face. 6, P. minuta, right hemelytra, paranota and lateral carina of the habitus. Scale = 1 mm.

VOLUME 102, NUMBER 4

821

Body length, body width, hood length, hood width and antennal segments measurements of the

New Caledonian species and P. approba. Last antennal segment measurements on P. approba is missing on the

Table 2.
type.
Body

Species Length Width Length
Parada majuscula 3.05 oJ 0.33
Parada paitae 2.98 7 0.31
Parada minuta 2.63 0.94 0.29
Parada approba 2.99 1.26 0.37

one or two areolae divided into two areolae
at level of apex, and discoidal area five to
six areolae broad. Antennal segments mea-
surements: I, 0.17: II, 0.12: III, 0.94: IV,
0.40.

Types.—Holotype: 2°, New Caledonia,
Mont Nondoué near Paita, sclerophyllous
forest, 16.X.1992, L. Bonnet de Larbogne,
J. Chazeau & E. Guilbert colls (fogging)
(MNHN). Paratype: 1 2, New caledonia,
Mont Nondoué near Paita, sclerophyllous
forest, 28.1V.1993, L. Bonnet de Larbogne,
J. Chazeau & E. Guilbert colls (fogging)
(MNHN).

Parada minuta Guilbert, new species
(Fig. 6)

Description.—Head and _ pronotum
brown, body, hemelytra, legs and antenna
clear to yellowish, last antennal segment
brown fuscous, some brown spots on hem-
elytra on discoidal and sutural areas. Length
2.63; width 0.94; hood length 0.29; hood
width 0.31.

Head short, with four tiny spines. Anten-
nal segments as two former species. Anten-
nal segments measurements: I, 0.15: II,
0.11: Il, 0.92: IV, 0.43. Bucculae short,
wide, three areolae broad. Labium short,
not extending beyond mesosternum, labial
sulcus widened between hind coxae.

Pronotum gibbose, areolate, tricarinate,
median carina uniseriate, elevated, lateral
carinae wide, four areolae broad. Paranota
four areolae broad.

Costal area completely uniseriate, sub-
costal area biseriate, discoidal area short,

Hood

Antennal Segments
Width I II Ii IV
0.38 0.19 OM 1.01 0.45
0.31 0.17 0.12 0.94 0.40
0.31 0.15 0.11 0.92 0.43
0.55 0.15 0.12 0.85 —

half length of hemelytra, five areolae broad
at widest part.

Types.—Holotype: 6, New caledonia, Ile
des Pins: Vao, 0-100 m, I. 1985, N.L.H.
Krauss coll. (BPBM). Paratype: 1 6, same
data as the holotype (BPBM).

KEY TO SPECIES OF PARADA

1. Head armed with 4 cephalic spines....... D
Head armed with 5 cephalic spines....... 4
2. Subcostal area triseriate at widest part .... 3

— Subcostal area biseriate at widest part (Fig. 6)

P. minuta Guilbert, n. sp.

3. Labium extending beyond hind coxae (Fig. 3)
Berend eo acee nines okra P. majuscula Guilbert, n. sp.

— Labium not reaching hind coxae (Fig. 5)...

P. paitae Guilbert, n. sp.

4 Paranotasthreesateolac wide ys acne es 5

Paranota four areolae wide
5. Median carina areolate and uniseriate (Fig.
IES) Loeieaeeai erica ieR Ceol P. solla Drake and Ruhoff
=) Medianvcarina not arcolates; - ere
el eevee eee he kas P. taeniophora (Horvath)
6. Lateral carinae five areolae wide (Fig. 10) . .
MERE SARS Cons RR Oto ore P. absona Drake
Lateral carinae less than five areolae wide .. 7
7. Third antennal segment 4 times longer than
last = .. yelePe MAE eee dae:s P. approba (Drake)
— Third antennal segment less than 4 times lon-
gerthan lasts. aa aS ioe w cee Bets 8
8. Costal area with areolae the same size than
that of subcostalearcay(sea 2) eee
Pace ina ema ld Spee a te iE P. hackeri Drake
— Costal area with areolae larger than that of
subcostalearearanan = cries on eee 9
9. Lateral carinae width of 2 areolae broad (Fig.
Aa Aa deas de one eck P. darlingtoni Drake
— Lateral carinae width of 3 areolae broad .. 10
10. Costal area largely biseriate at base (Fig.
P. popla Drake
— Costal area uniseriate except for two areolae

across) very base (Fis: I) S44 3 P. torta Drake

OWY
Gots) SLO «

S8seoosg © A iS Ss

Figs. 7-8.

CLADISTIC ANALYSIS

Choice of the taxa.—The eleven species
of the genus Parada are included in this
analysis as the ingroup. Three species are
included as outgroups: Cysteochila cubens
Guilbert, Physatocheila dissimilis Guilbert
and Nobarnus pilosus Guilbert. Cysteochila
cubens shows some affinities with the genus
Parada, while N. pilosus is clearly different.
Physatocheila dissimilis could be considered
as intermediate between these two genera by
its general morphology. Therefore, 14 spe-
cies were included in this analysis.

Character states.—Sixty-one morpholog-
ical characters are analysed. They are ana-
lysed in two differents ways. A first anal-
ysis concern all the characters. A second
analysis concern only fifty-one characters

% ootssoeO Ose

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Parada approba. 7, Profile. 8, Ventral face. Scale = 1 mm.

active and ten inactive. These ten characters
define the size of various structures in terms
of number of areolae. They are not used in
the analysis because the homology between
the areolae cannot be established with cer-
tainty, but they figure as attributes. When a
structure has two areolae, does it mean that
an areola divided into two areolae, or does
a second areola appears near the existing
one? The results of the two analyses are
compared. Inactive characters in the second
analysis are mentioned. Characters of the
genitalia are not taken into account because
they do not vary within the genus. Multi-
state characters are treated unordered. The
matrix of characters is given in Table 3.
Question marks represent characters lacking
or uncertainly stated.

VOLUME 102, NUMBER 4 823

; Sa
>)
We’
SS
iy
we
3

4 DOD AS)
fi Noo re is
MS) wee °0) Ves
; 4 A 7 rf} i
dj 0 \\ |
, ; go \ pg
Us 6 U ladl \
(y S D i
0); g @ 1h
i S toy \ |
) | \0
D 0) © \ Q
0 ic) \9 ) 0
© \ 0
\ 9 Ka Q 0
i) a ic ; a
0 5 \ ; ¢
Q 6)
by 4 " D
f ; Y \ :
a () Q Xo =: 9)
“0 C Se \
Q ° Xe
9 \S 10 11 12 Xe

Figs. 9-15. Parada spp. left costal area, paranota and lateral carinae. 9, P. popla. 10, P. absona. 11, P torta.
12, P. hackeri. 13, P. taeniophora. 14, P. darlingtoni. 15, P. solla. Scale = 1 mm.

0. Head black (0), brown (1), yellowish to 12. Occipital spines stout (0), slender (1).

white (2). 13. Median spine long (0), short (1), tuber-
1. Pronotum black (0), brown (1), yellow- cle-like (2), absent (3).
ish (2). 14. Median spine erect (0), prostrate (1).
2. Abdomen black (0), brown (1), yellow- 15. Median spine stout (0), slender (1).
ish (2). 16. Antennae densely pubescent (0), not
3. Hemelytra black (0), brown (1), yel- pubescent (1).
lowish to white (2), brown with colored 17. Last antennal segment longer than first
spots (3). and second together (0), shorter than
4. Body glabrous (0), pilose (1). first and second together (1).
5. Head convex (0), flat (1). 18. Last antennal segment longer than first
6. Frontal spines long (0), short (1). (0), shorter than first (1).
7. Frontal spines erect (0), prostrate (1). 19. Third antennal segment | time as long
8. Frontal spines stout (QO), slender (1). as the last one (0), 2 times (1), 3 times
9. Frontal spines in contact or crossed (0), (2);°4 times’ G).
not in contact (1). 20. Bucculae biseriate (0), triseriate (1),
10. Occipital spines long (0), short (1). quadriseriate (2), pentaseriate (3). In-
11. Occipital spines adpressed (0), pros- active character.
trate (1). The occipital spines are pros- 21. Bucculae open in front (0), closed in
trate when the spines do not rest onto front (1).

the head, otherwise they are adpressed. 22. Labium almost extending beyond the

824

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 3. Matrix of states of the 61 characters for 14 species.

Taxa Character states
Nobarnus pilosus 00001100000003?70111012100100?700?701?70004200010010030??202133
Physatocheila dissimilis 01110100000100001102101110101210?701??0111011111111001??110420
Cysteochila cubens 2223 00 OVO OOM2 0 Pal 702212 OOMOLOPS 05052703 20 Osa ema TOM 02 O a0 ee
Parada darlingtoni MOL SOO OOLOOOLOLLLOO2ZINLOPOOLU2112203 21 Ata Os WOM eA Aer OMe 2G
Parada hackeri Ala aL Sj{0)aL (0) aL (0)(0)(0)(ON0) LaalaE(O)(O)aL{o)sLak(o)tL (0) aa ieeya zat a ee MaE AL ALS} aL gL aL (0) iL ak (OAL aL (0) aL AU (0) al aL al at il (0)
Parada torta NOM ZO OLA COIN TOO2Z2 TNO LO OL NO 2 VA AAA te eae Oded Ones? AO) @ a2
Parada absona 0003000111001111100211100001122152110111211111110111110111220
Parada popla (ONY S4(O)(O) AL TL IL (OV(ON(0) ak iN aL aa) (0) 2A al A S}(0)(0)(0)(O)a al Ast Sy OVA Aral salsa UAL (O)al se (O}A ae ah SL AGO} aN TL AL 3),2,(0)
Parada solla HL ANS}(0) A110) a) 1 O)(ONCOVaL bea balo(oy Ayal al -A(0)sboj(ojabaal Aral ey zi 2a EAS) aA bs OAL aL LOO) AL aL?
Parada taeniophora il (0) a1 SOMO AL MAL LOVLONCOVAL wh Oak a7 AA AL Al ALCO) As a a2) A) A aL Sh aA AE ALCO) AL ALCO) 2A ALO) IL WL L200
Parada majuscula (0) LOS} OO)aL LALO ako) sk Sy aL O\O)IL-7AO) aL (0) ao) a AA) Aa Seay S ya EGE ICSE TE IL (O)IL aL a2) 2) AL aL Lat a
Parada paitae A ALAL aL (O)(O) AL TL A AL ALG) sb Sy aL (O)(O) ML LAL ICO) (O)(O) aL Aa LAA AL ara GE IETS IL LO) AE AL AL LO) aL IL 12) 2) AL
Parada minuta APASOO MLL MO MOU PP WO Os SAG OOM UA AO Aakala Wass Lala OAL aL LOO iL i 30
Parada approba MOA SOMO ssl Pojoy POO) ablOv(o)s} shal sya O)s@ak geal AAs al LNA ALA IL (aL wo) AL AL aL aL LO) AL AL 1. (0) 2,0

metasternum (0), extending to the mid-
dle of the metasternum (1), extending
little beyond the mesosternum (2),
reaching the posterior margins of the
mesosternum (3), extending to the mid-
dle of the mesosternum (4).

. Mesosternal laminae subparallel (0),

not subparallel (1).

. Metasternal laminae wide (Q), narrow

(1).

. Sternal laminae open behind (0), closed

behind (1).

. Pronotum strongly gibbose (0), slightly

gibbose (1).

. Pronotum pubescent (0), glabrous (1).
. Median carina not areolate (0), uniser-

iate (1).

. Median carina with large areolae (0),

small areolae (1), minute areolae (2).

. Lateral carinae ridge like (0), with one

or two rows of areolae (1), with 3 to 5
rows of areolae (2), with 6 or more
than 6 rows of areolae (3). Inactive
character.

. Lateral carinae erect (0), reflexed but

not resting onto the pronotum (1), re-
flexed and resting onto the pronotum

@):

. Lateral carinae 8 areolae long (0), 9

areolae long (1), 10 areolae long (2),
11 areolae long (3), 12 areolae long (4),

40.

41.

14 areolae long (5), 15 areolae long (6).
Inactive character.

. Hood absent (0), present but flat (1),

cyst-like and partly covering the head
(2). When the hood is present, it covers
most of the head but generally not the
eyes.

. Hood higher than posterior pronotal

lobe (0), not higher than posterior pron-
otal lobe (1).

. Hood 3 areolae long (0), 4 areolae long

(1), 5 areolae long (2), 6 areolae long
(3), 7 areolae long (4). Inactive char-
acter.

. Hood 1 areola high (0), 2 areolae high

(1), 3 areolae high (2). Inactive char-
acter.

. Collar triseriate (0), biseriate (1). In-

active character.

. Paranota 2 areolae broad (0), 3 areolae

broad (1), 4 areolae broad (2), 5 areolae
broad (3). Inactive character.

. Paranota with large areolae (0), small

areolae (1), minute areolae (2).
Paranota more than 14 areolae long (0),
13-14 areolae long (1), 11—12 areolae
long (2), 10 areolae long (3), 8—9 are-
olae long (4). Inactive character.
Paranota reflexed with free margins in
contact with pronotum (0), reflexed
with free margins not in contact with

VOLUME 102, NUMBER 4

42.

43.

44.

45.

46.

47.
48.

49.
50):
=
a2)

SS:

54.

SS)

56.
Si

58.

50)

60.

pronotum (1), not reflexed with free
margins not in contact with pronotum
2):

Paranota not covering pronotum (0),
covering part of pronotum (1).
Hemelytra wide (0), narrow (1). Hem-
elytra are considered narrow when not
extending far beyond the abdomen.
Hemelytra sharply widened at base (0),
not sharply widened at base (1).
Junction of RM and Cu veins raised
and forming a little swollen zone (0),
not raised and forming a little swollen
zone (1).

Hemelytra areolae hyaline (0), not hy-
aline (1).

Hemelytra pubescent (0), glabrous (1).
Hemelytral veins raised, forming ridges
(O), not raised and forming ridges (1).
Costal area wide (0), narrow (1).
Costal area plane (0), raised (1).
Costal area areolae very small (0),
small (1), large (2), very large (3).
Costal area 6 areolae broad (0), uni-
seriate (1).

Costal area uniseriate at base (0), bi-
seriate at base along 3-5 areolae (1),
biseriate at base along 7—10 areolae (2).
The costal area is mostly uniseriate and
shows in part two areolae for the same
width like if the original areolae were
divided in two smaller areolae (Figs.
9-15).

Costal area uniseriate at the middle (0),
biseriate at the middle along 2-3 are-
olae (1), biseriate at the middle along
5—6 areolae (2).

Subcostal area areolae very small (0),
small (1), large (2), very large (3).
Subcostal area large (0), small (1).
Discoidal area areolae very small (0),
small (1), large (2), very large (3).
Discoidal area 4 areolae wide (0), 5
areolae wide (1), 6 areolae wide (2), 7
areolae wide (3), 8 areolae wide (4), 9
areolae wide (5). Inactive character.
Sutural area areolae very small (0),
small (1), large (2), very large (3).
Sutural area 6—7 areolae wide (0), 8—9

825

areolae wide (1), 12 areolae wide (2),
less than 6 areolae wide (3). Inactive
character.

The phylogenetic analysis was performed
with Hennig86 (Farris 1988). The algorithm
ie* was used to build tree (s), using Fitch
parsimony.

RESULTS

First analysis.—Three trees of 119
steps, with ci = 61 and ri = 50 were gen-
erated. They differ by the relative position
of the three New Caledonian species
among themselves. Seventy-seven percent
of the active characters are not informa-
tive for the resolution of these conflicting
branches. Here is presented one of the
trees which corresponds to the following
interpretation (Fig. 16). For the 7 infor-
mative characters, Parada minuta and P.
paitae share the same state for four of
them, while P. majuscula and P. minuta
share the same state for one of them, and
P. majuscula and P. paitae for any of
them. Then, Parada minuta may be more
closely related to P. paitae than to P. ma-
juscula. The coefficients and number of
steps for each characters is given Table 4.

The genus Parada (including the species
approba) is monophyletic. The monophyly
of the genus is supported by the raised
veins on the hemelytra, the lateral carinae
and the paranota which are curved inwards
but not resting onto the pronotum.

The genus is divided in two groups, the
New Caledonian group which comprise
the three New Caledonian species, and the
Australian group comprising the other
eight species. The Australian species have
five cephalic spines, while New Caledon-
ian species have four. No unambigous
synapomorphy support the monophyly of
the Australian group, while the short oc-
cipital spine and the lack of a median
spine are the unambiguous synapomor-
phies of the New Caledonian group. Par-
ada approba is completely integrated in
the genus; however, it is the basal species

826 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

N. pilosus

P. dissimilis

N C. cubens
56
P. majuscula
i P. paitae
44 Gt) LUG):
13(3), 19(1) P. minuta
P. approba
19(3)
P. popla
6(1)
N P. absona
314), 37): 35
41(1), 48(0) P. hackeri
i 19(1)
33 5(1) N P. solla
32 43
P. darlingtoni
24(1) 30(2)
P sorta
51(2)
6(1) P. taeniophora
43
Fig. 16. One of the three cladograms obtained with the ie* command of Hennig86 on data show in table 3
(14 taxa, 61 active characters). Length = 119 steps, ci = 61, ri = 50. Paranota and lateral carinae width,

respectively, in number of areolae are indicated under an arrow on the cladogram.

of the Australian group. It is characterised have a labial sulcus broad like the New
by the third antennal segment that is four Caledonian species. Among the Austra-
times the length of the last. Parada popla, lian group, two subgroups can be distin-
P. absona and P. approba are the basal guished, the hackeri subgroup composed
species of the Australian group. They by P. hackeri and P. solla, and the dar-

VOLUME 102, NUMBER 4

Table 4. Number of steps, ci and ri for each char-
acter for the first analysis (10 characters inactive) and
the second analysis (all characters active).

ae Analysis 1 Analysis |
acters Steps ci ri Steps ci ri

1 5 40 40 5 40 40

2 5 40 40 5 40 40

3 5 40 0) 5 40 0)

4 | 75 O 4 WS 0)

5 1 100 100 1 100 100

6 2; 50 66 3 33 33

7 3) 33 60 3 33 60

8 2 50 50 2 50 50

9 2 50 50 2 50 50
10 2 50 0) 2 50 0)
11 1 100 100 1 100 100
12 1 100 100 1 100 100
13 2 50 50 2 50 50
14 5) 60 60 5 60 60
15 2 50 66 3 33 33
16 1 100 100 1 100 100
17 1 100 100 1 100 100
18 1 100 100 1 100 100
19 i 100 100 1 100 100
20 4 50 50 4 50 50
21 >) 40 0) 3) 40 0)
22 1 100 100 1 100 100
23) 7 5)// 40 8 50 20
24 3 33 66 3 33 66
25 Z 50 TS 4 25 25
26 3 33 0) 3 33} O
27 2 50 50 2 50 50
28 1 100 100 1 100 100
29 D, 50 0) 2 50 0)
30 4 50 0) 3 66 50
31 4 75 O 4 75 0)
32 1 100 100 1 100 100
33 U 85 50 7 85 50
34 3 66 50 2 100 100
35 2 50 0) 2 50 0)
36 7 42 0) 6 50 25
37/ 5 40 0) 3 66 66
38 1 100 100 1 100 100
59 5 60 50 3 100 100
40 2 100 100 2 100 100
41 7 57 P25) 6 66 50
42 2 100 100 2 100 100
43 1 100 100 1 100 100
44 1 100 100 1 100 100
45 l 100 100 100 100
46 4 25 25 3 33 50
47 1 100 100 1 100 100
48 1 100 100 1 100 100
49 i 100 100 1 100 100
50 1 100 100 1 100 100
31 1 100 100 1 100 100

Table 4. Continued.

Analysis 1 Analysis |

Char-

acters Steps ci ri Steps ci ri
Sy 4 75 Ws 5 60 50
53 1 100 100 1 100 100
54 5) 40 25 4 50 50
55 3 66 10) 3) 66 O
56 2 100 100 2 100 100
57 1 100 100 1 100 100
58 2 100 100 2. 100 100.
59 9 55 0) 8 62 DS
60 +4 50 0) 4 50 0)
61 5 60 33 4 TS 66

lingtoni subgroup composed by P. dar-
lingtoni, P. torta and P. taeniophora. The
monophyly of both subgroups together is
supported by a narrow labial sulcus. The
monophyly of hackeri subgroup is sup-
ported by a flat hood. The monophyly of
darlingtoni subgroup is supported by very
large areolae on costal area. The slightly
gibbose pronotum is autapomorphic for P.
hackeri, and the large areolae of the me-
dian carina is autapomorphic for P. torta.

Second analysis.—A single tree of 168
steps, with ci = 62 and ri = 48 is gener-
ated (Fig. 17). The conflicting branch be-
tween the New Caledonian species is re-
solved, and Parada minuta is more close-
ly related to P. paitae than to P. majus-
cula. As in the first analysis, the genus
Parada is monophyletic. The monophyly
being supported by the same characteris-
tics. Parada hackeri is the basal species
of the genus. It is the only species of Par-
ada with stout frontal and occipital
spines, biseriate bucculae and having the
shortest rostrum of the genus. It shares the
same hood shape and the same size of are-
olae on costal area than Cysteochila cub-
ens, the outgroup most related to the ge-
nus. The other species of the genus is di-
vided in two groups, the torta group, com-
prising P. torta (basal species of the
group), P. absona, P. popla, P. darling-
toni and P. approba, and the solla group,
comprising P. solla (basal species of the

828 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

A
56
8(1)
38(1)
33
Y

13@1), 31(1), 19(2)
STL) ool):
41(1), 48(0)

A
43 38(2)

6(1), 10(1),
13(3), 190) &

N. pilosus
P. dissimilis
C. cubens

P. hackeri

P. torta
29(0)
P. absona

P. popla
7(1), 30(2)
P. darlingtoni
32
13(0) P. approba
19(3)
P. solla
S(1)
P. taeniophora

P. majuscula
22(0)

P. paitae

P. minuta

Fig. 17. The single three cladograms obtained with the ie* command of Hennig86 on data show in table 3
(14 taxa, 51 active characters). Length = 168 steps, ci = 62, ri = 48. Paranota and lateral carinae width,
respectively, in number of areolae are indicated under an arrow on the cladogram.

group), P. taeniophora, P. majuscula, P.
paitae and P. minuta. The torta group is
characterized by the paranota three areo-
lae wide, that is shared with Parada hack-
eri and the outgroup Physatocheila dissi-

milis. The solla group is characterized by
the paranota four areolae wide, the only
autapomorphic character for the group.
The coefficients and number of steps vary
for 15 characters over 61 between the two

VOLUME 102, NUMBER 4

analysis (Table 4), 6 of them being inac-
tive in the first one.

DISCUSSION AND CONCLUSION

The species of Parada can easily be rec-
ognized by the combination of the paranota
(characters 41, 42), the lateral carinae (char-
acters 30, 31) and the costal area shapes
(characters 49 to 54).

Despite the different results of the two
analyses, some characters have the same
tendencies of evolution. The lateral carinae
and the paranota width in terms of number
of areolae used as attributes in the first ana-
lysis (because of their uncertain homolo-
gies) narrow from basal to terminal taxa.
However, these tendencies in the second
analysis are not as clear as in the first one.

In the first analysis, the width of the lat-
eral carinae narrow from the outgroup C.
cubens (lateral carinae 6 areolae broad) to
the New Caledonian group (4 areolae
broad), the Australian group (3 areolae
broad) and P. darlingtoni (2 areolae broad),
except for P. absona which has lateral ca-
rinae of 5 areolae broad (Figs. 9-15). The
same way, the paranota narrow from C.
cubens (5 areolae broad) to the New Cale-
donian group (4 areolae broad) and the
Australian group (3 areolae broad). There
are two reversals for P. solla and P. taenio-
phora which have the paranota 4 areolae
broad. The others pronotal and hemelytral
characters do not show such a tendency.
However, the lateral carinae and the par-
anota are longer in terms of number of are-
olae for C. cubens than for the Parada spe-
cies, and the lateral carinae of the New Ca-
ledonian species is shorter than the one for
the Australian species.

In the second analysis, the lateral carinae
narrow from C. cubens to Parada species
from 6 to 3 areolae wide. They still narrow
in the torta group for P. darlingtoni from 3
to 2 areolae wide, while they enlarge in the
solla group for the three New Caledonian
species from 3 to 4 areolae. There is also a
reversion for P. absona which lateral cari-
nae enlarge to 5 areolae wide. The paranota

829

narrow from C. cubens to Parada species
from 5 to 3 areolae wide but enlarge in the
solla group to 4 areolae.

If characters such as the width among the
different hemelytral areas and the various
pronotal expansions like the hood, the lat-
eral carinae and the paranota show a trend
of evolution, this is not the case of other
characters such as the length of the lateral
carinae and the paranota. They do not show
any extension or shortening through the
cladogram. Thus, wide lateral carinae and
wide paranota could be plesiomorphic. The
two major groups found by the first analy-
sis, are separated by a geographic barrier:
the sea between the Queensland and New
Caledonia, isolated from Australia since the
Triassic. Then, lateral carinae and paranota
still narrowed in Australia after the Triassic.
However, these characters show slight var-
iation among Parada species, in compari-
son with other genera. As a general rule,
Tingidae present traits that have no appar-
ent biological explanation and that could be
hypertelic by their disproportionate size.
But, these possibilities still require further
study and much more data such as behav-
ioral parameters, and host plants, need to
be added to test these evolutionary hypoth-
eses.

ACKNOWLEDGMENTS

I am most grateful to Gordon Nishida (B.
P. Bishop Museum, Honolulu, Hawaii) for
the loan of many specimens, and Thomas
Henry (Systematic Entomology Laboratory,
USDA) for his helpful support in the access
to the types at the National Museum of Nat-
ural History, Smithsonian Institution,
Washington, DC. I am also indebted to Cyr-
il d’Haese, Philippe Grandcolas, Laure De-
sutter-Grandcolas, and Thierry Bourgoin
(MNHN, Paris) for their fruitful comments.

LITERATURE CITED

Drake, C. J., 1942. New Tingidae (Hemiptera). Iowa
State College Journal of Science 17(1): 1-21.

. 1952. The genus Parada Horvath (Hemiptera:

Tingidae). Psyche 59(4): 143-147.

830 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

. 1961. Some Australian Tingidae (Hemiptera),
including new genera and new species. Records
of the Australian Museum 25(6): 107—114.

Drake, C. J. and FE A. Ruhoff. 1961. New genera and
new species of lacebugs from the Eastern Hemi-
sphere (Hemiptera: Tingidae). Procceeding of the
United States National Museum 113: 125-183.

. 1965. Lace bugs of the world: a catalog (He-

miptera: Tingidae). United States National Muse-

um Bulletin 243, 634 pp.

Farris, J. S. 1988. Hennig86, version 1.5, Port Jeffer-
son Station, New York.

Guilbert, E. In press. New data on New Caledonian
Tingidae (Hemiptera) with description of new spe-
cies. Zoologia Neocaledonica Vol. 5. Mémoires
du Museum National d’Histoires naturelles.

Horvath, G. 1925. Results of Dr. E. Mj6berg’s swed-
ish scientific expeditions to Australia 1910—
1913. 45. Tingitidae. Arkiv for Zoologi 17(24):
1-17.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 831-837

A NEW NEOTROPICAL SPECIES OF CLINODIPLOSIS
(DIPTERA: CECIDOMYIIDAE), AN IMPORTANT NEW PEST OF
CULTIVATED PEPPERS (CAPSICUM SPP.: SOLANACEAE)

RAYMOND J. GAGNE, HELGA BLANCO-METZLER, AND JEAN ETIENNE

(RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S.
Department of Agriculture, % National Museum of Natural History, Smithsonian Insti-
tution, Washington, DC 20560-0168, U.S.A. (e-mail: rgagne @sel.barc.usda.gov); (HB-M)
Centro de Investigacion en Proteccién de Cultivos, Estacién Experimental Fabio Baudrit,
University of Costa Rica, P.O. Box 183-4050, Alajuela, Costa Rica; (JE) Institut National
de la Recherche Agronomique, Station de Zoologie et Lutte Biologique, Domaine Duclos,
Petit Bourg, Guadeloupe

Abstract.—A new species of cecidomyiid, Clinodiplosis capsici Gagné, is reported
from cultivated sweet and hot pepper in Costa Rica, Guadeloupe, and French Guiana
where it appears to have potential as a serious pest. Adults, pupae, and larvae are de-

scribed, illustrated, and compared to other Clinodiplosis species.

Key Words:

In the mid-1990s, a new species of gall
midge belonging to Clinodiplosis was in-
dependently discovered on Guadeloupe and
in Costa Rica attacking sweet pepper, Cap-
sicum annuum L., and hot pepper, Capsi-
cum frutescens L. The white larvae of the
gall midge cause swellings on stems,
leaves, and bases of flowers (Figs. 1-3).
Galls can be found all year round and,
where locally abundant, the cecidomyiid is
a serious pest. Larvae live singly in cells
within the swellings. The full grown larva
pupates in the gall, the fully developed
pupa forms a circular exit hole with the
help of the antennal horns, emerging about
halfway out of the hole, and the adult im-
mediately emerges from the pupal skin. Af-
ter adult emergence, the damaged part of
the plant is susceptible to rot. Similar galls
were later found in 1999 in French Guiana.

In Costa Rica, galls were collected in dry
climates as in Guanacaste, wet climates as
in Pérez Zeled6n, and intermediate cli-

gall midge, sweet pepper, hot pepper

mates, such as Alajuela. They were found
in gardens, backyards and greenhouses of
Guanacaste, Alajuela, and Cartago Provin-
ces, but not in commercial plantations. This
absence in the larger plantations may be
due to chemical applications intended for
the control of the pepper weevil, Anthono-
mus eugenii Caro (Coleoptera: Curculioni-
dae).

The new species of gall midge will read-
ily run to Clinodiplosis in the key in Gagné
(1994). Clinodiplosis is a worldwide genus
of some 75 known valid species. Outside
the Neotropics, most species of this genus
appear not to be host specific and are usu-
ally associated with fungus growing in or
on plant tissue, such as spent flowers or old
galls. A few species, however, are associ-
ated with simple bud galls where they are
doubtless primary plant feeders (Gagné
1989). While Clinodiplosis spp. may also
be found associated with fungus in the Neo-
tropics, most known regional species in this

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.

and closely related genera peculiar to the
Neotropics, e.g., latrophobia and Schisma-
todiplosis, are phytophagous, host specific,
and associated with complex galls (Gagné
1989), as is the new species described be-
low.

METHODS

Galls with pupae or full-grown larvae
were placed in small containers until adults

Galls of Clinodiplosis capsici on stems and leaves of Capsicum sp.

emerged. Specimens of immature stages
and reared adults were preserved in 70%
isopropyl alcohol. Samples were mounted
on microscope slides using the method out-
lined in Gagné (1989). Terminology for
adult morphology follows usage in Mc-
Alpine et al. (1981) and for larval mor-
phology that in Gagné (1989). Galls were
obtained, adults reared, and other stages se-
cured by H. Blanco-Metzler in Costa Rica

VOLUME 102, NUMBER 4

and by J. Etienne in Guadeloupe. J. Etienne
also found similar galls in French Guiana.
The taxonomic investigation was the re-
sponsibility of R. J. Gagné.

Clinodiplosis capsici Gagné, new species
(Figs. 4—15)

Adult.—Head: Eyes connate, 11—12 fa-
cets long at vertex; facets mostly hexago-
noid, all closely adjacent. Occiput with dor-
sal protuberance with 2 apical setae. Frons
with 6-10 setae. Labella ellipsoid and
pointed apically, each with several lateral
setae. Palpus 4-segmented. Male antennal
flagellomeres (Fig. 4) binodal; one circum-
filum on basal node, two on the distal, the
loops of the three circumfila subequal in
length. Female flagellomeres (Fig. 5) cylin-
drical with long necks, surrounded by two
appressed circumfila connected by two lon-
gitudinal bands.

Thorax: Wing unmarked, 2.2 to 2.9 mm
long, R; curved toward apex, joining C pos-
terior to wing apex. Tarsal claws (Fig. 6)
untoothed, curved near basal third; empodia
very short, not attaining bend in claws.

Male abdomen: First through sixth ter-
gites entire, rectangular, with single poster-
ior row of setae, several lateral setae, scat-
tered scales, and 2 anterior trichoid sensilla;
seventh tergite unsclerotized posteriorly
and lacking the posterior row of setae and
scales, but lateral setae and anterior pair of
trichoid sensilla present; eighth tergite un-
differentiated, the only vestiture the anterior
pair of trichoid sensilla. First through eighth
sternites rectangular, covered with setae and
with 2 anterior trichoid sensilla; eighth ster-
nite similar to preceding except weakly
sclerotized anterolaterally. Genitalia (Figs.
9-10): cerci rectangular, with posterior se-
tae; hypoproct much longer than cerci, wid-
est at midlength, narrowing beyond mid-
length and widening again posteriorly, the
basal two-thirds expanded laterally to curve
slightly around aedeagus, and with strong
setae and several smaller ones posterolat-
erally; aedeagus elongate, narrower and
longer than hypoproct, with longitudinal

833

rows of sensory pits; gonocoxite elongate-
cylindrical with mesoposterior surface
forming nearly right angle; gonostylus
elongate-cylindrical, with setulae near base
and covered beyond with minute carinae
and widely scattered short setae.

Female abdomen (Figs. 7-8): First
through seventh tergites entire, rectangular,
with mostly single row of posterior setae,
several lateral setae, extensively covered
with scales, and with 2 anterior trichoid
sensilla. Eighth tergite unsclerotized, with
mostly single row of posterior setae and an-
terior pair of trichoid sensilla the only ves-
titure. Second through seventh sternites
quadrate, extensively covered with setae
and scales and with anterior pair of trichoid
sensilla. Ovipositor slightly protrusible,
venter of eighth segment and dorsum of
ninth and tenth segments without vestiture,
venter of ninth and tenth segments with se-
tae, cercus large, ovoid, with pair of apical
sensory setae and scattered setae elsewhere,
hypoproct short, narrow, with 2 posterior
setae.

Pupa.—Head (Figs. 11-12): Antennal
base pointed apically, the apex projecting
ventrad; cervical sclerite with two elongate
setae; face without ventral projections, with
1—2 papillae, one with seta, on each side of
base of labrum. Prothoracic spiracle elon-
gate, pointed apically. Abdominal tergites
covered dorsally with uniformly small spic-
ules, none enlarged and spiniform.

Third larval instar (Figs. 13—15).—
Length, 2.5—2.7 mm. White. Integument
with scattered spicules. Antenna about
twice as long as wide. Spatula with 2 wide-
ly separated anterior teeth, the space be-
tween weakly indented. Lateral thoracic pa-
pillae in 2 groups of 3 on each side of cen-
tral line, 2 papillae in each group each with
tiny seta. Dorsal and pleural papillae elon-
gate. Terminal segment rounded, with 8 pa-
pillae as follows: 1 pair as long as dorsal
setae of previous segment; 1 pair with setae
about half as long; the two posterior pairs
with short, corniform setae, those of inner
pair slightly narrower than outer pair.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

‘@

Figs. 2-3. 2, Galls of Clinodiplosis capsici on stems and base of flowers of Capsicum sp. 3, Detail of same,

one in section to show larva in larval cell.

VOLUME 102, NUMBER 4 835

?
\
\
\
‘ |
XX

< 10

Figs. 4-10. Clinodiplosis capsici. 4, Male third antennal flagellomere. 5, Female third antennal flagellomere.
6, Tarsal claws and empodium. 7, Female postabdomen, seventh segment to cerci (lateral). 8, Female cercus,

detail (lateral). 9, Male genitalia, only one gonopod shown (dorsal). 10, Same, only one gonocoxite shown
(ventral).

836 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 11-15. Clinodiplosis capsici. 11, Pupal head (ventral). 12, Pupal head, part, showing ventral projection
of antennal base (lateral). 13, Larval spatula and associated lateral papillae. 14, Larva (ventral). 15, Larval eighth
and terminal segments (dorsal).

VOLUME 102, NUMBER 4

Holotype.—¢d, from Capsicum frutes-
cens, Guadeloupe, French Antilles, Ravine
Chaude, 27-XII-1994, J. Etienne, GR 1575,
deposited in the National Museum of Nat-
ural History (USNM), Washington, DC.

Other material examined (all deposited in
USNM).—Same data as holotype, 1 d6, 2
2, 3 pupae, 3 larvae; from Capsicum sp.,
Guadeloupe, Ste. Rose, 3-II-1993, J. Etien-
ne, GR 1309, 2 6, 2 2, 5 pupae, 6 larvae;
from Capsicum sp., Guadeloupe, Lamentin,
1-VII-1992, J. Etienne, GR 1110, 1 d, 1 2,
1 pupa, | larva; from Capsicum annuum,
Costa Rica, Guanacaste Province, 3-I-1995,
H. Blanco, 1 3d, 1 @; from Capsicum an-
nuum, Costa Rica, Turialba, VIII-1995, H.
Blanco; 36; 22. 3 pupae.

Etymology.—The specific name, capsici,
means “of capsicum.”

Remarks.—The larva of C. capsici dif-
fers substantially from other Clinodiplosis
spp. in that the corniform setae of the ter-
minal segment are small and not each sit-
uated at the end of lobes, as is usual for the
genus. It cannot be said with confidence
whether in this species the lack of lobes is
primitive or the result of reduction. The
male and female of C. capsici have the gen-
eral habitus of the genus, the only pecu-
liarity being the shape of the male hypo-
proct, which is broadened at midlength, its
sides bending slightly around the aedeagus.
The pupal abdomen is without dorsal spines
and resembles in that way the species of
Clinodiplosis that feed on fungi. Unlike the

837

new species, several other Neotropical spe-
cies of Clinodiplosis that pupate in their
galls, e.g., Clinodiplosis eupatorii Felt in
conical leaf galls on Chromolaena (Aster-
aceae) (Gagné 1977), have enlarged dorsal
spines on the pupal abdomen that are pre-
sumably adaptations in those species for es-
caping from the galls.

ACKNOWLEDGMENTS

We thank P. Malikul for making the slide
preparations, Lucrecia Rodriguez for com-
puter assistance in making the plates, and,
for their comments on drafts of the manu-
script: P. Hanson, Universidad de Costa
Rica, San José; K.M. Harris, International
Institute of Entomology, CAB, London,
UK; P. Kolesik, The University of Adelaide,
SA, Australia; and A.S. Konstantinov and
A.L. Norrbom of the Systematic Entomol-
ogy Laboratory, USDA, Washington, DC.

LITERATURE CITED

Gagné, R. J. 1977. The Cecidomyiidae (Diptera) as-
sociated with Chromolaena odorata (L) K. & R.
(Compositae) in the Neotropical Region. Brenesia
12/13: 113-131.

1989. The Plant-Feeding Gall Midges of

North America. Cornell University Press, Ithaca,

New York, xiii & 355 pp. & 4 pls.

. 1994. The Gall Midges of the Neotropical Re-
gion. Cornell University Press, Ithaca, New York,
xv & 352 pp.

McAlpine, J. E, B. V. Peterson, G. E. Shewell, H. J.
Teskey, J. R. Vockeroth, and D. M. Wood, eds.
1981. Manual of Nearctic Diptera, Vol. 1. Re-
search Branch, Agriculture Canada, Monograph
27, vi & 674 pp.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 838-842

NEW SPECIES OF PUERTO RICAN PHYCITINAE
(LEPIDOPTERA: PYRALIDAE)

H. H. NEUNZIG

Department of Entomology, North Carolina State University, Raleigh, NC 27695-7613,

U.S.A.

Abstract.—Zamagiria borinquensis and Oryctometopia maricaoensis are described as
new species. Both were collected as adults in western Puerto Rico.

Key Words:

Post-Columbian Puerto Rico has been
subjected to major environmental changes.
Agriculture, mainly the extensive growing
for many years of sugarcane and coffee, re-
placed much of the native vegetation. Some
refugia exist, however, chiefly as parks or
reserves. These vary from tropical dry for-
est to tropical rain forests. Recently, ento-
mologists at the Carnegie Museum of Nat-
ural History (CMNH) have intensively
light-trapped Lepidoptera in these remain-
ing pockets of relatively undisturbed flora.
Two previously unknown species of phy-
citines were collected. These are described
below, and are added to the 20 species (Ta-
ble 1) previously recorded from Puerto
Rico by Heinrich (1956).

Zamagiria borinquensis Neunzig,
new species
(Figs. 1, 3-5)

Type locality.—Puerto Rico, Isabela,
Bosque Estatal de Guajataca, Montanas Ay-
mamon.

Diagnosis.—The male and female geni-
talia can be used to separate Zamagiria bor-
inquensis from other species in the genus.
Features peculiar to the new species are the
short, triangular inner clasper on the inner
base of the valva, the somewhat elongate,
spoonshaped apical gnathal process of the

moths, phycitines, Puerto Rico

male (Fig. 3), and the girdle of robust
spines in the corpus bursae of the female
(Fig. 5).

Description.—Forewing length 6.5—7.0
mm. Head brownish white. Labial palpus
white and fuscous. Maxillary palpus white
to brownish white, aigrettelike in male.
Dorsum of thorax brownish white suffused
with fuscous. Forewing brown dusted with
white; black ridge of raised, curled scales
preceding antemedial line; antemedial line
white, weakly developed, with associated
adjacent bands of dark scales; postmedial
line similar, in color and development, to
antemedial line; discal spots black, moder-
ately distinct, separate; patch of very pale
reddish brown scales at inner base of wing
and scattered patches of darker reddish
brown on outer % of wing. Underside of
male fore and hindwing without patches or
streaks of contrasting colored scales (fre-
quently seen in other Zamagiria). Male
genitalia (Figs. 3, 4): Uncus triangular, nar-
rowly rounded distally; apical process of
gnathos expanded into a somewhat elongate
spoon-shaped process; juxta a U-shaped,
thin plate with setiferous lateral arms; tran-
stilla absent; valva slender, unarmed except
for triangular, erect clasper on its inner
base; aedoeagus with single, short, stout,
well-sclerotized cornutus; vinculum about

VOLUME 102, NUMBER 4

Table 1. Phycitinae listed by Heinrich (1956) as
occurring in Puerto Rico!.

Crocidomera fissuralis (Walker)
Ectomyelois furvidorsella (Ragonot)
Fundella ignobilis Heinrich
Difundella distractor Heinrich
Anadelosemia texanella (Hulst)
Davara interjecta Heinrich
Davara rufulella (Ragonot)
Sarasota furculella (Dyar)
Caristanius pellucidellus (Ragonot)
Eurythmasis ignifatua Dyar
Paconius corniculatus Heinrich
Mescinia berosa Dyar

Phestinia costella Hampson
Laetilia portoricensis Dyar
Metephestia simplicula (Zeller)
Moodnopsis portoricensis Heinrich
Prosoeuzophera impletella (Zeller)
Caudellia clara Heinrich

Ribua contigua Heinrich

Varneria dubia Heinrich

! List does not include a few widespread neotropical
phycitines that Heinrich included in his checklist as
occuring in his category “‘West Indies,’ some of
which may be found in Puerto Rico.

as long as greatest width. Female genitalia
(Fig. 5): Ductus bursae short, about / as
long as corpus bursae, narrow, except
slightly broadened near ostium bursae,
without sclerotized plates common to most
other Zamagiria species, but generally
sclerotized anteriorly; corpus bursae elon-
gate, with well-developed, irregular comb
of many slender spines extending distally
from ductus bursae to about midway on
corpus, and with strongly formed band en-
circling corpus bursae at about middle and
bearing about 24 robust spines; ductus sem-
inalis attached to corpus bursae near junc-
tion of ductus bursae and corpus bursae.
Type material—Holotype: ¢: Puerto Rico,
Isabela, Bosque Estatal de Guajataca, Mon-
tanas Aymamon, 18°25'06N, 66°57'55’W,
forest, 210 m, 14—15 June 1996, J. Rawlins,
W. Zanol, R. Davidson, C. Young, M. Klin-
gler, S. Thompson; genitalia slide HHN
4427. Paratype: 1 2, same collection data
as for holotype; genitalia slide HHN 4428.

839

Jae, 15 2.
lotype. 2, Oryctometopia maricaoensis, holotype.

Males. 1, Zamagiria borinquensis, ho-

Holotype and paratype deposited in
CMNH.

Remarks.—Zamagiria borinquensis is
very similar externally to other species in
the genus. The character states of the male,
particularly the grooved head with its un-
usual compact tuft of scales attached to the
frons, and the unique tuft, composed of sev-
eral distinct types of scales, at the base of
the antenna, are easily recognized traits of
the species.

Etymology.—The specific epithet is de-
rived from the Taino name for Puerto Rico
(Borinquen), and the Latin suffix -ensis (de-
noting place).

Oryctometopia maricaoensis Neunzig,
new species
(Figs. 2, 6-8)

Type locality.—Puerto Rico, 3.3 km. sw
Maricao, Bosque Estatal de Maricao.

840

Figs. 3—5.
bursae and corpus bursae.

Diagnosis.—The male is readily identi-
fied by the elongate, medially constricted
vinculum (Fig. 6), and the female by the
large spined plate that covers most of one
side of the corpus bursae (Fig. 8).

Description.—Forewing length 10.5—
12.0 mm. Head of male brownish white to
pale reddish brown suffused with reddish
brown, and with vertex sulcate and white;
upper part of frons produced dorsally into
a ridge. Head of female simple, brownish
white to pale reddish brown. Labial palpus

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Zamagiria borinquensis. 3, Male genitalia (aedoeagus omitted). 4, Aedoeagus. 5, Female ductus

brown dusted with white and with a few
scattered brownish red scales. Maxillary
palpus of male aigrettelike, mostly pale
brown, in part pink. Maxillary palpus of fe-
male simple, pale brownish white. Dorsum
of thorax pale brown or pale purplish
brown. Forewing pale purplish brown in
basal 4%; antemedial line moderately distinct
to distinct, white, its inner % preceded by
black and pale reddish brown patch, and
followed by black line and associated white
patch; postmedial line moderately distinct

VOLUME 102, NUMBER 4

Figs. 6-8.
ductus bursae, corpus bursae and anterior part of ductus seminalis.

to distinct, white; medial area with patches
of red to dark red mostly along costa and
at discal spot; subterminal area with broad
brownish red patch following postmedial
line. Male genitalia (Figs. 6, 7): Uncus nar-
rowly triangular, rounded at apex; apical
process of gnathos a well-developed hook;
juxta platelike with thin, inwardly curved
lateral arms; transtilla absent; valva simple,
with short, stout lobes near inner base; ae-
doeagus, long, slender; vesica with cluster

841

Oryctometopia maricaoensis. 6, Male genitalia (aedoeagus omitted). 7, Aedoeagus. 8, Female

of small spines; vinculum long and slender
(about 2% times as long as basal width), and
constricted medially. Female genitalia (Fig.
8): Ductus bursae slightly shorter than cor-
pus bursae, sclerotized over % its length;
corpus bursae with large, many-spined
plate, covering most of left side (when
viewed ventrally) of bursae; plate, in part
perforate; ductus seminalis attached to cor-
pus bursae near junction of ductus bursae
and corpus bursae.

842

Type material.—Holotype: d¢: Puerto
Rico, Bosque Estatal de Maricao, 3.3 km
sw Maricao, 18°09'39"N, 67°00'05’W, for-
est, 550 m, 10-11 June 1996, J. Rawlins,
C. Young, R. Davidson, W. Zanol, S.
Thompson, M. Klingler; genitalia slide
HHN 4459. Paratypes: (4 ¢, 2 @, total);
same collection data as for holotype, geni-
talia slides HHN 4415, 4416 (3 6, 2 2);
Puerto Rico, Isabela, Bosque Estatal de Gua-
jataca, Montanas Aymamon, 18°25'06’N,
66°57'55"W, forest, 210 m, 14-15 June
1996, J. Rawlins, W. Zanol, R. Davidson,
C. Young, M. Klingler, S. Thompson (1 ¢).
All specimens from Maricao deposited in
CMNH,; Isabela paratype in North Carolina
State University Collection (NCSU).

Remarks.—The new species differs most
noticeably from Oryctometopia fossulatella
Ragonot, the only other species in the ge-
nus, in that the male genitalia lack the short,
thornlike spines projecting from the lower
margin of the sacculus, and have a cluster
of small spines on the vesica (rather than a
single spinelike cornutus), and in that the

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

female ductus bursae is more extensively
sclerotized. Nevertheless, males of both
species have an enlarged frons and sunken
vertex, tufted and otherwise modified an-
tennae, simple scale tufts on abdominal seg-
ment VIII, and generally similar genitalia.
Females of both have a characteristically
large, sclerotized plate in the corpus bursae
bearing numerous short spines.
Etymology.—tThe specific epithet is de-
rived from the locality (Maricao) where al-
most all the specimens were collected, and
the Latin suffix- ensis (denoting place).

ACKNOWLEDGMENTS

J. E. Rawlins generously made available
specimens for study, R. L. Blinn photo-
graphed the new species, and L. L. Deitz,
D. L. Stephan, and B. M. Wiegmann re-
viewed an initial draft of the manuscript.

LITERATURE CITED

Heinrich, C. 1956. American moths of the subfamily
Phycitinae. United States National Museum Bul-
letin 207: 1-581.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 843-851

TAXONOMY AND POLYTENE CHROMOSOMES OF
SIMULIUM PARNASSUM MALLOCH (DIPTERA: SIMULIIDAE)

Eric S. PAYSEN AND PETER H. ADLER

Department of Entomology, Clemson University, Clemson, SC 29634-0365, U.S.A.
(e-mail: padler@clemson.edu)

Abstract.—The taxonomic status of Simulium parnassum Malloch was investigated cy-
tologically and morphologically. Specimens were collected at 44 localities from the Gaspé
Peninsula of Quebec to northern Alabama. A previously unreported pupal form of S.
parnassum from two streams in South Carolina and one stream each in North Carolina
and Massachusetts lacked the typical rugose sculpturing on the dorsum of the head and
thorax. No additional structural characters in any life stage differentiated the two forms.
A standard map of the silk-gland chromosomes was constructed and compared with the
standard map for the subgenus Simulium. Simulium parnassum differed from the subge-
neric standard by six inversions in IL, one in IIL, three in IIS, and an unresolved number
in IL. Placement of S. parnassum in a separate species group, the S. parnassum species
group, in the subgenus Simulium is suggested on the basis of cytological evidence. No
fixed rearrangements were found among populations of S. parnassum from Quebec to
Alabama nor between populations of smooth and rugose pupal forms. A Y-linked poly-
morphism was observed in the centromere region of chromosome I in some males, but it
was not associated with either pupal form. Present evidence indicates that S. parnassum

is a single species throughout its range.

Key Words:

Simulium parnassum is one of the most
distinctive members of the subgenus Si-
mulium in the Nearctic Region (Stone
1964). The female is shiny black and each
claw is long and nearly sigmoidal, with a
small tooth near its midlength. The pupal
gill of six filaments and the simple, slipper-
shaped cocoon are similar to those of the S.
tuberosum and S. venustum species groups,
but the rugose head and thorax are unique.
The deeply incised, triangular postgenal
cleft of the larva is distinctive. Despite the
unique attributes of S. parnassum, this spe-
cies has been assigned to the S. tuberosum
species group (Crosskey and Howard
1997), ostensibly because of perceived sim-
ilarities in the male genitalia, notably the

Simulium parnassum, black flies, aquatic insects, polytene chromosomes

presence of a medially directed lobe on the
gonostylus. The homology of this lobe,
however, is questionable based on differ-
ences in location, size, and vestiture. Ac-
cordingly, a reevaluation of the phyloge-
netic placement of S. parnassum is war-
ranted.

Simulium parnassum was described from
females taken in northern New Hampshire
(Malloch 1914). Dyar and Shannon (1927)
described a male from eastern Virginia as
S. hydationis, which was synonymized with
S. parnassum by Stone and Jamnback
(1955). All life stages of S. parnassum have
been described and figured (Stone and
Jamnback 1955, Davies and Peterson 1956,
Davies et al. 1962, Wood et al. 1963, Stone

844

Table 1. Numbers of two pupal forms (rugose,
smooth) of Simulium parnassum collected at Smith
and Abner Creeks in Pickens County, South Carolina,
1998-1999.

Smith Creek Abner Creek
Ru- Ru
Year Date gose Smooth Date gose Smooth
1998 25 April D2 0) 2 June Ope
1998 5 May 0) + 9 June O 20
1998 14 May 7) 10) 11 June 1 23
1998 23 May! 4 23 June! 0) ]
1999 10 May ] 0) 1 June 1 12
1999 15 May 1 1 5 June Orel
1999 18 May 1 5) 10 June! QO 26
1999 21 May 30-36
1999 24 May 1 30
1999 28 May 0 17
1999 1 June! 0) 4

' No pupae found beyond these dates.

and Snoddy 1969, Adler and Kim 1986).
The polytene chromosomes have not been
studied, although Rothfels (1979) com-
mented briefly on the banding sequence of
two of the six chromosomal arms. Conse-
quently, S. parnassum has not been inves-
tigated for sibling species, which are com-
mon among black flies (Adler 1988).

This species occurs in eastern North
America from Canada to the southern end
of the Appalachian Mountains (Stone and
Snoddy 1969), with an isolated record from
Missouri (Doisey et al. 1986). The imma-
ture stages occupy cool, rocky, forest
streams (Davies et al. 1962, Adler and Kim
1986). Females are mammalophilic (Fuller
1940, Downe and Morrison 1957, Addison
1980) and can be pests of humans (Adler
and Kim 1986, Gibbs et al. 1986).

Our objectives were to screen S. parnas-
sum for sibling species, using morphology
and polytene chromosomes, and to resolve
its chromosomal banding sequence relative
to the subgeneric standard of Rothfels et al.
(1978), with the intent of gaining phylo-
genetic insight.

MATERIALS AND METHODS

Larvae and pupae of S. parnassum were
collected from 1998 to 1999 at 44 sites

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

from the Gaspé Peninsula of Quebec to
northern Alabama, and were fixed in Car-
noy’s solution (1 part glacial acetic acid: 3
parts 95% ethanol). Additional pupae were
reared individually to adults on moist filter
paper in petri dishes.

Seasonal sampling of larvae and pupae
was conducted at Smith Creek (35°0.17'N
82°49.01'W) and Abner Creek (35°4.27'N
82°47.11'W) in Pickens Co., South Caroli-
na. Streams were sampled every 10 days in
1998 and every three days in 1999. In 1998,
Smith Creek was sampled 3 March—11 Au-
gust and Abner Creek 13 May—11 August.
In 1999, both streams were sampled 17
March—26 June.

Polytene chromosomes were prepared
from larval silk glands, using the Feulgen
method of Rothfels and Dunbar (1953). A
standard chromosome map was constructed
for S. parnassum, following the procedures
and nomenclature of Rothfels et al. (1978).
The banding pattern of the standard map
was compared with that of the subgeneric
standard of Rothfels et al. (1978). For chro-
mosomal arms of S. parnassum that had
banding patterns identical to those of the
subgeneric standard, the subgeneric num-
bering was preserved. Chromosome arms
of S. parnassum that were rearranged rela-
tive to the subgeneric standard were given
new section numbers to provide a continu-
ous sequence. Slide-mounted chromosomes
of larvae from 15 of the 44 collection sites
(see Chromosomal Material Examined)
were compared band for band against the
standard map of S. parnassum in a search
for rearrangements within and between
populations of S. parnassum. Specimens
and chromosomal photographic negatives
and working maps were deposited in the
Clemson University Arthropod Collection,
Clemson, South Carolina.

Chromosomal material examined (num-
bers of each sex refer to larvae for which all
chromosomal bands were read).—ALA-
BAMA: Talladega Co., Cheaha State Park,
Cheaha Creek, 33°28.27'N 85°49.07'W, 27
March 1998 (1 3); 19 May 1998 (9 @);

VOLUME 102, NUMBER 4 845

Figs. 1-2. Cephalic plates of pupal exuviae of Simulium parnassum. 1, Rugose form (Smith Creek, Pickens
Co., South Carolina, 18 May 1999). 2, Smooth form (Abner Creek, Pickens Co., South Carolina, 11 June 1998).

846

pe

Figs. 3-5.

3 3)
ie! > \ >,
4] all
al 34 32 | - v35\32\ >

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Silk-gland chromosomes of Similium parnassum, with landmarks of Rothfels et al. (1978); C =

centromere region. 3, Chromosome arm IS. Numbers indicate section numbers for both S. parnassum standard
sequence and Simulium subgeneric standard of Rothfels et al. (1978). Male from South Carolina, Pickens Co.,
Oil Camp Creek, 8 May 1998 (sections 1 to center of section 13) plus female from Massachusetts, Berkshire
Co., 29 June 1998 (section 20 to center of section 13); Gl = glazed, 2 = two blocks, 3 = three heavy. 4,
Chromosome arm IL. Numbers on top of chromosome indicate S. parnassum standard banding sequence; bottom
numbers correspond with those of Simulium subgeneric standard of Rothfels et al. (1978). Brackets indicate
inversions relative to Simulium subgeneric standard of Rothfels et al. (1978); numbered brackets indicate se-
quence in which overlapping inversions occurred. Arrows indicate breakpoints for complex of three inversions
relative to subgeneric standard. Female from Georgia, Dade Co., 19 May 1998; Ma = marker, Sp = spongy. 5,
Centromere region of chromosome I. Male from Georgia, Dade Co., 19 May 1998; C’ = condensed, Y-linked

centromere band.

Cheaha State Park, Dry Creek, 33°28.23'N
85°48.72'N, 19 May 1998 (1 6, 6 Q);
GEORGIA: Dade Co, Johnson’s Crook,
Newsome Gap Road, 0.8 km west of Moore
Rd., 34°47.23'N 85°28.25'W, 19 May 1998
(2 6,7 2); MASSACHUSETTS: Berkshire
Co., Jug End Rd., 0.24 km west of Guilded
Hollow Road, 42°8.99'N 73°26.98'W, 29
June 1998 (2 36, 16 2); NEW HAMP-
SHIRE: Carroll Co., Bear Notch Rd., 3.2
km “north “ef “SR 112." “44° 1750N
71°19.13'W, 24 July 1998 (1 3); Grafton
Co., SR118, 10.5 km southwest of SR 112,

43°59.00'N 71°47.93'W, 24 July 1998 (2 6,
7 2); NEW JERSEY: Sussex Co., Tillman
Brook, 41°0.15'N 74°0.86'W, 22 June 1998
(1 36, 4 2); NORTH CAROLINA: Hay-
wood Co., US Rt. 276, 1.6 km north of Cru-
so, 35°26.4'N 82°48.8’W, 7 June 1998 (2
6); Madison Co., Silver Mine Creek
35°0.53'N 82°0.48'W, 14 May 1998 (3 gd,
2 2); SOUTH CAROLINA: Pickens Co.,
Abner Creek, SR 1105, 4.8 km east of Hwy
178, 35°4.27'N 82°47.11W’, 23 May 1998
(14 6,5 2); 2 June 1998 (10 2); Oil Camp
Creek, Oil Camp Creek Rd., 0.8 km west

VOLUME 102, NUMBER 4

847

Fig. 6. Chromosome arm IIS of Simulium parnassum, with landmarks of Rothfels et al. (1978). C = cen-
tromere region. Section numbers on bottom indicate S. parnassum standard banding sequence; those on top refer
to Simulium subgeneric standard sequence of Rothfels et al. (1978). Bracket indicates B inversion of Rothfels
et al. (1978). Arrows indicate breakpoints of two overlapping inversions relative to subgeneric standard. Female

from Massachusetts, Berkshire Co., 29 June 1998. Bu = bulge, Rb = ring of Balbiani, Tr = trapezoidal.

of River Falls Rd., 35°6.65'’N 82°34.08’W,
8 May 1998 (7 6, 1 2); TENNESSEE:
Monroe Co., Buckhorn Cr., 35°19.98'’N
84°9.83'W, 6 June 1998 (2 3); VIRGINIA:
Augusta Co., East Dry Branch, SR 688, 3.2
km: north of SR 42, 38°12.70'N
79°16.31'W, 25 June 1998 (8 2); NEW
BRUNSWICK: Restigouch Co., Collector
Hwy. 180, 37 km west of Bathurst, 27 July
1998 (3 3, 12 2); QUEBEC, Gaspé Pen-
insula, Hwy. 132, 5.6 km south of Routh-
ierville, 29 July 1998 (5 6, 19 @).

RESULTS

Two pupal forms of S. parnassum were
discovered among the 472 pupae examined.

Rugose pupae had a raised pattern of retic-
ulation on the dorsal surface of the head
and thorax (Fig. 1). Smooth pupae lacked
surface sculpturing, although the pattern of
rugosity could be seen, with substage light-
ing, as a vague outline on the head and tho-
rax of the pupal exuviae (Fig. 2). Interme-
diates between the smooth and rugose
forms were not found. No additional mor-
phological characters in larvae, pupae, or
adults were found that correlated with the
smooth and rugose pupal forms.

Smooth pupae were collected from four
streams. They were found with the rugose
form in streams in Pickens Co., South Car-

Fig. 7.

Chromosome arm IIIS of Simulium parnassum, with landmarks of Rothfels et al. (1978). C =

centromere region. Section numbers are those of both S. parnassum standard banding sequence and Simulium
subgeneric standard of Rothfels et al. (1978). Male from South Carolina, Pickens Co., Oil Camp Creek, 8 May
1998. Bl = blister, Ca = capsule.

848 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 8-9. Silk-gland chromosomes of S. parnassum, with landmarks of Rothfels et al. (1978). C = centro-
mere region. 8, Chromosome arm IIL. Section numbers refer to Simulium parnassum standard banding sequence.
Bracket indicates simple inversion relative to Simulium subgeneric standard of Rothfels et al. (1978). Female
from Tennessee, Monroe Co., 6 June 1998 (sections 55 through 70) plus female from North Carolina, Haywood
Co., 7 June 1998 (section 71). DNA = DNA puff, gB = gray band, Po = polar, St = saw tooth, Sy =

VOLUME 102, NUMBER 4

olina (Smith and Abner creeks), where they
were the predominant form, and in Berk-
shire Co., Massachusetts. In Transylvania
Co., North Carolina (South Prong Gladdy
Fork), one smooth pupa and no rugose pu-
pae were collected. Pupae from near the
type localities of S. parnassum and S. hy-
dationis were rugose. No differences in sea-
sonality were detected in streams where
both pupal forms occurred (Table 1), al-
though the rugose form was infrequent in
these streams. Seasonal differences, how-
ever, might have been apparent if popula-
tions of each form had been larger.

In Smith Creek, where a minimum-max-
imum thermometer had been placed, larvae
first appeared on 13 March 1998; water
temperature was 3.0—9.0°C for the three
days prior to collection. Pupae were first
found in Smith Creek on 25 April 1998;
water temperature was 11.0—12.0°C during
the 11 days prior to collection. In 1998, the
last larvae were found at Smith Creek on 2
June (water temperature for 9 days prior to
collection = 17.5—18.5°C) and in 1999 on
1 June (water temperature for 3 days prior
to collection = 17.5—18.0°C). Pupae were
first collected from Abner Creek on 2 June
1998 (temperature at time of collection =
18.0°C) and 1 June 1999 (16.0°C). The last
larvae and pupae were collected from Ab-
ner Creek on 23 June 1998 (18.0°C) and 10
Jone 19995(15:0°C),

Of 211 larvae prepared for chromosomal
analysis, 72% were read, band for band, in
their entirety. The banding sequences of the
short arms of chromosomes I and III (Figs.
3, 7) matched those of the subgeneric stan-
dard, although we were unable to match all
of the fine bands in the base of IS (sections
19 and 20). The remaining four arms had
fixed rearrangements relative to the subge-

849

neric standard. IL had one central inversion,
two overlapping basal inversions, and a
complex of three inversions distally (Fig.
4). ILS carried the B inversion of Rothfels
et al. (1978), plus two additional inversions
(Fig. 6). IfL had one central inversion (Fig.
8). The ITIL arm, with the nucleolar orga-
nizer displaced more distally relative to the
subgeneric standard, was highly rearranged
(Fig. 9). Many bands could be recognized,
but we conservatively matched only the
bands in the base of IIIL, relative to the
subgeneric standard, and indicated the char-
acteristic “‘marker’’? (sensu Rothfels et al.
1978), which was partitioned by rearrange-
ments into two pieces.

We found no fixed or floating inversions
in our material relative to the S. parnassum
standard sequence. Chromosome I is impli-
cated as the sex chromosome. Two males
from Georgia (Newsome Gap Road) had
differentiated centromere regions in chro-
mosome I; the centromere band of one ho-
mologue was expanded (standard) more
than the other (Fig. 5). About 52% of males
(n = 44) from other sites showed a failure
to pair on either side of the centromere of
chromosome I. This failure to pair was
found at sites with and without the two pu-
pal forms. One male from Abner Creek (13
May 1998) was heterozygous for expres-
sion of the nucleolar organizer. Simulium
parnassum was otherwise chromosomally
monomorphic.

DISCUSSION

A new pupal form of S. parnassum was
discovered in which the reticulation of the
head and thorax, long used as a diagnostic
character (e.g., Stone and Jamnback 1955),
was absent. No intermediates between
smooth and rugose pupae were found; how-

ee

symmetrical, 3 = three sharp. 9, Chromosome arm IIIL. Section numbers on bottom indicate S. parnassum
standard banding sequence; those on top refer to Simulium subgeneric standard sequence of Rothfels et al.
(1978). Arrows indicate breakpoints of inversions. N.O. = nucleolar organizer, M = marker (divided in two

pieces). Female from South Carolina, Pickens Co., Oil Camp Creek, 14 May 1998.

850

ever, we discovered no additional informa-
tion, either morphological, chromosomal,
distributional, or seasonal, to suggest that
two species are present. The synonymy of
S. hydationis with S. parnassum (Stone and
Jamnback 1955) is justified, both morpho-
logically and cytologically, because mate-
rial from near the two type localities was
morphologically and chromosomally ho-
mogeneous. However, the possibility that
the two pupal forms of S. parnassum rep-
resent homosequential species or that hom-
osequential sibling species exist, as they do
in other black flies (e.g., Henderson 1986),
cannot be excluded.

Relative to the Simulium subgeneric stan-
dard of Rothfels et al. (1978), we found
only two inversions that are shared with
other taxa. The IIS-B inversion of Rothfels
et al. (1978) is shared with most species in
subgenus Simulium (Adler et al. 1999), as
is an inversion in IIIL that has one of its
breakpoints at the 91/94 junction (Fig. 9)
(Adler, unpublished). Because IIIL is not
fully resolved, shared inversions with other
taxa could be present. The remaining in-
versions in IL, IIS, and IIL are apparently
autapotypic for S. parnassum. Chromosom-
al evidence does not suggest a relation with
the S. tuberosum species group, which is
defined by at least eight unique rearrange-
ments, including four in IIL (Adler and Ku-
usela 1994), nor does it provide resolution
of relationship with other species groups.
Tentative placement of S. parnassum in a
separate species group, the S. parnassum
species group, is therefore more appropri-
ate.

Simulium parnassum was univoltine in
South Carolina. In Pennsylvania, it has
been considered univoltine but with over-
lapping cohorts (Tessler 1991). Other au-
thors (e.g., Stone and Snoddy 1969, Cupp
and Gordon 1983) suggest that S. parnas-
sum completes more than one generation
per year, although conclusive evidence sup-
porting these claims is lacking.

Based on the results of our study, we
consider S. parnassum a single species, al-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

beit polymorphic in pupal surface texture.
We suggest, however, that additional (e.g.,
molecular) evidence be brought to bear to
corroborate or falsify this hypothesis.

ACKNOWLEDGMENTS

We thank D. C. Currie, J. C. Morse, and
A. G. Wheeler for reviewing the manu-
script, and C. E. Beard for technical assis-
tance. This work was supported, in part, by
grant no. DEB-9629456 from the National
Science Foundation to PHA. This is Tech-
nical Contribution No. 4545 of the South
Carolina Agriculture & Forestry Research
System.

LITERATURE CITED

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The black flies (Diptera: Simuliidae) of Ontario.
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cation of blood meals of blackflies (Diptera: Sim-
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Zibilske. 1986. Experimental applications of B.t.i.
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PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 852-861

TYPES OF SAWFLIES DESCRIBED IN THE GENUS PONTANIA A. COSTA
(HYMENOPTERA: TENTHREDINIDAE) IN THE ILLINOIS
NATURAL HISTORY SURVEY

ALEXEY G. ZINOVJEV AND DAVID R. SMITH

(AGZ) Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Rus-
sia (e-mail: zag@zisp.spb.su); (DRS) Systematic Entomology Laboratory, PSI, Agricul-
tural Research Service, U.S. Department of Agriculture, % National Museum of Natural
History, MRC-168, Washington, DC 20560-0168, U.S.A. (e-mail: dsmith @sel.barc.usda.
gov)

Abstract.—The types of ten sawfly species described in the genus Pontania by
MacGillivray and Ross are examined. This study has resulted in the following taxonomic
changes: Pontania daedala MacGillivray is a new synonym of Pontania proxima (Ser-
ville); Pontania demissa MacGillivray is a new synonym of Eupontania gracilis (Mar-
latt); Pontania devincta MacGillivray is a new synonym of Pontania populi (Marlatt);
Phyllocolpa dedecora (MacGillivray), n. comb.; Phyllocolpa derosa (MacGillivray), n.
comb.; and Nematus dotata (MacGillivray), n. comb. Notes are given on the types and

hosts where known.

Key Words:

This study of ten sawfly types described
in the genus Pontania A. Costa deposited
in the Illinois Natural History Survey,
Champaign, Illinois (INHS) focuses on
some that were reared by Yuasa and treated
by Yuasa (1922) in his work on sawfly lar-
vae and concurrently or later described by
MacGillivray (1921, 1923); two species de-
scribed by Ross (1929) are also included.
Placement of these species by Smith (1979)
was based on the generic concepts of that
time. Study of these specimens is prelimi-
nary to further work on eastern North
American gall-forming sawflies and an up-
dated catalog of Nearctic gall-forming saw-
flies of the genera Euura Newman, Pontan-
ia A. Costa, Eupontania Zinovjev, and
Phyllocolpa Benson. Generic concepts have
changed since Smith (1979) and the gall-
forming sawflies of North America are
poorly known. As a result of extensive

galls, leaf rolls, Phyllocolpa, Eitelius, Nematus

work on the Palearctic and world fauna by
Zinovjev (1993) and Zinovjev and Vikberg
(in press), the types of Nearctic species
need to be reexamined and incorporated
into the current classification.

We give some notes on each species, il-
lustrate the types, and attempt to associate
the MacGillivray species with species treat-
ed in Yuasa (1922). We propose three new
synonymies and three new combinations.
The other four species have been treated
correctly in previous literature. Species
headings are in the original combinations.

SPECIES DESCRIBED BY MACGILLIVRAY

MacGillivray (1921) described 19 spe-
cies of Nematinae, including seven species
of Pontania “‘as a result of an extended se-
ries of collecting and breeding of sawfly
larvae by Dr. H. Yuasa at Ithaca, New York,
and by the Maine Agricultural Experiment

VOLUME 102, NUMBER 4

Station at Orono, Maine.” The larvae of all
these species were supposed to be described
by Yuasa (1922). We examined part of Yu-
asa’s collection from Maine (Orono) which
is deposited in the National Museum of
Natural History, Smithsonian Institution
(USNM). It consists of a set of vials with
larvae or remnants of leaves with galls kept
in alcohol.

MacGillivray (1923) described 22 sawfly
species in a paper on sawflies collected
from ‘“‘the Katmai Expedition to Alaska.”
Only one of them was described under the
genus Pontania and currently (after Benson
1960) it is treated as a synonym of Phyl-
locolpa excavata (Marlatt 1896) (= mega-
cephala Rohwer 1908; destricta Mac-
Gillivray 1923; apicifrons Malaise 1932;
carinifrons Benson 1940) (Benson 1960,
Smith 1979).

Pontania daedala MacGillivray 1921: 33
(Ergs2 ie 105418)

Type locality.—Ithaca, New York.

Lectotype.—Designated by Frison (1927),
female, “‘7-6’’; “‘cocoon; Ithaca N. Y. 21
Aug. 11 [or 17? ]”; “Type of female Pon-
tania daedala A.D. MacGillivray [red la-
bell]; “INHS TYPE #1592.”

Valid name.—Pontania proxima (Ser-
ville 1823) (= P. daedala MacGillivray), n.
syn.

Notes.—The species was reared by Yu-
asa (1922), and he described the larvae of
P. proxima (as Pontania hyalina Norton)
from the material numbered Y-7-1, -7-4-1,
-8.8, Cu-cu 201, and M-92. The remnants
of host-plant leaves in the vial (M-92) de-
posited in the USNM cannot be identified
to species, but the sawfly is undoubtedly P.
proxima. We did not find the number ‘‘7-
6” in Yuasa (1922).

The female holotype has asymmetric
mandibles; ovipositor (sawsheath + oblong
plate) as long as the hindtibia (Fig. 10);
short cercus, about half the length of the
sheath; antennal hollow glabrous and shin-
ing with only few hairs on the lower part
(Fig. 18), on the upper part the border be-

853

tween the inner orbit and antennal hollow
convex and shining; and supraclypeal area
glabrous. This combination of characters
places it in the proxima group. Mac-
Gillivray mentioned the black color of the
pronotum which may separate this species
from P. hyalina Norton (a synonym of P.
proxima), but in the holotype, the margins
of the pronotum are pale, exactly as in spec-
imens of P. proxima from Europe.

Pontania decrepita MacGillivray 1921: 33
(Fig. 11)

Type locality.—Ithaca, New York.

Holotype female.—‘“Ithaca, N. Y. 21
July °17; 35-2-5”; “‘Type of female Pon-
tania decrepita A.D. MacGillivray [red la-
bel]”; “INHS TYPE #1594.”

Valid name.—Eitelius gregarius (Marlatt
1896) (= Pontania decrepita Mac-
Gillivray). Synonymy by Ross (1951).

Note.—This species (under the name Mi-
cronematus gregarius Marlatt) was treated
by Yuasa (1922) as being collected by him-
self (““Y’’), but without mentioning its rear-
ing number, and he did not mention the
name Pontania decrepita MacGillivray.
The holotype has claws of the same shape
as characteristic for Eitelius and also the
short, broad ovipositor in the holotype (Fig.
11) is typical for free-feeding Nematinae.

Pontania dedecora MacGillivray 1921: 32
ig: 7)

Type locality—Ithaca, New York.

Lectotype.—Designated by _ Frison
(1927), female, designated here; “*185a2,
May 24, 1919”; *“‘Type of Pontania dedcora
[sic!] A.D. MacGillivray, female [red la-
bel]; “INHS TYPE # 1595.”’

Valid name.—Phyllocolpa dedecora
(MacGillivray), n. comb.

Note.—This species was described from
two females (““Nos. 185a2, 8.51 (?)-1-1’’).
Yuasa (1922) mentions none of these num-
bers.

854

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Type specimens of Pontania in lateral view. 1, P. daedala. 2, P. demissa. 3, P. devincta.

VOLUME 102, NUMBER 4 855

Figs. 4-6. Type specimens of Pontania in lateral view. 4, P. dotata. 5, P. destricta. 6, P. pepii.

856

Figs. 7-9. Type specimens of Pontania dorsal view.

Pontania demissa MacGillivray 1921: 33
(Figs. 2, 12)

Type locality.—Ithaca, New York.

Lectotype.—Designated by Frison (1927),
female, “‘191-1-1"; “Ithaca N. Y.’’; ‘““Type
of female Pontania demissa A.D. Mac-
Gillivray [red label]’’; “INHS TYPE
FIST.”

Valid name.—Eupontania gracilis (Mar-
latt 1896) (= Pontania demissa Mac-
Gillivray), n. syn.

Notes.—Larvae and galls were described
by Yuasa (1922) as P. demissa Mac-
Gillivray 1921 (Y-191-1-1). We have not
seen larval material with this number.

Pontania demissa could be treated either
as synonym of Eupontania gracilis (Marlatt
1896) or E. petiolaridis (Rohwer 1917).
Both these species are associated with dif-
ferent willow species: Salix sericea Marsh.,
and Salix petiolaris Smith, respectively.
These sawflies are very similar, and they
might prove to be conspecific. Both willow
species occur in the vicinity of Ithaca, and,
at least in herbarium material studied by
AGZ, the galls are known also from both
host plants. However, Salix sericea seem to
be more common there and galls only on
this willow were found in the fall of 1997

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

7, P. dedecora. 8, P. derosa. 9, P. mariana.

around Ithaca by AGZ and T. Carr. The type
of Pontania demissa fits best the type ma-
terial of E. gracilis, and we consider them
as synonymous.

Pontania derosa MacGillivray 1921: 34
(Fig. 8, 19)

Type locality.—Ithaca, New York.

Holotype.—Female, *‘142-1-1 May 13,
1919”; “Ithaca, N. Y¥.’’; Dy pexor female
Pontania derosa A.D. MacGillivray. [red la-
bell]; “INHS TYPE #1599.”

Valid name.—Phyllocolpa
(MacGillivray), n. comb.

Notes.—The leaf-rolls and larvae were
described by Yuasa (1922) in his key, but
the host plant is unknown. The species is
very similar to Phyllocolpa nigrita Marlatt,
and these species may be conspecific.

derosa

Pontania devincta MacGillivray 1921: 34
(Figs. 3, 13)

Type locality—Orono, Maine.

Lectotype.—Designated by Frison (1927),
female: ‘““Me. Exp. Sta. Lot. 1694 Sub. 9, 1
Aug. 13”; “Type of female Pontania de-
vincta A.D. MacGillivray [red label]”’;
“INHS TYPE #1601.”

Valid name.—Pontania populi Marlatt

VOLUME 102, NUMBER 4

Figs. 10-13.
12, P. demissa. 13, P. devincta.

1896 (= Pontania devincta MacGillivray),
n. syn.

Notes.—MacGillivray described this spe-
cies from two (or more) females, with num-
bers Subs. 9 and 226. Yuasa (1922) men-
tioned the number 226, but he did not men-
tion 9. The galls described by Yuasa, kept
in the USNM (Maine: Bangor Bay, “*M-
226’), belong to the gall-making Eupon-
tania consors (Marlatt 1898). The plant
species is Salix humilis Marsh. with com-
paratively large, glabrous leaves (which is
typical of plants growing in shadowy for-
ests). We have not seen reared specimens
with Nr. 226, but according to the galls, the
paralectotype with this number belongs to
Eupontania consors. However, the lecto-
type belongs to the leaf-rolling species
group of Pontania s. str. and is conspecific
with Pontania populi, a leaf-rolling species

857

Apex of abdomen and sheath in lateral view of Pontania types. 10, P. daedala. 11, P. decrepita.

on Populus grandidentata Michx. It is con-
specific also with a specimen identified as
P. devincta by R. B. Benson (kept in The
Natural History Museum, London) who
(1960) incorrectly synonymized Pontania
devincta under P. excavata Marlatt.

Pontania dotata MacGillivray 1921: 34
(Fig. 4)

Type locality.—Ithaca, New York.

Lectotype.—Designated by Frison (1927),
male, “8-48 (?)-1-1”; “Ithaca N.Y. 25 Aug.
18”; “‘Type of male Pontania dotatus A.D.
MacGillivray [red label]; ““INHS TYPE
#1603.”

Valid name.—Nematus dotatus (Mac-
Gillivray), n. comb.

Note.—This species belongs to the genus
Nematus Panzer (= Pteronidea Rohwer)
but cannot be placed more precisely at pre-

858 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 14-17. 14-15, Apex of abdomen and sheath in lateral view of Pontania types. 14, P. destricta. 15,
P. excavata. 16-17, Head and thorax in lateral view of Pontania types. 16, P. destricta. 17, P. excavata.

VOLUME 102, NUMBER 4 859

Figs 18-19. Head in frontal view of Pontania types. 18, P. daedala. 19, P. derosa.

860

sent. It seems that this specimen was reared
by Yuasa, but he did not mention this name
or the number ‘‘8-48 (?)-1-1” in his keys.
However, the larvae with a similar number
(Y-8-48(?)-1) were described by Yuasa as
Pteronidea sp. 6.

Pontania destricta MacGillivray 1923:
168
(Figs. 5, 14, 16)

Type locality —Alaska: Katmai

Holotype.—Female (glued on the card-
board triangular point): ““Katmai Alaska
June ’17”’; “Jas S Hine Collector’; Type of
Pontania destricta A.D. MacGillivray fe-
male”; “INHS TYPE #1600.”

Valid name.—Phyllocolpa excavata
(Marlatt 1896) (= Pontania destricta
MacGillivray). Synonymy after Benson
(1960).

Note.—This species belongs to the Phyl-
locolpa leucapsis group, characterized by
glabrous and shining antennal hollows and
the hairs of the upper posterior part of the
mesepisternum directed upwards. We are
not sure that it is conspecific with Phyllo-
colpa excavata (Marlatt 1896), described
from ‘California, Colorado” and “*Veta
Pass, Colo.” The type of P. destricta differs
from that of Phyllocolpa excavata Marlatt
1896 (Figs. 15, 17) by a less rounded saw-
sheath with a distinct emargination beneath
in lateral view (Figs. 14—15), by the shape
of the inner orbits in lower part (Figs. 16—
17), and by the darker coloration (while
type of P. exavata is paler colored than it
is typical for this species).

SPECIES DESCRIBED BY ROSS

H. H. Ross (1929) described four species
in his paper revising Marlatt’s “Group I’
of Pontania (= Phyllocolpa Benson). Types
of two of them are deposited in INHS.

Pontania mariana Ross, 1929: 91
(i2..9)
Type locality.—Canada, B.C., Vancou-
ver.
Holotype.—Female, reared from _leaf-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

curl on Populus balsamifera; ‘‘Vancouver.
B. @) Larva coll; x24. 1927 HOH: Ross”:
‘““emerged VI. 4. 1928”’; [red label] ““HO-
LOTYPE Pontania mariana Ross female’’;
“INHS TYPE #1606.”

Valid name.—Phyllocolpa mariana
(Ross).

Host plant.—Populus balsamifera L.

Additional material examined.—Female
(paratype), Vancouver. B. C. Larv. Coll. IX.
4. 1928 H. H. Ross; Reared from Populus
balsamifera \eaf-curl; emerged VI. 4. 1928;
[red label] Paratype No 42755; PARATYPE
Pontania mariana H. H. Ross. Deposited in
the USNM.

Pontania pepii Ross, 1929: 95
(Fig. 6)

Type locality—NMontana, Florence.

Holotype.—Female, “‘Mont. Exper. Sta-
tion Coll. Florence Mont. June 1, 1912”;
‘“‘“HOLOTYPE Pontania female peppii Ross
[red label]; ““INHS TYPE #1608.”

Valid name.—Phyllocolpa pepii (Ross).

ACKNOWLEDGMENTS

We thank Kathy Zeiders, Illinois Natural
History Survey, Champaign, Illinois, for al-
lowing examination of the MacGillivray
and Ross types. We extend our appreciation
to Terry Nuhn for taking the photos and
Cathy Anderson for arranging the plates.
Thanks to the following who reviewed the
manuscript: H. Goulet, Agriculture and
Agri-Food Canada, Ottawa, and S. Scheffer,
Systematic Entomology Laboratory, U. S.
Department of Agriculture, Beltsville, MD.

LITERATURE CITED

Benson R. B. 1960. Studies in Pontania (Hym., Ten-
thredinidae). Bulletin of the British Museum (Nat-
ural History) Entomology 8(9): 367—384.

Frison, T. H. 1927. A list of the insect types in the
collections of the Illinois State Natural History
Survey and the University of Illinois. State of I-
linois Department of Registration and Education,
Division of the Natural History Survey, Bulletin
16, Article 4, pp. 137-309.

MacGillivray A. D. 1921. New species of Nemati-
nae—(Hymenoptera). Journal of the New York
Entomological Society 29(1): 27-35.

VOLUME 102, NUMBER 4

. 1923. Saw-flies of the Katmai expedition to
Alaska. Journal of the New York Entomological
Society 31(4): 163-171.

Marlatt, C. L. 1896. Revision of the Nematinae of
North America, a subfamily of leaf-feeding Hy-
menoptera of the family Tenthredinidae. United
States Department of Agriculture, Technical Se-
ries No. 3, 133 pp.

. 1898. Some new nematids. Canadian Ento-
mologist 30: 302-316.

Rohwer, S. A. 1917. Pontania petiolaridis, new spe-
cies, p. 19. In Cosens, A., ed., Reports on Insects
of the Year, Division No. 3, Toronto District. For-
ty-Seventh Annual Report of the Entomological
Society of Ontario, 1916, pp. 18-19.

Ross H. H. 1929. A study of Marlatt’s group I of the
genus Pontania with descriptions of four new spe-
cies (Hymenoptera: Tenthredinidae). Proceedings
of the Entomological Society of Washington
31(5): 91-97.

. 1951. Symphyta, pp. 4-87. In Muesebeck, C.

861

FE W., K. V. Krombein, and H. K. Townes, eds.,
Hymenoptera of America North of Mexico, Syn-
optic Catalog. United States Department of Agri-
culture, Agriculture Monograph 2, 1420 pp.

Smith D. R. 1979. Symphyta, pp. 3-137. Jn Krombein,
K. V., P. D. Hurd, Jr., D. R. Smith, and B. D.
Burks, eds., Catalog of Hymenoptera in America
North of Mexico, Vol. 1, Smithsonian Institution
Press, Washington, D.C.

Yuasa H. 1922. A classification of the larvae of the
Tenthredinoidea. Illinois Biological Monographs
7(4), 172 pp.

Zinovjev, A.G. 1993. Subgenera and Palaearctic spe-
cies groups of the genus Pontania, with notes on
the taxonomy of some European species of the
viminalis-group (Hymenoptera: Tenthredinidae).
Zoosystematica Rossica 2: 145-154.

Zinovjev A. G. and V. Vikberg. In press. The sawflies
of the Pontania crassispina-group with a key for
genera of the subtribe Euurina (Hymenoptera,
Tenthredinidae, Nematinae). Entomologicae Scan-
dinavicae.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 862-868

THE LARVA AND PUPA OF SCHIZONYXHELEA FORATTINI WIRTH AND
GROGAN (DIPTERA: CERATOPOGONIDAE) WITH A DISCUSSION OF THE
PHYLOGENETIC RELATIONSHIPS OF THE GENUS

ART BORKENT

Research Associate, Royal British Columbia Museum and the American Museum of
Natural History, 1171 Mallory Road, R1-S20-C43, Enderby, British Columbia, VOE 1V0O,

Canada (e-mail: aborkent @jetstream.net)

Abstract.—The larva and pupa of Schizonyxhelea forattinii Wirth and Grogan are de-
scribed for the first time. Larval, pupal and adult character states suggest that this genus
is closely related to Stilobezzia Kieffer and may even be most closely related to only

some species of Stilobezzia.

Resumen.—Se describen por primera vez larva y pupa de Schizonyxhelea forattinii
Wirth y Grogan. El estado de los caracteres de larva, pupa y adulto, sugiere que este
género se halla muy relacionado a Stilobezzia Kieffer, y puede atin, ser mas cercano a

solo algunas especies de Stilobezzia.

Key Words:
Costa Rica, Neotropical

The immatures of the Ceratopogonidae
remain as one of the most poorly known
life stages of all nematoceran families. As
part of a project to generate descriptions,
keys and phylogenetic interpretation of
these stages (Borkent, in preparation), this
paper provides a description of the larva
and pupa of Schizonyxhelea forattinii Wirth
and Grogan and interprets available char-
acter stages of the genus Schizonyxhelea
Clastrier from a cladistic perspective.

Schizonyxhelea includes two Neotropical
species and both are very small in size, with
wing lengths of 0.7—0.8 mm. When the ge-
nus was first proposed by Clastrier (1984),
he suggested that the single known species,
S. guyana, was similar to species of Nan-
nohelea Grogan and Wirth and Bothahelea
Grogan and Wirth. Wirth and Grogan
(1988) described a second species, S. for-
attinii, including the first male of the genus
and hypothesized that Schizonyxhelea was

Schizonyxhelea, Stilobezzia, immatures, taxonomy, phylogeny, systematics,

related to a group of genera with reduced
features: Baeohelea Wirth and Blanton,
Baeodasymyia Clastrier and Raccurt, Nan-
nohelea Grogan and Wirth and Rhynchoh-
elea Wirth and Blanton. Since then Borkent
(1992, 1995), Borkent and Craig (1999) and
Grogan and Borkent (1992) have shown
that Baeohelea and Baeodasymyia, as sister
genera, are likely an early lineage within
the Ceratopogonini and that Nannohelea
and Rhynchohelea belong to a group which
also includes the genera Brachypogon Kief-
fer, Ceratoculicoides Wirth and Ratanawor-
abhan and Sinhalohelea Grogan and Bor-
kent. The genus Schizonyxhelea has re-
mained an enigma. Discovery of the im-
matures and examination of fresh adult
material of S. forattinii provided an oppor-
tunity to further investigate the genus.

MATERIALS AND METHODS

Terms for larval and pupal structures fol-
low Lawson (1951). The single live larva

VOLUME 102, NUMBER 4

was discovered by placing a mud sample in
water and shaking the container gently. The
swimming larva was then captured with an
eye-dropper and reared in a petri dish (5 cm
in diameter) with a small amount of sub-
strate (mud and detritus) from the original
habitat. The larva and subsequent pupa
were kept at ambient temperature (approx-
imately 25°C) and checked daily. When
needed, a small amount of water was added
to the petri dish to keep the sample from
drying. The larval and pupal exuviae and
partially emerged adult were preserved in
70% ethanol and slide-mounted using the
method described by Borkent and Bissett
(1990).

Adults were also captured in the field by
sweeping vegetation with an aerial net;
these were examined as cleared specimens
in glycerine and on slides. Specimens were
compared to the holotype of S. forattinii.

The specimens are deposited in the Ca-
nadian National Collection in Ottawa, On-
tario, Canada.

DESCRIPTION OF LARVA AND PUPA

Fourth instar larva.—Body length un-
known. Head capsule (Figs. 1A, B) rela-
tively short (111 pm), blunt (head length/
width = 1.28), uniformly medium brown.
Collar slightly darker than rest of head cap-
sule, well-developed ventrally with ventro-
lateral anterior apodeme, separated ventro-
medially, terminating dorsolaterally. Frontal
suture extending to near anterior margin of
labrum. Ventral suture short. Most setae
simple with the following divided or plu-
mose: s, v, posterior 0; arrangement as in
Figs. 1A, B; bases of sensilla more or else
equal in size; following sensilla not visible:
j, t, Z, xX, n. Antenna short, further details
not visible. Eye unknown. Labrum short,
wide, details not visible. Mandible elon-
gate, basal portion straight, apical third
curved, sensilla not visible. Maxilla not
clearly visible, palpus short. Hypostoma a
broadly rounded projection. Epipharynx
with 1 pair of serrate combs, divided me-
dially (Fig. 1B). Thorax, abdomen with

863

more or less uniform light pink pigmenta-
tion. Caudal segment, anal papillae un-
known.

Pupa.—Length 1.29 mm. General color-
ation uniformly light brown. Body surface
generally smooth with only a few tubercles
and spicules associated with setae, a few
restricted to the medioanterodorsal and me-
dioanteroventral areas of abdominal seg-
ments 3—8 and anal segment. Operculum
(Fig. 1C) with well-developed anteromar-
ginal seta located on tubercle, with pore at
tubercle base; dorsolateral margin with row
of well-developed spines. One well-devel-
oped anterodorsal seta on moderately elon-
gate, rounded tubercle (Fig. 1G). One long,
slender and two shorter dorsolateral setae
(Fig. 1H). Two short dorsomedial seta, one
thicker than other (Fig. 1F). Five dorsal
sensilla, 1 short thick seta, 11, iv short, slen-
der setae, v, Vi pores; i, ii, 1v on single,
rounded, short, separate tubercles (Fig. 2A).
Ventromedian setae not visible (if present,
they are very minute). Two ventrolateral se-
tae slender, bases somewhat separate (Fig.
1E). Respiratory organ (Fig. 1D) somewhat
curved, smooth; four spiracles situated
along length, with 5—6 spiracles situated
apically, opening at surface; tracheal tube
of more or less equal diameter for length.
Metathorax not divided medially, with me-
dial protuberance from scutum protruding
to near posterior margin. Abdomen with se-
tae separate from one another (none on
common tubercle), each bordered by comb-
like or single elongate spicules. Segment 4
setal pattern (Fig. 2C) with 2 d.a.s.m., 4
dp.nisw ls Laisa. 3. pam: 3: yng setae
d.p.m. i—ili not present or very small; setae
dipantiv,, Lasimeelipam: 1:, ii, yall thick;
stout. Anal segment (Fig. 2B) with caudal
spine moderately elongate, somewhat
curved apically, directed posterolaterally.

Bionomics.—The single larva was col-
lected on December 17, 1993, 2 km north-
east of Tarcoles, Costa Rica from wet mud
in a small seep which flowed into the out-
flow of a larger spring on private property
directly west of Reserva Carara. The larva

864 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Mandible

ZZ, Leppup) Pl) I

Fig. 1. Structures of Schizonyxhelea forattinii. A, Larval head capsule in dorsal view. B, Larval head capsule
in ventral view. C, Pupal operculum. D, Pupal respiratory organ. E, Pupal mouthparts in ventral view. E Pupal
dorsomedial setae. G, Pupal anterodorsal seta. H, Pupal dorsolateral setae.

swam with the same rapid serpentine mo- two male and 11 female adult S. forattinii.
tion as do so many Ceratopogoninae. Dur- Each of the sites was at, or very near, a first
ing eight months in Costa Rica (July, 1993— order seep or spring, or a small stream and
Feb. 1994) I collected only one larva and specimens were collected from August to

VOLUME 102, NUMBER 4 865

7
|
IV «

>
ii

. fo)

B
vi o A
Dorsal Ventral
dasm. ts i
|.a.S.m.

|.p.m. hw hw

Fig. 2. Pupal structures of Schizonyxhelea forattinii. A, Dorsal setae. B, Anal segment in ventral view. C,
Setae of fourth abdominal segment.

866

December (3 sites at 2 km NE Tarcoles
from Aug. 3 to Dec. 17, 1993; a first order
spring in Atenas Oct. 1 to Dec. 7, 1993).
This suggests that immatures of this species
are restricted to small lotic habitats.

DISCUSSION

There are two newly discovered charac-
ter states which provide cladistic informa-
tion regarding the phylogenetic position of
Schizonyxhelea. The first is the presence of
plumose setae on the head capsule of the
larva, which merely indicates the unsur-
prising conclusion that this taxon belongs
within that group of Ceratopogoninae
which is the sister group of Ceratopogon
Meigen and, possibly Baeodasymyia +
Baeohelea. This character is discussed fur-
ther by Borkent and Craig (1999). The sec-
ond character state concerns the nature of
the male aedeagus. Borkent (1995: 97—98)
discussed (as his character 39) the uniquely
divided aedeagus of species of Stilobezzia
Kieffer. Wirth and Grogan (1988) pointed
out that the male aedeagus of Schizonyxhe-
lea was difficult to interpret. However, re-
examination of fresh material here, espe-
cially from a posterior aspect, shows that
the aedeagus of S. forattinii is divided me-
dially, indicating at least a sister group re-
lationship to, or within, Stilobezzia.

Further character states suggest that Schi-
zonyxhelea is actually closely related to a
species group within Stilobezzia and they
are discussed below. Although these fea-
tures are not yet understood cladistically,
continuing studies show promise for the in-
terpretation of most of these (Borkent, in
preparation).

—Larval head capsule sensilla q, s, k are
all close to each other (present in Stilobez-
zia flavirostris (Winnertz) and S. papillata
Remm, the only two species of Stilobezzia
for which this character state has been de-
scribed). The character state appears to be
unique within the Ceratopogonini but is
also present in some Sphaeromiini and Pal-
pomyiini.

—The following features are shared with

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

the pupa of Stilobezzia bulla Thomsen: pu-
pal respiratory organ shape and distribution
of spiracles virtually identical; presence of
thick spines on margin of operculum (also
present in many other genera); same num-
ber and similar size of thoracic setae; spe-
cific distribution of sensilla and spines on
segment 4 especially including the anterior
lateral sensilla and the presence of comb-
like spines associated with most sensilla
very similar; presence of two medial, low
projections on the dorsum of the anal seg-
ment; presence of spines on posterior mar-
gin of apicolateral processes of anal seg-
ment.

—AlIl Stilobezzia pupae are missing one of
the dorsal setae (probably # 3) and this ap-
pears to be nearly unique within the Cera-
topogonidae. It is also known in Baeoda-
symyia; Borkent and Craig (1999) misiden-
tified sensilla v as iii and vi as v.

—The male antenna of Schizonyxhelea
have flagellomeres 5—10 fused. Flagello-
meres 5—9 are fused in S. bulla and flagello-
meres 5—10 in S. thomsenae Wirth. The
males of most species of Stilobezzia have
separate flagellomeres.

—A few genera of Ceratopogonini (includ-
ing Stilobezzia) and most genera of Heter-
omyiini + Sphaeromiini + Palpomyiini +
Stenoxenini have a secondary row of pali-
sade setae (defined in detail by Borkent, in
press). This is also present in Schizonyxhe-
lea.

—Adult eyes of Schizonyxhelea have mi-
nute interfacetal pubescence. Very small in-
terfacetal spicules are also present in S. bul-
la (otherwise rare in Stilobezzia).

—Wing with radial cells reduced. Although
Wirth and Grogan (1988) suggested that
Schizonyxhelea lack radial cells, the male I
have had a narrow second radial cell. A
number of Stilobezzia species have a small
or completely reduced first radial cell.
However, loss of radial cells is known in a
number of other genera of Ceratopogonidae
and the character state cannot be interpreted
at present. It was likely the reduced radial
cells which led Wirth and Grogan (1988) to

VOLUME 102, NUMBER 4

suggest that Schizonyxhelea was related to
other genera with reduced wing venation
but, as noted in the introduction, these taxa
with reduced character states are not closely
related to one another. It is apparent that
several lineages of Ceratopogonidae have
produced small species which have inde-
pendently reduced their wing venation and,
for some, their mouthparts, antennae and
parts of the male genitalia.

—Wing with a narrow apical band of ma-
crotrichia. This character state is shared
with S. bulla and S. thomsenae and appears
to be unique with the subgenus Stilobezzia.
Members of the subgenera Debenhamia
Wirth and Grogan and Acanthohelea also
have macrotrichia on the wing membrane
but this is nearly always more generally dis-
tributed on the apex of the wing. Stilobezzia
(Acanthohelea) insolita Das Gupta and
Wirth from Malaysia also has a narrow
band of macrotrichia and the female has a
distinctively (but not unique within the Cer-
atopogonidae) curved single spermathecae
very similar to that present in female Schi-
zonyxhelea.

—Aedeagus very reduced in size. This
character state is shared with S. bulla, S.
thomsenae and a few other Stilobezzia. The
condition is otherwise rare within the Cer-
atopogonidae.

—Male cerci are closely approximated.
This is present in S. thomsenae, some other
Stilobezzia and some other genera of Cer-
atopogonini.

The striking similarity of the pupae of
Schizonyxhelea forattinii and Stilobezzia
bulla and the presence of fusion of male
flagellomeres 5—9 or 10, the narrow band
of apical of macrotrichia on the wing and
the reduced aedeagus shared with S. bulla
and S. thomsenae probably indicates that
the two species of Schizonyxhelea are ac-
tually small Stilobezzia with a somewhat re-
duced wing venation and with equal claws
on each leg of the female. A reversion from
a single claw to double claws is known in
other genera such as Serromyia Meigen
(Borkent and Bissett, 1990) and Alluaudo-

867

myia Kieffer (personal observation of new
species) and it would not be surprising to
find the same within Stilobezzia; indeed, I
have a small Stilobezzia from Costa Rica
with equal claws but with typical Stilobez-
zia-like wing venation (i.e., a small first and
a long second radial cell is present).

It is yet possible that the pupal character
states shared by Schizonyxhelea forattinti
and Stilobezzia bulla are plesiomorphic
within Stilobezzia and that Schizonyxhelea
is therefore the sister group of Stilobezzia.
Further study is required of character state
polarities to resolve this question (Borkent,
in preparation). For the present it is best to
continue to recognize Schizonyxhelea as a
valid genus.

ACKNOWLEDGMENTS

My wife Annette and brother and sister-
in-law, Herman and Pieta Borkent, financed
part of the collecting trip to Costa Rica and
this is deeply appreciated. Annette also pro-
vided the resources to work up the results
of that expedition and I thank her for her
incredible support.

El Servicio de Parques Nacionales kindly
provided permission to collect in the nu-
merous National Parks and Reserves in
Costa Rica. I express my appreciation to
Alvaro Castro, who helped organize our
lives in Costa Rica and directed me to pris-
tine habitats on the west coast of Costa
Rica. Finally, Alvaro Vargas permitted col-
lecting on his farm, just west of Reserva
Carara, which included an exceptional vir-
gin rainforest and some beautiful springs
from which the larva of Schizonyxhelea for-
attinii was collected.

I acknowledge the support of the Global
Environment Facility of the World Bank
through the “Biodiversity Resources De-
velopment Project’? developed by the Na-
tional Institute of Biodiversity (INBio), the
National System of Conservation Areas
(SINAC), Ministry of the Environment and
Energy (MINAE) in Costa Rica.

Steve Murphree provided helpful criti-
cisms as a referee of this paper which was

868

much appreciated. My thanks to Nancy E.
Adams and David G. Furth for the loan of
the holotype of S. forattinii from the Smith-
sonian Institution in Washington, D.C.

LITERATURE CITED

Borkent, A. 1992. A new key to some genera of Cer-
atopogonini in the Holarctic (Diptera: Ceratopo-
gonidae). Entomologica Scandinavica 22: 433-
436.

. 1995. Biting Midges in the Cretaceous Amber

of North America (Diptera: Ceratopogonidae).

Backhuys Publishers, Leiden, The Netherlands,

237 pp.

. In press. Biting Midges (Ceratopogonidae:
Diptera) from Lower Cretaceous Lebanese Amber
with a Discussion of the Diversity and Patterns
Found in Other Ambers. Backhuys Publishers,
Leiden, The Netherlands.

Borkent, A. and B. Bissett. 1990. A revision of the
Holarctic species of Serromyia Meigen (Diptera:

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Ceratopogonidae). Systematic Entomology 15:
153-217.

Borkent, A. and D. A. Craig (1999). A revision of the
Neotropical genus Baeodasymyia Clastrier and
Raccurt (Diptera: Ceratopogonidae) with a discus-
sion of their phylogenetic relationships. American
Museum Novitates 3274: 1—26.

Clastrier, J. 1984. Schizonyxhelea guyana n.g., n.sp. de
la Guyane Francaise (Diptera, Ceratopogonidae).
Revue Francaise d’Entomologie 6: 1—4.

Grogan, W. L. and A. Borkent. 1992. Sinhalohelea, a
new genus of predaceous midge from Sri Lanka
(Diptera: Ceratopogonidae). Proceedings of the
Entomological Society of Washington 94: 314—
Bi9%

Lawson, J. W. H. 1951. The anatomy and morphology
of the early stages of Culicoides nubeculosus Mei-
gen (Diptera: Ceratopogonidae = Heleidae).
Transaction of the Royal Entomological Society
of London 102: 511—570, pl. 1.

Wirth, W. W. and W. L. Grogan. 1988. The predaceous
midges of the world (Diptera: Ceratopogonidae;
Tribe Ceratopogonini). Flora and Fauna Hand-
book 4, xv + 160 pp., E.J. Brill.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 869-877

DESCRIPTION OF THE LAST LARVAL INSTAR AND PUPA OF
LUCIDOTA ATRA (G. A. OLIVIER 1790) (COLEOPTERA: LAMPYRIDAE),
WITH A DISCUSSION OF ABDOMINAL SEGMENT HOMOLOGY
ACROSS LIFE STAGES

Marc A. BRANHAM AND MIGUEL ARCHANGELSKY

(MAB) The Ohio State University, Museum of Biological Diversity, 1315 Kinnear Rd.,
Columbus, OH 43212-1192, U.S.A. (e-mail: branham.24 @osu.edu); (MA) Centro Regional
de Investigaciones Cientificas y Transferencia Tecnologica, CRILAR (CONICET—UNLaR)
Entre Rios y Mendoza s/n, 5301, Anillaco, La Rioja, Argentina (e-mail: marchangelsky @
crilar.com.ar)

Abstract.—The last larval and pupal stages of the widespread and common North Amer-
ican firefly, Lucidota atra (G. A. Olivier 1790), are described and illustrated. Last instar
larvae were collected in rotting logs in the early spring and fed terrestrial snails from the
same logs until the larvae pupated. The larva of L. atra was misidentifed in the literature
and has subsequently been misidentified as a species in the genus Photinus. A discussion
of the homology of abdominal sclerites in larval, pupal, and adult fireflies is provided.

Key Words: Lampyridae, Lucidota atra, Photinus, larva, pupa, morphology, firefly,

lightningbug, glowworm, ventrite

The genus Lucidota, as defined by La-
porte (1833) and fixed by Motschulsky
(1853), is restricted to the New World and
contains some 64 described species. The
genus ranges from the United States to Ar-
gentina. Lucidota atra (G. A. Olivier) oc-
curs from the northeastern United States to
Central America (McDermott 1966). The
larva of this species was first described by
H. E Wickham (1895), but because larvae
in the tribe Photinini are difficult to distin-
guish (LaBella and Lloyd 1991), a more de-
tailed larval description is required. Peter-
son (1951) apparently incorrectly identified
the larva on which he based his drawing of
*“Photinus sp.” in his book “‘Larvae of In-
sects.’ Upon examining “‘Photinus sp.” in
Peterson’s larval collection at The Ohio
State University and comparing it with lar-
vae reared by one of us (MAB), it was dis-
covered that Peterson’s larva is actually L.
atra.

For this study, last instar larvae were col-
lected in early spring and kept until eclo-
sion, thus allowing a positive identification
from the adult. No larval descriptions exist
for other species of this genus, most likely
due to difficulties in rearing firefly larvae
(Archangelsky and Branham 1998).

MATERIALS AND METHODS

Seven last instar larvae were collected in
a rotting log on April 6, 1993 outside of
Lawrence, KS, and kept in a glass jar with
damp wood from the log along with some
terrestrial snails collected in the same
wood. Empty snail shells were removed
from the jar every few days. The wood in-
side the jar was inspected for moisture con-
tent periodically. When the wood appeared
to be drying out, it was moistened with dis-
tilled water. To further simulate the inside
of the log, the jar was wrapped with paper

870 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

to reduce light entering the jar. No special
requirements were necessary for pupation.

Three larvae and one pupa were fixed in
boiling water and transferred to 7O%EtOH.
In order to study the larval morphology, a
specimen’s head, mouthparts and antennae
were dissected, cleared in lactic acid, and
mounted on microscope slides using Hoy-
er’s as the mounting medium. The descrip-
tions and drawings were done using a Wild
MS dissecting microscope and a Zeiss Ax-
ioscope 20 compound microscope, both
with a camera lucida.

RESULTS
Lucidota atra (G. A. Olivier)

Description of last larval instar.—
Length: 13.0 to 15.0 mm. Body elongate,
fusiform, slightly flattened dorsoventrally
(Fig. 1). Whitish ventrally with pink along
sides of thorax and abdomen. Sclerotized
regions uniformly light to dark brown and
granulose. All tergites, except abdominal
tergites 8 and 9, bearing 3 light colored
stripes that are more or less parallel to the
longitudinal axis of body.

Head capsule: Prognathous, subquadra-
te, dorsoventraly flattened, and robust (Fig.
2); retractable within thorax. Labrum and
clypeus fused. Epicranial suture present as
well as frontal sutures that extend to bases
of antennae. One pair of lateral stemmata,
posterior to base of antennae. Head capsule
not fused ventrally (Fig. 3).

Antenna: 3-segmented, partially retract-
able within membranous base (Fig. 4); orig-
inating on latero-apical edges of head cap-
sule. Basal segment widest, attached to
membranous base, median portion of dorsal
surface covered with medium length setae
pointing anteriorly, lateral pointing setae on
anterior third of segment approximately 2
to 3 times a long as setae in medial region. Fig. 1. Lucidota atra, fifth instar larva, habitus.
Second segment shorter than third, narrow- Seale han Senn
er, evenly covered by long setae, carrying
a large globular sensorium slightly longer
than third antennomere. Third segment very
Short, stout with several short setae, an api-

VOLUME 102, NUMBER 4 871

Figs. 2-5. Lucidota atra, head of fifth instar larva. 2, Dorsal view. 3, Ventral view. Scale bar = 1 mm. 4,
Right antenna, dorsal view. 5, Right mandible, dorsal view. Scale bars = 0.2 mm.

\

SS

Bil 3 SN
Zip WN | 4
ie, ae 14

Figs. 6-7.
dorsal view. Scale bar = 0.23 mm.

cal spine, and a small globular sensorium
on inner surface just below the antennal
apex.

Mandible: Symmetrical, strongly falcate,
with an inner channel opening subapically
on outer edge (Fig. 5). Retinaculum present,
forming 2 inner teeth on the apical third of
mandible. Basal third of the retinaculum
covered with a dense brush of setae. Medial
region of mandible covered by a single row
of long setae pointing inward toward the
retinaculum, perpendicular to the inner
channel of the mandible; 1 long seta par-
allel to apical point of the mandible, just
anterior to row of setae located medially.
One 4-pronged seta or sensory appendage
on outer margin of mandible, just before
channel opening; outermost prong of this
seta longer than the other 3.

Labium: Closely attached to maxilla,
formed by a short and strongly sclerotized
prementum, mentum (distally membranous)
and submentum (fused to mentum). Pre-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Lucidota atra, fifth instar larva. 6, Labium, dorsal view. Scale bar = 0.15 mm. 7, Left maxilla,

mentum heart shaped, in both dorsal and
ventral views with distal apical cleft (Fig.
6); in dorsal view, bearing 2 basal regions
of very fine setae with longer setae present
on the segments of the palp; 2 brushes of
fine cuticular spines present on each side of
prementum. Palpus 2-segmented; basal seg-
ment short, bearing several spines, second
segment twice as long as first, pointed and
somewhat forked with a single spine (Fig.
6).

Maxilla: Apical region (Fig. 7). Basal re-
gion (Fig. 3). Long and robust, closely at-
tached to labium. Cardo (Fig. 3) irregularly
shaped, bearing no setae. Stipes (Fig. 7)
very broad, ventral surface covered with se-
tae and bearing a single long seta; dorsal
surface bearing 2 long setae. Galea large,
2-segmented, basal segment very long, 3
times as long as second segment and lack-
ing setae; distal segment short, conical and
bearing several short setae with 1 seta on
distal apex of segment. Lacinia large, twice

VOLUME 102, NUMBER 4

as long as first segment of the galea, inner
surface covered with a thick brush of cutic-
ular spines. Palpus 3-segmented, basal seg-
ment largest, subquadrate, longer than other
2 segments combined, distal two-thirds
covered with medium to long setae; second
segment wider than long and bearing me-
dium length setae; distal segment subconi-
cal without setae, bearing a globular sen-
sorium-type structure.

Thorax: Prothorax subcircular, wider at
base, containing retracted head when larva
is in repose. Meso- and metathorax sub-
rectangular. Thoracic tergites subdivided by
sagittal line. Each segment with pleural area
formed by an upper laterotergite, below it
an epimeron and episternum separated by
pleural suture; mesothoracic laterotergite
subdivided, anterior plate smaller, carrying
mesothoracic spiracle. Prosternum medium
sized; meso- and metasterna smaller, nar-
row, subdivided into an anterior basister-
num and a posterior sternellum. | pair of
biforous spiracles present on mesopleuron.

Legs: 5-segmented, coxae long and cy-
lindrical, robust; trochanters small, sub-
triangular in lateral view; femora long and
cylindrical, widening slightly apically, with
a single long seta in medial inner portion;
tibiotarsi as long as femora, tapering to-
wards distal end; pretarsi strong, simple,
with a pair of stout setae at base. Double
row of strong setae on inner margin of ti-
biotarsi, lacking on inner margin of femora.

Abdomen: 10-segmented, segments 1 to
8 similar in shape, tapering toward end;
each tergite subrectangular, tergites 1
through 8 divided by a sagittal line and 2
lighter colored lines parallel to sagittal line;
lateral portions of tergite 8 lightly colored;
lateral portions of tergite 9 lightly colored
and without sagittal line; segment 10 a nar-
row ring surrounding anal region, carrying
holdfast organ. Pleural areas well devel-
oped, segments 1 to 7 subdivided, upper
plate large, suboval, carrying spiracles,
lower plate small, narrowly subtriangular;
pleuron 8 with only 1 suboval plate carry-
ing a spiracle; pleural areas of segments 9

873

and 10 reduced. Abdominal sterna large,
subquadrate, narrowing towards end of ab-
domen. Postero-lateral corners of sternite 8
bearing a twin spotted photic organ. Color
pattern similar to that of thorax. Biforous
spiracles present on pleurites 1 to 8.

Description of pupa.—Female, one day
old. Slightly curved, ventrally concave;
young pupa white, older pupa approaching
charcoal in color. Length: 10.0 to 11.0 mm.

Head: Completely covered by pronotum
in dorsal view (Fig. 9), white. Eyes small,
on sides of head; antennae inserted in front
of eyes, serrate with 11 obvious segments,
extending in length to metacoxae; antenna
and mouthparts white.

Thorax: Pronotum large, subtriangular,
slight emargination on either side of ante-
rior apex, covering head; white or cream.
Meso- and metanotum shorter than prono-
tum, subrectangular, carrying wing pads on
sides; posterior medial portion of mesono-
tum coming to a point, point lacking on me-
tanotum. First and second pair of legs fully
visible in ventral view; third pair of legs
almost completely covered by wingpads,
only metatarsus visible.

Abdomen: Segments wider than long,
white. Tergite 1 with postero-lateral corners
pointing perpendicular to sagittal axis of
pupa; postero-lateral corners of tergites 2
through 8 coming to a point and directed
posteriorly. Pleurites fused to the sternites
(except for pleurite 1 which bears spiracle
1) thus forming lateral margins of abdom-
inal “‘ventrites’’ (see Discussion.) First ster-
nite lacking, first ventrite (sternite 2) par-
tially visible, remaining ventrites fully vis-
ible, 7 total in female pupa, (male pupa
with 8 ventrites total.) Medial-lateral cor-
ners of ventrite 7 (sternite 8) bearing a twin
spotted photic organ.

Spiracles: 9 pairs; First on pleuron of
mesothorax, remaining 8 on abdominal seg-
ments | to 8.

DISCUSSION

Biology

The activity period of L. atra adults
range from early June to July, and the larval

874

—

8

Figs. 8-9. Lucidota atra, pupa. 8, Ventral view. 9,

life is suspected to be approximately two
years long, since both large and small lar-
vae have been found together during mid-
summer (Balduf 1935). However, as this
species has not been successfully reared
from egg to adult, the two year life cycle
remains speculative. Because L. atra larvae
are typically found in rotten logs and
stumps from the fall through early spring,
it is assumed that these are the larvae that
overwinter (Williams 1917; MacDermott

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Dorsal view. Scale bar = 4 mm.

1964; MAB, personal observation). It
should be noted, however, that stumps and
logs are places where many coleopterists
typically look for beetle larvae. Both larvae
and pupae produced a glow from a two-
spotted photic organ when disturbed. The
two-spotted photic organ is located on the
eighth sternite of the larvae and seventh
ventrite (eighth sternite) of the female pupa.
Therefore, it is expected that male pupa
would also have a similar organ on its sev-

VOLUME 102, NUMBER 4

enth ventrite as the seventh ventrite of adult
males of this species bare such an organ,
though the ability to luminesce seems to di-
minish shortly after eclosion (MAB, per-
sonal observation). Since no L. atra larvae
have been found foraging in the open, they
may be subterranean in habit. Adults usu-
ally fly during the day, and males follow
pheromone plumes to the females (Lloyd
1972). The female is typically up to a third
larger than the male.

Traditional Perspective, and Modern View

Peterson (1951) included a side view
drawing of a lampyrid larva along with a
mandible and antennae which was labeled
“Photinus sp.”’ This same drawing was in-
cluded by LaBella and Lloyd (1991) and
has thereafter been used as an example of
a Photinus larva. Upon comparing our L.
atra larvae with the actual specimen upon
which Peterson based his drawing (in the
Peterson Larval Collection, The Ohio State
University, Columbus, Ohio), they are iden-
tical matches. Additionally, Peterson’s de-
termination label in the vial with this spec-
imen reads “‘Photinus sp.? .”’ This question
mark on the determination label, evidently
put there by Peterson himself, was most
likely accidentally overlooked, and, thus for
some 49 years, the specimen has been mis-
identified as Photinus sp., rather than Lu-
cidota atra. This mistake is very easy to
make due to the great morphological simi-
larity between genera in the tribe Photinini,
in which both Photinus and Lucidota are
assigned. The only definitive method to as-
sociate larvae and adults is to rear the lar-
vae.

Abdominal Sclerites in Lampyridae

Considerable confusion has occurred
concerning the number of abdominal scler-
ites in discussions of adult firefly abdomen
morphology. We believe that this confusion
has been caused largely by the fact that
lampyrids posses a varying number of vis-
ible ventral abdominal sclerites and there is
a lack of accuracy in defining the terms

875

used in descriptions and discussions of the
abdomen. Without both an understanding of
the homology among abdominal sclerites
and the use of accurate terminology, mor-
phological investigations of lampyrids are
bound to remain confused.

The description of both larval and pupal
states of L. atra is instructive for following
the reduction and fusion of various abdom-
inal segments and sclerites from the larval
stage, where all abdominal sclerites are pre-
sent and obvious, to the pupal stage, where
the effect of internalization, reduction and
fusion can be first detected. In all firefly
larvae currently known, there are ten ab-
dominal segments, with the tenth being
quite small and, therefore, commonly over-
looked. Each abdominal segment bears a
tergite, distinct pleurites that bear the spi-
racles (segments one through eight) and a
sternite. Identification of abdominal seg-
ments and sclerites in the larvae is not dif-
ficult. Reduction of abdominal segments
and the internalization of sclerites in the
adults however, can make it difficult to de-
termine homology among abdominal seg-
ments.

In Coleoptera, the adult abdomen is usu-
ally composed of ten segments in the male
(with the tenth often being highly reduced
or fused with the ninth), and nine in the
female (with the ninth being modified to
form the genital segment) (Lawrence and
Britton 1991). As was pointed out by Green
(1956), the first abdominal segment in adult
Lampyridae is indicated only by the first
abdominal tergite, except females of Pho-
tinus granulatus (Green, p. 597). However,
investigation concluded that the pleurite
bearing the first abdominal spiracle is also
present, though in reduced form, in pupal
L. atra and the adults of some lampyrids.
In the adult, the first visible ventral sclerite
actually is of the second abdominal seg-
ment, as the ventral portion of the first ab-
dominal segment is usually so internalized
and reduced, it is not visible ventrally. This
condition is termed a “hologastrous type
abdomen” (Nichols 1989). Additionally, in

876

adult lampyrids the abdominal pleurites are
fused to the sternites, thus forming a con-
tinuous ventral plate. The median half of
this ventral plate was the larval sternite and
the lateral regions on each side were the
pleurites. Green (1956), therefore suggested
**... it would be incorrect to refer to the
ventral segments of the abdomen as ster-
nites.”’ Lawrence and Britton (1991) use the
term “‘ventrites’’ to denote sternites that are
externally visible.

In light of these two situations: sternite
one lacking and the fusion of sternites with
pleurites, while also keeping with Green
(1956) and Lawrence and Britton (1991),
we adopt the term “‘ventrites’’ to denote the
visible ventral sclerites in the L. atra pupa,
which like other firefly species, has the
same abdominal morphology as the adult.
In most firefly species, the female has one
ventrite fewer than the male, with species
in the Luciolinae being the exception, the
male having six and the female having sev-
en ventrites. McDermott (1964) stated that
*‘The Lampyridae may be defined as that
family of the Cantharoidea having usually
seven visible ventral abdominal segments in
the male.’’ Apparently, McDermott did not
count the ninth abdominal segment when
visible, as a visible ventral segment. This
may be due to the small size if the ninth
ventral sclerite in relation to the other ven-
tral sclerites and the fact that it is usually
the terminal ventral sclerite. It is our present
conclusion that the sclerite of the ninth ab-
dominal segment (ventrite eight) needs to
be counted as a “‘ventrite’’? when visible.
Depending upon whether the eighth ventrite
is concealed under ventrite seven or ex-
posed, adult males will have either seven or
eight ventrites (MAB, personal observa-
tion), with eight ventrites being found in the
majority of genera family-wide, examined
by MAB. Therefore, in males of most fire-
fly species, ventrites one through eight cor-
respond to abdominal segments two
through nine. The only known exception is
for members of the subfamily Luciolinae,
which bear only six ventrites (McDermott

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1964; Ballentyne 1987a, b) and the paedo-
morphic brachypterous male of the Euro-
pean species Phosphaenus hemiperus La-
porte (MAB, personal observation). In the
Luciolinae, the last segment exposed in the
male is ventrite six (abdominal segment
seven), with segment eight apparently re-
duced, or altogether lost, with segment nine
forming part of the aedeagal sheath which
is retracted into the abdomen (Ballentyne
1992). The male of Phosphaenus hemiperus
more or less retains a larviform type ab-
domen. Therefore, it is no surprise that Tor-
re-Bueno’s (Nichols 1989) definition of
‘“‘sternite = ventrite” is insufficient in con-
veying the homology of ventral abdominal
segments in adults of Lampyridae.

Even though ‘“‘segmental fusion’’ does
not seem to occur in the firefly abdomen,
the use of the term ‘“‘ventrite’’ should be
used with care to avoid confusion of the
homology of various abdominal segments.
However, the use of the term “‘ventrite”’ to
denote only visible abdominal segments in
the adult, while also keeping in mind (and
mentioning a point of reference) that ““ven-
trite one”’’ is actually the ventral sclerite of
the second abdominal segment (in almost
all cases), is simply good nomenclature and
serves to avoid confusion concerning which
adult abdominal segment is being referred
to.

CONCLUSION

The firefly larva labeled as ‘‘Photinus
sp.’ by Peterson (1951) is actually the larva
of Lucidota atra, which is herein rede-
scribed in greater detail than the original
description (Wickham 1895) in order to fa-
cilitate larval identification. Through rear-
ing this species from larva to adult, it was
possible to investigate the homology of ab-
dominal segments and track possible fusion
or reduction events that lead to a decrease
in number of visible ventral abdominal
sclerites in the adult. Fusion of both abdom-
inal segments and ‘“‘ventrites’’ are not
known to occur in currently studied lam-
pyrid taxa.

VOLUME 102, NUMBER 4

ACKNOWLEDGMENTS

We thank Lesley A. Ballentyne for dis-
cussions concerning Pteroptyx abdominal
segmentation, and James E. Lloyd, Andrey
Sharkov, David R. Smith, and John W.
Wenzel, for their helpful comments on the
manuscript.

LITERATURE CITED

Archangelsky, M. and M. A. Branham. 1998. Descrip-
tion of the Preimaginal Stages of Pyractomena bo-
realis (Randall, 1838) (Coleoptera: Lampyridae)
and Notes on its Biology. Proceedings of the En-
tomological Society of Washington 100(3): 421—
430.

Balduf, W. V. 1935. The Bionomics of Entomophagous
Coleoptera. John S. Swift, St. Louis, 220 pp.
Ballentyne, L. A. 1987a. Further Revisional Studies on
the Firefly Genus Pteroptyx Olivier (Coleoptera:
Lampyridae:Luciolinae). Transactions of the

American Entomological Society 113: 117-170.

. 1987b. Lucioline Morphology, Taxonomy and

Behaviour: A Reappraisal (Coleoptera: Lampyri-

dae). Transactions of the American Entomological

Society 113: 171-188.

. 1992. Revisional Studies on Flashing Fireflies
(Coleoptera: Lampyridae), Ph.D. Dissertation,
University of Queensland, St. Lucia, Brisbane,
Australia.

Green, J. W. 1956. Revision of the Nearctic Species of
Photinus (Lampyridae: Coleoptera). Proceedings
of the California Academy of Science XXVIII
(15): 561-613.

877

LaBella, D. M. and J. E. Lloyd. 1991. Lampyridae, pp.
427—428. In Stehr, E W., ed., Immature Insects,
Vol. 2, Kendall Hunt Publishing Company, Du-
buque, Iowa, 974 pp.

Laporte, EL.N. (Comte de Castelnau). 1833. D’une
Révision du genre Lampyre. Annales de la Société
Entomologique de France II: 122-153.

Lawrence, J. F and E. B. Britton. 1991. Coleoptera,
pp. 543-695. In C.S.I.R.O., ed., The Insects of
Australia, Second Edition, Vol. 2, Cornell Uni-
versity Press, Ithaca, New York, 1,137 pp.

Lloyd, J. E. 1972. Chemical communication in fire-
flies. Environmental Entomology 1(2): 265-266.

McDermott, FE A. 1964. The Taxonomy of the Lam-
pyridae (Coleoptera). Transactions of the Ameri-
can Entomological Society 90: 1—72.

. 1966. Lampyridae. Jn Steel, W. O., ed., Co-
leopterorum Catalogus Supplementa. Pars 9 (edi-
tio secunda). W. Junk, ‘s-Gravenhage, 149 pp.

Motschulsky, V. 1853. Etudes Entomologiques I: 24—
58.

Nichols, S. W. 1989. The Torre-Bueno Glossary of En-
tomology. Revised Edition of ‘“‘A Glossary of En-
tomology” by J.R. de la Torre-Bueno. New York
Entomological Society, American Museum of
Natural History, New York, New York, 840 pp.

Peterson, A. 1951. Larvae of Insects. Part II, Edwards
Brothers, Inc., 416 pp.

Wickham, H. F. 1895. On the Larvae of Lucidota, Si-
noxylon and Spermophagus. Bulletin from the
Laboratories of Natural History of the State Uni-
versity of Iowa III, pp. 28-35.

Williams, E X. 1917. Notes on the Life-History of
some North American Lampyridae. Journal of the
New York Entomological Society 25: 11—12.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 878-891

LIFE HISTORY AND DESCRIPTION OF IMMATURE STAGES OF
NEASPILOTA PUBESCENS FREIDBERG AND MATHIS (DIPTERA:
TEPHRITIDAE) ON LESSINGIA FILAGINIFOLIA (HOOKER AND ARNOTT)
M. A. LANE (ASTERACEAE) IN SOUTHERN CALIFORNIA

RICHARD D. GOEDEN

Department of Entomology, University of California, Riverside, CA 92521, U.S.A.
(e-mail: rgoeden @ucrac1l.ucr.edu)

Abstract.—Neaspilota pubescens Freidberg and Mathis is a univoltine, monophagous
fruit fly (Diptera: Tephritidae) developing solely in the flower heads of Lessingia filagin-
ifolia (Hooker and Arnott) M. A. Lane (Asteraceae) belonging to the subtribe Solidagi-
ninae of the tribe Astereae in southern California. The egg, second- and third-instar larvae,
and puparium are described and figured. The anterior thoracic spiracle of the second instar
has five papillae, reduced to two papillae in the third instar. The second instar has seven
oral ridges and the third instar eight oral ridges, which, except for the most ventral, eight
oral ridge in the latter instar, are ventrally toothed. The arrangement of these oral ridges
in a vertical series lateral to the oral cavity is a distinguishing generic character. The
larvae feed mainly on the ovules and soft achenes as first and second instars; however,
as third instars, they may extend their feeding into the receptacle and supplement their
diet with sap. The nonfeeding prepuparium overwinters in a protective cell that occupies
much of the excavated flower head and is formed of ovule-, achene-, chaff-, pappus-, and
corolla-fragments impregnated with excess sap and liquid feces that harden when dry. A
few prepuparia pupate and emerge from their cells in the late summer and probably
Overwinter as adults, but most pupariate during the next year in late spring, and emerge
as adults that aggregate on preblossom host plants to mate and subsequently oviposit.
Pteromalus sp. (Hymenoptera: Pteromalidae) was reared as a solitary, larval-pupal en-
doparasitoid from a puparium of N. pubescens.

Key Words: Insecta, Neaspilota, Lessingia, Asteraceae, nonfrugivorous Tephritidae, bi-
ology, taxonomy of immature stages, flower-head feeding, monophagy, seed

predation, parasitoid

Revision of the genus Neaspilota (Dip-
tera: Tephritidae) by Freidberg and Mathis
(1986) facilitated determination of speci-
mens reared from California Asteraceae
(Goeden 1989) and stimulated several life-
history studies, including those on N. viri-
descens Quisenberry (Goeden and Headrick
1992), N. wilsoni Blanc and Foote (Goeden
and Headrick 1999), N. signifera (Coquil-
lett) (Goeden 2000a), N. aenigma Freidberg
and Mathis (Goeden 2000b), and N. appen-
diculata Freidberg and Mathis (Goeden
2000c). This paper describes some imma-

ture stages and the life history of a sixth
species from California, N. pubescens
Freidberg and Mathis.

MATERIALS AND METHODS

The present study was based in large part
on dissections of flower heads of Lessingia
filaginifolia (Hooker and Arnott) M. A.
Lane (Asteraceae) collected during 1990—
1997 mainly from the following two loca-
tions in the South and North Sections, re-
spectively, of the San Bernardino National

VOLUME 102, NUMBER 4

Forest: Bautista Canyon at 1,100 m eleva-
tion, Riverside Co. and North of South Fork
Campground at 1,550 m, SW San Bernar-
dino Co. One-liter samples of excised, im-
mature and mature flower heads containing
eggs, larvae, and puparia were transported
in cold-chests in an air-conditioned vehicle
to the laboratory and stored under refrig-
eration for subsequent dissection, photog-
raphy, description, and measurement. Eight
second- and 12 third-instar larvae and five
puparia dissected from flower heads were
preserved in 70% EtOH for scanning elec-
tron microscopy (SEM). Additional prepu-
paria and puparia were placed in separate,
glass shell vials stoppered with absorbant
cotton and held in humidity chambers at
room temperature for adult and parasitoid
emergence. Specimens for SEM were hy-
drated to distilled water in a decreasing se-
ries of acidulated EtOH. They were osmi-
cated for 24 h, dehydrated through an in-
creasing series of acidulated EtOH and two,
1-h immersions in hexamethyldisilazane
(HMDS), mounted on stubs, sputter-coated
with a gold-palladium alloy, and studied
and photographed with a Philips XL-30
scanning electron microscope in the Insti-
tute of Geophysics and Planetary Physics,
University of California, Riverside.

Most adults reared from isolated prepu-
paria and puparia were individually caged
in 850-ml, clear-plastic, screened-top cages
with a cotton wick and basal water reser-
voir and provisioned with a strip of paper
toweling impregnated with yeast hydroly-
zate and sucrose. These cages were used for
studies of longevity and sexual maturation
in the insectary of the Department of En-
tomology, University of California, River-
side, at 25 + 1°C, and 14/10 (L/D) photo-
period. Two pairs of virgin males and fe-
males obtained from emergence cages also
were held in each of six, separate, clear-
plastic, petri dishes provisioned with a flat-
tened, water-moistened pad of absorbant
cotton spotted with honey (Headrick and
Goeden 1994) for observations of their
courtship and copulation behavior.

879

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

RESULTS AND DISCUSSION
Taxonomy

Adult.—Neaspilota pubescens was de-
scribed by Freidberg and Mathis (1986: 55—
57), who pictured the unpatterned wing,
along with drawings (p. 56) of the lateral
aspect of the head; male right foretarsus,
epandrium, distiphallus, epandrium and cer-
ci, aculeus and its apex enlarged, and sper-
matheca.

Immature stages.—The first-instar larva
remains undescribed, but the egg, second-
and third-instar larvae and puparium are de-
scribed below, as the only stages available
at this writing.

Egg: Only three intact eggs were found
and measured in situ within separate, im-
mature, preblossom flower heads. These
eggs were white, opaque, smooth, elongate-
ellipsoidal, and averaged 0.71 + 0.03
(range, 0.64—0.74) mm long and 0.18 +
0.003 (range, 0.17—0.18) mm wide, tapered
and smoothly rounded at both ends. As no
eggs were examined by scanning electron
microscopy, the egg of N. pubescens could
only be generally compared with the eggs
of N. viridescens, N. wilsoni, and N. appen-
diculata, which were described in detail by
Goeden and Headrick (1992, 1999) and
Goeden (2000c).

Second instar: White, elongate-cylindri-
cal, rounded anteriorly, truncated dorsopos-
teriorly (Fig. 1A), body segments well-de-

880 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Det WD Exp 200 jum A... Det WD

142 64 SE 144 4

Acc.V Spot MaG

00k 0 2 g a2 ” -

Fig. 1. Second instar of Neaspilota pubescens: (A) habitus, anterior to left; (B) gnathocephalon, ventrolateral
view, |1-minute acanthae, 2-anterior sensory lobe, 3-stomal sense organ, 4-mouthhook, 5-median oral lobe, 6-
integumental petal, 7-oral ridge; (C) anterior sensory lobe, 1-dorsal sensory organ, 2-terminal sensory organ, 3-
lateral sensory organ, 4-supralateral sensory organ, 5-pit sensory organ, 6-stomal sense organ; (D) anterior
thoracic spiracle; (E) caudal segment, l-rima, 2-interspiracular process, 3-intermediate sensory complex; (F)
intermediate sensory complex, 1-stelex sensillum, 2-medusoid sensillum.

VOLUME 102, NUMBER 4

fined, circumscribed anteriorly with few
minute acanthae (Figs. 1B-1); dorsal sen-
sory organ well-defined, dome-shaped (Fig.
1C-1); anterior sensory lobe (Fig. 1B-2, C)
with terminal sensory organ (Fig. 1C-2),
lateral sensory organ (Fig. 1C-3), suprala-
teral sensory organ (Fig. 1C-4), and pit sen-
sory organ (Fig. 1C-5); stomal sense organ
(Figs. 1B-3, C-6) ventrolaterad of anterior
sensory lobe; mouthhook bidentate (Fig.
1B-4); median oral lobe (Fig. 1B-5), fla-
belliform, laterally compressed (not
shown); about six papilliform, integumental
petals dorsal to each mouthhook (Fig. 1B-
6); seven oral ridges toothed ventrally, in
vertical series lateral to oral cavity (Fig. 1B-
7); prothorax, at least, circumscribed ante-
riorly by posteriorly-directed, minute acan-
thae (Fig. 1B-1); anterior thoracic spiracle
with five, cuboidal papillae (Fig. 1D); lat-
eral spiracular complexes not seen; caudal
segment with two stelex sensilla (not
shown) dorsolaterad and ventrolaterad of
posterior spiracular plate (Fig. 1E); poste-
rior spiracular plate bears three ovoid rimae
(Fig. 1E-1), ca. 0.015 mm long, and four
interspiracular processes (Fig. 1E-2), each
with one to four, simple or forked branches,
longest measuring 0.01 mm; intermediate
sensory complex (Figs. 1E-3, F) with a ste-
lex sensillum (Fig. 1F-1) and a medusoid
sensillum (Fig. 1F-2).

The habitus of the second instar of N.
pubescens (Fig. 1A) is more like N. wilsoni
(Goeden and Headrick 1999), N. signifera
(Goeden 2000a), N. aenigma (Goeden
2000b), and N. appendiculata (Goeden
2000c) than the barrel-shaped second instar
of N. viridescens (Goeden and Headrick
1992). The dorsal sensory organ of N. pu-
bescens is well defined in the second instar
(Fig. 1C-1), as with N. signifera (Goeden
2000a) and N. appendiculata (Goeden
2000c), but is not well defined in N. viri-
descens (Goeden and Headrick 1992), N.
wilsoni (Goeden and Headrick 1999), and
N. aenigma (Goeden 2000b). The integu-
mental petals of the second instars of all six
species are papilliform and six in number

881

in N. pubescens (Fig. 1B-6), like N. viri-
descens (Goeden and Headrick 1992), but
number four in N. signifera (Goeden
2000a), seven in N. wilsoni (Goeden and
Headrick 1999) and N. appendiculata (Goe-
den 2000c) and eight in N. aenigma (Goe-
den 2000b). In the first instars of all six
congeners examined to date, though not
available in the present study, the integu-
mental petals are broad, flattened, and
paired (Goeden and Headrick 1992, 1999;
Goeden 1999a, b). An apparent difference
in N. pubescens is the five papillae on the
anterior spiracle of the second instar (Fig.
1D), compared to eight in N. appendiculata
(Goeden 2000c), and three to four papillae
in second instars of N. viridescens (Goeden
and Headrick 1992), N. wilsoni (Goeden
and Headrick 1999), N. signifera (Goeden
2000a), and N. aenigma (Goeden 2000b).
Finally, the interspiracular processes of N.
pubescens each bear one to four branches
like N. aenigma (Fig. 1E-2, Goeden 2000b),
not two to four branches like WN. signifera
(Goeden 2000a), nor five to nine branches
like those of N. viridescens (Goeden and
Headrick 1992), nor two to six branches
like those of N. wilsoni (Goeden and Head-
rick 1999), nor four branches like those of
N. appendiculata (Goeden 2000c). How-
ever, it is recognized that most specimens
of N. pubescens with branch numbers at the
high end of the range will not differ from
most other Neaspilota in this character.
Third instar: Pale yellow, ellipsoidal,
with posterior spiracular plate dark brown
to black, tapering anteriorly; posterior spi-
racular plate on caudal segment flattened
and upturned dorsally ca. 60° (Fig. 2A), mi-
nute acanthae circumscribe anterior fifth of
thoracic and anterior abdominal segments,
but more common posteriorly (Fig. 2B-1);
gnathocephalon conical (Fig. 2B); dorsal
sensory organ an elliptical, flat, poorly de-
fined pad (Fig. 2C-1) punctured centrally
and peripherally by pore sensilla (Fig. 2C-
2); anterior sensory lobe (Fig. 2C) bears ter-
minal sensory organ (Fig. 2C-3), lateral
sensory organ (Fig. 2C-4), supralateral sen-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

spot Magn §=Det WD Exp HJ 1mm A

fal } f 14

AccV SpotMagn Det WD Exp
p.00 kV 3.0_174x__ SE_13.4 39

eas ;
c= Mar

Acc.V Spot Magn Det WD. Expl
0:0 kV 3.0 1309x SE "13.4 47

Fig. 2. Third instar of Neaspilota pubescens: (A) habitus, anterior to left; (B) gnathocephalon, lateral view,
1-minute acanthae, 2-oral ridge, 3-stomal sense organ, 4-mouthhook; (C) anterior sensory lobe, 1-dorsal sensory
organ, 2-pore sensillum, 3-terminal sensory organ, 4-lateral sensory organ, 5-supralateral sensory organ, 6-pit
sensory organ, 7-integumental petal, 8-stomal sense organ; (D) anterior thoracic spiracle; (E) metathoracic
(left) and abdominal (right) lateral spiracular complexes, 1-spiracle, 2-verruciform sensilla on metathorax,

VOLUME 102, NUMBER 4

sory organ (Fig. 2C-5), and pit sensory or-
gan (Fig. 2C-6); eight, papilliform (above)
or spatulate (below), integumental petals in
two rows above each mouthhook (Fig. 2C-
7); eight oral ridges (Fig. 2B-2), all but
most ventral ridge, toothed ventrally and
lateral to oral cavity; stomal sense organ
(Figs. 2B-3, C-8) ventrolaterad of anterior
sensory lobe; mouthhook (Fig. 2B-4) tri-
dentate (not shown); median oral lobe lat-
erally flattened, apically pointed (not
shown); prothorax circumscribed by minute
acanthae (Fig. 2B-1); verruciform sensilla
circumscribe prothorax posteriorad of mi-
nute acanthae (not shown); anterior thoracic
spiracle on posterior margin of prothorax
bears two oblong papillae (Fig. 2D); meta-
thoracic lateral spiracular complex with a
spiracle (Fig. 2E-1) and three verruciform
sensillae, one above (not shown) and two
below the spiracle (Fig. 2E-2); abdominal
lateral spiracular complex with a spiracle
(Fig. 2E-3) and two verruciform sensilla
(Fig. 2E-4) dorsoposteriorad of, and two
verruciform sensilla ventroposteriorad of,
the spiracle, these sensilla arranged as two
vertical pairs; caudal segment circum-
scribed dorsally by minute acanthae (Fig.
2F-1); a stelex sensillum dorsolaterad (Fig.
2F-2), laterad (Fig. 2F-3), and ventrolaterad
(Fig. 2F-4) of posterior spiracular plate
(Fig. 2G); each posterior spiracular plate
(Fig. 2G) bears three ovoid rimae (Fig. 2G-
1), ca. 0.03 mm in length, and four inter-
spiracular processes (Fig. 2G-2), each with
one to three, simple, pointed or forked
branches, longest branch measuring 0.013
mm; intermediate sensory complex (Fig.
2H) with a medusoid sensillum (Fig. 2H-1)
and a stelex sensillum (Fig. 2H-2). Each
stelex sensillum surrounding the posterior
spiracular plate apparently has a single,

ce

883

hemispherical, minute acanthus at its base
(Fig. 2H-3).

The habitus of the third instar of N. pu-
bescens generally is like that reported for
N. viridescens (Goeden and Headrick
1992), N. wilsoni (Goeden and Headrick
1999), N. signifera (Goeden 2000a), and N.
aenigma (Goeden 2000b), and N. appendi-
culata (Goeden 2000c). Like N. signifera
(Goeden 2000a) and N. appendiculata
(Goeden 2000c), only the anterior part of
each body segment of N. pubescens is cir-
cumscribed by minute acanthae, whereas, in
N. aenigma the anteriors, pleura, and pos-
teriors of each segment are thus circum-
scribed (Goeden 2000b); in N. wilsoni, all
intersegmental areas and all abdominal seg-
ments except the pleura are circumscribed
(Goeden and Headrick 1999); and in N. vir-
idescens, the intersegmental areas are free
of acanthae (Goeden and Headrick 1992).
Like N. viridescens (Goeden and Headrick
1992) and N. wilsoni (Goeden and Headrick
1999), but not like N. signifera (Goeden
2000a), N. aenigma (Goeden 2000b) and N.
appendiculata (Goeden 2000c), the dorsal
sensory organ is not well defined, and flat-
tened, and not dome-shaped, in the third in-
star of N. pubescens (Fig. 2C-1). In the sec-
ond instar of N. wilsoni (Goeden and Head-
rick 1999), N. signifera (Goeden 2000a), N.
appendiculata (Goeden 2000c), and N. pu-
bescens (Fig. 1C-1) the dorsal sensory or-
gan is both prominent and dome-shaped, as
it is in the first instar of all congeners ex-
cept N. signifera, where it was hidden in
my specimens and could not be examined
for comparison (Goeden 2000a), and in the
present study where first instars were not
available.

Additional similarities involved the inte-
gumental petals in the third instars of all

3-spiracle, 4-verruciform sensilla on first abdominal segment; (F) anal segment, 1-minute acanthae, 2-dorsolateral
stelex sensillum, 3-lateral stelex sensillum, 4-ventrolaterad stelex sensillum; (G) posterior spiracular plate, 1-
rima, 2-interspiracular process, 3-ecdysial scar; (H) intermediate sensory complex, 1-medusoid sensillum, 2-

stelex sensillum, 3-basal, conical, minute acanthus.

884

five congeners examined to date, all of
which are papilliform and arranged in a
double row above each mouthhook (Goe-
den and Headrick 1992, 1999; Goeden
2000a, b, c). The stomal sense organ of the
third instar of N. pubescens bears one or
two verruciform sensilla, two pore sensilla,
and one or two, cone-shaped or short pa-
pilliform sensilla (Fig. 2C-8). Thus, it ap-
pears similar in complexity to the stomal
sense organ of the second instar (Figs. 1B-
3, C-6); however, the stomal sense organs
of the third instars of four congeneric spe-
cies appear especially well-developed com-
pared to earlier instars and each bears dif-
ferent combinations of sensory structures,
variously described as several cone-shaped
sensilla in N. viridescens (Goeden and
Headrick 1992); as papilliform and pit-type
in N. wilsoni (Goeden and Headrick 1999);
as verruciform or “‘compound verruciform”’
in N. signifera (Goeden 2000a), and ver-
ruciform and pit-type in N. aenigma (Goe-
den 2000b).

The third instars of all six species of
Neaspilota examined to date have oral ridg-
es with dentate ventral margins character-
istically arranged in vertical series ventro-
laterad of the dorsal sensory organ and lat-
erad of the oral cavity. Similar oral ridges
also were described in the second instars of
N. viridescens (Goeden and Headrick
1992), N. wilsoni (Goeden and Headrick
1999), N. signifera (Goeden 2000a), and N.
appendiculata (Goeden 2000c). The oral
ridges number eight in N. pubescens (Fig.
2B-2), seven or eight in the third instar of
N. aenigma (Goeden 2000b), seven in N.
appendiculata (Goeden 2000c), but six in
the second and third instars of the other
three congeners examined to date. The ap-
pearance and arrangement of these oral
ridges appears to be a generic character;
however, the present study and Goeden
(2000c) confirm that the oral ridges vary in
number among third instars of Neaspilota
species. Also, the most ventral, eighth oral
ridge of N. pubescens is not ventrally
toothed (Fig. 2B-2). The third instars of

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Trupanea imperfecta (Coquillett), T. jonesi
Curran, 7. nigricornis (Coquillett), T. pseu-
dovicina (Hering), 7. signata Foote, and T.
wheeleri Curran also bear serrated oral ridg-
es (Goeden and Teerink 1997b, 1998,
1999a; Goeden et al. 1998; Knio et al.
1996; Teerink and Goeden 1999), but these
oral ridges appear to be fewer in number,
and are not arranged in a more or less reg-
ular, vertical row lateral to the mouth hook,
as in Neaspilota.

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

The number and appearance of the stelex
sensilla surrounding the posterior spiracular
plate differ among the Neaspilota species
examined to date. These number only four
in the first instars of N. wilsoni (Goeden and
Headrick 1999), N. aenigma (Goeden
2000b), and N. appendiculata (Goeden
2000), but, unfortunately, were not ob-
served with N. signifera (Goeden 2000a) or
in the present study. This count of stelex
sensilla remains at four in the second instars
of N. aenigma (Goeden 2000b), N. appen-
diculata (Goeden 2000c), and N. pubes-
cens, increases to six in third instars of N.
wilsoni (Goeden and Headrick 1999), N.
aenigma (Goeden 2000b), and N. appendi-
culata (Goeden 2000c). These stelex sen-
silla also show inter-instar (intraspecific)
and interspecific differences in the inci-
dence and appearance of the minute acan-
thae that may ring them basally, but this
was not recognized, studied or recorded by
my coworkers and me until recently (Goe-
den 2000b, c; Fig. 2H-2, -3).

VOLUME 102, NUMBER 4

pot Magn
J 3) 2bhy

wed Det WD Exp -——————__ 50 um Cc

Acc.V Spots
RLOLO KS) 475x SE

Fig. 3. Puparium of Neaspilota pubescens: (A)
habitus, anterior to left; (B) anterior end, 1-invagina-
tion scar, 2-anterior thoracic spiracle; (C) caudal seg-
ment, l-rima, 2-interspiracular process, 3-intermediate
sensory complex.

13.2 59

Puparium: Mostly pale yellow, with pos-
terior two-three segments grayish to black-
ened posteriorly, ellipsoidal, and smoothly
rounded at both ends (Fig. 3A); anterior end

885

bears the invagination scar (Fig. 3B-1) and
anterior thoracic spiracles (Fig. 3B-2); cau-
dal segment circumscribed by minute acan-
thae; three stelex sensilla, dorsolaterad, lat-
eral, and ventrolateral of posterior spiracu-
lar plates; posterior spiracular plate bears
three broadly elliptical rimae (Fig. 3C-1),
and four interspiracular processes, each
with one to three branches (Fig. 3C-2); in-
termediate sensory complex (Fig. 3C-3)
with a medusoid sensillum and a stelex sen-
sillum. Seven puparia averaged 2.79 + 0.05
(lange, 255—2:95) mmeainveneth 1.31
0.06 (range, 1.10—1.60) mm in width.

DISTRIBUTION AND Hosts

Freidberg and Mathis (1986) described
the distribution of N. pubescens as ‘‘South-
em California south of 35° north latitude
and west of the Sierra Nevada Mountains.”
Freidberg and Mathis (1986) and Foote et
al. (1993) mapped the distribution to in-
clude only California.

The only reported and confirmed host
plant of N. pubescens (reported as appen-
diculata) is Lessingia (reported as Coreth-
rogyne) filaginifolia by Goeden (1989),
which belongs to the subtribe Solidagininae
of the tribe Astereae in the family Astera-
ceae (Hickman 1993, Bremer 1994). This
perennial subshrub has at least two distinct
varieties and itself is widely distributed in
coastal scrub, oak woodlands, and grassland
below 2,600 m throughout California and
into adjacent southwestern Oregon and
northern Baja California, Mexico (Hickman
1993, Shreve and Wiggens 1964), where N.
pubescens also probably occurs.

Host records in Freidberg and Mathis
(1986) present a quandary, as only N. brun-
neostigmata Doane is reported from Lessin-
gia (reported as Corethrogyne) filaginifolia,
including both varieties filaginifolia and
californica (deCandolle) M. A. Lane (re-
ported as C. californica). Neither of these
records obtained from Wasbauer (1972) had
been checked by Freidberg and Mathis
(1986). Instead, both varietal records prob-
ably should refer to N. pubescens, not N.

886

brunneostigmata. On the other hand, I have
only reared N. brunneostigmata from Les-
singia glandulifera A. Gray and L. lem-
monii A. Gray, never N. pubescens, which
is easily distinguished from the former te-
phritid using the keys in Freidberg and
Mathis (1986) and Foote et al. (1993). The
latter host record for N. brunneostigmata
(as aenigma) was first reported by Goeden
(1989), and was corrected in Foote et al.
(1993). The host record for L. glandulifera
is new. Both represent the only confirmed
host records for N. brunneostigmata. The
seven other host records for N. brunneo-
stigmata listed by Wasbauer (1972) and re-
peated by Freidberg and Mathis (1986) re-
main unconfirmed and refer either to un-
documented records or to probable mis-
identifications listed in Foote and Blanc
(1963), which, of course, predated Freid-
berg and Mathis (1986). In summary, I be-
lieve that N. pubescens is a true mono-
phage, and I suspect, the better plant tax-
onomist in the case whereby C. filaginifolia
was renamed L. filaginifolia (Hickman
1993):

BIOLOGY

Egg.—lIn each of three, closed, preblos-
som, immature flower heads of L. filagini-
folia, a single egg was inserted pedicel-last;
two of these eggs had their long axes per-
pendicular to the receptacle, one rested at a
45° angle to the receptacle. The last egg had
been inserted through the phyllaries and
was embedded for half its length in the co-
rolla of a peripheral floret; whereas, the oth-
er two eggs were placed between an inner
phyllary and a peripheral floret. The diam-
eters of the receptacles of these flower
heads containing eggs averaged 1.7 + 0.17
(range, 1.40—1.99) mm.

Larva.—Upon eclosion, the only two
first instars found feeding in separate, pre-
blossom flower heads either tunneled into
an ovule, or into a corolla before entering
the ovule to which the corolla was basally
attached. The receptacles averaged 1.4 mm
in diameter and an average of 1.5 ovules

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

was damaged in these two flower heads.
Neither receptacle was abraded or pitted by
feeding.

Second instars continued feeding on
ovules in preblossom flower heads or on
soft achenes in open, blossom and post
blossom flower heads. All fed within a se-
ries of adjacent ovules/soft achenes with
their bodies more or less perpendicular to
and their mouthparts directed towards the
receptacles, but always above the recepta-
cles. Receptacles of 12 flower heads con-
taining second instars averaged 1.92 + 0.17
(range, 1.42—3.42) mm in diameter. These
flower heads each contained a single larva
that had destroyed an average of 3.8 + 0.6
(range, 1—8) ovules/soft achenes. Based on
23 (range, 17—34) as the average total num-
ber of ovules. and achenes respectively
counted in 40 preblossom to postblossom
flower heads, about 9.5% (range, 2.5—20%)
of the ovules/soft achenes in the 12 flower
heads were damaged by second instars.

Third instars (Fig. 4A) initially continued
to feed mainly on soft achenes in blossom
or postblossom flower heads. Twenty-four
flower heads that averaged 1.95 + 0.14
(range, 1.10—2.56) mm in diameter each
contained a single third instar. An average
of 15 + 2.1 (range, 4-34) of the soft
achenes therein were damaged, or about
38% (range, 10—100%) of the average total
of 40 ovules/soft achenes per flower head.
These percentages of seed predation per lar-
va per flower head, like those reported for
N. aenigma (Goeden 2000b) and N. appen-
diculata (Goeden 2000c), are on the high
side among florivorous tephritids studied by
us to date (Headrick and Goeden 1998).
This percent seed destruction per flower
head is exceeded only by gregarious flori-
vorous species like Trupanea_ conjuncta
(Adams) (Goeden 1987) and T. pseudovi-
cina Hering (Goeden and Teerink 1998) or
by species with large larvae that develop in
immature or small flower heads like Para-
cantha cultaris (Coquillett) (Cavender and
Goeden 1982) and Xenochaeta dichromata
Snow (Goeden and Teerink 1997a). Also, it

VOLUME 102, NUMBER 4

887

Fig. 4.
achenes in open flower head, (B) late third instar feeding on soft achenes in open flower head, (C) puparium
formed in spring by overwintered prepuparium in flower head, (D) mating adults, dorsal view, (E) mating adults,
ventral view, (F) mating adults lateral view. Lines = 1 mm.

should be noted that the rates of flower
head infestation per sample for N. pubes-
cens was very low, e.g., mean of 4.5%
(range, 1.5%—10%) for five samples, like
most Neaspilota in nature that we have
studied (unpublished data).

Third instars in flower heads fed with
their long axes oriented perpendicular to
and mouthparts directed towards the recep-
tacles (Fig. 4B). Only five (21%) of the
third instars in the 24 infested heads scored
or pitted the receptacles; however, most
presumably supplemented their diet with

Life stages of Neaspilota pubescens in Lessingia filaginifolia: (A) early third instar feeding on soft

sap. Goeden and Headrick (1992, 1999) and
Goeden (2000c) described and discussed
this similar type of feeding by N. virides-
cens, N. wilsoni, and N. appendiculata, re-
spectively. And, as also reported for all
three of these congeners that overwinter as
prepuparia (Goeden and Headrick 1992,
1999; Goeden 2000c), most third instars be-
came surrounded for about 90% of their
lengths by cells, which occupied most of
the interior of the flower heads and con-
sisted of ovule-, achene-, chaff-, pappus-,
and corolla-fragments cemented together by

888

liquid feces and sap that hardened when dry
(Fig. 4C). These protective cells were
slightly larger than the mature larvae, ex-
ternally incorporated the outer walls of
achenes and the few uneaten achenes, and
were blackened and smooth inside (Fig.
4C). Upon completing feeding and cell con-
struction, the larvae oriented with their an-
terior ends towards the receptacles, retract-
ed their mouthparts, and formed prepuparia
(Headrick and Goeden 1998). Most individ-
uals overwintered in diapause as prepuparia
(Goeden and Headrick 1992, 1999; Goeden
2000c; Headrick and Goeden 1998), but a
few individuals pupariated early and
emerged in summer (July—August). Prior to
pupariation the prepuparia reversed their
orientation within their cells and turned
180° such that their heads were directed
away from the receptacles (Fig. 4C).

Adult.—Adults emerged from overwin-
tered, mature flower heads, and probably
are long-lived. Under insectary conditions,
five unmated males averaged 67 + 23
(range, 23-152) days, but only two virgin
females were available for study and aver-
aged 27 (range, 24—30) days. Such lengthy
longevities for males compare favorably
with average adult longevities reported for
adults of N. viridescens (Goeden and Head-
rick 1992), N. wilsoni (Goeden and Head-
rick 1999), N. signifera (Goeden 2000a), N.
aenigma (Goeden 2000b), and N. appendi-
culata (Goeden 2000c).

The premating and mating behaviors of
N. pubescens were not studied in the field,
but were observed in petri dish arenas
found to be so useful with many other non-
congeneric, nonfrugivorous, tephritid spe-
cies (Headrick and Goeden 1994). Premat-
ing behaviors observed with paired N. pu-
bescens were abdominal pleural distension
and side-stepping by males while tracking
females (Headrick and Goeden 1994) and
rapid wing hamation, sometimes combined
with lofting about 20° by both sexes (Head-
rick and Goeden 1994). No trophallaxis or
nuptial gift presentation was noted as re-
ported with N. viridescens (Goeden and

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Headrick 1992). Seven matings, six by one
pair during a 2-week period (Figs. 4D, E,
F) were observed that began during late af-
ternoon at dusk and lasted an average of
1,032 + 73 (range, 850—1,440) min (or 17
h and 12 min on average). This was three
to five times longer than the average dura-
tions of 190 min reported for N. aenigma
(Goeden 2000b), 235 min reported for UN.
wilsoni (Goeden and Headrick 1999), 238
min reported for N. signifera (Goeden
2000a), 285 min reported for N. appendi-
culata (Goeden 2000c), and 318 min re-
ported for N. viridescens (Goeden and
Headrick 1992). It also was among the lon-
gest mating durations reported by my col-
laborators and me (Headrick and Goeden
1994), exceeded only by a pair of Tephritis
baccharis (Coquillett) that mated for 2 days
and nights under similar conditions (Goe-
den and Headrick 1991). A pair of Dioxyna
picciola (Bigot) was reported to stay to-
gether in the field for 28 h, during which
time they copulated and oviposited three
times in one day, remained in copula on
one flower head overnight, and copulated
and oviposited again three times during the
following day (Headrick et al. 1996), but
this involved mate guarding (Headrick and
Goeden 1994, Headrick et al. 1996), also
observed with N. signifera (Goeden 2000a).
As far as could be determined, the extended
mating by N. pubescens involved continu-
ous copulation.

The only precopulary behavior observed
for the female was an elevation of the ovis-
cape 20° to 30° coupled with rapid wing
hamation and lofting, which may have sig-
naled receptivity (but see postcopulatory
behavior below). A male was observed to
initiate mating by rapidly pursuing and
grabbing the head of the female from the
front with his forelegs and -tarsi, and wres-
tling her into submission while he mounted
her. No copulatory induction behavior was
noted with N. pubescens and intromission
was gained rapidly. Most matings were ob-
served only after copulation had begun,
suggesting that precopulatory behavior was

VOLUME 102, NUMBER 4

perfunctory. The mating position (Figs. 4D,
E, F) was such that the wings of the male
were parted about 20°, while the wings of
the female were parted at about 60°, with
both pairs of wings centered over their re-
spective body midlines (Figs. 4D, E). The
body of the female was parallel to the sub-
strate, while the body of the male was el-
evated about 20° anteriorly (Fig. 4F). The
hind- and midtarsi of the male usually rest-
ed on the substrate, but sometimes his mid-
tarsi grasped the base of the oviscape, while
his foretarsi hooked onto the abdominal
pleura of the female midway along the ab-
domen (Figs. 4D, F). The male’s mouth-
parts were positioned above the second ab-
dominal tergite of the female (Figs. 4D, FP).
In this position the male tenaciously held
onto the female as copulation continued,
while defeating her increasingly vigorous
efforts to dislodge him. These dislodgment
efforts by the female consisted of walking
rapidly, then stopping to strongly arch her
dosum at the juncture of her thorax and ab-
domen while rapidly hamating and lofting
her wings (Fig. 5A), or bending her ovi-
scape upward, while kicking at his head,
ventrum, and mid- and hind legs with her
hind legs and tarsi. In between these bouts
of female “restlessness,” the pair largely
remained quiescent with only their mouth-
parts pumping, or sometimes both forming
regurgitation droplets (Headrick and Goe-
den 1994), or while the female groomed her
head and fore legs. Besides tightly clinging
to the female, the male sometimes reposi-
tioned his foretarsi so as to gain better pur-
chase or rub his midtarsi alternately along
her oviscape to induce passivity as a vari-
ation on copulatory induction behavior
(Headrick and Goeden 1994). These bursts
of activity by the female always preceded
disengagement, but only infrequently re-
sulted in disengagement. What action(s) fi-
nally triggered or caused the pair to sepa-
rate, remain unanswered.

Two mating terminations and disengage-
ments were observed, which involved each
male turning in place and walking away

Figs 5:
mating by Neaspilota pubescens: (A) female arching
dorsum and kicking at male with hind legs and tarsi,
(B) male turning and stepping off female, (C) male
pulling genitalia from female as he walks away. Lines
= 1 mm.

Disengagement sequence at termination of

from the female while pulling his genitalia
from within her, a process that lasted less
than 10 s in both cases (Figs. 5A, B. C).
Postcopulatory behavior by N. pubescens
mainly consisted of storing of the genitalia
by males and cleaning and grooming by
both sexes (Headrick and Goeden 1994); al-

890

though, instances of a female strongly ele-
vating her oviscape was seen, as she ran
away from a pursuing male to avoid recou-
pling following disengagement. As noted
above, this same behavior often preceded
mating.

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

Natural enemies.—A single female of
Pteromalus sp. (Hymenoptera: Pteromali-
dae) was reared from a puparium of N. pu-
bescens as a solitary, larval-pupal endopar-
asitoid.

ACKNOWLEDGMENTS

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

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 892-900

A TAXONOMIC REVIEW OF THE GENUS DELPHINIOBIUM MORDVILKO
(HOMOPTERA: APHIDIDAE) IN CHINA

GE XIA QIAO AND GUANG XUE ZHANG

Institute of Zoology, The Chinese Academy of Sciences, 19 Zhongguancun Lu, Haidian,
Beijing 100080, People’s Republic of China (e-mail: leifm@panda.ioz.ac.cn)

Abstract.—The aphid genus Delphiniobium Mordvilko from China is reviewed. There
are four species in China, Delphiniobium gyamdaense Zhang, D. yezoense Miyazaki, D.
violisuctum Qiao and Zhang, n. sp., and D. aconitifoliae Zhang and Qiao, n. sp. A
key to species from China, morphological descriptions, distributions, host plants, and
diagnosis of new species are provided. In addition, the apterous viviparous female of D.
gyamdaense Zhang is described for the first time.

Key Words:

Mordvilko (1914) erected the genus Del-
phiniobium for the species Myzus junack-
ianum Karsch 1887. Hille Ris Lambers
(1947) studied the genus Delphiniobium
from Europe. At present, this genus is rep-
resented by eight species and one subspe-
cies, D. bogdouli Szelegiewicz 1969, D.
canadense (Robinson 1968), D. carpaticae
Mamontova 1966, D. hanla Paik 1971, D.
Junackianum (Karsch 1887), D. junackinum
sylvanae (Knechtel and Manolache 1944),
D. lycoctoni B6rner 1950, D. yezoense Mi-
yazaki 1971 and D. gyamdaense Zhang
1981 (Remaudiére and Remaudiére 1997).
In the course of the present study, two new
species, D. aconitifoliae Zhang and Qiao
and D. violisuctum Qiao and Zhang are de-
scribed from China, plus two other species,
D. gyamdaense Zhang and D. yezoense Mi-
yazaki. A key to species in China is pro-
vided. In addition, the apterous viviparous
female of D. gyamdaense Zhang 1981 is
described for the first time.

MATERIALS AND METHODS

All specimens studied including types are
housed in the Zoological Museum, Institute

Homoptera, Aphididae, Delphiniobium, new species, China

of Zoology, the Chinese Academy of Sci-
ences, Beijing.

Aphids were mounted on microscope
slides in Arabic gum, and observed under
phase contrast microscopy. Terminology
follows Miyazaki (1971). Drawings were
made by the first author, Dr. X. L. Chen,
and Mr. T. S. Zhong using microscopy.
Measurements are in millimeters.

Delphiniobium Mordvilko 1914

Delphiniobium Mordvilko 1914: 65.

Type-species: Myzus junackianum Karsch
1887 (= Rhopalosiphum aconiti van der
Goot 1912). By monotypy.

Generic diagnosis.—Body elliptical. Me-
dial frontal tubercle undeveloped or devel-
oped, lateral frontal tubercles with diverg-
ing inner sides present. Thoracic stigmal
pores much longer than abdominal stigmal
pores. Spiracular sclerites of thorax strong-
ly produced, with opening very large and
round. Mesosternal furca with short or long
stem. Antennal segment III with large and
small round secondary rhinaria. Ultimate
rostral segment stout. First tarsal segment
chaetotaxy: 3, 3, 3. Siphunculi slender,

VOLUME 102, NUMBER 4

slightly or distinctly swollen, with distinct
reticulation consisting of several rows of
hexagonal cells near apex, paler than cauda
basally. Cauda dark, long tapering. Mainly
on Aconitum and Delphinium.

In general aspects this genus resembles
Megoura Buckton, from which it can be
easily separated by the reticulated apices of
the siphunculi. On the other hand many
Macrosiphum-like species in America have
swollen siphunculi with a reticulated area,
but in none of them a black, sclerotic cauda
occurs. Other genera with swollen siphun-
culi have either no reticulated area (Ampho-
rophora Buckton), or (Rhopalosiphoninus
Baker) often no rhinaria on the antennal
segment III in apterous viviparous females.

Distribution.—Europe (Romania, Eng-
land, Netherlands, Germany, Russia), North
America (Canada), Asia (China, Japan, Ko-
rea, Mongolia).

KEY TO APTEROUS VIVIPAROUS FEMALES OF
SPECIES FROM CHINA

1. Medial frontal tubercle not developed, antennal
tubercles small (Fig. 22); abdominal tergite I
with 14 hairs; ultimate rostral segment with 2
pairs accessory hairs; China: Shanxi (Yang-
cheng County) Delphiniobium violisuctum
Qiao and Zhang, n. sp.
— Medial frontal tubercle developed, antennal tu-
bercles large (Figs. 1, 15); abdominal tergite I
with at most 10 hairs; ultimate rostral segment
with 3—5 pairs accessory hairs ........... 2
. Siphunculi slightly or distinctly swollen in
middle; ultimate rostral segment at most 1.30
times as long as second hind tarsal segment;
antennal segment III with either at most 25 sec-
ondary rhinaria or at least 44............. 3
— Sipunuculi not swollen; ultimate rostral seg-
ment 1.35 times as long as second hind tarsal
segment; antennal segment III with 21—39 sec-
ondary rhinaria; China: Beijing (Sanpu Dis-
trict, Xiaolongmen District)
Delphiniobium aconitifoliae
Zhang and Qiao, n. sp.
3. Antennal segment HI with 44—57 secondary
rhinaria, on basal %4 of segment; siphunculi dis-
tinctly swollen (Fig. 20); China: Qinghai (Hu-
zhu County), Xizang (Gyamda County)
Bre ori Bers Delphiniobium gyamdaense Zhang
— Antennal segment III with 10—25 secondary
rhinaria, on basal half of segment; siphunculus
slightly swollen; China: Sichuan (An County),

N

893

Hebei (Kuancheng County, Wulinshan Moun-
tain, Xiaowutai Mountain); Japan .......
Ae TE ee Meta Delphiniobium yezoense Miyazaki

KEY TO ALATE VIVIPAROUS FEMALES OF
SPECIES FROM CHINA

1. Siphunculi distinctly swollen; antennal seg-

ment III longer than segmentsIV+V..... 2
— Siphunculi indistinctly swollen; antennal seg-
ment III shorter than segments IV + V .... 3

N

. Antennal segment III with 77-81 secondary
rhinaria; yellow in life; siphunculus 2.00 times
as long as cauda; China: Qinghai (Huzhu
County), Xizang (Gyamda County) .....
Delphiniobium gyamdaense Zhang
— Antennal segment HI with 40—65 secondary
rhinaria; bluish green in life; siphunculus 1.60—
1.70 times as long as cauda; China: Hebei (Ku-
ancheng County, Wulinshan Mountain, Xia-
owutai Mountain), Sichuan (An County); Japan
Delphiniobium yezoense Miyazaki
3. Median frontal tubercle indistinct; ultimate ros-
tral segment 1.06 times as long as second hind
tarsal segment; siphunculi 1.74 times as long
as cauda; China: Shanxi (Yangcheng County)
Delphiniobium violisuctum
Qiao and Zhang, n. sp.
— Median frontal tubercle distinct; ultimate ros-
tral segment 1.40 times as long as second hind
tarsal segment; siphunculi 2.00 times as long
as cauda; China: Beijing (Sanpu District, Xiao-
longmen District) . .

Delphiniobium aconitifoliae
Zhang and Qiao, n. sp.

Delphiniobium aconitifoliae Zhang and
Qiao, new species
(Figs. 1-14)

Description.—Measurements: Apterous
viviparous female: Body 3.56 in length,
1.78 in width. Antenna 3.72, length of seg-
ments I-VI 0.15, 0.09, 1.20, 0.59, 0.50,
0.12+1.08, respectively. Ultimate rostral
segment 0.19 in length, 0.09 in basal width.
Hind femur 1.41, hind tibia 2.53, second
hind tarsal segment 0.16. Siphunculus 0.72
in length. Cauda 0.50 in length. Alate vi-
viparous females: Body 3.20 in length, 1.18
in width. Antenna 3.93, length of segments
EVE (O41 45. OOH WIAD) 0262; .0:58;
0.12+1.23, respectively. Ultimate rostral
segment 0.21 in length, 0.11 in basal width.
Hind femur 1.38, hind tibia 2.51, second
hind tarsal segment 0.15. Siphunculus 0.72
in length. Cauda 0.37 in length. Alate male:

894

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-14. Delphiniobium aconitifoliae. 1-9, Apterous viviparous female. 1, Dorsal view of head. 2, An-
tennal segments I-III. 3, Antennal segment VI. 4, Ultimate rostral segment. 5, Dorsal view of abdominal seg-
ments V—VIII. 6, Dorsal hair of body. 7, Ventral hair. 8, Siphunculus. 9, Cauda. 10—11, Alate viviparous female.
10, Antennal segment III. 11, Fore wing. 12-13, Alate male. 12, Antennal segments III-V. 13, Clasper. 14,
Oviparous female, hind tibia.

Body 2.64 in length, 1.13 in width. Ovip-
arous female: Body 3.61 in length, 1.83 in
width.

Apterous viviparous female: Body oval,
red and yellow in life. Mounted specimens:
Head grey, thorax and abdomen pale, with-
out patches. Antenna black except base of
antennal segment III pale; rostral segments
III-V, distal half of femora, tibiae and tarsi
black, siphunculi pale at base, brown at
middle, black distally, cauda and anal plate
black, genital plate pale. Abdominal seg-
ments VII and VIII with slight imbrications.
Spiracles large, closed, spiracular plates
slightly brown. Mesosternal furca with
short stem, pale. Dorsal hairs of body thick

and short, stout at apex. Ventral hairs sim-
ilar to but longer than dorsal hairs. Head
with 1 pair cephalic hairs, 1 pair antennal
tubercular hairs, 4—5 pairs dorsal hairs.
Pronotum with 1 pair spinal, 1 pair pleural
and 1 pair marginal hairs. Abdominal ter-
gites I-VI each with 4—6 spino-pleural
hairs, tergite VII with 2 or 3; tergites II-
VII each with 2-3 pairs marginal hairs, ter-
gite I with | pair marginal hairs, tergite VIII
with 2 pairs hairs. Length of cephalic hairs
0.04, length of marginal hairs on abdominal
tergite I 0.03, and length of dorsal hairs on
tergite VIII 0.05, 0.52 times, 0.39 times and
0.75 times as long as widest diameter of
antennal segment III, respectively. Median

VOLUME 102, NUMBER 4

frontal tubercle developed, antennal tuber-
cles developed and diverging, higher than
median front. Antenna 6-segmented, basal
part of segment III and segments IV—VI
with distinct imbrications, length 1.10 times
as long as body; length in proportion to seg-
ments I-VI: 13, 7, 100, 49, 41, 10+90, re-
spectively. Antennal hairs similar to dorsal
hairs, segments I-VI each with 6 or 7, 4,
25-31, 7-13, 7-10, 3 or 4 +3-8 hairs, re-
spectively, apex of processus terminalis
with 3 hairs. Length of hairs on antennal
segment III 0.04, 0.57 times as long as wid-
est diameter of segment. Antennal segment
III with 21—39 large and small round sec-
ondary rhinaria (Fig. 2) on basal % of seg-
ment. Rostrum thick and large, reaching
midcoxae, clypeus with 2 pairs hairs ante-
riorly; ultimate rostral segment 2.00 times
as long as its basal width, 1.20 times as
long as second hind tarsal segment, with 7
pairs hairs, 3—4 pairs accessory hairs
among them. Hind femur 1.20 times as long
as antennal segment III. Hind tibia 0.71
times as long as body. Length of hairs on
hind tibia 0.06, 0.93 times as long as middle
tibia width of segment. First tarsal segment
chaetotaxy: 3, 3, 3. Siphunculi long, taper-
ing, slightly constricted at distal %4, with
12-14 rows of reticulations at constricted
part; 0.20 times as long as body, 1.40 times
as long as cauda; with flange. Cauda taper-
ing, rough, with spinulose imbrications;
with 6-8 thick and long hairs. Anal plate
circular at apex, with 11—19 long and short
hairs. Genital plate semicircular, with 14—
16 hairs.

Alate viviparous female: Mounted spec-
imens: Head and thorax slightly brown,
head surrounding eyes black, abdomen
pale, without patches. Spiracles large and
opened, spiracular plates brown. Marginal
areas of body and some abdominal seg-
ments behind siphunculi with distinct im-
brications. Dorsal hairs of body thick and
short, stout at apex. Head with 1 pair ce-
phalic hairs, 2—3 pairs antennal tubercular
hairs, 4 pairs dorsal hairs. Abdominal ter-
gites I-VII each with 3 or 4 spinal, 1 pair

895

pleural and 4—8 pairs marginal hairs, tergite
VIII with 2-3 pairs hairs. Length of ce-
phalic hairs 0.04, length of marginal hairs
on abdominal tergite I 0.03, and length of
dorsal hairs on tergite VII 0.05, 0.61 times,
0.49 times and 0.72 times as long as widest
diameter of antennal segment III, respec-
tively. Antenna 6-segmented, 1.20 times as
long as body; length in proportion to seg-
ments: Vi: 12.18; 100: 55.51 LO aa07,
respectively. Antennal segment III with 29—
36 hairs, length 0.03, 0.46 times as long as
widest diameter of segment. Antennal seg-
ment III with 46—73 large and small round
secondary rhinaria, on entire segment. Ros-
trum reaching midcoxae, ultimate rostral
segment 1.40 times as long as second hind
tarsal segment, with 8—9 pairs hairs, 5—6
pairs accessory hairs among them. Hind fe-
mur 1.21 times as long as antennal segment
III. Hind tibia 0.79 times as long as body.
Veins normal. Siphunculi swollen at distal
half, constricted at apex, 0.23 times as long
as body, 2.00 times as long as cauda. Cauda
long, tapering, with 7 hairs. Anal plate with
17-19 hairs. Other characters similar to ap-
terous viviparous female.

Alate male: Body long oval. Mounted
specimens: Antenna, rostrum, distal half of
femora, tibiae, tarsi, siphunculi, cauda, anal
plate dark brown, other appendages slightly
brown. Abdominal tergites I-V each with 1
pair spino-pleural and 1 pair marginal
patches. Dorsal hairs of body with sclerites
at base. Dorsal hairs slightly short, ventral
hairs longer than dorsal hairs. Antenna 6-
segmented, 1.50 times as long as body. Sec-
ondary rhinaria small round, on segments
III-V; segment III with 53—63, segment IV
with 8-11, segment V with 7-11. Fore-
wing media two-branched, hindwing two
obliques. Clasper shown in Fig. 13. Other
characters similar to alate viviparous fe-
male.

Oviparous female: Body oval. Mounted
specimens: Antennal segments I-III, distal
part of segment IV, distal part of segment
V, basal part of segment VI, distal part of
rostrum, distal half of femora, tibiae, tarsi,

896

cauda, anal plate, distal / of siphunculi
dark brown, other appendages slightly
brown; basal Y; of siphunculi pale, rest of
siphunculi slightly brown. Genital plate
slightly brown. Antennal segment III with
12-18 small round secondary rhinaria, on
basal half of segment. Hind tibia swollen at
basal half, with about 250 small round
pseudo-sensoria. Other characters similar to
apterous viviparous female.

Diagnosis.—The new species differs
from D. yezoense Miyazaki as follows:
Clypeus with 4 hairs (yezoense, 2 hairs),
basal and distal of antennal segment III and
basal of segment IV pale (yezoense, black,
except basal of segment III), and yellow or
red in life (yezoense, green or bluish green).
It differs from D. bogdouli Szelegiewicz as
follows: Antennal segment V black (bog-
douli, basal part of segment pale), yellow
or red in life (bogdouli, shining brown in
life), and length of dorsal hairs of body 0.33
(bogdouli, 0.45).

Etymology.—The species name is based
on the host plant, Aconitum kusnezofii.

Holotype.—Apterous viviparous female,
No. 6804-1-2-3, 1978-VI-8, China, Beijing
(Sanpu District, 116.4°E, 89.9°N), Col. G.
X. Zhang and T. S. Zhong, on young twigs
and upper of leaves of Aconitum kusnezofii
Reichb.

Paratypes.—Two apterous viviparous fe-
males and 2 alate viviparous females, No.
6804, other data same as holotype; 2 alate
viviparous females, 5 apterous females and
2 alate males, No. 6502, 1976-X-5, Col. B.
L. Zhang, other data same as holotype; 2
apterous viviparous females, | alate vivip-
arous female, 3 alate males and 12 apterous
females, No. 6628, 1977-X-14, other data
same as holotype; 5 apterous viviparous fe-
males and 3 alate viviparous females, No.
6230, 1976-V-18, other data same as holo-
type; 1 apterous viviparous female, 1 alate
viviparous female and 2 apterous females,
No. 11535, 1997-IX-11, China, Beijing
(Xiaolongmen District, 116.0°E, 39.9°N),
Col. J. G. Xiangyu, on Aconitum sp.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Delphiniobium gyamdaense Zhang 1981
(Figs. 15-21)

Delphiniobium gyamdaense Zhang 1981:
264.

Description.—Apterous viviparous fe-
male: Body 3.04 in length, 1.50 in width.
Body elliptical, yellow in life, siphunculi,
antennae, legs and cauda black. Mounted
specimens: Head and prothorax slightly
brown, mesonotum, metanotum and abdo-
men pale, without patches. Antennal seg-
ments dark brown, except basal part of seg-
ment III and distal part of segment VI; dis-
tal part of rostrum, distal half of femora,
distal part of tibiae and tarsi dark brown;
distal ¥, of siphunculi, cauda and anal plate
dark brown, others pale. Mesosternal furca
with long stem. Median frontal tubercle
slightly developed, antennal tubercles de-
veloped, higher than median frontal tuber-
cle. Antenna 6-segmented, about as long as
body; length in proportion to segments I—
VI: 13, 8, 100, 40, 40, 10+115. Antennal
segment III with 44—46 round secondary
rhinaria, over the entire segment. Length of
hairs on segment III % as long as widest
diameter of segment. Rostrum exceeding
midcoxae, ultimate rostral segment 2.35
times as long as its basal width, with 8 ac-
cessory hairs. Siphunculi long barrel-
shaped, swollen at middle, width at swollen
part 1.90 times distal width, distal % with
reticulations, 0.16 times as long as body,
1.25 times as long as cauda. Cauda long
tapering, indistinctly constricted, 2.56 times
as long as its basal width, with 6 hairs. Anal
plate with 16 hairs.

Specimens examined.—Three apterous
viviparous females, No. 11391, 1997-VI-8,
China: Qinghai (Huzhu County, 101.9°E,
36.8°N), Col. X. L. Chen, on Aconitum sp.,
Xizang (Gyamda County, 93.1°E, 29.9°N).

Delphiniobium violisuctum Qiao and
Zhang, new species
(Figs. 22-31)

Description.—Measurements: Apterous
viviparous female: Body 3.13 in length,

VOLUME 102, NUMBER 4

897

Figs. 15—21.
segments I-IV. 17, Antennal segments V and VI. 18, Ultimate rostral segment. 19, Mesosternal furca. 20,
Siphunculus. 21, Cauda.

1.43 in width. Antenna 3.70, length of seg-
ments I-VI 0.14, 0.10, 1.10, 0.56, 0.46,
0.14+1.27, respectively. Ultimate rostral
segment 0.12 in length. Hind femur 1.37,
hind tibia 2.30, second hind tarsal segment
0.16. Siphunculus 0.67 in length. Cauda
0.45 in length. Alate viviparous female:
Body 3.40 in length, 1.58 in width.
Apterous viviparous female: Body large,
dark green in life, on upper side of leaves.
Mounted specimens: Head and prothorax
brown, antennal segments I and IJ, distal 7%
of segment III, distal % of segment IV, dis-
tal half of segment V, basal part of segment
VI and distal half of processus terminalis
dark brown, others brown; apex of rostrum,
distal ¥; of femora, distal ¥, of tibiae, tarsi,

Delphiniobium gyamdaense, apterous viviparous female. 15, Dorsal view of head. 16, Antennal

distal %4 of siphunculi, cauda and anal plate
dark brown, coxae, trochanters, basal /; of
femora, basal ”, of siphunculi and genital
plate slightly brown, others brown. Spira-
cles large, oval, opened; spiracular plate
long oval, slightly pale. Mesosternal furca
with long stem. Dorsal hairs of body thick,
short, stout at apex. Head with 1 pair of
cephalic hairs and 1| pair antennal tubercu-
lar hairs, 4 pairs dorsal hairs; pronotum
with 3 pairs spinal, 2 pairs pleural and 1
pair marginal hairs; abdominal tergite I
with 14 hairs, tergite VII with 4 hairs.
Length of cephalic hairs 0.04, length of
marginal hairs on tergite I 0.03, and length
of dorsal hairs on tergite VII 0.05, 0.73
times, 0.55 times and 0.91 times as long as

898

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 22-31.
Antennal segments I-IV. 24, Antennal segments V and VI. 25, Ultimate rostral segment. 26, Mesosternal furca.
27, Dorsal view of abdominal tergites II—VIII. 28, Spiracle on abdominal segment II. 29, Siphunculus. 30,
Cauda. 31, Alate viviparous female, antennal segment III.

widest diameter of antennal segment III, re-
spectively. Median frontal tubercle unde-
veloped, antennal tubercles distinct, higher
than median frontal tubercle, slightly di-
verging at inner margins. Antenna 6-seg-
mented, 1.18 times as long as body, length
in proportion to segments I-VI: 14, 10, 100,
54, 45, 14+123. Processus terminalis 9.11
times as long as base of segment. Antennal
hairs similar to dorsal hairs, antennal seg-
ments I-VI each with 7, 4, 30, 11, 6, 3+3
hairs, respectively, apex of preocessus ter-
minalis with 3 hairs. Length of hairs on an-
tennal segment III 0.04, 0.64 times as long
as widest diameter of segment. Antennal
segment III with 11 small round secondary

Delphiniobium violisuctum. 22—30, Apterous viviparous female. 22, Dorsal view of head. 23,

rhinaria, on basal half of segment. Rostrum
reaching midcoxae, ultimate rostral seg-
ment thick, 1.33 times as long as its basal
width, 0.87 times as long as second hind
tarsal segment; with 2 pairs accessory hairs.
Legs normal. Hind femur 1.33 times as long
as antennal segment III. Hind tibia 0.72
times as long as body. Hairs on legs slightly
longer than dorsal hairs of body. Length of
hairs on hind tibia 0.07, 1.40 times as long
as middle width of segment. First tarsal
segment chaetotaxy: 3, 3, 3. Siphunculus
long barrel-shaped, 3.42 times as long as
basal width, 8.13 times as long as distal
width, 0.21 times as long as body, 1.48
times as long as cauda, swollen at middle,

VOLUME 102, NUMBER 4

distal ¥, of siphunculi with reticulations.
Cauda long, tapering, slightly constricted at
middle, 2.44 times as long as its basal
width, with 7 hairs. Anal plate circular at
apex, with 18 hairs. Genital plate with 15
hairs, 2 long anterior hairs among them.
Gonapophyses three, each with 5 or 6 hairs.
Alate viviparous female: Body large.
Mounted specimens: Basal of antennal seg-
ment III pale, distal 4% of tibiae black, distal
half of femora, and other antennal segments
slightly brown, other appendages dark
brown; dorsum of head brown; distal of
rostrum, cauda, and anal plate greyish
brown; basal ’/,; of siphunculi slightly
brown, distal ¥; of siphunculi dark brown.
Abdominal dorsum without patches. Anten-
na 6-segmented, segment III with 42—47
small round secondary rhinaria, on entire
segment. Fore wing veins brown, media
two-branched. Other characters similar to
apterous viviparous females.
Diagnosis.—The new species is similar
to D. bogdouli Szelegiewicz from Mongolia
by the median frontal tubercle and antennal
tubercles, but differs as follows: antennal
segment III with 42—47 secondary rhinaria
in alatae (bogdouli, 58—62); ultimate rostral
segment 0.87 times as long as second hind
tarsal segment (bogdouli, longer than, 1.10
times); siphunculi slightly swollen at mid-
dle (bogdouli, indistinctly swollen); and
dark green in life (bogdouli, shining
brown).
Etymology.—The new species is named
based on the host plant, Viola verecunda.
Holotype.—Apterous viviparous female,
No. Y8070-1-1-1, China: Shanxi (Yangch-
ene County; 112.4°E, 35.5°N), Col-.. Fei
Zhao, 1996-VI-1, on Viola verecunda.
Paratypes.—One alate viviparous female
and 1 apterous larvae, No.Y8070, other
data same as holotype.

Delphiniobium yezoense Miyazaki 1971

Delphiniobium yezoense Miyazaki 1971:
34(1):40.

Distribution.—No.Y4347, China: Hebei
(Wulinshan Mountain, 117.4°E, 40.6°N),

899

1983-IX-12, Col. S.P. Tian; No. Y5718, Chi-
na: Hebei (Kuancheng County, 118.4°E,
40.6°N), 1983-IX-9, Col. S.P. Tian;
No.Y5597, China: Hebei (Xiaowutai
Mountain, 115.°E, 89.9°N), 1984-VI-22,
Col. S.P. Tian; No.Y1598, China: Sichuan
(An County, 104.4°E, 31.6°N), 1979-V-22,
Col. H.Y. Li; No.Y1436, China: Sichuan
(An County), 1978-IX, Col. H.Y. Li; Japan.

Host-plants.—Aconitum kusnezoffi and
A. comiichali (in China); A. chinense, A. ki-
tadakense, A. yesoense and A. sachalinense
(in Japan).

ACKNOWLEDGMENTS

We express our thanks to Mr. S. P. Tian,
Mr. T. S. Zhong, Mr. H. Y. Li, Mrs. E Zhao,
and Dr. X. L. Chen for collection of spec-
imens and preparing some illustrations. The
project was supported by the National Nat-
ural Sciences Foundation of China (a grant:
39700015) and a Biological and Technical
Innovation Grant from the Chinese Acade-
my of Sciences (A2999084).

LITERATURE CITED

Borner, C. 1950. Neue europdische Blattausarten.
Selbsverlag. Naumbury, Private publication. 19
PP-

Hille Ris Lambers, D. 1947. Contributions to a Mono-
graph of the Aphididae of Europe II. Temminckia
VII: 179-319.

Karsch, A. 1887. Myzus junackianus n. sp. Berliner
Entomologica Zoologici 31: 21-23.

Knechtel, W. K. and C. I. Manolache. 1944. Neue Blat-
tlause fiir Rumanien. V. Academic Rumanine,
Bulletin Section Science 25: 267-270.

Mamontova-Solukha, V. O. 1966. Aphids of the So-
vietic Karpats. Insects of Ukrainian Karpats and
Transkarpattia, ser. Prolems of Zoological Akad
Nauk Ukrainian RSR, Kiev, pp. 3—13.

Miyazaki, M. 1971. A revision of the tribe Macrosi-
phini of Japan (Homoptera: Aphididae, Aphidi-
nae). Insecta Matsumurana 34(1): 1—9, 40—42.

Mordvilko, A. K. 1914. [Insectes Hémiptéres (Insecta
Hemiptera), Vol. I. Aphidoidea. Livraison I. (Fati-
ne de la Russie et des pays limitrophes, fondée
principalement sur les collections du Musée Zool-
ogique de 1’Académie Impériale des Sciences de
Petrograd.)] Petrograd. 236 pp.

Paik, W. H. 1971. A new species of aphid from Mt.
Hanla. Korean Society Plant Protection 10(2): 75—
76.

900

. Insecta V. Illustrated Encyclopedia of Fauna
& Flora of Korea 13: 245, 354—356.

Remaudiére, G. and M. Remaudiére. 1997. Catalogue
of the World’s Aphididae. Homoptera Aphidoidea.
Institut National de la Recherche Agronomique.
147, rue de I’ Universite, 75338 Paris Cedex 07:
473 pp.

Robinson, A. G. 1968. Two new species of aphids

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(Homoptera: Aphididae) from Canada. The Ca-
nadian Entomologist 100: 275—279.

Szelegiewicz, H. 1969. Neue Blattlaus (Homoptera,
Aphididae) aus der Mongolei. Annales Zoologici
Warszawa 27: 169-194.

Zhang, G. X. and T. S. Zhong. 1981. Homoptera:
Aphidoidea, pp. 264—265. Jn Huang E S., ed., In-
sects of Xizang, Beijing: Science Press, 600 pp.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 901-911

NEW COSTA RICAN SPECIES OF PHYLLOPHAGA HARRIS
(COLEOPTERA: MELOLONTHIDAE: MELOLONTHINAE)

MIGUEL ANGEL MORON AND ANGEL SOLIS

(MAM) Departamento de Entomologia, Instituto de Ecologia, A.C. (SEP-CONACYT).
Apdo. Postal 63, Xalapa, Veracruz 91000, México (e-mail: moron.ma @ecologia.edu.mx);
(AS) Departamento de Entomologia, Instituto Nacional de Biodiversidad (INBio), Santo
Domingo de Heredia, Costa Rica

Abstract.—Four new species of Phyllophaga are described from Costa Rica, Phyllo-
phaga kohlmanniana and P. picadoana from the montane rain forest of the Monteverde
Biosphere Reserve, Puntarenas, and P. janzeniana and P. guanacasteca from the tropical
deciduous forests lowlands of Guanacaste. Drawings of male genital capsules, female
genital plates, and tarsal claws are provided.

Resuimen.—Se describen cuatro especies nuevas de Phyllophaga procedentes de tres
localidades en Costa Rica: P. kohlmanniana y P. picadoana del bosque lluvioso de mon-
tama de la Reserva de la Biosfera Monteverde, Puntarenas; P. janzeniana y P. guanacas-
teca de los bosques tropicales caducifolios de las tierras bajas de Guanacaste. Se incluyen
ilustraciones de las capsulas genitales masculinas, de las placas genitales femeninas y de

las unas tarsales.

Key Words:

During curatorial work at the collection
of INBio (1993-1999), we found specimens
that represent four undescribed species of
Phyllophaga from the provinces of Guan-
acaste and Puntarenas, Costa Rica. Males of
these species have deep cleft or bifid tarsal
claws, while the females have dentate tarsal
claws. Using the criteria of Saylor (1942),
Sanderson (1958), or Mor6n (1986), the
males of these species key to the subgenus
Phytalus, whereas the females key to the
subgenus Phyllophaga (s.str.). A similar
situation is present with other species from
Mexico, such as Phyllophaga ambigenus
(Bates) and Phyllophaga mesophylla Mo-
ron and Rivera (1992), both placed in the
subgenus Phytalus based on the structure of
the male tarsal claws and other characters.
Awaiting for new advances in the supras-
pecific classification of the genus, we place

Phyllophaga, May beetles, new species, tropical forests, Costa Rica

the presently described species as incerta
sedis. In this paper, we describe males and
females, address variation within the spe-
cies, and present the precise distribution
data for four new species of Phyllophaga
from the montane rain forests and decidu-
ous tropical forests in Costa Rica.

Acronyms used in the text are as follows:
CAS, California Academy of Sciences, San
Francisco, USA; INBio, Instituto Nacional
de Biodiversidad, Costa Rica; MXAL, pri-
vate collection M.A. Moron, Xalapa, Méx-
ico.

Phyllophaga kohlmanniana Moron and
Solis, new species
(Figs. 1-5)

Description.—Holotype male: Head and
pronotum shiny dark brown, with dense
vestiture of short whitish setae; elytra, ster-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

3

Figs. 1—5.

Phyllophaga kohlmanniana. 1, Male protarsal claw, lateral view. 2, Genital capsule, lateral view.

3, Paramera, distal view. 4, Female protarsal claw, lateral view. 5, Female genital plates, ventral view. Scale

lines = 1 mm, except figs. 1, 4 = 0.5 mm.

nites and pygidium reddish dark brown,
sligthly pruinose, completely covered with
dense vestiture of uniform, whitish, short
setae, that offer a macroscopic grayish, vel-
vety appearance; mouth parts, tibiae and
tarsi shiny reddish brown; coxae and fem-
ora partially pruinose. Clypeus 3.7 * wider
than long, anterior border curved, not sin-
uate, with elevated margin, surface slightly
concave, with many uniformly distributed,
deep, round punctures each provided with
short, erect seta. Frontoclypeal suture wide-
ly sinuate and deeply impressed. Frons 1.6
x wider than long, convex, finely punctate
rugose, with short erect setae on entire sur-
face. Antenna 10-segmented, with three
segmented club, lamellae of 8th to 10th
segments 1.6 X longer than length of pre-
ceeding six segments combined; segments
4 or 5 shorter than 3, with rounded promi-
nences on anterior sides; segments 6 and 7
wider than long, with semiconical promi-
nences on anterior sides. Frons 3.7 X wider

than dorsal diameter of eye. Eye canthi long
and wide, with 13-14 setae. Labrum bi-
lobed, widely sinuated, with scattered slen-
der setae. Mentum slightly concave, im-
punctate, with scarce slender setae, anterior
border briefly sinuated, nearly straight.
Pronotum 1.7 X wider than long and 2.9 X
wider than frons. Pronotal disk shiny, but
with basi-central area pruinose, total sur-
face with deep, round punctures regularly
separated by 1—2 diameters; lateral borders
widely angulate, lateral marginal bead cren-
ulate, with long, curved setae; basal bead
complete, indicated by wide, deep sulcus;
anterior angles straight, slightly prominent;
posterior angles nearly straight, not promi-
nent. Scutellum 1.5 X wider than long, with
many small punctures. Elytron 2.3 X longer
than wide, pruinose, densely and regularly
punctate-setose; epipleural border narrow,
extended along complete margin, slightly
widened at % and ¥ of its length, provided
with scarce slender setae at anterior third,

VOLUME 102, NUMBER 4

dull or pruinose at middle and posterior
thirds; humeral calla rounded, prominent;
apical calla rounded. Metathoracic wings
completely developed. Propygidium prui-
nose, with dense setiferous punctures. Py-
gidium scarcely convex, pruinose, with
round shaped, shallow rounded seitiferous
punctures, regularly distributed; apical mar-
gin with 16 long, slender setae; basal mar-
gin effaced. Pterosterna with long, dense,
yellowish setae. Visible abdominal sternites
II to IV slightly depressed at midline, with
a noticeable shallow sulcus; sternite V
sligthly convex, pruinose, with dense seti-
ferous punctures at middle; anal plate nar-
rowed, transversely excavated, with elevat-
ed anterior and posterior margins, and 10
setae on apical border. Protibia nearly as
long as protarsus (1:1.1), with external bor-
der tridentate, preapical spur acute, straight,
longer than 2nd protarsomerus (1.2:1). Me-
sotibia with one oblique, well marked, se-
tiferous carina on external side; upper api-
cal spur straight, narrow, and 1.4 xX longer
than lower spur. Metatibia slightly shorter
than metatarsus (1:1.2), with one oblique
setiferous carina on external side; upper
apical spur articulated, straight, sharply
pointed, as long as basal metatarsomere,
and 1.5 X longer than lower spur; lower
apical spur articulated, apex acute. Tarso-
meres semicylindrical, elongate, with en-
larged apex, some setae apically and two
lines of setae along ventral side. Tarsal
claws symmetrical, similar in all legs, deep-
ly cleft, upper tooth shorter than lower
tooth (Fig. 1). Genital capsule with short,
narrowed parameres, dorsal and ventrally
fused, ring shaped, apex with medium size,
tooth-like projections (Fig. 3). Aedeagus
very long, with preapical patches of spi-
nules at sides and sclerotized dorsal support
with sinuose flagellum (Fig. 2). Tectum (=
phallobase) uniformly convex. Length of
genital capsule from apex of parameres to
border of basal piece: 6.8 mm. Total body
length: 21.2 mm. Humeral width: 10.0 mm.

Allotype female: Similar to the male ex-
cept as follows: antenna with segments 5

903

and 6 fused, and appearing 9-segmented.
Visible abdominal sternites II to VI convex,
with dense setiferous punctures; anal plate
1.8 X longer than male anal plate, very con-
vex, punctate, with 12 slender setae at the
apical border. Meso- and metatibiae each
with one oblique, strong, setiferous carina
near middle and other vague carina on basal
third of external side. Both apical spurs of
metatibia articulated, wide, lanceolate and
curvated. Tarsal claws dentate, with long
tooth near middle of ventral border (Fig. 4).
Ventral genital plates well sclerotized, near-
ly symmetrical, elongated, with short setae
at distal process; dorsal genital plates fused
with ventral ones toward lateral sides, with
acure apices directed mesad and setae on
distal borders (Fig. 5). Total body length:
20.6 mm. Humeral width: 9.8 mm.

Variation.—Male: Similar to holotype
except as follows: total body length: 19.8—
21.5 mm, humeral width: 9.6—10.3 mm,
pronotum of some specimens more reddish
than holotype, other specimens with anten-
nal club as long as preceeding six segments
combined. Female similar to allotype ex-
cept as follows: pygidium with more seti-
ferous punctures; total body length: 19.5—
20.4 mm; humeral width: 9.6—9.8 mm.

Type material.—Described from 5 6, 4
?. Holotype 6 INBio: Costa Rica: Puntar-
enas, Monteverde, La Casona, IV-92, 1,520
m, N. Obando. Allotype 2 INBio: Costa
Rica: Guanacaste, SW Volcan Cacao, Est.
Mengo, II-89, 1,100 m, GNP Biodiversity
Survey. Paratypes INBio, MXAL: same
data as holotype (2 ¢ and 2 ¢); same data
as allotype (2 6, 1 @).

Type locality.—La Casona, Estaci6n
Monteverde, province of Puntarenas, Costa
Rica (aprox. 10°30’N; 85°10’ W).

Biological data.—Specimens of P. kohl-
manniana were collected at lights near
cloud forests and tropical premontane for-
ests located between 1,100—1,520 m. Phe-
nology: February (4), April (5). Other spe-
cies of Phyllophaga flying at the same time
were P. (Phyllophaga) tapantina Mor6n

904

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

8

Figs. 6-11.

Phyllophaga janzeniana. 6, Male protarsal claw, lateral view. 7, Genital capsule, lateral view.

8, Paramera, distal view. 9, Paramera, ventro-apical view. 10, Female tarsal claw, lateral view. 11, Female genital
plates, ventral view. Scale lines = 1 mm, except figs. 6, 10, 11 = 0.5 mm.

and Solis, P. (Phyllophaga) tilarana Mor6n
and Solis and P. picadoana, n.sp.
Remarks.—Phyllophaga kohlmanniana
does not belong to any species group de-
scribed by Moron (1986). The structure of
the male tarsal claws suggests placement in
the subgenus Phytalus, but the form of the
female tarsal claws indicates placement in
the subgenus Phyllophaga (s.str.), as is
known for P. ambygenus (Bates) and some
other Mexican and Central American spe-
cies. By the color, vestiture, body size,
length of tarsi, and form of clypeus is sim-
ilar to P. (Phytalus) zeteki Saylor, 1942, de-
scribed from Barro Colorado Island, Pana-
ma, but the shape and details of the male
genital capsule (Fig. 2), sculpture and ves-

titure of the sternites, shape of tarsal claws
(Fig. 1) and proportions of antennal seg-
ments will aid in the recognition of this new
species.

Etymology.—This new species is dedi-
cated to our friend, Dr. Bert Kohlmann,
whose enthusiastic devotion to the study of
the scarab beetles of Mexico and Central
America is well known to coleopterists and
also to people interested in insect ecology
and biogeography.

Phyllophaga janzeniana Moron and
Solis, new species
(Figs. 6-11)

Description.—Holotype male: Clypeus,
frons and pronotum shiny dark reddish

VOLUME 102, NUMBER 4

brown; elytra and pygidium shiny reddish
brown, without macroscopic vestiture;
mouth parts, sterna and legs shiny testa-
ceous reddish brown. Clypeus 3.6 X wider
than long, anterior border rounded, not sin-
uate, with elevated margin, surface slightly
concave, with some wide, deep, coarse,
round punctures without microscopic setae.
Frontoclypeal suture sinuate and vaguely
impressed. Frons 2.0 X wider than long,
convex, coarsely punctate rugose, with
transverse keel on anterior half and some
medium size setae at sides and minute setae
scattered on the posterior half of disk. An-
tenna 9-segmented, with 3 segmented club,
lamellae as long as the length of preceeding
5 segments combined; 5th segment longer
than 4 or 3; segment 6 compresed with
semiconical anterior process. Frons 3.3 X
wider than dorsal diameter of eye. Eye can-
thi long and wide, with 9—10 setae. Labrum
widely concave, sligthly bilobed, with scat-
tered slender setae. Mentum slightly con-
cave, impunctate, with scarce, slender setae,
anterior border briefly notched. Pronotum
1.6 X wider than long and 2.9 X wider than
frons. Pronotal disk shiny, with deep,
coarse umbiliform punctures irregularly
separated by 1-3 diameters; lateral borders
widely angulate, lateral marginal bead
widely crenulated, with scattered, long,
slender setae; basal bead vaguely indicated
by punctures, mainly toward the sides; an-
terior angles straight, prominent; posterior
angles widely obtuse, rounded. Scutellum
1.3 X wider than long, with some irregular
punctures. Elytron 2.8 < longer than wider,
shiny, densely rugo-punctate; epipleural
border very narrow, extended along com-
plete margin, provided with some scattered
short setae; humeral calla rounded, promi-
nent; apical calla rounded. Metathoracic
wings completely developed. Propygidium
glabrous, with distal half nearly shiny and
basal half slightly pruinose, with sparse um-
biliform punctures. Pygidium moderately
convex, shiny, glabrous, coarsely rugo-
punctate; apical margin with 14 long, slen-
der setae; basal margin effaced medially.

905

Pterosterna with medium size, moderately
dense, yellowish setae. Visible abdominal
sternites II and IV slightly convex, nearly
polished and glabrous toward the middline;
sternite V convex, shiny, irregularly rugo-
punctate with scarce, scattered setae at mid-
dle; anal plate narrowed, transversely con-
cave, irregularly punctate, anterior and pos-
terior borders elevated, with some slender
setae. Protibia slightly shorter than protar-
sus (1:1.2), with external border tridentate,
preapical spur acute, nearly straight, as long
as 2nd protarsomerus. Mesotibia with one
oblique, sharp setiferous carina on external
side; upper apical spur straight, narrow, and
1.1 X longer than lower spur. Metatibia
nearly as long as metatarsus, with one
oblique, sharp setiferous carina on external
side; upper apical spur articulated, angulat-
ed, sharply pointed, 1.2 < longer than basal
metatarsomere, and 1.1 X longer than lower
spur; lower apical spur articulated, with
acute apex. Tarsomeres semicylindrical,
elongate, with enlarged apex, some setae
apically and two lines of setae on ventral
side. Tarsal claws symmetrical, similar in
all legs, widely and deeply cleft, upper
tooth nearly as long as lower tooth, poste-
rior ventral border slightly serrated (Fig. 6).
Genital capsule with medium size paramer-
es, dorsally fused, elongated with apex ex-
panded and curvated toward the middle line
(Figs. 7-8); ventrally the paramera are not
fused (Fig. 9). Aedeagus long, with a preap-
ical dorsal sclerotized plate that support a
pair of long curved spine-like sclerotized
structures (Fig. 7). Tectum uniformly con-
vex. Length of genital capsule from apex of
parameres to border of basal piece: 3.7 mm.
Total body length: 11.9 mm. Humeral
width: 5.0 mm.

Allotype female: Similar to the male ex-
cept as follows: anterior angles of clypeus
more wide and rounded, expanded; anten-
nal club as long as the length of four pre-
ceeding segments; pronotum with anterior
angles more prominent and lateral bead no-
ticeably crenulate. Visible abdominal ster-
nites II to IV convex, with scattered seti-

906

ferous punctures; sternite V more long and
convex; anal plate 2 X< longer than male
anal plate, convex, punctate, with 10 slen-
der setae near the posterior border. Pygidi-
um with central part of disk slightly flat-
tened. Both apical spurs of metatibia slight-
ly curvated with the apex rounded. Tarsal
claws with upper tooth slightly shorter than
ventral one (Fig. 10). Ventral genital plates
with basal half well sclerotized and distal
part membranous, sclerotized only at bor-
ders, sligthly asymmetrical, basally round-
ed, elongated to apex, with some setae on
border; dorsal genital plates poorly sclero-
tized, fused medially, with some setae at
each side (Fig. 11). Total body length: 11.2
mm. Humeral width: 5.2 mm.

Paratype variation.—Male similar to ho-
lotype except in total body length: 10.8—
11.8 mm, humeral width: 4.8—5.2 mm,
pronotum and elytra of some specimens
darker than holotype, other specimens with
frontal transverse carina more or less ac-
centuated. Female similar to allotype except
as follows: pronotum, elytra and pygidium
with more or less punctures; total body
length: 10.8—11.5 mm; humeral width: 5.0—
5.4 mm.

Type material.—Described from 8 6, 10
?. Holotype 6 INBio: Costa Rica: Guan-
acaste, 12 km SE La Cruz, Cerro El Hacha,
Casa Oeste, VI-87, 300 m, GNP Biodiver-
sity Survey. Allotype 2°: same data as ho-
lotype (INBio). Paratypes CAS, INBio,
MXAL: same data as holotype (4 6, 1 2);
Guanacaste, PN. Rincon de La Vieja, Est.
Las Pailas, 1/22-VII-92, 800 m, D. Garcia
GB. #8-2));

Type locality—Cerro El Hacha, 12 km
SE La Cruz, province of Guanacaste, Costa
Rica (approx. 10°58'N; 85°32'W).

Biological data.x—Specimens of P. jan-
zeniana were collected at lights in tropical
deciduous forests located between 300—800
m. Phenology: June (7), July (11). Other
species of Phyllophaga flying at the same
time were P. (Chlaenobia) scabripyga
(Bates), P. (Phytalus) obsoleta (Blanchard),

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

P. (Phyllophaga) elenans Saylor and P.
(s.str.) hondurasana Moser.

Remarks.—Phyllophaga janzeniana does
not belong to any species group proposed
by Mor6én (1986). It may be related to P.
soctona Mor6n from Chiapas, México,
which is also a small species with frontal
transverse carina, 9-segmented antenna and
similar tarsal claws, but the shape and de-
tails of the male genital capsule (Figs. 7—8)
and female genital plates (Fig. 11) are very
different. Each paramere of P. soctona have
two large, apical teeth, and dorsal genital
plates of female are not fused medially.

Etymology.—This new species is dedi-
cated to Dr. Daniel H. Janzen, whose inter-
est in the study and conservation of tropical
forests, mainly in the Guanacaste area, is
well known.

Phyllophaga picadoana Moré6n and
Solis, new species
(Figs. 12-16)

Description.—Holotype male: Head,
pronotum and pygidium shiny dark reddish
brown; with sparce vestiture of short to
long setae; mouth parts, sterna and legs
shiny reddish brown. Clypeus 3.2 * wider
than long, anterior border shallowly and
widely sinuated, with margin elevated, sur-
face nearly flat, with large, shallow, round
punctures and some scattered short setae.
Frontoclypeal suture fine, vaguely indicated
by a wide, shallow sinuate sulcus. Frons 1.7
x wider than long, convex, coarsely punc-
tate rugose, covered with short to medium
size slender setae. Antenna 10-segmented,
with 3-segmented club, lamellae 2 < longer
than length of preceeding 5 segments com-
bined. Frons 4.1 X wider than dorsal di-
ameter of eye. Eye canthi long and wide,
with 7-8 setae. Labrum widely sinuate,
slightly bilobed, with scattered slender se-
tae. Mentum slightly convex, impunctate,
with few slender setae, anterior border shal-
lowly and widely sinuated. Pronotum 1.7 X
wider than long and 3 X wider than frons.
Pronotal disk shiny, with shallow, round
shaped, large setiferous punctures irregular-

VOLUME 102, NUMBER 4

907

16

Figs. 12-16. Phyllophaga picadoana. 12, Male protarsal claw, lateral view. 13, Genital capsule, lateral view.
14, Paramera, distal view. 15, Female protarsal claw, lateral view. 16, Female genital plates, ventral view. Scale

lines = | mm, except figs. 12, 15 = 0.5 mm.

ly separated from one another by 1-3 di-
ameters; lateral borders widely angulate,
lateral marginal bead slightly crenulated,
mainly toward basal half, with scattered,
long, slender setae; basal bead erased, only
with shallow sulcus, mainly toward sides;
anterior angles acute, prominent; posterior
angles slightly obtuse, not prominent. Scu-
tellum 1.6 X wider than long, with 10
rounded punctures. Elytron 2.4 X longer
than wide, densely punctate-rugose, mostly
shiny, except the preapical area and distal
half of the lateral borders that have vague
pruinose vestiture; epipleural border mod-
erately narrowed, extended along complete
margin, provided with scattered short, slen-
der setae; humeral calla rounded, promi-
nent; apical calla rounded. Metathoracic
wings completely developed. Propygidium
shiny, with dense setiferous punctures. Py-
gidium moderately convex, shiny, rugose,
with round, shallow setiferous punctures ir-
regularly distributed; apical margin with 18

long, slender setae; basal margin effaced
medially. Pterosterna with long, dense, yel-
lowish setae. Visible abdominal sternites II
and IV slightly depressed, polished and
nearly glabrous at the middline; sternite V
convex, with a patch of granules and erect
short setae on middle and with a wide, shal-
low transverse sulcus before posterior bor-
der; anal plate sligthly concave, vaguely
furrowed at middle with some erect, medi-
um size setae and anterior and posterior
borders slightly elevated. Protibia slightly
shorter than protarsus (1:1.2), with external
border tridentate, preapical spur acute, near-
ly straight, slightly longer than 2nd protar-
somerus (1:1.2). Mesotibia with one
oblique, sharp, setiferous carina on external
side; upper apical spur straight, narrowed,
as long as the lower spur. Metatibia slightly
shorter than metatarsus (1: 1.2), with one
oblique, sharp, setiferous carina on external
side; upper apical spur nearly lanceolate,
curved, with rounded apex, as long as basal

908

metatarsomere, and 1.4 X longer than lower
spur; lower apical spur articulated, with
rounded apex. Tarsomeres semicylindrical,
elongate, with enlarged apex, some setae
apically and two rows of setae with a fine
longitudinal keel in middle of ventral side.
Protarsomeres 1—3 with subapical short
spines. Tarsal claws symmetrical, similar in
all legs, deep cleft, upper tooth as long as
lower tooth, with middle of ventral border
finely serrate (Fig. 12). Genital capsule with
proportionately short parameres, dorsally
fused, apical thirth canaliculated by inner
side (Fig. 14). Aedeagus large, with wide
apical patches of spinules and dorsal preap-
ical tuft of strong setae on a wide sclero-
tized support (Fig. 13). Tectum uniformly
convex. Length of genital capsule from
apex of parameres to border of basal piece:
4.1 mm. Total body length: 17.1 mm. Hu-
meral width: 7.8 mm.

Allotype female: Similar to male except
as follows: elytra with long erect setae near
scutellum and toward apex. Pygidium less
convex and rugose. Visible abdominal ster-
nites II to V convex, with scattered setifer-
ous punctures; anal plate convex, punctate,
with many slender setae. Apical spurs of
metatibia wide, lanceolate and curved. Tar-
sal claws with preapical tooth on ventral
border, nearly as long as apical tooth, with
part of ventral border serrate (Fig. 15). Ven-
tral genital plates slightly sclerotized, nearly
symmetrical, with apical border rounded
(Fig. 16). Total body length: 17.3 mm. Hu-
meral width: 8.1 mm.

Paratype variation.—Male similar to ho-
lotype except in total body length: 17.0—
17.4 mm, humeral width: 7.6—7.9 mm, dor-
sal vestiture of some specimens less dense
than holotype, other specimens with anten-
nal club 1.8—2.1 X longer than length of
preceeding five segments combined. Spec-
imens from Chirripo have a darker color,
distal half of paramera less canaliculated
and have a wide shallow concavity on the
dorsal part of parameral base. Female sim-
ilar to allotype except as follows: elytra and
pygidium with more or less setiferous punc-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tures; total body length: 16.8—17.3 mm; hu-
meral width: 7.9—8.1 mm.

Type material—Described from 10 6
and 8 2. Holotype ¢ INBio: Costa Rica:
Puntarenas, R.B. Monteverde, La Casona,
I-92, 1,520 m, N. Obando. Allotype 2 IN-
Bio: same data as holotype except III-1994.
Paratypes CAS, INBio, MXAL: same data
as holotype (5 ¢); same data as allotype (2
3,3 2); Cartago, Turrialba, Chirripo, Gra-
no de Oro, I[X-92, 1,120 m, P. Campos (2
6a4Ae®):

Type locality—La Casona, Reserva Biol-
6gica Monteverde, province of Puntarenas,
Costa Rica (approx. 10°30'N; 85°10'W).

Biological data.—Specimens of P. pica-
doana were collected at lights near montane
tropical forests and coffee plantations lo-
cated at 1,120—1,520 m. Phenology: March
(12), September (6). Other species of Phyl-
lophaga flying at the same time were P.
(Phyllophaga) tilarana Moron and Solis, P.
(s.str.) tapantina Moroén and Solis and P.
kohlmanniana Moron and Solis.

Remarks.—Phyllophaga picadoana does
not belong to any species group proposed
by Moron (1986). Externally this species
resembles P. (Phyllophaga) brevidens
Bates and allies, but the male and female
tarsal claws of P. brevidens have a small
tooth at the middle of ventral border, each
paramere is deeply bifurcate and the aedea-
gus do not present wide apical patches of
spinules.

Etymology.—The name of this new spe-
cies is dedicated to the memory of the for-
mer Costa Rican biologist, Dr. Clodomiro
Picado Twight, who devoted much of his
life to the study of animals, tropical ecology
and natural richess of that country.

Phyllophaga guanacasteca Moron and
Solis, new species
(Figs. 17-22)

Description.—Holotype male: Head,
pronotum and elytra dark reddish brown
with dense pruinose greyish vestiture;
mouth parts, pygidium and legs shiny red-
dish brown; sternites light reddish brown

VOLUME 102, NUMBER 4

909

Figs. 17-22.

Phyllophaga guanacasteca. 17, Male protarsal claw, lateral view. 18, Male hind tarsal claw,

lateral view. 19, Genital capsule, lateral view. 20, Paramera, distal view. 21, Female protarsal claw, lateral view.

22. Female genital plates, ventral view. Scale lines =

with fine pruinose vestiture. Clypeus 3.4 X
wider than long, anterior border wide and
deeply sinuate, with elevated margin, sur-
face slightly convex, with many uniformly
distributed, shallow large punctures, with-
out setae. Frontoclypeal suture sinuate and
deeply impressed. Frons 1.4 < wider than
long, convex, densely punctate, without se-
tae. Antenna 10-segmented, with 3-seg-
mented club, lamellae 1.3 xX longer than
length of preceeding 6 segments combined.

1 mm, except figs. 17, 18, 21 = 0.5 mm.

Frons 2.5 X wider than dorsal diameter of
eye. Eye canthi long and wide, with 9-11
setae. Labrum bilobed, widely sinuated,
with scattered slender setae. Mentum slight-
ly convex, impunctate, with few slender se-
tae, anterior border nearly straight. Prono-
tum 1.6 X wider than long and 2.9 X wider
than frons. Pronotal disk pruinose, with
shallow, round small punctures regularly
separated from one another by 2—4 diame-
ters; lateral borders widely rounded, lateral

910

marginal bead entire, with 2-3 scattered
slender setae; basal bead indicated by punc-
tures, mainly toward sides, where shallow
sulcus are also indicated; anterior angles
straight, slightly prominent; posterior an-
gles obtuse, slightly prominent. Scutellum
1.5 X wider than long, scattered small
punctures. Elytron 2.6 X longer than wide,
pruinose, uniformly punctate with some in-
terstriae slightly elevated; epipleural border
narrow, extended along complete margin,
provided with some scattered short setae;
humeral calla rounded, prominent; apical
calla rounded. Metathoracic wings com-
pletely developed. Propygidium pruinose,
with uniform round, small punctures. Py-
gidium convex, shiny, with round, shallow
punctures irregularly distributed; apical
margin with 14 long, slender setae; basal
margin effaced medially. Pterosterna with
medium size, moderately dense, yellowish
setae. Visible abdominal sternites II and III
slightly depressed, pruinose, with small,
black, granules at midline; sternite [V with
dentiform irregular black granules near pos-
terior border; sternite V with rounded area
scarcely prominent at middle, covered with
fine rugosities, granules and short setae;
anal plate narrowed, sligthly convex, with
some short setae near midline and 6 slender
setae on apical border. Protibia shorter than
protarsus (1:1.8), with external border tri-
dentate, but basal tooth very reduced;
preapical spur acute, nearly straight, shorter
than 2nd protarsomerus (1:2.2). Mesotibia
with one oblique, setiferous carina on ex-
ternal side; upper apical spur straight, nar-
rowed to the apex, and 1.8 X longer than
lower spur; lower spur with truncated apex.
Metatibia shorter than metatarsus (1:1:3),
with one oblique, setiferous carina on ex-
ternal side; upper apical spur straight,
sharply pointed, slightly longer than basal
metatarsomere (1.2:1), and 1.9 X longer
than lower spur; lower apical spur articu-
lated, curved, with rounded apex. Protar-
someres 1—4 slightly concave by ventral
sides, with lateral rows of setae. Meso- and
metatarsomeres semicylindrical, elongate,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

with enlarged apex, some setae apically and
two lines of setae on ventral side. Protarsal
claws slightly asymmetrical, deeply cleft,
upper tooth longer and narrower than lower
tooth (Fig. 17). Meso- and metatarsal claws
symmetrical, deeply cleft, with upper tooth
slightly longer than lower tooth (Fig. 18).
Genital capsule with long parameres, dor-
sally fused, with preapical tooth-like pro-
jections on the external and internal borders
(Figs. 19-20). Aedeagus short, with preap-
ical, large, sclerotized asymmetrical plate
(Fig. 19). Tectum with two large latero-dis-
tal rounded prominences (Fig. 19). Length
of genital capsule from apex of parameres
to border of basal piece: 5.9 mm. Total
body length: 19.1 mm. Humeral width: 7.9
mm.

Allotype female: Similar to male except
as follows: clypeal disk with more deep
punctures; antennal club shorter than length
of five preceeding segments (1:1.3). Pygid-
ium smaller, slightly convex, with more
punctures, mainly toward apex. Visible ab-
dominal sternites II to TV convex, with scat-
tered setae; sternite V slightly longer than
the IV segment, with more setae; anal plate
convex, punctate, with 30 slender setae.
Meso- and metatibiae each with one
oblique, strong, setiferous carina on exter-
nal side. Both apical spurs of metatibia ar-
ticulated, wide, lanceolate and curved. Pro-
tarsal segments 1—4 semicylindrical, elon-
gated with enlarged apex, but not concave
on ventral surface. Tarsal claws clearly den-
tate, with apical tooth nearly as long as pos-
terior one (Fig. 21). Ventral genital plates
well sclerotized, symmetrical, rounded with
wide apical borders; dorsal genital plates
ovate, with slightly projected rounded apex
(Fig. 22). Total body length: 20.0 mm. Hu-
meral width: 8.4 mm.

Paratype variation.—Male similar to ho-
lotype except in total body length: 18.6—
19.3 mm, humeral width: 7.6—8.0 mm, dor-
sal color of some specimens darker than ho-
lotype or with the pruinose vestiture more
or less dense. Female similar to allotype ex-
cept as follows: frons and pygidium with

VOLUME 102, NUMBER 4

more punctures; total body length: 19.5—
20.3 mm; humeral width: 8.3—8.4 mm.

Type series.—Described from 4 d, 3 9.
Holotype ¢ INBio: Costa Rica: Estacion
Pitilla, 9 km S Sta. Cecilia, PN. Guana-
caste, Guanacaste, Costa Rica, V-94, 700
m, C. Moraga. Allotype 2 INBio: same
data except III/IV-93, Malaise. Paratypes
INBio, MXAL: same data as holotype (3 6,
ey):

Type locality—Estacion Pitilla, 9 km S
Santa Cecilia, Guanacaste National Park,
province of Guanacaste, Costa Rica
(approx. 10°59'26"N; 85°25'40"W).

Biological data——Males and females of
P. guanacasteca were collected at lights
near deciduous tropical forest located at
700 m. Phenology: March—April (1), May
(6). Other species of Phyllophaga flying at
the same time were P. hondurasana Moser
and P. guapiles Saylot.

Remarks.—Phyllophaga guanacasteca
does not belong to any species group de-
fined by Moron (1986). It is general ap-
pearance suggest a relationship with P.
(Phytalus) pruinosa (Blanchard), but the ar-
ticulated lower metatibial spur in the male,
and the dentate tarsal claws of the female
do not match with the diagnostic characters
of the group pruinosa. The shape and de-
tails of the male genital capsule (Figs. 19—
20) and female genital plates (Fig. 22),
length of lower metatibial spur of male,
sculpture of the pygidium and sternites and
shape of protarsal claws (Fig. 17) will aid
in the recognition of this new species.

Etymology.—Specific epithet derived

OV

from the name of the province of Guana-
caste, to which this species appears to be
restricted.

ACKNOWLEDGMENTS

Study of the collections of INBio made
by Miguel A. Moron was made possible by
the support of Instituto de Ecologia, Xalapa
and Programa Diversidad Biolégica, CYT-
ED-México (1993), and by the support of
INBio (1998). Research done at Instituto de
Ecologia, Xalapa by Angel Solis was made
possible with the financial aid of INBio
(1997) and CONACYT (1999). We appre-
ciate corrections and suggestions of two
anonymous reviewers for better English ex-
pression. This paper is a contribution to the
project ‘““Sistematica y Biologia del género
Phyllophaga en México y América Cen-
tral’? (225260-25723-N CONACYT Méxi-
CO).

LITERATURE CITED

Moron, M. A. 1986. El género Phyllophaga en Méx-
ico. Morfologia, Distribuci6n y Sistemdatica Su-
praespecifica (Insecta: Coleoptera). Publ. 20, In-
stituto de Ecologia, México, 341 pp.

Moron M. A. and L. E. Rivera-Cervantes. 1992. Dos
especies nuevas de Phyllophaga (Phytalus) (Co-
leoptera: Melolonthidae) de la Sierra de Manan-
tlan, Jalisco, México. Anales Instituto de Biologia
Universidad Nacional Autonoma de México, serie
Zoologia 63(1): 79-87.

Sanderson, M. W. 1958. Faunal affinities of Arizona
Phyllophaga, with notes and descriptions of new
species. Journal of the Kansas Entomological So-
ciety 31: 158-173.

Saylor, L. W. 1942. Notes on beetles related to Phyl-
lophaga Harris with descriptions on new genera
and subgenera. Proceedings of the United States
National Museum 92(3145): 157-165.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 912-918

TWO ADDITIONAL SPECIES OF ROBBER FLIES OF THE GENUS
OMMATIUS WIEDEMANN (DIPTERA: ASILIDAE) FROM THE BAHAMAS
AND WITH REPLACEMENT NAMES FOR TWO OTHER SPECIES

A. G. SCARBROUGH

Department of Biology, Towson University, Baltimore, MD 21252, U.S.A. (e-mail:

ascarbrough @ towson.edu)

Abstract.—Two species of robber flies of the genus Ommatius Wiedemann are reported
from the Bahamas, increasing the number of species to six. Ommatius mariae, Nn. sp.,
from Eleuthera and Nassau, is described, illustrated, and contrasted with O. hanebrinki
Scarbrough and Rutkauskas. Ommatius mariae, O. hanebrinki, and O. hispidus Scar-
brough from the hanebrinki species group. A Cuban species, O. lineolatus Scarbrough is
reported for the first time from Long Island in the Bahamas. Two new replacement names
are proposed: Ommatius dignus new name for Ommatius dimidiatus Scarbrough and
Ommatius fimbrillus new name for Ommatius fimbriatus Scarbrough and Poinar.

Key Words:

The robber flies of the genus Ommatius
Wiedemann from the Bahama Islands are
poorly known. To date four species are re-
ported from single localities, 1.e., O. abana
Curran (1953) from Bimini, O. hanebrinki
Scarbrough and Rutkauskas (1983) from
San Salvador, O. membranous Scarbrough
(1985a) from Rum Cay, and _ setiferous
Scarbrough (1988) from Mayaguana. Fur-
thermore, only one sex is known for all spe-
cies except for O. hanebrinki. This paper
reports a new species of Ommatius Wie-
demann from Eleuthera and Nassau, and the
discovery of a Cuban species, O. lineolatus
Scarbrough (1988), from Long Island. Two
replacement names are proposed to correct
nomenclatural errors.

METHODS

General methods and terminology follow
that described by Scarbrough (1997) and
McAlpine (1981), respectively. Ratios used
in the text are as follows: face:head width

nomenclature, Asilidae, Ommatius, new species, new records, Bahamas

ratio (FHWR) is the greatest width of the
head in front profile divided by the width
of the face at the base of the antennae; cell
m1 width ratio (m1WR) is the ratio of the
width of the cell at the basal third and api-
cal two-thirds divided by the width of the
base; hind femoral width:length ratio
(HFWLR) is the greatest dorsal length di-
vided by the greatest width.

Acronyms of museums used in the text
are as follows: TUMZ, Museum of Zoolo-
gy, Towson University, Baltimore, MD;
AMNH, American Museum of Natural His-
tory, NY; CMNH, Carnegie Museum of
Natural History, Pittsburgh, PA; USNM,
National Museum of Natural History,
Smithsonian Institution, Washington, DC;
FSCA, Florida State Collection of Arthro-
pods, Gainesville, FL; GPAC, George Poin-
ar Collection of Amber, Department of En-
tomology, Oregon State University, Cor-
vallis, OR.

VOLUME 102, NUMBER 4

Ommatius mariae Scarbrough,
new species
(Figs. 1-9)

Male.—Brown. Body 11.1—12.3 mm;
wing 8.3-8.6 mm. Head: Dull yellowish
gray tomentose; frons, vertex, and narrow
margin of occiput dorsally more yellow;
vestiture largely whitish to pale yellow;
FHWR 1.0:6.3—1.0:6.9. Antenna mostly
brown setose, whitish setae present below
scape; flagellum broadly oval, slightly lon-
ger than wide, and slightly longer than
scape. Frons with margins parallel. Occiput
with 3-5 short, thick, brown postocular
bristles; bristles slightly curved forward,
apices of bristles above eye.

Thorax: Pronotum yellowish gray tomen-
tose with 2 brown and 2 whitish bristles.
Mesonotum with wide median stripe divid-
ed medially by a thin, light yellowish to-
mentose line; 2 lateral spots brown tomen-
tose spots present; tomentum otherwise yel-
low to yellowish gray; setae sparse, limited
to lateral margins; 3 posterior dorsocentral
and 4 lateral stout, dark brown bristles. Scu-
tellum yellowish tomentose with scattered
whitish setae; 2 short, brown, marginal se-
tae sometimes present, each about half as
long as dorsal setae; preapical groove ab-
sent. Pleura dull yellowish gray tomentose
anteriorly, more grayish posteriorly; vesti-
ture whitish; halter yellow.

Wing: Margin anteriorly straight, without
a costal bulge. Cell r4 narrow and long,
sides only slightly divergent apically; base
before apex of cell d. Crossvein r-m just
beyond middle of cell d, slightly shorter
than or as long as vein CuA1+M3. Cell m1
Vass 120:1.9:1-4-1.0:2.7:2.1.

Legs: Coxae brown with grayish tomen-
tum and pale yellow to white vestiture; fore
coxa with numerous stout bristles anterior-
ly. Trochanters brown. Femora mostly yel-
low with yellowish bristles; fore femur
mostly brown anteriorly with a narrow yel-
low band at apical third, and narrow apex
dorsally and posteriorly brown; middle fe-
mur with apical half to two-thirds anteriorly

913

and apical fourth posteriorly brown; hind
femur with apical half to two-thirds ante-
riorly and posteriorly and apical third to
half dorsally and ventrally brown. Middle
femur with 2 anterior, 2 anteroventral, and
1 preapical, dorsoposterior bristles. Hind fe-
mur with 3 anterior and 5 anteroventral
bristles, all long and yellowish, most stout;
anteroventral bristles at middle two-thirds;
1 posteroventral bristle basally plus 6 bris-
tles on a raised posterior tubercle at middle
of hind femur; all posteroventral bristles
quite thick from base to apex, only slightly
flattened with round apices, most or all
brown; HFWLR 1/2.8—1/3.0. Tibiae mostly
yellow with yellowish bristles laterally and
brown bristles apically; narrow apices and
anterior surfaces of all tibiae brown; fore
tibia with only narrow brown stripe anteri-
orly. Hind tibia short, two-thirds as long as
hind femora and four-fifths as long as hind
tarsus; thick, constricted preapically, apex
abruptly wide, clublike. Basal tarsomere of
fore and mid tarsi yellow with narrow apex
brown; basal tarsomere of hind tarsus yel-
lowish basally grading to brownish yellow
apically; apical 4 tarsomeres of all tarsi
brown; fore and middle tarsi with 5 and 4
yellow bristles respectively; hind tarsus
with 2 yellow bristles.

Abdomen: Yellow brown basally, brown
apically; mostly yellowish gray tomentum
and yellowish setae present; terga brown to-
mentose with brown setae dorsally.

Terminalia (Figs. 1-5): Epandrium with
a short, asymmetrical, apical hook; apical
third with abundant short, bristly setae.
Ventral lamella with a short, sclerotized,
preapical, tongue-like process, sides slight-
ly curved downward; dense, long, yellow
setae present. Gonostylus with prominent
wide base and a short, narrow dorsal pro-
cess. Aedeagus with broad hooded sheath,
distiphallus strongly arched downward with
only apex exposed. Gonocoxite with long,
thin, erect flanges, one along inner margin
and another more lateral a long, horizontal
process with pointed apex present. Hypan-

914 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

2D

if il

0.5 mm

1 0.5 mm D)

ee as

3 0.5 mm

0.3 mm

SD

Ommatius mariae, male terminalia. 1, Dorsal view. 2, Lateral view. 3, Ventral view. 4, Gonostylus.

Figs. 1—S.
5, Aedeagus, lateral view.

VOLUME 102, NUMBER 4

8 0.2 mm

Figs. 6-9.
fork.

drium strongly produced apically, apical
margin subtruncate, with rounded corners.

Female.—As male, differing as follows:
Body 13.6 mm; wing 10.0 mm; FHWR 1/
6.9; cell ml WR 1.0:2.1:1.4; HFWLR 1/
5.6. Head: Face with 2 brownish yellow to
brownish bristles. Occiput with 1-2 brown

915

9 0.2 mm

Ommatius mariae, female terminalia. 6, Dorsal view. 7, Ventral view. 8, Spermatheca. 9, Genital

postocular bristles. Thorax: Tomentum
largely dull yellowish. Wing: Crossvein r-
m longer than vein CuAl+Ms3. Base of cell
r4 at apex of cell d. Legs: Fore coxa with
several, long, thin bristles. Middle and hind
femora black on apical half or slightly less
and with all or most anterior bristles black.

916

Hind femur slender, not especially swollen,
without a mound-like tubercle, and bristles
acutely pointed; all ventral bristles yellow
and distributed in well defined rows. Tibiae
with narrow apex and anterior surface
black; hind tibia black except yellow on
basal half anteriorly. Hind tibia slender,
only gradually wider apically, apex not un-
usually wide or flat. Tarsi largely or entirely
black with black bristles; fore and middle
tarsi with basal tarsomere mostly yellow,
narrow apex black; only 2—3 yellow bristles
present.

Abdomen: Apical margin of tergite 7
with 2 midlateral black bristles, 1 of these
contrastingly long and stout. Tergite 8
shiny, jet black, apical corner somewhat an-
gular, oblique; a long, midlateral, black
bristles present posterior to apical margin
and 2-3 shorter bristles more basally, these
black or yellowish. Tergite 9 dorsally with
narrow base sclerotized, less than one-fifth
as long as cercus, corners contrastingly
wide, extending around base of ventral la-
mella forming a wide sclerotize bridge in
dry specimens. Cercus unusually narrow
with apex slightly emarginate. Ventral la-
mella narrowly sclerotized along inner mar-
gin. Sternites 5—7 with 3—4 long, stout bris-
tles laterally, mostly yellowish, those in api-
cal corner of sternite 7 black and contrast-
ingly long. Sternite 8 shiny jet black with
a transverse row of yellow setae and bris-
tles; 4 bristles medially, darker, light amber,
contrastingly long and thick, extending for-
ward to subapex of ventral lamella. Sper-
mathecae obovate; ducts long and thin with
bases of ducts separate, each originating
from very short, common duct between
base of arms of genital fork. Genital fork
largely membranous, apical third of each
arm, narrow, slightly sclerotized.

Types.—Holotype 6, BAHAMAS:
Eleuthera, Rainbow Bay, 21.x.85, J. R. Wi-
ley (FSCA). Allotype 2, same data as ho-
lotype except: 29.vi-6.vii.1990, J. R. & S.
C. Wiley (FSCA). Paratype: 1 6, same data
as holotype (FSCA); 1 6, BWI, Nassau,
3.xii.61, N. A. Roeff (USNM).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Etymology.—Latin mariae, for Mary, in
memory of a loving, devoted wife and
mother.

Remarks.—Ommatius mariae and O. ha-
nebrinki (holotype and allotype, USNM;
paratypes, AMNH, TUMZ; others CMNH/
FSCA) are very similar species, almost
identical in general habitatus, and differ
markedly from other known New World
species. They are readily separated from
each other by the uniquely modified struc-
tures of the terminalia. In addition, the
whitish ocellar setae and the flat, mostly
brown bristles with round tips, clustered on
a raised, posterior tubercle on the hind fem-
ora further characterize the male. In O. ha-
nebrinki, these bristles are flatter, yellow,
and distributed more in a linear row on a
lower tubercle. The anterior surface of all
tibiae in the female of O. mariae is mostly
black whereas the tibiae are yellow with
narrow dark apices in O. hanebrinki. The
basal tarsomere of the hind tarsus is black
in O. mariae whereas it is much lighter,
brownish yellow in O. hanebrinki. Further-
more, in O. mariae segment 8 is shiny, jet
black; tergite 8 has scattered bristly setae
dorsally and stout bristles laterally, none are
present along the apical margin. Sternite 8
medially has a transverse row of numerous,
short setae and four contrastingly long bris-
tles. In O. hanebrinki, segment 8 is much
lighter brown; tergite 8 has mixed bristly
and abundant, short, recumbent setae dor-
sally, and several, stout bristles are present
along the apical margin. The transverse row
of vestiture on sternite 8 medially consists
of 8—10 stout bristles and sparse, short se-
tae.

Ommatius hanebrinki (1983), O. hispidus
Scarbrough (1985a), and O. mariae are
similar in several regards, and differ from
all known species from Cuba and the Ba-
hamas. This group, i.e., hanebrinki group,
is readily recognized by the following com-
bination of characters of the male: 1)
Strongly inflated hind femur; 2) a series of
flat bristles with rounded tips on the pos-
terior margin of the hind femur; 3) an api-

VOLUME 102, NUMBER 4

cally clubbed [hanebrinki and mariae] or
long digitate process [hispidus] on the hind
tibia; 4) the unusually long, blade-like pro-
cesses of the gonocoxite in males; 5) the
unusually stout bristles on the apical ab-
dominal segments; and 6) the dorsally
membranous tergite 9 which wraps below
the bases of the ventral lamella, and the un-
usually strongly, apically pointed sternite 8
with stout bristles posteriorly in the female.

Ommatius lineolatus Scarbrough

Ommatius lineolatus Scarbrough 1988: 90—
94. 5 Holotype, 2 allotype. Type locality
Cuba (USNM).

Specimens examined.—BAHAMAS: 1
2 of O. lineolatus, Bimini (FSCA); 1 6 of
O. lineolatus, Long Island (FSCA). CUBA:
Holotype 6 and 2 of O. lineolatus Scar-
brough (USNM).

Remarks.—Ommatius lineolatus, a Cu-
ban species, is reported here from Long Is-
land in the Bahama Islands and increases
the number of species of Ommatius to 6
from this region. The species is readily rec-
ognized by its small size (S—6 mm), yellow-
ish brown color of the legs, 6 ventral setae
distributed the length of the antennal style,
and combined characters of the terminalia.
It is otherwise quite similar to O. abana
(types series, AMNH), and may prove
eventually to be that species as more ma-
terial is located and studied. This type series
of O. abana consist of only 3 females, i.e.,
males are unknown. The species is large (9
mm), the body is jet black, the legs are am-
ber or reddish with black markings, and the
style has only four long setae near the tip.

Ommatius dignus Scarbrough,
new name

Ommatius dimidiatus Scarbrough 1985b:
647-650. Figs. 9-14 (primary junior
homonym of Ommatius dimidiatus Mac-
quart 1850: 394, Asilidae); (types
GPAC).

The binomen Ommatius dimidiatus Mac-
quart (Macquart 1850) was first used for a

Ov

species in the Australian region. It is now
reported from Tasmania and Australia
(Daniels 1989). Later (Scarbrough 1985b),
the same binomen was applied to a new
Neotropical species from Dominica, Lucia,
and Martinique in the Lesser Antilles. I pro-
pose dignus (L, meaning worthy or fit) as
a new replacement name.

Ommatius fimbrillus Scarbrough,
new name

Ommatius fimbriatus Scarbrough and Poin-
ar 1992: 13-16. Figs 1—4 (primary junior
homonym of Ommatius fimbriatus Hardy
1949: 301, Asilidae); (types GPAC).

Unfortunately, Scarbrough and Poinar
(1992) were unaware that the binomen Om-
matius fimbriatus had already been used for
a species of an asilid fly from Australia
when they used it for a fossil species in
Dominican amber. I propose fimbrillus (L,
meaning a fringe) as a new replacement
name for it.

ACKNOWLEDGMENTS

My sincere thanks to David Grimaldi,
AMNH, New York; Chen Young, CMNH,
Pittsburgh; E Christian Thompson, System-
atic Entomology Laboratory, USDA, Wash-
ington, DC; and Gary Stark, FSCA, Gaines-
ville, FL, for the loan of specimens. I also
thank FE Christian Thompson and Gary
Steck for their help and cooperation during
my visit to examine specimens; and to Drs.
William Grogan, Department of Biological
Sciences, Salisbury State University, Salis-
bury, MD, and E Geller-Grimm, Museum
Wiesbaden (MWNH), Department of Nat-
ural Sciences, Germany, for their invaluable
reviews of this manuscript.

LITERATURE CITED

Curran, C. H. 1953. The Asilidae and Mydaidae of the
Bimini Islands, Bahamas, British West Indies (Dip-
tera). American Museum Novitates 1644: 5—6.

Daniels, G. 1989. Family Asilidae, pp. 326-349. In
Evenhuis, N. L., ed., Catalog of the Diptera of the
Australasian and Oceanian Regions. Bishop Mu-
seum Press & E. J. Brill, 1,155 pp.

Hardy, G. H. H. 1949. Miscellanous notes on Austra-

918 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

lian Diptera. XIV. Venation and other notes. Pro-
ceedings of the Linnean Society of New South
Wales 73(1948): 298-303.

Macquart, P. J. M. 1850. Diptéres exotiques nouveaux
ou peu connus. 4. Supplément. - Mémoires de la
Société Royale des Sciences, de |’Agriculture et
des Artes (1849): 309—479, 14 plates. (365—400,
plates. 6—9). Lille.

McAlpine, J. E 1981. Morphology and terminology—
Adults, pp. 9-63. In McAlpine, J. FE et al., Manual
of Nearctic Diptera, Vol. 1 Ottawa. Monograph
No. 28, 674 pp.

Scarbrough, A. G. 1985a. New Ommatius Wiedemann
(Diptera: Asilidae) from Cuba and the Bahamas.
Journal of the New York Entomological Society
93: 1226-1239.

. 1985b. Ommatius (Diptera: Asilidae) in the

lesser Antilles. Proceedings of the Entomological

Society of Washington 87(3): 641—655.

. 1988. New robber flies (Diptera: Asilidae)

from Mayaguana Island, Bahamas, and Cuba. En-

tomological News 99(2): 90—94.

. 1997. New and old species of Ommatius Wie-
demann (Diptera: Asilidae) from Hispaniola. In-
secta Mundi 11: 9—24.

Scarbrough, A. G. and R. Rutkauskas. 1983. A new
species of Ommatius Wiedemann (Diptera: Asili-
dae) from San Salvador, the Bahamas. Proceed-
ings of the Entomological Society of Washington
85: 144-151.

Scarbrough, A. G. and George O. Poinar. 1992. Upper
Eocene robber flies of the genus Ommatius (Dip-
tera: Asilidae) in Dominican Amber Insecta Mun-
di. 6: 13-18.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 919-923

A NEW SPECIES OF CHLORONIA BANKS (MEGALOPTERA:
CORYDALIDAE) FROM SOUTHEASTERN BRAZIL,
WITH A KEY TO THE SPECIES OF BRAZIL

ATILANO CONTRERAS-RAMOS

Corregidora 460-7, Amp. Miguel Hidalgo, 14410 Tlalpan, México, D.F, México
(e-mail: acontreras_ramos @ hotmail.com)

Abstract.—The Neotropical dobsonfly genus Chloronia (Megaloptera: Corydalidae:
Corydalinae) contains 14 previously described species. In this paper, Chloronia pennyi,
a new species from Minas Gerais, Brazil, is described and illustrated. In general appear-
ance C. pennyi resembles most the Andean C. bogotana Weele, especially in the small
fuscous markings on the head and the pair of fuscous bands on the pronotum. In the new
species, the antennae are entirely yellow (apically infuscate in C. bogotana), the wings
are mostly pale (patterned in C. bogotana), and the dark bands on the pronotum are
continuous (nearly meeting in middle in C. bogotana). A key for adult males of the four
currently recognized Brazilian species is included. New distribution records are given for
C. corripiens (Walker) in southeastern Brazil.

Key Words:
sonfly, taxonomy, key

The Neotropical genus Chloronia is dis-
tributed from northwestern and northeastern
Mexico, southward into western and south-
eastern South America, including some of
the Lesser Antilles. Individuals of this ge-
nus are rather small dobsonflies (forewing
length 24—50 mm) with a characteristic col-
or (yellow with black spots), which allows
for easy recognition. After the revision by
Penny and Flint (1982), Flint (1991) clari-
fied the identity of Chloronia bogotana
Weele and added two new species from
Costa Rica (Flint 1992), and Contreras-Ra-
mos (1995) described two new species each
from Ecuador and Guatemala. In all, 15
species of Chloronia, including the one de-
scribed in this paper, are currently recog-
nized. Larvae of Chloronia have been di-
agnosed by Penny and Flint (1982) and
Contreras-Ramos and Harris (1998). Ac-

Chloronia, new species, Megaloptera, Brazil, Neotropics, Corydalidae, dob-

counts on habitat of a few species have
been given by Penny and Flint (1982), Ge-
ijskes (1984), and Contreras-Ramos (1999).

In this paper, a new species of Chloronia
from southeastern Brazil is described and
illustrated. In 1998, five specimens of the
new species were collected in two localities
in Minas Gerais during caddisfly survey
work by Ralph W. Holzenthal (UMSP) and
colleagues. General similarity of the distinct
Brazilian series with the distantly distrib-
uted C. bogotana (Andes of Bolivia, Co-
lombia, Ecuador, and Peru). indicated high
possibilities for the Brazilian specimens to
belong in a new species. This was corrob-
orated after careful examination of the male
genitalia of the Brazilian series. Additional
distributional records for C. corripiens
(Walker) in southeastern Brazil, and a key
for the identification of the four Chloronia

Rig e

Habitus of Chloronia pennyi, holotype.

species currently recorded in Brazil also are
presented in this paper.

The genitalic and venational terminology
used here follows that of Glorioso (1981),
as modified by Contreras-Ramos (1998).
Specimens were originally preserved in
80% ethanol. Three males were spread and
pinned for dry preservation. Genitalia were
dissected, cleared, and stored by standard
methods, as explained by Contreras-Ramos
(1998). Specimens will be deposited at the
entomological collections of the Museu de
Zoologia, Universidad de Sao Paulo, Brazil
(MZSP); Department of Entomology, Uni-
versity of Minnesota, St. Paul (UMSP); and
the National Museum of Natural History,
Smithsonian Institution, Washington, DC
(NMNH).

Chloronia pennyi Contreras-Ramos,
new species
(Figs. 1—6)
Diagnosis.—This species, together with
Chloronia corripiens (Walker) and C. plau-
manni Penny and Flint, appears to have a

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

phylogenetically basal position in the ge-
nus. All three have an unmodified ninth
sternite, a sparsely setose ninth tergum
(lacking small clusters of spinous setae pre-
sent in most species), as well as conspicu-
ous dorsolateral pregenital sacs between the
eighth and ninth abdominal segments.
However, on the basis of general appear-
ance, the new species is most similar to C.
bogotana. Both have small posterolateral
spots on the head (Flint 1991, figs. 5, 6)
and elongate dark spots on the pronotum.
In C. bogotana, nevertheless, the pronotal
spots are discontinuous, not meeting in the
middle (Flint 1991, figs. 5, 8), whereas in
the new species they form a pair of contin-
uous lateral bands (Fig. 6). Forewing col-
oration in C. bogotana is patterned, with
most crossveins dark (Flint 1991, figs. 7—
9). In the new species, forewings are mostly
clear with few crossveins slightly dark (Fig.
1). With respect to genitalic characters, the
10th sternite of C. pennyi resembles that of
C. convergens Contreras-Ramos. In both,
10th sternite lobes are sclerotized and su-

Figs. 2-6.
eral. 6, Pronotum, dorsal.

bquadrate, but in the new species lobes lack
apical spines and the 10th sternite has only
slightly developed anterolateral projections
(Fig. 4).

Description of adult (Figs. 1, 5—6).—
Head width, ¢ 3.6—4.1 mm (average 3.8
mm, n = 3), 2 4.1-4.3 mm (average 4.2
mm, n = 2); forewing length, 6 27.4—29.4
mm (average 28.3 mm, n = 3), 2 34.3-

ee . ae

25
Senet

7

Chloronia pennyi. 2, Male genitalia, dorsal. 3, Same, ventral. 4, 10th sternite. 5, Head, dorsolat-

34.7 mm (average 34.5 mm, n = 2). Color
generally pale yellow with fuscous spots
and bands. Head pale yellow, mandible yel-
lowish brown with teeth and outer side dark
reddish-brown. Labrum subquadrate with
pair of long, flattened setae. Clypeal margin
nearly straight. Compound eyes and base of
ocelli dark. Postocular spine blunt, colored
as head. Pattern of fuscous wide-elongate

922

spot at postocular plane, thin spot at adja-
cent ridge, pair of posterior spots, and
round spot at occiput, on each side of head
(Fig. 5). Antenna 37 to 44-segmented, fili-
form, yellow, with at most last segment in-
fuscate. Maxilla yellow with 5-segmented
palp, last two segments pale brown. Labial
palp 4-segmented, yellow.

Pronotum yellow, with pair of dark, con-
tinuous, longitudinal bands (Fig. 6). Meson-
otum without fuscous spots. Legs yellow,
tarsal claws brown. Forewing pale yellow,
hyaline, with 26—27 costal crossveins, 0—2
forked, a few in proximal % of wing with
dark ends. Veins mostly yellow, R,-Rs
crossveins, forking of M, and few basal
crossveins finely fuscous. Posterior margin
of wing with subtle grayish maculations.
Anterior 2A cell variably with a spot to
only a slight maculation. Hindwing pale
yellow, hyaline, but 2nd r brown.

Male genitalia (Figs. 2—4): Ninth tergum
subtriangular, finely and sparsely setose,
without patches of spinous setae; V-shaped
internal inflection reaching midlength of
tergum. Tenth tergites about 2.5 times as
long as ninth tergum, subcylindrical, blunt-
ly tapering, basal 7, divergent, apical /, di-
rected posteriorly, finely and evenly setose
(Fig. 2). Ninth gonostylus incurved, fusi-
form, anteroventral margin slightly more
convex, with sharp apical point. Ninth ster-
num moderately sclerotized, subquadrate,
with well developed posterolateral lobes,
slightly convex posteromesally (Fig. 3).
Membrane between 9th and 10th sternites
eversible, broadly bilobate, thickened.
Tenth sternite convex, with small, sharply
pointed anterolateral projections; lobes
sclerotized, papilliform, subequal in width
and length, sparsely and conspicuously se-
tose (Fig. 4).

Material examined.—Holotype 3: BRA-
ZIL. Minas Gerais: Serra do Cip6, Rio Cip6
in Cardeal Mota (Cach. Baixo), 19°20.553’S,
43°38.531'W, el. 750 m, 10—15.ii.1998, Hol-
zenthal, Paprocki, Huisman [head width =
3.6 mm, forewing length = 28.0 mm]
(MZSP).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Paratypes: Same data as holotype, 1 @
(MZSP), 1 do, 1 2 (UMSP); Minas Gerais:
confluence Rio Peixe & Rio Preto do Itam-
bé,, 19°17.525'S, 43°15.457'W,, ely S00mm:
4.11.1998, Holzenthal & Paprocki, 1 ¢d
(NMNH).

Etymology.—This new species is gladly
dedicated to Norman D. Penny from the
California Academy of Sciences, in recog-
nition of his extensive contributions to Neo-
tropical neuropterology, as well as ac-
knowledging his support for the author’s
graduate research projects.

KEY TO KNOWN BRAZILIAN SPECIES OF
CHLORONIA
(MODIFIED FROM PENNY AND FLINT 1982)

1. Head with pair or few small fuscous spots pos-
teriorly
— Head unicolorous or only with lateral margin
INfUSCAIE: 2 sc)y5 a8 Yolo aca ee 2
2. Lateral margin of head fuscous (Penny and
Flint, 1982. fig. 44). ates Se
C. plaumanni Penny and Flint
— Lateral margin unmarked (Penny and Flint
OS Dh oe AD) eee gta C. corripiens (Walker)
3. Pronotum with two anterior and two posterior
fuscous spots, wings patterned (Penny and
Flint 1982, fig. 48) .. C. hieroglyphica (Rambur)
— Pronotum with two fuscous, longitudinal bands
(Fig. 6), wings mostly clear (Fig. 1) .....
bo, Sasa tY ans eg USS C. pennyi, new species

ADDITIONAL MATERIAL EXAMINED

Chloronia corripiens (Walker).—BRA-
ZIL. Minas Gerais: Serra do Cip6, Cardeal
Mota, Cachoeira Veu da Noiva, 19°18.912'S,
43°36.260'W, el. 800 m, 12.11.1998, Holzen-
thal & Paprocki, 1 ¢6 (UMSP); Parana: Rio
Mae Catira, 10 km N Porto de Cima,
25°21.821'S,. 48°52.473'W, el. 200iimies—
9.xii.1997, Holzenthal & Huisman, 3 °&
(UMSP).

ACKNOWLEDGMENTS

Thanks to Ralph W. Holzenthal (UMSP)
for calling my attention to a series of dob-
sonfly specimens collected through his cad-
disfly survey work in Brazil, among which
the new Chloronia species was found.
Thanks also to Philip J. Clausen for curat-

VOLUME 102, NUMBER 4

ing dobsonflies (UMSP) and processing a
loan to me. A brief research visit to the
University of Minnesota was funded by the
Instituto de Biologia of the Universidad Na-
cional Aut6noma de México. The hospital-
ity of Fernando Mufioz-Quesada (UMSP)
and family is greatly appreciated. Finally,
thanks to David E. Bowles (Texas Parks
and Wildlife Department) and an anony-
mous reviewer for improving the quality of
the manuscript.

LITERATURE CITED

Contreras-Ramos, A. 1995. New species of Chloronia
from Ecuador and Guatemala, with a key to the
species in the genus (Megaloptera: Corydalidae).
Journal of the North American Benthological So-
ciety 14: 108-114.

1998. Systematics of the dobsonfly genus

Corydalus (Megaloptera: Corydalidae). Thomas

Say Publications in Entomology: Monographs.

Entomological Society of America, Lanham,

Maryland, 360 pp.

1999. Mating behavior of Platyneuromus

(Megaloptera: Corydalidae), with life

history

923)

notes on dobsonflies from Mexico and Costa Rica.
Entomological News 110: 125-135.

Contreras-Ramos, A., and S. C. Harris. 1998. The im-
mature stages of Platyneuromus (Corydalidae),
with a key to the genera of larval Megaloptera of
Mexico. Journal of the North American Bentho-
logical Society 17: 489-517.

Flint, O. S., Jr. 1991. On the identity of Chloronia
bogatana [sic] Weele (Neuropterida: Megaloptera:
Corydalidae). Proceedings of the Entomological
Society of Washington 93: 489-494,

1992. A review of the genus Chloronia in Cos-
ta Rica, with the description of two new species
(Neuropterida: Megaloptera: Corydalidae). Pro-
ceedings of the Biological Society of Washington
105: 801-809.

Geijskes, D. C. 1984. Notes on Megaloptera from the
Guyanas, S. Am., pp. 79-84. In Gepp, J., H. As-
pock, and H. Holzel, eds., Progress in World’s
Neuropterology; Proceedings of the Ist Interna-
tional Symposium on Neuropterology. Graz, Aus-
tria.

Glorioso, M. J. 1981. Systematics of the dobsonfly
subfamily Corydalinae (Megaloptera: Corydali-
dae). Systematic Entomology 6: 253-290.

Penny, N. D., and O. S. Flint, Jr. 1982. A revision of
the genus Chloronia (Neuroptera: Corydalidae).
Smithsonian Contributions to Zoology 348: 1—27.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 924-928

A NEW AFROTROPICAL SPECIES OF ALLOGNOSTA OSTEN SACKEN
(DIPTERA: STRATIOMYIDAE)

NORMAN E. WOODLEY

Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture, % National Museum of Natural History, Smithsonian Institution,
Washington, DC 20560-0168, U.S.A. (e-mail: nwoodley @sel.barc.usda.gov)

Abstract.—Allognosta njombe, new species, is described from localities around the
north end of Lake Nyassa in Tanzania (type locality) and Malawi. A key to the four
known species of Afrotropical Allognosta is provided. Additional locality records are
given for A. stuckenbergae Lindner and A. tessmanni Enderlein.

Key Words: Beridinae, Tanzania, Malawi

The genus Allognosta Osten Sacken is
the only genus of the stratiomyid subfamily
Beridinae in the Afrotropical Region. When
I reviewed the Afrotropical species (Wood-
ley 1987) I had available only 12 specimens
representing three species, although about
seven additional specimens of A. stucken-
bergae Lindner were known at that time. A
small number of additional specimens have
come to light, including some of a new spe-
cies, that are reported upon in this paper.
Since my review of the African species, I
have published a revision of the world gen-
era of Beridinae (Woodley 1995) which can
be consulted for general information and
cladistic relationships at the generic level
within the Beridinae as well as an overview
of the genus Allognosta.

Terminology and methodology follow
that used in my previous paper (Woodley
1987) except that the aedeagal complex is
now called the phallic complex (Sinclair et
al. 1994). Specimens studied are from the
Carnegie Museum of Natural History, Pitts-
burgh, Pennsylvania, USA (CMNH); Natal
Museum, Pietermaritzburg, Natal, South
Africa (NMP); and Naturhistoriska riks-
museet, Stockholm, Sweden (NRS). The
sections on material examined under al-

ready described species treat only speci-
mens seen since my 1987 paper.

KEY TO AFROTROPICAL SPECIES OF
ALLOGNOSTA

1. Pilosity of eyes long and dense, easily visible

at low magnifications; occurring only in South
AtiTiCa wes Sane one A. stuckenbergae Lindner

— Pilosity of eyes short and sparse, visible only

at higher magnifications; occurring north of
South Africa
. Apical one-third to one-fourth of middle and
hind femora darkened, strongly contrasting
with whitish-yellow basal portions; second
segment of palpus yellow

Nw

— Middle and hind femora yellowish, without
strongly contrasting darker areas; second seg-
ment of palpus black

3. Female frons strongly produced, the antennae
inserted below the prominence; male with me-
dian lobe of phallic complex similar in diam-
eter and equal in length to lateral lobes; pleura
wholly yellowish, only vaguely darker beneath
wing base A. bwamba Woodley

— Female frons moderately produced (Fig. 2), the
antennae inserted at the apex of the promi-
nence; male with median lobe of phallic com-
plex in dorsal view (Fig. 5) more slender and
shorter than lateral lobes; pleura usually
brownish to brownish-black, nearly concolo-
rous with scutum, but sometimes paler .....
Seas EEN Aare Ae Se eee A. njombe, n. sp.

VOLUME 102, NUMBER 4

9,

y

\

v

\
yh

i
1 loner

>

Figs. 1-2.

Allognosta njombe Woodley, new species
(Figs. 1-7)

Diagnosis.—This species is in a group
easily distinguished from A. stuckenbergae
by having nearly bare eyes. It differs from
A. tessmanni Enderlein in having all femora
wholly dark yellow. It is most similar to A.
bwamba Woodley, but differs in having
darker pleura, a less strongly produced
frons in females, and different male geni-
talia.

Description.—Male: Head (Fig. 1)
black, 1.5 time higher than long; eye large,
upper ommatidia larger than lower ones,
but size transition not sharply delimited;
ocellar tubercle moderately prominent; face
short, receding; frons very small, slightly
convex; head mostly whitish-gray tomen-

O25

Ly Meth.
wee Dp,

{

SSS
Nash

—

SS

A=
\\ \
Mines

/

==
/
hg

LZ

EZ Wy;
icp fi fl) yy
pe

NO

Left lateral views of heads of Allognosta njombe. 1, Male. 2, Female.

tose, sparse on frons, face, and upper oc-
ciput, almost absent on ocellar tubercle; pi-
losity sparse, pale on lower occiput and
gena, more brownish on face; eye with ex-
tremely short, sparse pilosity, visible only
at high magnification; antenna 0.8 length of
head, ratio of segments 4:4:[5:2:3:3:3:3:2.5:
3.5], scape and pedicel brownish-yellow,
flagellomeres 1—3 yellow, those beyond
three brownish; hairs on scape and pedicel
moderately short, dark; first flagellomere
with usual subapical dark hairs, other fla-
gellomeres with a few scattered hairs; pal-
pus slender, the two segments subequal in
length, the second slightly clavate; both pal-
pal segments dark brownish with pilosity
mostly pale, but with a few dark hairs at
apex of second segment; proboscis dark

926

brownish. Thorax with scutum and scutel-
lum blackish-brown with slight bronzy re-
flections, postpronotal lobe slightly paler,
postalar callus dark yellowish; pleura dark
brownish, but posterior half of anepister-
num and katepisternum and entire anepi-
meron orangish-brown; pleura with pale to-
mentum but with noticeable bare area on
ventral half of katepisternum; scutum and
scutellum entirely, densely pilose with
short, semi-erect brownish hairs, posterior
part of scutum and scutellum with some
longer, more erect hairs intermixed; pleura
with pale pilosity except anterior half of
anepisternum and entire meron and katepi-
meron apilose; coxae dark yellow, anterior
pair slightly suffused with brown; femora
dark yellowish, posterior pair a little more
brownish; tibiae brownish; tarsi dark
brownish, basal portions of first tarsomeres
vaguely more yellowish; pilosity on legs
mostly dark, some pale hairs intermixed on
femora, entirely pale on coxae, scattered
longer, erect hairs present on femora and
tibiae; wing moderately infuscated with
grayish-brown, pterostigma and veins
brownish, entire wing set with dense micro-
trichia; halter dark brownish, basal portion
of stem paler. Abdomen brownish, tergal
grooves slightly darker; entire abdomen
with sparse tomentum; pilosity of tergites
mostly very short, dark, but longer laterally
and basally, some hairs on first tergite pale,
pilosity of sternites pale; terminalia with
gonocoxites (Fig. 3) moderate in size, mar-
gins diverging posteriorly; apex of hypan-
drium moderately produced, the apex of the
process feebly bilobed; gonostylus with lat-
eral margin slightly arcuate, otherwise sim-
ple; phallic complex (Figs. 5—6) arcuate in
profile, median lobe shorter and more slen-
der than lateral lobes; epandrium (Fig. 4)
small, unmodified. Length 5.3 mm.
Female: Differs from male as follows:
Head (Fig. 2) 1.2-1.3 times higher than
long, strongly dichoptic, eye with ommatid-
ia uniform in size; frons 0.4—0.5 width of
head at ocellar tubercle, produced at lower
portion to meet upper facial margin, the an-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tennae being inserted at anterior-most point
of head in profile; upper frons with margins
very slightly diverging ventrally, lower part
appearing vaguely depressed, overall shiny
and finely punctate; lower frons very short,
slightly convex; face short, strongly reced-
ing; upper occiput margins produced pos-
terodorsally with sharply rounded margins;
upper frons and upper occipital margins
without tomentum, lower frons and face
with dense, pale grayish tomentum; upper
frons and occipital margins with moderately
dense pale golden pilosity; palpus more ro-
bust than in male, second segment more
strongly clavate. Thorax with scutum and
scutellum as in male but with very slight
metallic purplish reflections; pilosity of scu-
tum and scutellum shorter than in male,
uniform in length, entirely pale golden col-
ored; pleura brownish to brownish-black,
vaguely paler below wing (uniformly paler
in specimen from Malawi; see remarks be-
low); hind coxa sometimes darkened; all
femora uniformly dark yellowish; tibiae
brownish-black, approximately basal one-
third of fore and middle femora yellow,
hind femur only vaguely pale at extreme
base; tarsi uniformly brownish-black. Ab-
domen with tergites with vague purplish re-
flections; sternites slightly paler in color
than tergites; furca (Fig. 7) with median ap-
erture large, emarginate anteriorly; postero-
medial margin triangularly emarginate me-
dially; spermathecal ducts unsclerotized
posteriorly; cerci short, second segment
about 0.7 length of first. Length 5.6—6.0
mm.

Type material.—Holotype ¢ (NRS) from
Tanzania is labeled: “S TANGANYIKA
Melando Forest 30 mi S of Njombe 2,450
m. 10.1962 leg. G. Heinrich(HOLOTY PE
3 Allognosta njombe N. E. Woodley
1999.”” The locality of collection is in Ir-
inga Province in Tanzania. The specimen is
missing the right front tarsus and the left
middle tibia and tarsus, otherwise it is in
excellent condition. The terminalia are
stored in glycerin in a microvial on the
specimen pin. Allotype 2, 3 2 paratypes

VOLUME 102, NUMBER 4

927

Figs. 3-7.
and postgenital segments, dorsal view. 5, Phallic complex, dorsal view. 6, Phallic complex, left lateral view. 7,
Female furca, ventral view. Abbreviations: ep, epandrium; gc, gonocoxites; gs, gonostylus; hyp, hypandrium;
llb, lateral lobe of phallic complex; ma, median aperture of female furca; m/b, median lobe of phallic complex;
pmar, posteromedial margin of female furca.

(NRS, USNM): Tanzania: Mbeya Province,
Rungve Mt., 2,600 m, 8—11.1962, G. Hein-
rich. 3 2 paratypes (NRS): Tanzania:
Mbeya Province, Rungve Mt., 20 mi. SSE
Mboya [probably = Mbeya], 2,600 m,
1962, G. Heinrich. 1 2 paratype (CMNH):
Malawi: Chitipa District, Jembya Reserve,
18 km SSE Chisenga, 10-08S, 33-27E,
1,870 m, 1-10 January 1989, J. Rawlins, S.
Thompson.

Distribution.—Known only from high-

Male and female terminalia of Allognosta njombe. 3, Genital capsule, dorsal view. 4, Epandrium

land localities in Malawi and Tanzania
around the north end of Lake Nyassa.

Etymology.—The species name is a noun
in apposition based on the name of the type
locality.

Remarks.—This species is most similar
to Allognosta bwamba Woodley. The elon-
gation of the anterior region of the head in
females is similar in the two species, al-
though in A. njombe the antennae are in-
serted at the apex of the anterior most por-

928

tion of the head, not below it. The overall
structure of the male terminalia is similar in
the two species, especially the shape of the
posterior margin of the hypandrium. Al-
though no phylogenetic analysis has been
done on Allognosta, it seems probable that
these two species are closely related. None
of the other Afrotropical species of Allog-
nosta have been collected in the vicinity of
Lake Nyassa.

Allognosta stuckenbergae Lindner

Allognosta stuckenbergae Lindner 1961: 1.

Material examined.—mSOUTH AFRICA:
1 6, 1 &, Natal, Pietermaritzburg, Town
Bush, xi.1976, R. Miller (NMP); 1 6,
Transvaal, 12 km. S Sabi, 2530BB, indig-
enous bush, 3.xii.1976, R. Miller (NMP); 1
2, Cape Province, Hogsback, 3226DB, for-
est and forest margins, 13—16.xu1.1985, J. &
B. Londt (NMP); 2 2, Cape Province, Tsit-
sikama Coastal National Park, Stormsriv-
ermond, 34°02'S, 23°53’E, moist medium
high coastal forest with Podocarpus, 15—
19.x.1994, Michael Séderlund, Malaise trap
(NRS); 1 6, Cape Province, Tsitsikama
Forest Reserve, 33°58'S, 23°54'E, moist
high indigenous forest with Podocarpus,
14—19.x.1994, Michael Sdderlund, Malaise
trap (NRS); 1 2, Cape Province, Bloukrans
Pass at Varkrivier, 33°57'S, 23°38’E, coastal
rainforest in ravine, 14—19.x.1994, Michael
Séderlund, Malaise trap (NRS).

Remarks.—All of the new locality data
fit within the known range of A. stucken-
bergae in the eastern half of South Africa.

Allognosta tessmanni Enderlein

Allognosta tessmanni Enderlein 1921: 182.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Material examined.—UGANDA/ZAIRE:
12, North of Lake Edward, Gyldenstolpe
(NRS).

Remarks.—This specimen was collected
on an expedition headed by Nils Gylden-
stolpe along the northern parts of Lake Ed-
ward from 4 April to 6 May 1921 (Thomas
Pape, personal communication).

ACKNOWLEDGMENTS

I thank Chen Young, Carnegie Museum
of Natural History, Pittsburgh, Pennsylva-
nia, USA; David Barraclough, Natal Mu-
seum, Pietermaritzburg, Natal, South Afri-
ca; and Thomas Pape, Naturhistoriska riks-
museet, Stockholm, Sweden, for loaning
specimens on which this paper is based.
Thomas Pape also supplied additional in-
formation on some collection localities.
Stephen D. Gaimari, Smithsonian Institu-
tion and Thomas J. Henry and Allen L. No-
rrbom, both of the Systematic Entomology
Laboratory, provided comments on the
manuscript.

LITERATURE CITED

Enderlein, G. 1921. Uber die phyletisch alteren Stra-
tiomyiidensubfamilien (Xylophaginae, Chiromy-
zinae, Solvinae, Beridinae, und Coenomyiinae).
Mitteilungen aus dem Zoologischen Museum in
Berlin 10: 151-214.

Lindner, E. 1961. Athiopische Stratiomyiiden (Diptera)
V. Stuttgarter Beitrage zur Naturkunde 68: 1—13.

Sinclair, B. J., J. M. Cumming, and D. M. Wood. 1994.
Homology and phylogenetic implications of male
genitalia in Diptera-lower Brachycera. Entomolo-
gica Scandinavica 24: 407—432.

Woodley, N. E. 1987. The Afrotropical Beridinae
(Diptera: Stratiomyidae). Annals of the Natal Mu-
seum 28: 119-131.

. 1995. The genera of Beridinae (Diptera: Stra-

tiomyidae). Memoirs of the Entomological Soci-

ety of Washington No. 16, 231 pp.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 929-956

A NEW GENUS OF THE OROBITIDINAE AND DISCUSSION OF ITS
RELATIONSHIPS (COLEOPTERA: CURCULIONIDAE)

Boris A. KOROTYAEV, ALEXANDER S. KONSTANTINOV, AND CHARLES W. O’ BRIEN

(BAK) Zoological Institute, Russian Academy of Sciences, 199034 St. Petersburg, Rus-
sia; (ASK) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S.
Department of Agriculture, c/o National Museum of Natural History, Smithsonian Insti-
tution, Washington, DC 20560-0168, U.S.A. (e-mail: akonstan @sel.barc.usda.gov);
(CWO) Center for Biological Control, Florida A&M University, Tallahassee, FL 32307-
4100, U.S.A.

Abstract.—The new genus Parorobitis of the subfamily Orobitidinae and two new
species, Parorobitis gibbus (Paraguay: Alto Parana) and Parorobitis minutus (Brazil:
Curitiba, Parana), are described. The subfamily Orobitidinae is redescribed based on char-
acters of Orobitis and Parorobitis, and diagnoses of the two genera are provided. Several
characteristics of the head, thorax, wings and elytra are used to compare the Orobitidinae
with the supposedly related taxa of Zygopinae, Ceutorhynchinae, and Baridinae. A key

to these taxa is provided.

Key Words:

The subfamily Orobitidinae is comprised
of two strikingly distinct genera. The Pa-
learctic Orobitis Germar includes the trans-
palearctic O. cyaneus Linnaeus which de-
velops in seed capsules of Viola spp. in
broad-leaved and mixed forests and O. ni-
grinus Reitter from the Balkans. A new ge-
nus of this subfamily is described herein in-
cluding a new species from Paraguay and
another from Brazil.

These new taxa are described and an at-
tempt is made to place them in the context
of present-day weevil classification. In the
last section of the paper, the close affinity
of the newly described genus with the ge-
nus Orobitis and preliminary conclusions
regarding the relationships of the Orobiti-
dinae are presented. It is clear that exami-
nation of a much larger sample of taxa and
characters in combination with rigorous
analysis may be necessary to propose ro-
bust hypotheses of relationships.

Curculionidae, Orobitidinae, new taxa, South America

As in many other beetle families, termi-
nology of structures in weevils is fairly in-
consistent and often is used in a misleading
way (Thompson 1989). Although the sub-
ject of this paper is not comparative mor-
phology or analysis of terminology, it is
useful to provide a brief summary of the
sources of terms for the most critical and
disputed parts of the weevil body.

To describe metathoracic structures, a
combination of terms proposed by Matsuda
(1970) and McHugh et al. (1997) and the
terms previously adapted for leaf beetles
(Konstantinov and Vandenberg 1996) is
used.

General wing terminology follows Ku-
kalova-Peck and Lawrence (1993). Some
unique structures of weevils were named by
Zherikhin and Gratshev (1995). These
terms are used for structures which were
not named by Kukalovd-Peck and
Lawrence (1993). Terms of internal elytral

930

structures are those used by Lyal and King
(1996).

Terms proposed by Thompson (1989) are
used for the spermatheca. The term
‘“‘pump,”’ widely accepted to describe the
same part of the spermathecal body at-
tached to the lobe of the duct and lobe of
gland (Smith 1979, Konstantinov 1998), is
used instead of “‘tail.”’ As for the other parts
of the female genitalia, commonly accepted
terms are used (Morimoto 1962, O’Brien
and Askevold 1995, Howden 1995). Mori-
moto (1962), Lyal (1993), and O’Brien and
Askevold (1995) are the main sources of
the names used for different parts of the
male genitalia.

Subfamily Orobitidinae

Description.—Body small, 2—3.5 mm
long, globose or transversely-subrhomboi-
dal, prothorax and elytra strongly conjointly
convex (Figs. 1—4). Black; vestiture dense,
tightly appressed and inconspicuous or sub-
erect, composed of white and dark scales.

Rostrum as (Figs. 3—6, 8) long as or
slightly longer than prothorax, basally
weakly curved, apically straight; dorsal
margin more or less angular in lateral view
over antennal base. Apical part of rostrum
in female (Fig. 5) much narrower than basal
part. Ventral surface of head without pos-
terior tentorial pits and occipital sutures.
Subgenal sutures fused (Figs. 10, 11). An-
tenna inserted at basal third of rostrum
(Figs. 3, 4, 8, 9). Antennal scrobe oblique,
ventral margin reaching venter of rostrum
less than half way between antennal inser-
tion and eye. Scape stout, short, less than
half as long as rather long and slender fu-
nicle with 7 antennomeres (Figs. 16, 17).
Eye medium-sized, weakly convex. Frons
as broad as base of rostrum or slightly nar-
rower, sometimes weakly narrowing in mid-
dle.

Mandible (Figs. 12, 13, 15) ventrally
with 2 visible denticles, third denticle cov-
ered with apex of rostrum; mandible dor-
sally separated from maxilla by long ven-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

trolateral appendage of rostrum (Figs. 12,
15). Labial palpi 2- (Parorobitis) or 3-
(Orobitis) segmented. Mentum long.

Prothorax subtrapezoidal with base very
broadly rounded or nearly straight and
slightly produced posteriorly for most of its
length. Sides slightly rounded or nearly
straight. Basal margins of prothorax and el-
ytra tightly coupled, sometimes jointly
raised, but not crenulate. Apical margin of
pronotum not raised. Postocular lobes well
developed to absent. Lateral tubercles ab-
sent. Disc of pronotum flat or moderately
convex in apical half, without median sul-
cus. Prosternum excavated, not keeled in
front of fore coxae. Distance between coxae
nearly equal to width of rostrum. Mesepi-
meron not visible from above, flat, deeply
inserted between base of prothorax and el-
ytra (Figs. 18, 23). Rostral furrow formed
by rather high, lamelliform keels behind
fore coxae on prosternum, sometimes lower
keels also present on mesosternum.

Mesoscutellum with tall stalk (Fig. 18).
Mesosternum between middle and hind
coxae with broad fold projecting over hind
coxae. Mesosternum and metasternum
fused. Internal ridge and border between
these structures barely visible. Mesendos-
ternite situated between internal wall of
coxal cavity and mesosternal ridge. Plate
nearly as long as wide. Basal and apical ap-
pendages slender (Fig. 19).

Scutoscutellar ridges of metanotum not
connected, directed anteroventrally. Alo-
cristal ridges not connected in middle (Fig.
20).

Metendosternite with moderately narrow
stalk. Furcal arm slender, apically widening.
Lateral arm short, situated nearly in middle
of furcal arm. Anterior and posterior trans-
verse ventral processes of equal length,
leaving narrow furrow between (Figs. 21,
22).

Elytra slightly longer than broad, with
strongly convex or obsolete humeral prom-
inences. Elytra with locking sutural struc-
ture composed of deep longitudinal groove

VOLUME 102, NUMBER 4

Fig. 1.

Parorobitis gibbus, 3.

on left elytron (Fig. 18) and longitudinal
ridge on right elytron (Fig. 27). Basal part
of suture with two callosities and depres-
sions between on left elytron. Right elytron
with two depressions, and with ridge be-
tween them (Figs. 27, 28, 31, 32). Submar-
ginal ridge of each elytron also complicated
(Fig. 29, 30). Apex of elytron with longi-
tudinal stridulatory file. Preapical groove
absent (Figs. 35, 36).

Wings well-developed or slightly re-

$33)

duced, with well developed R, radial fis-
sure, and first radial scelerite (Figs. 25, 26).!

Legs long. Femora broad from base, un-
armed, subulate to moderately clavate, shal-
lowly grooved for reception of tibiae. Tib-
iae (Figs. 39, 42, 43) in female not unci-
nate; in male, with short apical mucro (Figs.

! Zherikhin and Gratshev (1995) state that the Orobi-
tini are wingless; however, Orobitis cyaneus has short-
ened but fairly well-developed wings (Fig. 26).

982

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2—4.
lateral view.

2, Orobitis cyaneus, 3, dorsal view. 3, O. cyaneus, 3, lateral view. 4, Parorobitis gibbus, d,

VOLUME 102, NUMBER 4

40, 41). Apical comb of setae not or slightly
extending onto outer margin of tibia. Third
tarsomere bilobed. Claws separate, moder-
ately divergent, with long median paired
contiguous process (Figs. 46—48).

Abdomen with sterna in one plane. Ster-
na 1—4 of subequal length, 5 almost as long
as sterna 3 and 4 together. Sternum | nearly
completely divided by hind coxae. Hind
margins of sterna 2—4 parallel, moderately
curving backwards on sides (Fig. 23). Pleu-
rite 1 long, separated from rest of tergum 1
by groove, attached by wide membrane
with granulated microsculpture to metepis-
ternum (Figs. 24). Pleura 4—7 combined
with corresponding tergites. Tergum 8 ex-
posed in males (Fig. 1).

Diagnoses of the two genera of the
subfamily Orobitidinae

1 (2). Body subspherical, rounded. Humeral
prominences obsolete. Basal margins of
prothorax and elytra not raised (Figs. 2, 3).
Dorsum almost uniformly covered with
black or bluish black tightly appressed
scales, shining. Dorsum of rostrum evenly
curved over antennal base (Figs. 3, 9).
Frons flat. Pronotum evenly convex, with-
out median or lateral depressions. Basal
margin of pronotum almost straight, imper-
ceptibly notched opposite scutellum. Post-
ocular lobes absent. Mesoscutellum round-
ed, as long as broad, weakly and evenly
convex. Elytra longer than broad, all inter-
vals flat, striae linear. Lateral margin of el-
ytron slightly and almost evenly rounded,
inconspicuously emarginate near base over
mesepimera. Mesosternum without keels.

and dark brown spots on pronotum and
contrasting basal band on elytra, with back-
ground vestiture composed of dense sub-
erect to subappressed, linear to lanceolate,
white scales. Rostral dorsum angular over
antennal insertion (Figs. 4, 8). Frons mod-
erately depressed, anterior part sloping to
base of rostrum and posterior part, to ver-
tex. Pronotum with two submedian obtuse
prominences in apical half; laterally un-
even. Basal margin of pronotum notched in
middle and bisinuate laterally. Postocular
lobes from well- to poorly developed. Me-
soscutellum longer than wide, large, tu-
berculiform, steeply sloping anteriorly and
gradually sloping to elytra. Elytra broader
than long. Odd-numbered intervals weakly
to moderately convex, costiform near base.
Striae consisting of medium-sized, deep,
remote punctures, space between latter un-
dulate. Intervals very densely, finely punc-
tate, nearly matt. Lateral margin of elytron
moderately deeply emarginate over mese-
pimeron, very shallowly bisinuate behind.
Mesosternum with well-developed keels
reaching middle coxae. Fore coxa with ob-
tuse prominence at apex, separated laterally
by narrow sulcus. Femora distinctly cla-
vate, with shallow grooves on ventral side,
all uniformly covered with scales. Claws
long, length greater than width of last-tar-
somere at apex (Figs. 46, 47); median pro-
cess slightly shorter than claws. Abdominal
suture 1 with lateral pore under brown
scaly spot. Pleural area of thorax with
brown spots. 2.9—3.5 mm.. Neotropical

Shae tame Carats eam airs Parorobitis, new genus

Parorobitis Korotyaev, O’Brien, and

Konstantinov, new genus

eS)

2 (1).

Sides of meso- and metathorax evenly con-
vex, without prominences or depressions.
Fore coxa without prominence on inner
side of apex. Femora nearly parallel-sided,
with glabrous grooves ventrally. Claw
short, length less than width of tarsomere
5 at apex (Fig. 48); median process ca. half
as long as claw. Abdominal suture | with-
out lateral pore. Sterna uniformly covered
with yellowish scales. 2-2.8 mm. Palearc-
tic. Larvae in seed capsules of Viola spp.

3 6.6 BMS LCH ORES, ONG. Geer eae oer Orobitis Germar

Body subrhomboidal, angular humeral
prominence convex, forming broad, mod-
erately curved callus (Figs. 1, 4). Basal
margins of both prothorax and elytra both
finely conjointly raised. Dorsum with pale

Type species.—Parorobitis gibbus, new
species.

Description.—Small; body short, su-
brhomboidal, dorsally strongly convex;
prothorax and elytra conjointly convex.
Body black, densely clothed with suberect
to subappressed, elongate scales, dorsally
with contrasting pattern of white and brown
scales. Black, funicle and club of antennae
and apical third of rostrum brown, tarso-
mere 3 dark brown.

Head capsule small. Rostrum angularly
curved at antennal base, ventrally weakly
arcuate, with dorsum more (in female) or
less (in male) sharply angular (Figs. 4, 8).

934 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

*

ae '

Fig. 5—7. Head. 5, Parorobitis gibbus, 3, frontal view. 6, Orobitis cyaneus, ¢, frontal view. 7, P. gibbus,
view through occipital opening, tentorium and foregut with proventriculus.

VOLUME 102, NUMBER 4

10 11

Fig. 8-11. Head, ¢. 8, 9, Lateral view. 10, 11, Ventral view. 8, 10, Parorobitis gibbus. 9, 11, Orobitis
cyaneus.

936

Basal part of rostrum broader than apical
(Fig. 5), nearly as broad as fore tibia. Api-
cal part narrower (Fig. 5); in female,
straight, cylindrical, shining. Antenna in-
serted at basal third of rostrum. Scape short,
stout. Funicle long, antennomeres progres-
sively shorter apically; club oval, short
(Figs. 5, 16). Eyes strongly anteriorly di-
rected. Frons at anterior margin as broad as
base of rostrum, widened posteriorly, weak-
ly depressed, slightly, transversely ridged in
middle. Vertex weakly convex, not carinate.
Sculpture fine. Tentorium well-developed,
with moderately broad long stalk attached
to ventral surface of head and two lateral
branches attached to anteromedial corner of
eye (Fig. 7). [It is unclear if these branches
are homologous with anterior or dorsal
arms.] Foregut with 8-laminate chitinized
proventriculus (Fig. 7, 14). [According to
Crowson (1955) this structure may well be
plesiomorphic for curculionids. It is also
known to occur in Cossoninae (Caulophilus
latinasus Say) (Crowson, 1955), Baridini
(Pachybaris porosus Lec.) (in the latter spe-
cies it is much longer and all the laminae
are lower), and Zygopini (Lobotrachelus
subfasciatus Motschulsky) (it is generally
shorter with every lamina being taller pos-
teriorly and shorter anteriorly). ]

Left mandible with ventral denticles clos-
er together than on right mandible. Labial
palpus 2-segmented. Sides of mentum con-
vex (Fig. 12).

Prothorax transverse, trapezoidal. Base
sinuate, noticeably produced posteriorly
and enclosed by humeral prominences of
elytra, shallowly notched anterior to scutel-
lum and bisinuate laterally. Apical margin
of pronotum raised. Basal margin raised to
meet raised basal margin of elytra, finely
carinate. Hind angles weakly projecting,
sides weakly convex and strongly converg-
ing to apical prominences immediately
above postocular lobes. Apical constriction
moderately deep on dorsum and gradually
disappearing towards prominences, which
slope to anterior margin of prothorax. Sides
near anterior margin with two angular

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

prominences visible in dorsal view. Punc-
tures on disc dense, uniform, medium-
sized, rather deep; intervals between punc-
tures shining, narrow. Ocular lobe well-de-
veloped, angular. Fore coxae separated by
width of rostrum. Prosternum in front of
coxae not keeled, as long as width of ros-
trum at base, deeply excavated, with mod-
erately high lamelliform keels behind cox-
ae.

Mesothorax short, transverse (Figs. 18,
19). Mesosternum with keels reaching an-
teromedial margin of middle coxae. In lat-
eral view, mesosternum rather steeply de-
clivous to metasternum. Scutellum large,
strongly convex, broadly rounded on top.
Mesepimeron separated from mesocoxa by
mesosternal appendage (Fig. 23). Mesepim-
eron not visible from above, although en-
tering deeply between bases of prothorax
and elytron. Distance between mesocoxae
greater than width of coxa.

Metathorax much longer than mesotho-
rax (Fig. 18). Metanotal prescutal mem-
brane thin and vertical. Alocrista moderate-
ly narrow, without well-developed lateral
ridge. Scutum well separated by scutoscu-
tellar groove forming large “‘pocket”’ lat-
erally, nearly as large as scutellum. Ven-
trally, scutoscutellar ridge forming wide
plate directed anteroventrally and dividing
metanotal cavity into two almost equal-
sized compartments (Fig. 20). Scutellar
groove ending at basal margin of notum.
Allocristal part of notum containing dorsal-
ly and ventrally separated cavity with two
elongate openings. Axillary part of metan-
otum situated anterolaterad of scutum. Part
of scutum connected with axillary area
forming nearly parallel-sided, wide, weakly
sclerotized appendage. Metasternum form-
ing thick fold over hind coxa. Hind coxae
separated from elytra by slightly less than
width of coxa. Metepisternum convex.

Elytra much broader than pronotum,
transverse, strongly narrowing apically,
with unusually convex humeral prominenc-
es, separately and rather narrowly rounded
at apices, and extending over base of py-

VOLUME 102, NUMBER 4

Fig. 12-17. 12-14, 16, Parorobitis gibbus, S.
13, Tip of head, lateral view. 14, Proventriculus. 1

15, 17, Orobitis cyaneus, 3. 12, Mouth parts, ventral view,
5, Mouth parts. 16, 17, Antenna.

938 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

mstr

msc

Fig. 18-22.

Thoracic structures of Parorobitis gibbus. 18, Lateral view with right elytron removed. 19,

Mesothorax and metasternum (metanotum removed), view of internal structures. 20, Metanotum, ventral view.
21, 22, Metendosternite (21, dorsal view; 22, ventral view). Abbreviations: alr = allocrista; aly = anterior part
of longitudinal ventral process; amt = furcal arm; ana = anterior notal appendage; atv = anterior part of
transverse ventral process; elec = elytral lock; lamt = lateral arm; m = membrane; mesn = mesonotum:; mest
= stalk of mesoscutellum; msc = mesocoxal cavity; msds = mesendosternite; msm = mesepimeron; msst =
mesepisternum: mst = mesosternum; mstr = merosternal ridge; mtn = metanotum; mts =

= = metepisternum; mtt
posterior part of longitudinal ventral process; pma = postmedial appendage; psm =

= metasternum; ply =

VOLUME 102, NUMBER 4

gidium. Lateral margin of elytron emargin-
ate at base and very shallowly bisinuate
posteriorly. Internal surface with four lon-
gitudinal striae of tiny denticles (Fig. 27).
Right elytron with locking structure con-
sisting of narrow ridge and two nearly
round lateral impressions. Elytral apex cov-
ered with multidentate median callosities.
Middle of apex with patch of medially di-
rected setae (Figs. 27, 31). Striae composed
of medium-sized, rather deep, sparse, round
punctures. Intervals 3—4 times as broad as
striae; odd-numbered intervals weakly to
strongly convex, densely covered with
small shallow punctures.

Wing elongate, with apex slightly darker
than middle. Humeral field with short C not
touching Sc. R straight, widening apically.
Radial fissure indistinct. Radial fold weak.
First radial sclerite well-developed and
large, second small. Rr absent. Anal field
with two weakly developed anal veins (Fig.
D5):

Legs long, stout. Trochanters short. Fem-
ora broad from base, moderately clavate,
with deep ventral constriction near apex.
Shallow ventral groove in apical half cov-
ered with scales as elsewhere. Hind femur
more swollen in apical part than fore and
middle femora. Tibiae weakly broadening
and curved outward apically; fore and mid-
dle tibiae slightly flattened, hind tibia nearly
round in cross-section, medially and fron-
tally compressed apically (Fig. 31). In fe-
male, tibiae mutic; in male, all armed with
small curved mucro (Figs. 39—41).

Apical fringe of setae slightly oblique
only to outer margin of tibia. Setae dense,
fine and short. Tarsi of medium proportions,
tarsomere 3 bilobed, nearly twice as broad
as 2. Tarsomere 5 moderately widening api-
cally, slightly more than half length extend-
ing beyond lobes of tarsomere 3. Claws

ee

939

large, with transversely flattened, longitu-
dinally curved, apically subacute to acute,
median process, latter slightly shorter than
claws (Figs. 46, 47).

Venter weakly concave in female and
more strongly so in male. Sterna 1—4 short,
nearly equal in length. Sternum 5 nearly
twice as long as sternum 4 (Fig. 23). Small
indentation between sterna | and 2, slightly
smaller indentations between other sterna,
with latter indentations slightly more me-
dial. Abdominal terga 1—5 short, nearly
equal in length. Tergum 6 much shorter in
middle. Tergum 7 slightly longer than terga
4, 5 and 6 together (Fig. 24). Tergite 8 of
male with long basal projections (Fig. 57,
60).

Male genitalia: Apodemes longer than
upper part of median lobe. Apical [third?
fourth] of median lobe bent ventrally, grad-
ually narrowing apicad. Spiculum gastrale
long, attached to membrane of tegmen be-
tween sclerotized parts of sternum 8 (Figs.
57-62).

Female genitalia: Tergum 8 elongate,
weakly sclerotized medially, with moder-
ately well-sclerotized stripe along border.
Sternum 8 with two patches of long setae
apically and short setae between sclerotized
arms (Fig. 51). Vagina narrow. Coxite and
stylus narrow, at rest situated inside vagina
(Figs. 53, 55). Spermatheca with well-de-
veloped gland and ductal lobes, receptacle
more than 4X as long as wide (Figs. 52,
56).

Etymology.—This masculine generic
name is the result of combining the generic
name Orobitis and the Greek prefix para-
meaning “‘close by, or similar to” and re-
flects the morphological similarity and
close relationship of these two genera.

prescutal membrane; ptv = posterior part of transverse ventral process; scg = scutellar groove; scmt = meta-
scutellum; sct = metascutum; sctm = mesoscutellum; ssr = scutoscutellar ridge; stmt = furcal stalk; tmt =

tendons of meso- metafurcal muscles.

940 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

msst

<> EEE Ce mts

26

Fig. 23-26. 23-25, Parorobitis gibbus. 23, Meso- and metasternum and abdominal sternites. 24, Abdominal
tergites, pleurites, and metepisternum. 25, Wing. 26, Orobitis cyaneus, wing. Abbreviations: Irs, 2rs = radial

sclerites; m = membrane; msm = mesepimeron; msst = mesepisternum; p11, 2, and 3 = abdominal pleurites;
rfi = radial fissure; strm = medial stripe.

VOLUME 102, NUMBER 4

Parorobitis gibbus Korotyaev, O’Brien,
and Konstantinov, new species
(Figs. 1, 4, 5, 7, 8, 12-14, 16, 18-25, 27,
29, 31, 35, 39-41, 46, 47, 51-53, 57-59)

Types.—Holotype d, Paraguay, Puerto P.
Stroessner, 26—28.XII.1965, Nr. BR 9 (Ma-
hunka), HMNH. Paratypes. 2 6, 2 2, same
data as holotype (¢6, 2 HMNH, d USNM,
2 ZMAS). Paratypes. 6, °, Paraguay: Alto
Parana, 6 km W. Pto. Pres. Stroessner: 25—
28.1.1983, leg. E. G. Riley (CWOB)

Description.—Male: Rostrum nearly as
long as pronotum, with maximum width at
antennal insertion, tapering basally and api-
cally in dorsal view. Dorsal surface with
sharp median carina disappearing before
apex (Fig. 5). Lateral carina better devel-
oped between antennal insertion and head.
Space between median and lateral carinae
with row of deep punctures. Weak carinae
converging on sides above antennal inser-
tion, but vanishing separately near apex.
Antenna inserted 0.35 from base of rostrum
(Fig. 8). Scape shorter than funicular anten-
nomeres | and 2 together, not reaching eye.
Funicle rather slender and long, antennom-
eres progressively becoming shorter, anten-
nomeres 5 and 6 slightly longer than wide,
antennomere 7 globose. Club oval, short.
Setae on flagellomeres moderately long,
suberect.

Prothorax 1.65 as broad as long. Pos-
tocular lobe large, angular. Disc moderately
convex, with two broad obtuse prominences
in apical half, separated by shallow longi-
tudinal depression not extending to basal
half of disc. Sides of disc with rather deep
round fovea postero-lateral to apical discal
prominences, and with shallower fovea be-
hind anterolateral prominences. Scutellum
large, 1.4 as long as broad, rounded, very
strongly convex, with steep anterior slope
and gentle posterior slope to level of elytral
suture. Surface of scutellum concealed by
pale, sand-brown, imbricate scales.

Elytra 1.2X as broad as long, 1.6% as
broad as base of prothorax, very strongly
narrowing apically, with very strongly con-

94]

vex rounded humeral prominences. Disc
strongly convex, more steeply sloping to
base than to apex. Base of elytra deeply fo-
veate along basal half of scutellum. Elytral
striae with medium-sized, deep, sparse
punctures. Stria 1 at base parallel to antes-
cutellar margin of elytron and meeting in-
curved base of stria 2. Striae 3—6 nearly
straight at base, stria 7 vanishing near the
posterior margin of swollen interval 7, stria
8 reaching basal half of humeral promi-
nence. Intervals 2 and 4 rather strongly
convex, interval 6 depressed at base, be-
coming gradually more convex apically. In-
terval 8 in apical two-thirds slightly more
convex than others. Preapical prominence
obtuse and not very convex, stria beneath
prominence deepened. Intervals matt,
densely and finely punctate, lacking gran-
ulations.

Femora moderately swollen apically.
Tibiae inconspicuously widening and api-
cally curved outward. Mucro on middle and
hind tibiae slightly shorter than tarsal claws;
on fore tibia, half as long and much finer
(Figs. 39-41). Protarsomere one about
1.7X as long as broad, protarsomere two
1.3X as long as broad, protarsomere three
0.8X as long as broad, 1.1X as long as 2.
Tarsomere 5 weakly broadened apically, by
two-thirds of length extending beyond
lobes of tarsomere 3. Punctures on femora
dense, deep, medium-sized, round; on tib-
iae, more or less elongate. Median claw
process ca. twice as broad as claw (Figs.
46, 47).

Anal sternum flattened in middle one-
third and covered with suberect to erect
white hairs, sterna 3 and 4 with long, nar-
row, curved, suberect white scales along
hind margin. Pygidium rounded, weakly
transverse and raised along median line,
coarsely punctate, and moderately densely
covered with elongate suberect white
scales.

Genitalia (Figs. 57-59): Median lobe
narrowing apically, abruptly widening hor-
izontally. Apical third with lateral patches

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 27-28. Internal surface of right elytron. 27, Parorobitis gibbus. 28, Orobitis cyaneus. Abbreviations:
ag = apical groove of elytron; ape = apical area of elytron; coe = concave area of elyton; f = file; g =

submarginal ridge; rm = rim; sf = sutural flange.

VOLUME 102, NUMBER 4

of setae. Apodeme curved laterally, wid-
ening before base.

Head capsule densely covered with par-
allel-sided or weakly rounded, broad scales;
vertex with few narrower brown scales in
middle, and piceous scales on hind margin,
otherwise with white scales. Basal part of
rostrum with moderately dense white
scales, latter half as broad as on vertex, rap-
idly thinning to antennal insetion; rostrum
distally with sparse hair-like scales along
sides to slightly beyond middle. Pronotum
with moderately dense, narrow-lanceolate,
brown scales in middle of disc; median de-
pression with paler scales, often with few
white scales on bottom. Brown spots on
discal prominences and near base surround-
ed by narrow gray lines moderately densely
covered with hair-like white scales; latter
also covering sides of prothorax to angular
line formed by broad white scales, nearly
concealing pleural area and fore coxa. An-
terior slope of discal prominences densely
covered with narrower white scales; apical
constriction with two gray spots, formed by
narrow white scales. Sides also with small
spot of brown scales between hind angles
and lateral depression. Scutellum dull pale
brown apically, with white scales at base.
Base of elytra with band of brown scales;
intervals 1 and 2, preapical prominences
and sides in apical half densely covered
with broad white scales; middle of each el-
ytron with sparser vestiture of white, linear
to narrow-lanceolate scales, latter broader
in striae. Legs mottled with spots of narrow
and broad white scales, sternum | and base
of sternum 2 with brown spot on sides. Py-
gidium with scales not extending from
punctures, broadly oval to round in basal
half and elongate, slightly raised near apex;
brown scales predominant in basal half;
with white scales, along margins, median
line, and in apical half.

Body length 3.1—3.3, width 3 mm.

Female: Rostrum 1.25 as long as pro-
thorax, moderately widened and angularly
curved at antennal insetion, with dorsal out-
line angular in lateral view; in apical part,

943

straight, cylindrical, slender, about *4 width
of fore tibia. Basal half of rostrum matt,
densely covered with small shallow punc-
tures. Median carina well developed, ex-
tending considerably beyond antennal base;
lateral carinae finer. Sides of rostrum with
pair of carinae converging apically from an-
tennal base to middle of apical part of ros-
trum. Antennae inserted 0.3 from base of
rostrum.

Tibiae without mucro. Protarsomere 1
about 1.6 as long as broad, protarsomere 2
nearly as long as broad, protarsomere 3
twice as long as 2.

Anal sternum flat. Pygidium slightly lon-
ger than broad, weakly and rather evenly
convex, slightly raised along median line in
apical third, narrowly rounded at apex,
matt, moderately densely covered with me-
dium-sized, fairly deep punctures.

Scutellum dull pale brown.

Female genitalia: Arms of sternum 8
moderately long (Fig. 51). Coxite and sty-
lus slender (Fig. 53), latter nearly cylindri-
cal. Spermathecal pump and ductal lobe
forming broad loop. Apex of spermathecal
pump bent toward ductal lobe (Fig. 52).

Remarks.—Parorobitis gibbus shares
most character states with P. minutus. It can
be separated by the whiter scale pattern of
the head, larger size, longer arms of ster-
num 8 (Fig. 51), the slender coxite and sty-
lus (Fig. 53), the stylus nearly cylindrical,
the spermathecal pump and ductal lobe
forming a moderately wide loop, the apex
of the spermathecal pump bent toward the
ductal lobe (Fig. 52), the median lobe nar-
rowing apically with abrupt widening at the
horizontal part, the apical third with lateral
patches of setae, and the apodeme widening
apically (Fig. 57-59).

Etymology.—This specific epithet is
based on the Latin adjective gibbus =
‘*humpbacked, protuberant’”’ and refers to
the humpbacked appearance and the two
well-developed pronotal protuberances or
swellings.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 29-30.

Parorobitis minutus O’Brien, Korotyaev,
and Konstantinov, new species
(Figs. 54-56, 60—62)

Types.—Holotype ¢, Brazil, Curitiba,
Parana, I.13.1969, Araucaria forest, leg. C.
W. & L. B. O’Brien (OZUP). Paratype °,
same data as holotype (CWOB).

Description.—Male: Rostrum slightly
longer than pronotum, with maximum
width at antennal insertion, tapering basally
and apically in dorsal view. Dorsal surface
with sharp median carina disappearing in
front of apex. Lateral carina better devel-
oped between antennal insertion and head.
Space between median and lateral carinae

Internal surface of right elytron, anterolateral part. 29, Parorobitis gibbus. 30, Orobitis cyaneus.

without punctures, covered with longitudi-
nal wrinkles. Lateral carinae above antennal
insertion parallel to each other, vanishing
near apex. Antenna inserted 0.37 from base
of rostrum. Scape shorter than funicular an-
tennomeres 1 and 2 together, not reaching
eye. Funicle rather slender and long, anten-
nomeres progressively becoming shorter,
antennomeres 5 and 6 slightly longer than
wide, antennomere 7 globose. Club oval,
short. Setae on flagellomeres long, suberect.

Prothorax 1.67X as broad as long. Pos-
tocular lobe weakly developed. Disc mod-
erately convex, with single obtuse promi-
nence in middle. Lateral areas of disc with-

VOLUME 102, NUMBER 4

Fig. 31-34. Lock structures of elytron. 31, Parorobitis gibbus. 32, Orobitis cyaneus. 33, Pachybaris porosa.
34, Lobotrachelus subfasciatus (left elytron).

out deep fovea, with laterobasal promi-
nence.

Scutellum large, 1.67 as long as broad,
very strongly, roundly convex, with steep
anterior slope and posterior slope leveling
to elytral suture. Surface of scutellum con-
cealed by brownish (in male) or pale sand-
brown (in female) imbricate scales.

Elytra together 1.2 as broad as long,
1.69X as broad as base of prothorax,
strongly narrowing apically, with very
strongly convex, rounded humeral promi-
nences. Disc strongly convex, more steeply
sloping to base than to apex. Base of elytra
deeply foveate along basal half of scutel-
lum. Strial punctures small, moderately
deep, sparse, nearly entirely covered with

scales. Stria 1 parallel at base to antescu-
tellar margin of elytron, meeting incurved
base of stria 2. Striae 3—6 nearly straight at
base, stria 7 vanishing in front of preapical
callosity, stria 8 reaching basal half of hu-
meral prominence. Intervals 2—6 equally
convex; intervals 5 and 6 flattening apical-
ly; interval 8 in apical two-thirds nearly as
convex as adjacent intervals. Preapical
prominence convex, stria beneath promi-
nence deepened. Surface of intervals matt,
densely and finely punctate, lacking gran-
ulation.

Femora moderately swollen apically.
Tibiae inconspicuously widening and api-
cally curved outward. Mucro on middle and
hind tibiae slightly shorter than tarsal claw;

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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VOLUME 102, NUMBER 4

on fore tibia, mucro tiny, scarcely visible.
Protarsomere one about 2X as long as
broad, protarsomere two 1.2 as long as
broad, protarsomere three 0.72 as long as
broad, 1.3 as long as protarsomere two;
protarsomere five weakly widened apically,
extending two-thirds beyond lobes of pro-
tarsomere 3. Punctures on femora dense,
deep, medium-sized, round; on tibiae, more
or less elongate. Tarsal claws with median
process ca. as broad as claw.

Anal sternum flattened in middle and
covered with suberect to erect white hairs,
ventrites 3 and 4 with long, narrow, curved,
suberect white scales along hind margin.
Pygidium rounded, weakly transverse and
raised along median line, coarsely punctate,
and moderately densely covered with elon-
gate suberect white scales.

Male genitalia (Figs. 57-59): Median
lobe narrowing apically without abrupt
widening horizontally. Apical third without
lateral patches of setae. Apodeme at base
nearly as broad as at apex.

Head capsule densely covered with par-
allel-sided or weakly rounded broad scales;
vertex with two stripes of narrow brown
scales in middle, and piceous scales at hind
margin, otherwise with white scales. Basal
part of rostrum with moderately dense
white scales half as broad as those on ver-
tex, rapidly thinning to antennal insertion;
rostrum distally with sparse hair-like scales
along sides to slightly beyond middle.
Pronotum with moderately dense, narrow-
lanceolate, brown scales on disc and paler
scales in middle, with narrow transverse
stripe of white scales behind middle. An-
terior slope of discal prominence densely
covered with wider white scales. Sides also
with small spot of brown scales above fore
coxae. Scutellum dull pale brown on top,
with white scales at base. Outer corner of
mesepisternum with spot of black scales.
Base of elytra with band of brown scales,
intervals 1 and 2, preapical prominences,
and sides in apical half, densely covered
with broad white scales, middle of each el-
ytron with sparser vestiture of white, linear

947

to narrow-lanceolate scales, broader in stri-
ae. Legs mottled with spots of narrow and
broad white scales, sternum 1 and base of
sternum 2 with brown spot on sides. Pygid-
ium with scales not extending beyond punc-
tures, broadly oval to round on basal half
and elongate, slightly raised near apex;
brown scales predominant on basal half,
white scales along margins, median line,
and on apical half.

Body length 2.9—3.0, width 2.9 mm.

Female: Rostrum 1.25 as long as pro-
thorax, moderately widened and angularly
curved at antennal insertion, with dorsal
outline angular in lateral view; apically
straight, cylindrical, slender, about *%4 width
of fore tibia. Median carina shorter than in
male; lateral carinae well developed. An-
tenna inserted 0.3 from base of rostrum.

Tibiae without mucro. Protarsomere one
about 2.2 as long as broad, protarsomere
two 1.2X as long as broad, protarsomere
three 1.17X as long as protarsomere two.

Female genitalia: Arms of sternum 8
moderately short (Fig. 54). Coxite and sty-
lus robust (Fig. 55). Stylus tapering apical-
ly. Spermathecal pump and ductal lobe
forming moderately narrow loop. Apex of
spermathecal pump bent in direction op-
posite of ductal lobe (Fig. 56).

Remarks.—Parorobitis minutus shares
majority of the character states exhibited by
P. gibbus. It can be distinguished by the
darker scale pattern of the head, smaller
size, shorter arms of sternum 8 (Fig. 54),
robust coxite and stylus (Fig. 55), stylus ta-
pering apically, spermathecal pump and
ductal lobe forming rather narrow loop,
apex of spermathecal pump bent in direc-
tion opposite of ductal lobe (Fig. 56), me-
dian lobe narrowing apically without abrupt
widening horizontally, apical third without
lateral patches of setae, and apodeme at
base nearly as narrow as at apex (Fig. 60—
62).

Etymology.—This specific epithet is
based on the Latin adjective minutus =
‘“‘small’’ and refers to the relatively small
size of the body.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ali

Fig. 37-38. Elytral apex with stridulatory file. 37, Pachybaris porosa. 38, Lobotrachelus subfasciatus. Ab-
breviations: ag = apical groove; pag = preapical groove.

VOLUME 102, NUMBER 4 949

Fig. 39-43. Legs. 39-41, Parorobitis gibbus. 42, 43, Orobitis cyaneus. 39, Right metatibia, dorsal view.
40, 41, Tibial apex. 42, Right metatibia, dorsal view, 43, Right metatibia, ventral view.

950

DISCUSSION

In spite of the sharp differences in ap-
pearance, Orobitis Germar and Parorobitis
share many important characters including
the following: unusual, extremely convex
shape of body; rostrum weakly curved ba-
sally, apically straight; ventral surface of
head without posterior tentorial pits and oc-
cipital sutures; antennal scrobe oblique,
with ventral margin reaching venter of ros-
trum less than half way between antennal
insertion and eye; mandibles with two den-
ticles ventrally; labial palpi 2-segmented;
prothorax subtrapezoidal, with base ex-
tremely wide and broadly rounded; mese-
pimera not visible from above, flat, deeply
inserted between base of prothorax and el-
ytra; prosternal sulcus formed by rather
high lamelliform keels behind fore coxae;
mesoscutellum with tall stalk; mesosternum
and metasternum fused; elytra with strongly
convex humeral prominences and sutural
locking mechanism; apex of elytron with
longitudinal stridulatory file (Figs. 35, 36);
wings with well-developed R and radial fis-
sure; legs long; claws with well developed
median process (Figs. 46—48); abdominal
sternum 1 nearly completely divided by
hind coxae; pleurum 1 long, separated from
tergum 1 by groove, attached to metepis-
ternum by wide membrane with granulated
microsculpture (Figs. 24). One of the most
convincing pieces of evidence for the close
relationships between Orobitis and Paro-
robitis is the structure of the stridulatory
device. According to Lyal and King (1996),
Orobitis has a unique file isolated from sur-
rounding sculpture and placed on a differ-
ent part of the elytron (Figs. 28, 36). Based
on these observations, Lyal and King
(1996) suspected that the file of Orobitis is
non-homologous with files of other weevils
(Figs. 37, 38). Parorobitis shares all the
distinctive characters of the stridulatory file
of Orobitis (Figs. 27, 35).

The differentiation between the two gen-
era 1S most evident in the gestalt. The
strongly developed relief of the exoskeleton

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

in Parorobitis readily distinguishes it from
species of Orobitis. Perhaps this indicates
greater development of thoracic muscles,
possibly due to a much more active flight
behavior in the species of the new genus.

The systematic relationships of the Orob-
itidinae are uncertain. Colonnelli (1984:
207) placed Orobitis in the Ithyporinae
[previously considered a tribe of the Cryp-
torhynchinae (Hustache 1936)], and cited
Hustache (1936) as justification for the
placement. This is actually incorrect as
Hustache does not deal with Orobitis in the
Coleopterorum Catalogus, but only cites the
original placement of Cleogonus nuculus
(Germar) in the genus Orobitis, which has
nothing to do with the systematic position
of Orobitis. Zherikhin and Egorov (1990:
113) treat Orobitini as a tribe of the Bari-
dinae (which according to them also in-
cludes Ceutorhynchini, Zygopini, and Tri-
gonocolini).

Zherikhin and Gratshev (1995) elevated
Baridinae to family, but there are some
problems with this taxonomic decision.
Among the synapomorphies of Barididae
sensu Zherikhin et Gratshev, the presence
of the median keel on the inner surface of
metasternum seems the primary character.
However, this is shared by other apparently
very different groups and its distribution in
combination with other characters should
be studied. The first character in the wing
structure listed by Zherikhin and Gratshev
(1995: 773), among the most important
synapomorphies of the Barididae, is the
strongly curved basal wing margin. How-
ever this character state is more developed
in the Zygopinae and Ceutorhynchinae
(Zherikhin and Gratshev 1995, Figs. 108—
121), than in Baridinae (see Figs. 122—132
in the same work). Trigonocolinae do not
possess a critical diagnostic character which
distinguishes Ceutorhynchinae, Zygopinae,
and Baridinae from the rest of Curculioni-
dae, i.e., the dorsally visible mesepimera,
and it is not clear that this character state
should be considered to be a secondary loss
in Trigonocolinae as presumed by Zheri-

VOLUME 102, NUMBER 4

Fig. 44-50. Legs. 44, Pachybaris porosa, right metatibia, ventral view. 45, Lobotrachelus subfasciatus, right
metatibia, ventral view. 46—S0. Claw. 46, 47, Parorobitis gibbus. 48, Orobitis cyaneus. 49, Pachybaris porosa.
50, Lobotrachelus subfasciatus.

952

khin and Egorov. The large to very large
scutellum, the presence of the wax powder
secretion, and the often carinate elytral in-
tervals of Trigonocolus do not fit the di-
agnosis of the Barididae. Also Zherikhin
and Gratshev (1995) give no evidence from
the wing structure of close affinity of Tri-
gonocolinae with the other 4 subfamilies.
We prefer to consider Trigonocolinae as
short-bodied representatives of the phyletic
branch also including Mecyslobini, Mag-
dalini, and Carciliini (i.e., Molytinae in re-
cent classifications) rather than to place
them close to Orobitidinae, Ceutorhynchi-
nae, Baridinae, or Zygopinae. Each of the
last four groups can be clearly identified
and is very different from each other. Ex-
cept for the Orobitidinae, the other taxa
have very large numbers of genera and spe-
cies and are distributed worldwide. Thus it
seems that additional data is needed for
treating Baridinae, Ceutorhynchinae, Zyg-
opinae and Orobitidinae as members of a
single natural group. Until the presence of
a longitudinal keel on the metasternum is
shown to be a true synapomorphy, through
rigorous character analysis, there is no rea-
son to consider Barididae sensu Zherikhin
and Egorov (1990) and Zherikhin and
Gratshev (1995) to be a natural group.
Detailed comparative analysis of the taxa
considered is far beyond the aim of this pa-
per and needs examination of much greater
number of taxa and structures. However, we
would like to summarize here some char-
acters of the Orobitidinae and their sup-
posed relatives. We believe that revealing
clear distinctions will allow better under-
standing of the groups, rather than lumping
them together based on characters whose
synapomorphic value has not been proven.

KEY FOR DIFFERENTIATION OF THE
OROBITIDINAE AND WEEVIL SUBFAMILIES
WITH DORSALLY VISIBLE MESEPIMERA:

1 (2). Mesepimera not clearly visible from above.
Meso- and metasterna fused. Abdominal
sternum | not longer than 2, nearly com-
pletely divided by hind coxae. Rostrum

Pal):

3 (4).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

more or less bent at antennal insertion,
straight or nearly straight in apical part, al-
ways narrower than basal part; not separat-
ed from frons by sulcus. Claws with ap-
pendages fused into entire flat process half
as long or nearly as long as claws them-
selves. Body globose or subrhomboidal,
very strongly convex dorsally. 1.8—3.5 mm.
Palearctic and Neotropics .....
Mesepimera usually clearly visible from
above (not visible in Palearctic species of
Baris Germar developing on crucifers and
in some southern Asian and South African
Ceutorhynchinae with subconical, strongly
apically narrowing prothorax). Meso- and
metasterna not fused, separated by distinct
suture. Abdominal sternum | longer than
2, never nearly completely divided by hind
coxae. Rostrum not conspicuously bent at
antennal insertion. Claws simple, toothed,
or appendiculate, in latter case appendages
not fused in an entire flat median process.
Body shape variables aa) see
Prosternum with median sulcus for recep-
tion of rostrum, often prolonged on meso-
or metasternum, and with more or less de-
veloped keels in front of fore coxae. Males
always without horn-like projections on
prosternum before coxae and without fovea
between them. Head capsule not spherical,
but transverse or slightly flattened dorso-
ventrally. Rostrum not separated from head
capsule by sulcus, neither conspicuously ta-
pering apically nor dilated to base. Anten-
nal funicle often with fewer than 7 flagel-
lomeres. Eyes usually more or less convex,
always separated on frons (usually by not
less than twice width of antennal scape).
Pronotum often with raised anterior margin
and with lateral (and often also discal) tu-
bercles. Basal margin of pronotum often
raised conjointly with basal margins of el-
ytra and crenulate. Scutellum small to mi-
nute, never transverse or subcordate. Elytra
usually broad, slightly longer than broad,
in tropical species often broader than long.
Lateral margin of elytron more or less an-
gularly emarginate above anterior part of
metepisterna, latter projecting dorsally.
Many species apterous in temperate zones.
Hind femora broadest, often saltatorial.
Tibiae mutic (except in some Zacladus
Reitter and Scleropterus Schoenherr), mu-
cro usually developed on middle and hind
tibiae in males, but often also on fore tib-
iae; in xerophilous species with narrow tar-
si, females may have longer mucro. Claws
usually toothed or appendiculate, often

Orobitidinae

955

VOLUME 102, NUMBER 4

Fig. 51-56. Female genitalia. 51-53, Parorobitis gibbus. 54—56, Parorobitis minutus. 51, Sternite and tergite
= sternite 8;

8. 52, 56, Spermatheca. 53, 55, Vagina with coxites and styli. 54, Sternite 8. Abbreviations: 8 st
arm of sternite 8; apd = apodeme of sternite 8; cx = coxite; d = duct: dlb = ductal

gland lobe; sty = stylus; vg = vagina.

8 t = tergite 8; arm
lobe; g = gland; glb

954 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

mdl

Fig. 57-62. Male genitalia and tergite 8. 57-59, Parorobitis gibbus. 60—62, Parorobitis minutus. 57, 60,
Tergite 8 and aedeagus, ventral view. 58, 61, Dorsal view. 59, 62, Lateral view. Abbreviations: 8 st = sternite
8; 8 t = tergite 8; ap = apodeme; mdl = median lobe; spg = spiculum gastrale; tg = tegmen.

VOLUME 102, NUMBER 4

4 (3).

5 (6).

connate; rarely single. Pygidium exposed
except in Palearctic Trichosirocalus Colon-
nelli. Host plants mostly herbaceous or li-
anas, larvae develop in non lignified tissue.
1.5—7.0 mm. World-wide including Arctic
(but not Subantarctic) tundra

Be es CATS TEM ote Ceutorhynchinae

Prosternum usually without median sulcus
for reception of rostrum, without keels in
front of fore coxae (except in Lobotrach-
elini, Zygopinae). Prosternum in males
sometimes with horn-like projections be-
fore coxae and deeply foveate between
them. Head capsule spherical or transverse.
Rostrum usually separated from head cap-
sule by sulcus and/or more or less strongly
widening to base, often also tapering and
flattened apically. Antennal funicle usually
with 7 flagellomeres. Eyes often flat, broad-
ly separated or contiguous. Anterior margin
of pronotum never raised, sides without tu-
bercles, disc more or less evenly convex,
sometimes with median tubercle, but not
sulcate. Basal margins of pronotum and el-
ytra neither raised conjointly nor crenulate.
Scutellum small to large, often convex,
transverse or subcordate. Elytra often much
longer than broad. Lateral margin of ely-
tron shallowly sinuate, not conspicuously
emarginate above anterior part of metepis-
terna. Wings usually functional except in
herpetobiont species. Fore femur usually
broadest, often strongly enlarged and den-
tate- then fore tibia strongly bent; legs not
saltatorial. Tibiae uncinate; in males, often
also mucronate. Claws usually simple, free
or connate; rarely single. Pygidium ex-
posed or concealed. Host plants herbs, very
often lianas and trees, larvae often in lig-
nified tissue. 0.9-35 mm. Worldwide ex-
cept northern taiga and tundra ........
Eyes contiguous or subcontiguous, very
large, flat, limited to dorsal half on head
capsule, not visible ventrally. Head capsule
at least slightly transverse. Rostrum not
separated from head capsule by sulcus,
more or less widening to base, often taper-
ing and flattened apically. (In Lobotrach-
elini, Zygopinae, prosternum deeply sul-
cate, body polished, globose, legs subu-
late.) Elytra elongate to transverse, usually
flattened dorsally, never glabrous or metal-
lic, usually with scales. Wings usually
functional. Venter more or less strongly
oblique to apex in lateral view. Femora of-
ten dentate, fore femur often strongly en-
larged, then tarsus elongated, especially in

males. Claws usually free, simple or

toothed. Predominantly tropical and sub-
tropical #2. hs. c) Reena are eae ee Zy gopinae
6 (5). Eyes widely separated on frons, always ap-
proximate ventrally. Head capsule spheri-
cal. Rostrum usually separated from frons
by sulcus. Prosternum rarely deeply sulcate
for reception of rostrum. Elytra usually
elongate, in many tropical forms short or
subglobose, body then often glabrous or
metallic. Vestiture often reduced or absent.
Wings reduced in herpetobiont forms. Ven-
ter not conspicuously oblique apically in
lateral view. Femora usually unarmed, fore
femur rarely strongly enlarged. Claws often
connate, rarely toothed or single. World-
wide except northern taiga and tundra ...
spate ackah Fats MEPeteh aes Stee OTE Ac Baridinae

ACKNOWLEDGMENTS

We thank O. Merkl of the Hungarian
Museum of Natural History (HMNH) for
the opportunity to study material in his
care. The other depositories of the type ma-
terial are abbreviated as follows: C. W.
O’Brien collection, Florida A & M Univer-
sity, Tallahassee, FL (CWOB); National
Museum of Natural History, Smithsonian
Institution, Washington, DC (USNM); Zoo-
logical Institute, Russian Academy of Sci-
ences, St. Petersburg (ZIN); Federal Uni-
versity of Parana, Curitiba, Brazil (OZUP).
We are also grateful to T. L. Erwin (De-
partment of Entomology, National Museum
of Natural History, Washington, DC), A.
Norrbom, and N. Woodley (Systematic En-
tomology Laboratory, ARS, USDA, Wash-
ington, DC) for reviewing this manuscript
and providing valuable suggestions. Re-
search by B. Korotyaev was funded in part
by Russian Federation for Basic Research,
Grant # 98-04-49763. Research by C. W.
O’Brien was funded in part by the USDA's
Cooperative State Research, Education and
Extension Service (FLAX 910001).

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chinae de |’ Afrique tropicale (Coleoptera, Curcu-
lionidae). Annales Historico-Naturales Musei Na-
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Crowson, R. A. 1955. The Natural Classification of the
Families of Coleoptera. Lloyd, London, viii+ 187

956

pp. [Reprinted from Entomologist’s Monthly
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Howden, A. T. 1995. Structures related to oviposition
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Hustache, A. 1936. Curculionidae: Cryptorrhynchinae.
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PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 957-963

FIRST DOCUMENTED RECORD OF MONOMACHIDAE
(HYMENOPTERA: PROCTOTRUPOIDEA) IN NEW GUINEA,
AND DESCRIPTION OF TWO NEW SPECIES

LUCIANA MUSETTI AND NORMAN FE JOHNSON

Department of Entomology, The Ohio State University, 1315 Kinnear Road, Columbus,

OH 43212-1192 U.S.A.

Abstract.—The occurrence of the genus Monomachus Klug (Hymenoptera: Proctotru-
poidea, Monomachidae) in New Guinea is documented for the first time. Two new species,
M. cracens and M. comptus, are described. These are distinguished from the three known
Australian Monomachus as well as the numerous New World species.

Key Words:
asitoid

The family Monomachidae is a small
group of parasitic Hymenoptera (Procto-
trupoidea) with austral disjunct distribu-
tion. Only two genera have been recog-
nized, Monomachus Klug and Tetraconus
Szépligeti. The latter is known only from
a single female specimen from Brazil. We
are studying the family with the goal of
elucidating the phylogenetic relationships
among species and understanding their
biogeography. Nineteen species-group taxa
have been described from the New World
from Mexico to Chile and Argentina, and
these were last revised by Schulz (1911).
Three additional species of Monomachus
were recognized in the recent revision of
the Australian fauna (Naumann 1985).
Masner (1993) noted that the family is also
to be found in New Guinea, but no species
have been described. The Papuan material
in fact represents two species distinct from
those known in Australia. We describe
them here because they do not appear to
be closely related to the numerous Neo-
tropical species; these will be the focus of
a separate paper.

Monomachus, New Guinea, Australia, Proctotrupoidea, Hymenoptera, par-

MATERIALS AND METHODS

Specimens for this study from New
Guinea are found in the collections of the
American Entomological Institute, Gaines-
ville, FL (AEIC) and the Bishop Museum,
Honolulu, HI (BPBM). Australian material
is found in the Australian National Insect
Collection, Canberra; Canadian National
Collection of Insects, Ottawa; the Museum
of Comparative Zoology, Cambridge, MA;
The Natural History Museum, London; and
The Ohio State University, Columbus.

The mandibles in Monomachidae are re-
markably diverse in shape. We use the fol-
lowing terms to describe their structure.
The mandible is generally divided into two
areas, a basignath and distignath, separated
by a subbasal groove. The groove allows
for greater range of abduction of the man-
dibles. The distal margin of the groove is
sometimes sharply marked; this corre-
sponds to the mandibular fold of Naumann
(1985) and the Basalfalte of Schulz (1911).
The lowermost part of the subbasal groove
is sometimes clearly visible as a sharply de-

958

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1.

fined diagonal sulcus. The basignath is con-
vex and invaginated into the oral cavity
when the mandibles are widely opened. The
nomenclature for thoracic sulci follows
Huber and Sharkey (1993).

The label data for all specimens is avail-
able in electronic format in Hymenoptera
On-Line at the URL http://ris.biosci.ohio-
state.edu/hymenoptera (Johnson and Muset-
ti, in press). The OSUC numbers quoted are
the individual unique identifiers for the
specimens in this study.

Monomachus cracens Musetti and
Johnson, new species
(Fig. 1)

Female.—Measurements for the two fe-
males are reported in pairs separated by a
slash; first value refers to female bearing ID
number OSUC 117646 (holotype @), sec-
ond value refers to OSUC 117653 (paratype
2). Fore wing length 5.0/5.8 mm. Color:
Head, lateral margins of pronotum, central
disc of mesoscutum on either side of notau-
li, metanotum, upper portion of mesopleu-
ron, all of metapleuron, lateral and posterior
portions of propodeum, entire metasoma
brown to dark brown; mesosoma otherwise

Monomachus cracens, \ateral habitus. Scale line = 1 mm.

light brownish yellow; coxae and trochan-
ters brownish yellow, legs otherwise dis-
tinctly darker brown; wing membrane clear.

Head: Clearly transverse in frontal view,
width across eyes 2.10/1.96X greater than
length, head width across the compound
eyes (2.48 mm/2.69 mm) not differing sig-
nificantly from head width across gena
(2.40 mm/2.70 mm respectively). Eye
height 3.54/4.71 xX malar length. Vertex
posteriorly with moderately dense deep se-
tigerous punctures, each separated by ap-
proximately diameter of one puncture;
punctures sparser, more irregularly sized
and spaced near ocelli; punctures densely
packed, but irregularly shaped near anten-
nal insertions; vertex otherwise smooth,
without microsculpture; outer margins of
ocelli bordered by distinct furrows; vertex
immediately behind ocelli largely glabrous.
Occipital carina widely separated from oral
margin ventrally, occiput with dense setig-
erous punctures continuing from vertex,
continuing on to gena. Gena in frontal view
not swollen. Frons (above clypeus, below
antennal sockets) slightly convex, with
dense setigerous punctures nearly through-
out, i.e., extending to clypeus and beyond

VOLUME 102, NUMBER 4

level of lower margin of compound eyes;
with a central area immediately above clyp-
eus smooth and glabrous, extending dorsal-
ly % height of frons. Clypeus convex, pre-
dominantly smooth and glabrous, dorsal
margin marked by line of large setigerous
punctures, hairs long and erect, margin
broadly angled and raised medially, epis-
tomal sulcus narrow and distinct medially,
broad and poorly defined laterally. Malar
space mostly punctate and pilose; malar
sulcus present. Mandible with sparse setig-
erous punctures on distignath, hairs elon-
gate; gradually narrowed apically; biden-
tate, teeth broadly rounded, ventral tooth
larger, longer than dorsal; distignath broad-
ly convex; subbasal groove fairly broad,
margins rounded and somewhat indistinct,
bottom of groove angled, appearing as a di-
agonal sulcus extending from posterior
mandibular articulation to lateral clypeal
lobes; basignath distinctly widest near an-
terior articulation, distignath reaching pos-
terior articulation.

Mesosoma: Pronotum predominantly
smooth and glabrous dorsally; transition be-
tween pronotal neck and collar broadly
rounded in profile, marked by a weak trans-
verse fold, laterally with sparse hairs and
fine punctures. Mesoscutum predominantly
smooth, very sparsely setose. Notauli ar-
cuate, separated from transscutal articula-
tion by short distance subequal to their
width, without crenulae. Parapsidal furrows
very indistinct. Scutellar pit wide, extend-
ing laterally beyond notaulus, deep, without
crenulae, crescentic, lateral margins of pit
angled posteriorly; central convex portion
of scutellum quadrate, separated from axilla
by simple sulcus; posterior margin of scu-
tellum with single transverse row of small
foveae. Scutellum, axillae smooth, nearly
glabrous. Dorsellum transverse, slightly
bulging, anteriorly with a narrow crenulate
sulcus, posteriorly mostly smooth. Meta-
postnotum (anterior to transverse groove at
base of propodeum) with pair of indistinct
or broadly rounded posterior projections.
Mesopleuron punctate and pilose nearly

959

throughout, save for smooth, glabrous, con-
cave area adjacent to mesopleural suture.
Mesepisternal groove indicated by a row of
foveae extending ventrally from near base
of fore wing to scrobal groove, then turning
anteriorly and extending below fore coxa,
widely separated from the discrimen ven-
trally; scrobal groove indicated by trans-
verse line of deep foveae; mesepisternum
finely punctate and pilose, relatively flat
ventrally, broadly rounded toward medial
articulations, discrimen shallow, inconspic-
uous, widened posteriorly to form small de-
pression near coxae, with densely pilose,
distinctly fingerlike lobe projecting above
depression on each side. Metapleuron dis-
tinctly separated from propodeum by row
of deep, broad foveae; densely setose. Pro-
podeum moderately globose in dorsal view,
punctate along midline and around posterior
end, dorsal surface otherwise smooth; dor-
solaterally rugose, densely setose; anterior
margin without teeth opposite metapostno-
tal projections. Fore wing with radial cell
closed, length 4.5/4.0 X width, base of m-
cu only slightly displaced basad of bifur-
cation of Cu,; in hind wing M between Cu,
and Irm/Rs absent.

Metasoma: First segment (petiole) dis-
tinctly elongate, slender, strongly curved;
remaining segments elongate, cylindrical,
not laterally compressed; second segment
longer than third, length of second 1.3/1.2
length of third; second and third segments
with tergite loosely wrapped around the
sternite, not closely appressed and leaving
very visible separation between tergum and
sternum; length of metasomatic segments as
a percentage of total length: 1: 23.6/23.5;
2222319/22 42°32 18 3/1816; 4s 33:
8.9/9.3; 6: 8.0/8.1; 7 to apex: 6.2/5.8.

Male.—Other than characters of sexual
dimorphism in metasoma and antenna typ-
ical for family, differing from @ as follows:
Body color generally brown above, yellow-
ish brown below; base of legs including
femora brownish yellow, brown apically.
Fore wing length 4.2—4.9 mm (* = 4.54,
SD = 0.268, n = 6). Sculpture on body

960

other than frons generally with punctures
much less dense and largely smooth. Pro-
podeum with sculpture along midline more
extensive, extending from anterior to pos-
terior margins. Antenna with dense short
hairs, tyloid on A4—A8 small, with fine
seta. Fore wing with closed radial cell,
length 3.7—4.8 < width (* = 4.20, SD =
0.379). Hind wing sometimes with a short
stem of M arising from Irm/Rs.

Material examined.—Holotype 2°: PAP-
UA NEW GUINEA: Wau, 1,250 m,
3.ix.1965, malaise trap, J.& M. Sedlacek
(OSUC 117646, BPBM). Specimen in good
condition; lacking A3—A15 from right an-
tenna, A4—A15 from left. Paratypes: PAP-
UA NEW GUINEA: (NE) Wau, Morobe
Distr., 1,200 m, 5.x.1962, malaise trap, J.
Sedlacek, 2 (OSUC 117653), 26.x.1961, 3
(OSUC 117654), 23.x.1965, J. & M. Sed-
lacek, 6 (OSUC 117647); (NE), Wau, Mo-
robe Distr., 1,050 m, 11.ix.1961, malaise
trap, J. Sedlacek, gd (OSUC 117651),
30.1x.1961, ¢6 (OSUC 117652); ; NE Wau,
Little Wau Ck., 1,200—1,300 m, 3.xii.1965,
malaise trap, J. Sedlacek, 6 (OSUC
117650); NE Karimui, 1,080 m, 14.vii.1963,
M. Sedlacek (OSUC 117655). All speci-
mens in BPBM.

Etymology.—The specific epithet cra-
cens, Latin for neat, slender, graceful, refers
to the elongate petiole in the female of this
species and its overall graceful habitus.

Comments.—Monomachus cracens may
be most easily distinguished from M. comp-
tus by the remarkably long and curved pet-
iole in the female and, in both sexes, the
single row of foveae at the apex of the scu-
tellum. The ventral lobes found at the pos-
terior end of the mesepisternum are very
distinct in male and female specimens;
these lobes are visible only as weak raised
margins of the depression in front of the
mid coxae in M. comptus. Additionally, the
lateral profile of the pronotum is mostly
smooth and rounded, with only a weak line
marking the separation of the neck and the
collar dorsally.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Monomachus comptus Musetti and
Johnson, new species
(Figs. 2—3)

Female.—Fore wing length 5.9 mm. Col-
or: Head generally dark brown, clypeus
and mandibles brown; lateral portions of
pronotum, disc of mesoscutum on either
side of notauli (including prescutum), pro-
podeum, mesopleuron and metapleuron
dark brown; scutellum, axillae, dorsal pron-
otum, metanotum lighter in color, varying
from brownish yellow to brown; metasoma
brown; antenna brown; legs generally
brownish yellow, hind leg beyond trochan-
ter brown; wing membrane clear.

Head: Transverse in frontal view (Fig.
2), width across eyes 1.95 X wider than
long, head width across the compound eyes
(2.57 mm) slightly more than width across
gena (2.46 mm). Eye height 3.13 malar
length. Vertex and gena with same pattern
of sculpture as found in M. cracens, but
generally more smooth and shallow; outer
margins of ocelli bordered by distinct fur-
rows. Occipital carina short, widely sepa-
rated from oral margin, occiput with dense
setigerous punctures. Gena in frontal view
not swollen. Frons flatter than M. cracens,
with fine setigerous punctures extending be-
low level of the compound eyes; central
smooth, glabrous area above clypeus larger,
mostly flat, extending through *% distance to
antennal insertions; short raised carina pre-
sent between antennae. Clypeus convex,
smooth and nearly glabrous medially, shal-
lowly punctate laterally, medially differen-
tiated from frons by clear epistomal sulcus,
without distinct line of large setigerous
punctures, apical margin broadly angled
and raised medially, epistomal sulcus poor-
ly defined laterally. Malar area mostly
smooth, with few sparse setigerous punc-
tures near compound eyes, malar sulcus dis-
tinct. Mandible with sparse setigerous
punctures on distignath, hairs elongate;
gradually narrowed apically; bidentate,
teeth broadly rounded, ventral tooth wider,
longer than dorsal; distignath broadly con-

VOLUME 102, NUMBER 4

961

Figs. 2—4.
matic segments 1-3, lateral view. 4, M. australicus, metasomatic segments 1—3, lateral view. Stippled lines on
metasoma indicate lower margin of sterna visible through cuticle of terga. Scale line = 1 mm.

vex; subbasal groove fairly broad, margins
rounded and somewhat indistinct, bottom of
groove angled, appearing as a diagonal sul-
cus extending from posterior mandibular ar-
ticulation to lateral clypeal lobes; basignath
distinctly widest near anterior articulation,
distignath reaching posterior articulation.
Mesosoma: Pronotum predominantly
smooth and glabrous dorsally; transition be-
tween pronotal neck and collar abrupt,
marked by ruga; ventrally with sparse hairs
and fine punctures. Mesoscutum predomi-
nantly smooth, very sparsely setose. Notau-
li arcuate, separated from transscutal artic-
ulation by short distance subequal to their
width, finely crenulate. Parapsidal furrow
present, extending over half length of me-

Characters of Monomachus species. 2, M. comptus, head, frontal view. 3, M. comptus, metaso-

soscutum from transscutal articulation. Scu-
tellar pit slightly narrower, reaching later-
ally as far as notaulus, with fine longitudi-
nal crenulae, only slightly crescentic, lateral
margins of pit broadly rounded; central
convex portion of scutellum slightly widest
anteriorly, separated from axilla by finely
crenulate sulcus, posterior margin of scu-
tellum with a subapical row of large punc-
tures, apically with 2—3 rows of smaller fo-
veae. Scutellum, axillae nearly smooth, gla-
brous. Dorsellum less bulging than M. cra-
cens, transverse, bordered anteriorly with
sculptured sulcus half its length, posterior
margin with narrow finely punctate band.
Metapostnotum with pair of indistinct or
broadly rounded posterior projections. Me-

962

sopleuron anteriorly punctate, setose ante-
riorly and ventrally, with a wide smooth
and glabrous area adjacent to mesopleural
sulcus extending to intersection of scrobal
and mesepisternal groove. Mesepisternal
groove indicated by finely crenulate fold
near wing base and by line of foveae ex-
tending anteriorly from scrobal groove,
these two sections widely separated, mes-
episternal groove broadly separated from
discrimen ventrally; scrobal groove indicat-
ed by transverse line of deep foveae; mes-
episternum finely punctate, sparsely setose,
protuberant ventrally, closely abutting cox-
ae so as to hide medial articulations, dis-
crimen indicated by shallow longitudinal
invagination, widened posteriorly to form
small fusiform pit near mid coxae; margin
of pit slightly produced and raised laterally.
Metapleuron distinctly separated from pro-
podeum by row of small foveae; densely
setose. Propodeum moderately globose in
dorsal view, coriaceous throughout, with
scattered setigerous punctures, with vague
indication of median longitudinal carina,
posteriorly with longitudinal rugulae; dor-
solaterally densely setose, longitudinally ru-
gose; anterior margin without teeth opposite
metapostnotal projections. Fore wing with
radial cell closed, length 4.24 * width, base
of m-cu only slightly displaced basad of bi-
furcation of Cu,; in hind wing base of M
present arising from Irm/Rs.

Metasoma (Fig. 3): First segment slen-
der, fairly straight; remaining segments
elongate, cylindrical, not laterally com-
pressed; length of second segment 1.3 X
length of third; length of segments as a per-
centage of total metasoma length: 1: 22.3;
22 VOT 33: N83 As TAA Se 8; 62/947
to apex: 6.7.

Male.—Other than characters of sexual
dimorphism in metasoma and antenna typ-
ical for family, differing from female as fol-
lows. Body color displaying same pattern
as female except sometimes with less con-
trast between dark brown and brownish yel-
low areas. Fore wing length 4.8—6.2 mm (x
= 5.46, SD = 0.622, n = 4). Vertex with

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

more extensive smooth areas. Propodeum
with spiracle bulging, dorsolaterally some-
times with well defined longitudinal carina.
Antenna with dense short hairs, tyloid on
A4—A8 small, with fine erect setae. Fore
wing with radial cell closed, length 3.1—4.0
x width (« = 3.75, SD = 0.295). Hind
wing sometimes with M absent between
Cu, and Irm/RS.

Material examined.—Holotype 2: PAP-
UA NEW GUINEA: SE Mount Giluwe,
2,500—2,750 m, 30.v.1963, J. Sedlacek
(OSUC 117648, BPBM). Specimen in fair-
ly good condition; lacking A5—A15 from
right antenna, and A1l3—A15 from left. Ab-
domen broken, segments four and beyond
in gelatin capsule attached to pin. Paraty-
pes: PAPUA NEW GUINEA: Daulo Pass,
2,450 m, 22.xii.1978-8.1.1979, J. Sedlacek,
6 (OSUC 117079, AEIC); Mount Otto,
2,000 m, 22.xii.1978-9.1.1979, J. Sedlacek,
6 (OSUC 117077, AEIC); Tari Gap, nr.
Mount Hagen, 2,600 m, 29.i-4.11.1979, J.
Sedlacek, 6 (OSUC. 117080; “AEBI@);
Mount Giluwe, 2,800 m, 3.1-8.11.1979, J.
Sedlacek, ¢ (OSUC 117078, AEIC).

Etymology.—The specific epithet comp-
tus, Latin for ornamented, refers to the
more elaborate sculpture on the scutellum.

Comments.—See diagnosis under de-
scription of Monomachus cracens for the
most useful characters to distinguish the
two Papuan species.

DISCUSSION

Schulz (1911) distinguished the New
World species of Monomachus from those
in Australia by means of the curvature of
the petiole: strongly bowed in specimens
from America, straight in those from Aus-
tralia. Monomachus cracens (Fig. 1) clearly
does not conform to this rule. Both species
from New Guinea are very similar to M.
australicus Girault in terms of the structure
of the mandibles, the sculpture on the body,
the shape of the head, and the shape of the
clypeal margin. The color patterns observed
differ from that ‘‘yellow form’ described
by Naumann for specimens of M. australi-

VOLUME 102, NUMBER 4

cus from northern Queensland most notice-
ably in that the lateral lobes of the meso-
scutum are the same color, dark brown, as
the medial lobe. The ventral mesepisternal
lobes are also shared with all Australian
Monomachus. The Australian species are
immediately distinguishable on the basis of
the wing venation: the base of m-cu is
strongly displaced basad of the bifurcation
of Cu, in the fore wing. The vein is inserted
nearly in the middle of cell 2Cu (first sub-
discal cell). All of these Old World species
have the second and third metasomatic seg-
ments elongate, with the second longer than
the third. Neotropical Monomachus typical-
ly have the second segment much shorter
and apically widened, and the metasoma
beyond the petiole is laterally compressed.

ACKNOWLEDGMENTS

Thanks to D. Wahl (AEIC), G. Nishida
(BPBM), L. Masner (Ottawa), S. Lewis
(London), P. Perkins (Cambridge), and J. C.
Cardale (Canberra) for the loans of speci-

963

mens. This material is based upon work
supported by the National Science Foun-
dation under Grant No. DEB-9521648.

LITERATURE CITED

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

Johnson, N. E and L. Musetti. In press. Data ware-
housing architecture and tools for Hymenoptera
biodiversity informatics. Jn Austin, A. D., ed.,
Proceedings of the 4 International Hymenoptera
Conference. Canberra, Australia.

Masner, L. 1993. Superfamily Proctotrupoidea, pp.
537-557. In Goulet, H. and J. T. Huber, eds., Hy-
menoptera of the World: An Identification Guide
to Families. Agriculture Canada. Research
Branch. Publication 1894/E. Ottawa, Canada, 668
Pp-

Naumann, I. D. 1985. The Australian species of Mon-
omachidae (Hymenoptera: Proctotrupoidea), with
a revised diagnosis of the family. Journal of the
Australian Entomological Society 24: 261-274.

Schulz, W. A. 1911. Systematische Uebersicht der
Monomachiden. Memoirs of the International
Congress of Entomology 2: 405—422.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 964-968

DESCRIPTION OF THE PUPA OF ARMIGERES (LEICESTERIA) OMISSUS
(EDWARDS) AND A KEY TO THE LARVAE AND PUPAE OF THE
ARMIGERES OCCURRING IN NEPAL (DIPTERA: CULICIDAE)

RICHARD FE. DARSIE, JR.

Florida Medical Entomology Laboratory, University of Florida, 200 9th Street SE, Vero
Beach, FL 32962, U.S.A. (e-mail: rfd@ gnv.ifas.ufl.edu)

Abstract.—The pupa of Armigeres omissus (Edwards) is described for the first time.
Keys to the larvae and pupae of Armigeres species occurring in Nepal are added.

Key Words:

The pupal stage of species of the sub-
genus Leicesteria, genus Armigeres occur-
ring in Nepal were described by Darsie
(1998). Subsequently, a female of Armiger-
es (Leicesteria) omissus (Edwards) was dis-
covered in the collection of Nepal mosqui-
toes at the Florida Medical Entomology
Laboratory, Vero Beach, FL, a new country
record. Its pupa was briefly described by
Delfinado (1966) and Baisas (1974). A
more detailed description follows, based on
specimens from Thailand since none are
available from Nepal.

With this account, the pupae of all spe-
cies of Armigeres from Nepal have now
been described (Ramalingam 1987; Toma et
al. 1994; Darsie 1998, 2000). Inasmuch as
a detailed study of the larvae was a neces-
sary part of the pupal verification, a larval
key is also included. This is a revision of a
key by Darsie and Pradhan (1990). Since
then, four species have been added to the
fauna, i.e., Ar. (Arm.) theobaldi Barraud
(Pradhan and Darsie 1990), Ar. (Lei.) in-
choatus Barraud and Ar. (Lel.) digitatus
(Edwards) (Darsie et al. 1992) and Ar.
(Lei.) omissus, herein.

The pupa of Ar. omissus possesses the
generic and subgeneric characters given be
Darsie (2000). It is readily distinguished

Armigeres omissus, pupa, keys, Nepal

from pupae of the Nepal Armigeres species
by a combination of: the absence of seta 1-
P, the paddle fringe extending to near the
base, seta 6-II-V with thin branches and
seta 1-II with 17 or fewer branches.

METHODS AND MATERIALS

For procedures used in this study refer to
Darsie (1998). No pupae of Ar. omissus
were found in my collection from Nepal,
but specimens were borrowed from the
Walter Reed Biosystematic Unit, National
Museum of Natural History (NMNH),
Smithsonian Institution, with accompany-
ing larval exuviae for species verification.
In the description below br means branches
and Le and Pe mean exuviae of the fourth
instar larva and pupa, respectively.

DESCRIPTION

Armigeres (Leicesteria) omissus (Edwards)
(Fig. 1)

Position and size of setae as figured,
range and modal number of branches in Ta-
ble 1. Cephalothorax: Setae 1,3-CT long to
very long, thin, usually single (1,2); 6-CT
0.53—-0.82, * 0.66 length of 7-CT; trumpet
brown, reticulate, length 0.5—0.6 mm, index
1.54-2.50, x 2.05. Abdomen: Seta 1-Il
moderately long, with 10—17 br; 2-V-VII

VOLUME 102, NUMBER 4 965

PRorlde Toye

Fig. 1. Pupa of Armigeres (Lei.) omissus. A, Cephalothorax (left side). B, Metanotum and abdomen (dorsal
left, ventral right). Abbreviations: CT = cephalothorax; GL = genital lobe; Mr = paddle midrib; Pa = paddle;
T = respiratory trumpet.

966 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Pupa chaetotaxy of Armigeres omissus.
Abdominal segments
Cephalo

Seta thorax I II II IV Vv VI Vil Vill
0) - = 1 1 1 1 l

l i 2DY 5-10 (?) 10-17 (14) 2-8 (3) 25 (3) 2-6 (2) 1—3 (2) De 2) (C2) -

2 1 (2) 1 | l 1 1 1 _

3 ey 2D} l 1 N=7/ (6) =s)(G)) 1-3 (1) 27a) =

4 Don) 24 (3) 2-6 (4) 1-3 (1) IES Gls) 122) 1-3 (1) 1, 2 (1) We 22, ((Ib))
5 1-3 (2) l 2-5 (2) 1-5 (3) 1-3 (1) ee Gy) 122) 2 (GD) -

6 ] 1—4 (?) 2-7 (2) 35 (3) 14 (2) ils ZA) ls 2 (Ub) 14 (1) _

7 12119) 1 2@)* 2-5: (2)" 2, 32) 133) 24@€4) 1-3 (2) 1-3 (1) -

8 1-3 (2) = = BIC) BrayQy 2x 3@)y 220 @) 2-8 (?) 2-5 (5)
9 1 ] 1 1 1 1 Is) (2) =
10 1 (2) - 12) 2n@) 1-3 (2) 1-3 (1) 1 1 _

11 1 - = 1—3 (1) 1-3 (1) 1-3 (1) 1-3 (1) 1-3 (2) -
12 1 - - - - ~ = - ~
14 _ — - 1 1 l 1 tS 7 (Ob) 1

' Range followed in parenthesis by mode.

short, single, 0.22—0.40, « 3.0 length of 1-
V-VII; 3-ILIII stout, 0.62-0.91, x 0.75
length of following tergum; 5-IV,V stout,
more than 0.5 length of following tergum;
6-IJ]-V short, with thin branches, usually
double (1-5); 6-VI long, stout, sparsely
aciculate, single, seldom double; 9-VII
long, stout, aciculate, with 2—5 br, seldom
single; 9-VIII long, stout, aciculate, usually
with 5 br (2-5). Seta 12-IV was found in
only one pupa. Paddle: Length 1.10—1.27
mm, index 1.05—1.29, 1-P absent, except
one pupa with a seta-like spicule without
alveolus, fringe long, 0.19 mm, extending
to near base.

The description is based on the follow-
ing specimens, all from Thailand, depos-
ited in the NMNH: Chiang Mai Province,
Huey Muang Ban Ay, V-15-64, 12 LePe,
ex bamboo pot; Nan Province, Ban Pha
Hang, elev. 400 m, VIII-19-66, 22 LePe,
ex bamboo stump; Lampang Province, Doi
Pha Huat, elev. 420 m, V-21-68, 12 LePe,
ex bamboo stump. The Nepal specimen
was collected in Jhapa District, Kanchan-
bari, VIII-2-91, 1°, resting outdoors on
vegetation in primary forest (coll. no. 111-
x28):

KEY TO PUPAE OF ARMIGERES SPECIES OF
NEPAL

Ihe Seta 1-P small or absent; seta 5-IV-V at
least 0.5 length of following tergum; seta
6-CT shorter than 7-CT (subg. Leicester-

TGV. bbe odode ors eee 2
- Seta 1-P long, rather stout, if not, seta 5-
IV,V less than 0.5 length of following ter-
gum (theobaldi); seta 6-CT as long as or
longer than 7-CT (subg. Armigeres) 8
2(1) Paddle with fringe of long spicules on
outer margin extending to near base ... 3
~ Paddle with fringe of long spicules on
outer margin in apical 0.75 or less .... 6
3(2) Seta 6-II-V short, usually with 2 or more
thin branches’). «2 32.55 See eee 4
= Seta 6-II-V long, stout, single ....... 3
4(3) Seta 1-II with 28 or more branches; seta
1-I with thick unbranched base, 0.27 of
totallenctheepses eee digitatus (Edwards)
— Seta 1-II with 17 or fewer branches; seta
1-I with smaller unbranched base, 0.18 of
totaljlenothiesress eee omissus (Edwards)
5(3). Seta 3-IV usually 5- or 6-branched; seta
I= sparsely aciculate == 0-3-1)
SPS eM ees aca ahs, ‘ool OZ annulitarsis (Leicester)
— Seta 3-IV usually with 4 or fewer branch-
es; seta 1-II densely aciculate ......
aaah) OTE en fy Witte Veg Z wellh magnus (Theobald)
6(2). Seta 1-II with 24 or more branches; pad-

dle with large external lobe; seta 1-II-VII
subequal toxsetal 2a oe
dolichocephalus (Leicester)

VOLUME 102, NUMBER 4

- Seta 1-II with 21 or fewer branches; pad-
dle without large external lobe; seta 1 at
least 2.0 length of seta 2 on V-VII.... 7
Seta 3-CT with thin branches; seta 6-VI
single inchoatus Barraud
- Seta 3-CT with stout branches; seta 6-VI
usually with 2 or more branches ... .
sic Joust SD DESORBED TOR TLCe SP eRe dentatus Barraud
Setae 3-II, III and 5-IV, V shorter than
0.25 length of following tergum . .
theobaldi Barraud
- Setae 3-II, III and 5-IV, V longer than 0.5
length of following tergum
Seta 1-CT with thin branches, usually
double; seta 9-VII with 9 or more branch-
CS eee ae eee ICE. Pe aureolineatus (Leicester)
- Seta 1-CT stout, single or double; seta 9-
VII with 8 or fewer branches ....... 10
Seta 1-II small, usually with 5 or fewer
thin branches, 0.36 or less length of 3-II
od cages en ot DED U EERE kuchingensis Edwards
- Seta 1-II much thicker, with 6 or more
branches, 0.5 or more length of 3-II .. 11
1110). Seta 1-II pedunculate, brush-like,
branched in apical 0.7, with with 17 or
more branches ........ durhami Edwards
- Seta 1-II pedunculate or not, with 6-14
branches
12(10). Trumpet short and broad, index 1.2—1.7:
seta 3-VII closer to seta 4 than to seta 1
Seer a ts oes ciin subalbatus (Coquillett)
- Trumpet long, index 2.3 or greater; seta
3-VII closer to seta | than to 4

7(6).

8(1).

9(8).

10(9).

KEY TO FOURTH INSTAR LARVAE ARMIGERES
SPECIES OF NEPAL

(Partially adapted from Macdonald 1960)

1. Abdominal segment X with dorsal saddle
and very small ventral sclerotized plate
Sie Sask eae eee ne magnus (Theobald)
= Abdominal segment X with dorsal scler-

alized saddle Onllys .\3. ots She annie acs 2
2(1) Comb scales fringed with subequal spi-
TUES. ta i eatin Uae Sy ee ae at ae 3
— At least some comb scales with apical
SJOVCIYES | onrn pe ery GEE SARC Phan ee Aaah eee eee 8
3(2) Comb with more than 25 scales
55: 5 6 a OND ORO ROR eRe: inchoatus Barraud
= Comb with fewer than 25 scales...... 4
4(3) Comb with 18=25 scales =~... 4 62.
A Mate ke, Be ES Re annulitarsis (Leicester)
= Comb with fewer than 18 scales...... 5
5(4). Most comb scales rather pointed apically,
point fringed with subequal spinules ... 6

= Comb scales rounded apically, fringed
with subequal spinules.............. 7

967
6(5). Seta 1-X on saddle or close to it .....
Bede Tea ae Ea durhami Edwards
= Seta 1-X distinctly removed from saddle
BS PERS ROMEO fo “opis subalbatus (Coquillett)
7(5) Seta 1-III-VI reaching posterior margin of
following segment; seta 6-I with 3 or 4
[RINENES, 9.666 cielo + kuchingensis Edwards
- Seta 1-III-VI only reaching basal 0.25 or
less of following segment; seta 6-I with
D9) MONS. 255-55. kesseli Ramalingam
8(2) Comb with 11 or fewer scales ....... 9
= Comb with 12 or more scales....... 10
9(8) Seta 1-S very small, with 2 or 3 branches;
1-X small, not inserted on saddle .....
SOY ree eee aureolineatus (Leicester)
= Seta 1-S long, rather stout, single; seta 1-
X strong, inserted on saddle .......
Ny ae.’ bee syn cee oy. ER theobaldi Barraud
10(9). Abdominal segments I-VII with promi-
nent tubercles bearing setae .......
Bs gas ea isie sod cae 6 dolichocephalus (Leicester)
~ Abdominal segments I-VII without tu-
bercles ws: 4: Aaa vats a eee oes i
11(10). At least abdominal sterna I-IV with

large patch of fine spicules .......
digitatus (Edwards)
= Abdominal sterna I-IV without spicules

12(11). Seta 5-VIII with 3 or more fine branches:
comb scales with 2 or more apical spines

ae es sence ae omissus (Edwards)
= Seta 5-VIII stout, single or double; most
comb scales with single apical spine . . .

SERS s RR RY chanetined open dicws dentatus Barraud

ACKNOWLEDGMENTS

The author is indebted to the National
Geographic Society for their support of the
work, to T. Gaffigan, Walter Reed Biosys-
tematics Unit, for providing the specimens
used in this study, to B. Bower-Dennis for
the illustration, and to J. R. Rey and J. K.
Nayar for reviewing the manuscript. This is
Florida Agricultural Experiment Station
Journal Series No. R-06543.

LITERATURE CITED

Baisas, F E. 1974. The mosquito fauna of Subic Bay
Naval Reservation, Republic of the Philippines.
U.S. Navy, Headquarters First Medical Service
Wing Technical Report 72-2, 1-170.

Darsie, R. F, Jr. 1998. Descriptions of the pupae of six

968

species of Armigeres Theobald, subgenus Leices-

teria Theobald (Diptera: Culicidae) from Nepal.

Proceedings of the Entomological Society of

Washington 100: 234—246.

. 2000. Description of the pupae of five species
in subgenus Armigeres Theobald, genus Armiger-
es Theobald, with a key to species of the known
pupae of the subgenus (Diptera, Culicidae). Pro-
ceedings of the Entomological Society of Wash-
ington 102: 108-119.

Darsie, R. F, Jr. and S. P. Pradhan. 1990. The mos-
quitoes of Nepal their identification, distribution
and biology. Mosquito Systematics 22: 69-130.

Darsie, R. F, Jr., S. P. Pradhan, and R. G. Vaidya. 1992.
New species records from 1991 collections. Mos-
quito Systematics 24: 23-28.

Delfinado, M. D. 1966. The culicine mosquitoes of the
Philippines, tribe Culicini (Diptera, Culicidae).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Memoirs of the American Entomological Institute
(Ann Arbor) 7: 1—252.

Macdonald, W. W. 1960. Malaysian parasites
XXXVIII. On the systematics and ecology of Ar-
migeres subgenus Leicesteria (Diptera, Culicidae).
Studies of the Institute for Medical Research 29:
110-153.

Pradhan, S. P. and R. FE Darsie, Jr. 1990. New additions
to the mosquito fauna of Nepal. Journal of the
Institute of Medicine (Nepal) 12: 225-228.

Ramalingam, S. 1987. On the restriction of Armigeres
durhami Edwards and the description of Armiger-
es kesseli n.sp. (Diptera: Culicidae). Tropical Bio-
medicine 4: 55—65.

Toma, T., I. Miyagi, and N. Benjaphong. 1994. Rede-
scription of Armigeres (Armigeres) theobaldi
(Diptera: Culicidae). Mosquito Systematics 26:
11-18.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 969-973

NEW RECORDS OF FLEAS (SIPHONAPTERA) FROM
EASTERN WEST VIRGINIA

RALPH P. ECKERLIN AND HARRY EF PAINTER

Natural Sciences Division, Northern Virginia Community College, Annandale, VA

22003 U.S.A. (e-mail: reckerlin@nv.cc.va.us)

Abstract.—Nine species of fleas are reported for the first time from the boreal envi-
ronment in the mountains of eastern West Virginia. The new state records are Cerato-
phyllus gallinae (Schrank), Conorhinopsylla stanfordi Stewart, Epitedia faceta (Roths-
child), Hystrichopsylla tahavuana Jordan, Megabothris asio asio (Baker), M. quirini
(Rothschild), Nearctopsylla genalis genalis (Baker), Rhadinopsylla media Smit, and Tam-

iophila grandis (Rothschild).
Key Words:

We report nine flea species found in the
higher elevations of the Appalachian Moun-
tains of West Virginia along the eastern bor-
der with Virginia where boreal conditions
are found. There has been considerable
study of the biota of the Appalachian balds
(Mark 1958) and spruce and northern hard-
wood forests (Handley 1971, Stephenson
and Adams 1984, and references therein)
demonstrating plants and animals charac-
teristic of the boreal environments of more
northern regions. Results of these studies
suggested to us that fleas common to more
northern environments might be present in
West Virginia because of the presence of
appropriate mammal hosts and the exis-
tence of boreal conditions suitable for their
non-parasitic larval stages. Benton (1971,
1980) added seven new species to the West
Virginia list bringing the state total to 24
and further noted that an additional 20 fleas
not then known from the state should be
present. We add nine new flea species to the
West Virginia list with all but one expected
by Benton.

Appalachian Mountains, fleas, distribution, West Virginia

MATERIALS AND METHODS

Collections were made in the eastern
mountains of the Ridge and Valley Province
of West Virginia as often as could be ar-
ranged during all months of the year from
1984 to 1996. Bird and mammal nests were
placed in a Berlese funnel overnight in the
laboratory to extract fleas. Small mammals
were trapped using Sherman live traps and
snap traps. Live animals were placed in pa-
per bags containing paradichloro-benzene
crystals for a few minutes and then released
into plastic bags for identification, sexing
and a check for fleas still in the pelage. The
animals were then released in the vicinity
of their capture. Collection data were writ-
ten on the paper bag and the contents were
examined for fleas in the laboratory. Mam-
mals found dead were brushed over a white
enameled pan and fleas were placed in eth-
anol. Five nest boxes were placed about 3
m high in a stand of red spruce (Picea rub-
ens Sargent) on Alleghany Mountain in Po-
cahontas County at an elevation of 1,300 m
to monitor the fleas of flying squirrels.

970

White polyester fiber was provided as nest-
ing material and sampled as nearly monthly
as weather would permit for a year. Sam-
pling involved removal of only half of the
fiber from a nest box, placing it in a plastic
bag and replacing it with fresh fiber, thereby
permitting the maintenance of succeeding
generations of parasites. Fleas were re-
moved from the retained nesting material in
the lab and stored in ethanol. All fleas were
decolorized in 10% KOH, dehydrated in
ethanol, cleared in xylene and mounted in
Canada balsam. Fleas were identified using
keys developed by Benton (1983) and Hol-
land (1985). Mammal names follow Wilson
and Reeder (1993).

RESULTS

The following nine flea species new to
the state of West Virginia were collected.

Ceratophyllidae
Ceratophyllus gallinae (Schrank)

3 2 from the nest of Sialia sialis (L.), 26
Sept 1993, Pocahontas County, Top of Al-
legheny.

This European hen flea is a wide-ranging
species from Alaska and Canada (Holland
1985) as well as northeastern United States
(Benton 1980). Domestic chickens and na-
tive birds are the usual hosts. This is the
southernmost reported occurrence of this
species in the eastern United States.

Megabothris quirini (Rothschild)

1 2 ex Clethrionomys gapperi (Vigors),
15 Nov 1984, 1 2 ex C. gapperi, 20 April
1985, 1 2 ex Microtus pennsylvanicus 26
Sept 1993, all from Pocahontas County, Al-
leghany Mountain. 1 6, 2 2 ex C. gapperi
30 June 1985, 1 ex Peromyscus leucopus
(Rafinesque), 30 June 1985, Randolph
County, Spruce Knob Lake. 4 2 ex C. gap-
peri, 29 Aug 1985, 1 3 ex M. chrotorrhinus
(Miller), 9 Dec 1985, 3 2 ex C. gapperi 18
May 1986, all from Pendleton County,
Spruce Knob.

Hopla (1965) stated that M. quirini “‘is a
flea that has migrated into the boreal re-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

gions from the south following the retreat
of the Pleistocene glaciers.” It is found on
mice in seven genera (Holland 1985). Our
collections were primarily from microtines.
Five of 72 C. gapperi, 1 of 9 M. chrotor-
rhinus and 1 of 27 P. leucopus were in-
fested. In the eastern United States this flea
had been reported only from New York and
New England (Benton 1980). The nearest
record to those reported here is from south-
ern New York, over 500 km to the north-
east.

Megabothris asio asio (Baker)

1 3, 26 2 ex 9 Microtus pennsylvanicus
and 1 6 ex Peromyscus maniculatus (Wag-
ner), 18 Nov 1984; Pocahontas County, Al-
leghany Mountain; 2 2 ex 2 M. pennsyl-
vanicus, 1 3 ex P. maniculatus and 1 3 ex
C. gapperi, 20 Oct 1985 from Tucker
County, Dolly Sods; 1 6 ex M. chrotor-
rhinus, 9 Dec 1985 Pendleton County,
Spruce Knob; 1 6, 4 2 ex M. pennsylvan-
icus 30 Jun 1985, Randolph County, Spruce
Knob Lake.

Numerous collections were made at these
four sites during the Fall, Winter and Spring
months. This is another flea found primarily
on microtine hosts with a boreal distribu-
tion in eastern Canada and northeastern
United States. The previous nearest record
of this flea is in Pennsylvania. Thus, our
Appalachian records from West Virginia
extend the range about 300 km to the south.

Ctenophthalmidae
Conorhinopsylla stanfordi Stewart

1 36,1 2 ex Glaucomys volans (L.) nest,
10 Nov. 1985, Pocahontas County, Alle-
ghany Mountain.

This flea is blind and is primarily a nest
flea on flying squirrels. Other hosts include
squirrels in the genera Sciurus and Tamias-
ciurus (Holland 1985). It is known from
Ontario to Maryland so this is the south-
ernmost occurrence in the East. This flea is
not plentiful in collections, probably due to
a dearth of nest examinations of the flying
squirrel in winter. It was found in larger

VOLUME 102, NUMBER 4

numbers in more extensive nestbox arrays
in neighboring Virginia.

Tamiophila grandis (Rothschild)

2 2 ex Tamias striatus (L.), 25 Sep 1993,
12 Oct 1996, Pocahontas County, Top of
Allegheny.

As the name implies, this is a large flea
and a parasite of the chipmunk, of which 2
of 15 were parasitized. This flea is rare in
collections because it is a nest flea and is
rarely taken on the host. It occurs from On-
tario to southwestern Virginia where it was
recently reported by Eckerlin and Painter
01995):

Epitedia faceta (Rothschild)

2 2 from nest of G. volans, 19 Jan 1986,
Pocahontas County, Alleghany Mountain.

This nest flea is most abundant in the late
fall and early winter (Benton 1980). Both
northern and southern flying squirrels are
hosts but the flea is restricted to eastern
Canada and eastern United States, corre-
sponding closely with the range of G. vo-
lans (Holland 1985). The southernmost re-
ported occurrence of E. faceta is eastern
Tennessee in the Appalachians at high ele-
vation (Durden and Kollars 1997).

Nearctopsylla genalis genalis (Baker)

3 2 ex B. brevicauda, 9 Dec 1985, Pen-
dleton County, Spruce Knob; 1 @ ex B.
brevicauda, 3 Nov 1995, Pocahontas Coun-
ty, Alleghany Mountain, 4 2 ex Sorex fu-
meus Miller, 18 Nov 1984, Pocahontas
County, Top of Allegheny.

Benton (1980) refers to this flea as a fall
and winter flea and indeed our records are
all in the fall months. With a range from
Ontario and Illinois eastward to the Atlantic
seaboard states, N. genalis was previously
collected no further south than Pennsylva-
nia. The Pocahontas County site extends the
range of this species 300 km southward at
high elevations in the Appalachians. Two of
33 B. brevicauda and 1 of 2 S. fumeus were
infested.

O71

Rhadinopsylla media Smit

1 2 from nest of Neotoma floridana
(Ord), 17 Sep 1950, Pendleton County,
Franklin, collected by Price and Tipton.

The single damaged specimen is depos-
ited in the Monte L. Bean Life Sciences
Museum at Brigham Young University and
was kindly loaned us by Michael Hastriter.

Hystrichopsyllidae
Hystrichopsylla tahavuana Jordan

1 @ ex Blarina brevicauda (Say), 18
May 1986, Randolph County, Spruce Knob
Lake, and 1 2 from nest of Microtus penn-
sylvanicus (Ord), 25 Oct 1986, Pocahontas
County, Top of Allegheny.

This large flea was described from the
two host species we list. Scattered records
exist from Ontario to Tennessee. Only 1 of
33 B. brevicauda was infested so we regard
this as an uncommon flea.

DISCUSSION

Eight of the nine new records are species
Benton (1971) predicted would be found in
West Virginia. The only unexpected species
was Rhadinopsylla media whose nearest re-
ported occurrence is in Michigan and Min-
nesota (Benton and Timm 1976). Addition-
al comment is warranted by the consider-
ably greater physical separation between
this record and other specimens of this sel-
dom collected flea. Pfitzer (1950), in his un-
published master’s degree thesis reported
collecting a number of the closely related
Rhadinopsylla orama Smit 1957 from Wise
County, Virginia just north of the Tennes-
see/Virginia line. The number and location
of genal spines (of the R. media specimen,
Fig. 1) confirm placement of the West Vir-
ginia specimen in Rhadinopsylla and the
long suture between the metanotum and
metepimeron (Fig. 2) separates R. media
from the only other eastern species, R. or-
ama, which is found in the adjacent states
of Virginia, Pennsylvania, and Maryland
(Benton 1980), but was not observed in this
study. We agree with Benton and Timm

972

Figs. 1-2
eron.

Rhadinopsylla media. 1, Head. 2

(1976) that more specimens are required to
determine distribution of these poorly
known fleas.

Epitedia faceta, Hystrichopsylla tahavu-
ana, Megabothris a. asio, M. quirini,
Nearctopsylla g. genalis and Tamiophila
grandis have extensive distributions north
of West Virginia, but are restricted to high
elevations southward in the Appalachian
mountains. We regard these species and
their environments in West Virginia as bo-
real. Significant extensions of ranges were
demonstrated for N. g. genalis and M. a.
asio, previously not collected south of
Pennsylvania and M. quirini, not collected
south of New York. Conversely, Conorhin-
opsylla stanfordi and Ceratophyllus galli-
nae are widespread and not ecologically
bound to the Appalachians. Conorhinopsyl-
la stanfordi adults in the nests of flying
squirrels have a pronounced cold weather
peak of abundance and are absent for most
of the year (Benton and Day 1980). Many
species of birds serve as hosts of C. galli-
nae including many cavity nesters such as
bluebirds (Holland 1985) which are found
throughout the study area. The total number

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

“))))))) yo

, Metathorax showing suture between metanotum and metepim-

of species reported from West Virginia now
stands at 33. We believe that more of Ben-
ton’s (1971) hypothesized species will be
discovered in West Virginia with additional
collecting.

ACKNOWLEDGMENTS

We acknowledge with gratitude the tu-
telage and help of the late Robert Traub in
our studies. He made records available to
us, permitted us use of his reference collec-
tion and literature, taught us flea anatomy,
and constantly urged us to get out and do
more collecting. We thank the West Virgin-
ia Department of Natural Resources for
granting collection permits. We thank the
many students who helped us collect or
who donated specimens. Finally, we thank
the reviewers for comments and sugges-
tions that have greatly improved this paper.

LITERATURE CITED

Benton, A. H. 1971. An annotated list of the fleas (Si-
phonaptera) of West Virginia. Proceedings of the
West Virginia Academy of Science 40: 35-39.

. 1980. An atlas of the fleas of the eastern Unit-

ed States. Marginal Media, Fredonia, New York,

177 pp.

VOLUME 102, NUMBER 4

. 1983. An illustrated key to the fleas of the
eastern United States. Marginal Media, Fredonia,
New York, 34 pp.

Benton, A. H. and J. F Day. 1980. Seasonal changes
in the flea fauna of nests of the southern flying
squirrel, pp. 401—403. Jn Traub, R. and H. Starke,
eds., Fleas. Proceedings of the International Con-
ference on Fleas, Ashton Wold, Petersborough,
UK 21-25 June, 1977. A. A. Balkema, Rotterdam

Benton, A. H. and R. M. Timm. 1976. Description of
the female of Rhadinopsylla media Smit (Siphon-
aptera: Hystrichopsyllidae). Journal of Medical
Entomology 13: 473—475.

Durden, L. A. and T. M. Kollars, Jr. 1997. The fleas
(Siphonaptera) of Tennessee. Journal of Vector
Ecology 22: 13-22.

Eckerlin, R. P. and H. E Painter. 1995. First record of
Tamiophila grandis (Insecta: Siphonaptera) from
Virginia. Banisteria No. 6: 24—25.

Handley, C.O., Jr. 1971. Appalachian mammalian ge-
ography—Recent epoch, pp. 263-303. Jn Holt, P.
C., R. A. Paterson, and J. P. Hubbard, eds., The
Distributional History of the Biota of the Southern
Appalachians Part III. Vertebrates. Research Di-

973

vision Monograph 4, Virginia Polytechnic Insti-
tute and State University, Blacksburg, Virginia.

Holland, G. W. 1985. The fleas of Canada, Alaska and
Greenland (Siphonaptera). Memoirs of the Ento-
mological Society of Canada No. 130, Entomo-
logical Society, Ottawa, Canada, 631 pp.

Hopla, C. E. 1965. Alaskan hematophagous insects,
their feeding habits and potential as vectors of
pathogenic organisms. I. The Siphonaptera of
Alaska. Arctic Aeromedical Laboratory, Fort
Wainwright, Alaska, 267 pp.

Mark, A. E 1958. The ecology of the southern Appa-
lachian grass balds. Ecological Monographs 28:
293-336.

Pfitzer, D. W. 1950. A manual of the fleas of Tennes-
see, M.S. Thesis, University of Tennessee, Knox-
ville, 103 pp.

Stephenson, S. L. and H. S. Adams. 1984. The spruce-
fir forest on the summit of Mount Rogers in south-
western Virginia. Bulletin of the Torrey Botanical
Club 111: 69-75.

Wilson, D. E. and D. M. Reeder, eds. 1993. Mammal
species of the world, A taxonomic and geographic
reference, 2d Edition., Smithsonian Institution
Press, Washington, D.C., 1206 pp.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 974-990

SAWFLIES (HYMENOPTERA: TENTHREDINIDAE) DESCRIBED BY
BENJAMIN D. WALSH, WITH NOTES ON THEIR HOSTS AND BIOLOGY

ALEXEY G. ZINOVJEV AND DAVID R. SMITH

(AGZ) Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Rus-
sia (e-mail: zag@zisp.spb.su); (DRS) Systematic Entomology Laboratory, PSI, Agricul-
tural Research Service, U.S. Department of Agriculture, % National Museum of Natural
History, MRC-168, Washington, DC 20560-0168, U.S.A. (e-mail: drsmith@sel.barc.
usda.gov)

Abstract.—The 15 species of sawflies described by Benjamin D. Walsh in 1866 are
identified. Lectotypes are designated for six species: Euura salicisovum, Euura salicis-
nodus, Nematus salicispomum, Nematus salicisdesmodioides, Nematus salicispisum, and
Nematus quercicola. Neotypes are designated for four species: Pristiphora grossulariae,
Euura perturbans, Nematus inquilinus, and Nematus hospes. Most species described by
Walsh are the gall-forming Nematinae on willows. Host plants, types of galls, and bio-

logical notes are presented where known.

Key Words:
Walsh

Benjamin D. Walsh described 15 species
of sawflies, all of the subfamily Nematinae
of the Tenihredinidae: Pristiphora grossu-
lariae (Walsh 1866a) from Davenport,
Iowa, and 14 species from near Rock Is-
land, Illinois, in his paper treating insects
associated with galls on willows (Walsh
1866b). Three of the latter species, Nematus
mendicus, N. fur, and Pristiphora_ syco-
phanta, have free-feeding larvae which
were only incidental in galls using them as
a pupation site. All other species described
in that paper are gall formers. The interpre-
tation and fixation of the identity of the
Walsh species by designation of types are
especially important in the study of North
American gall-forming sawflies because
they are among the oldest names available.
Walsh was the first in North America to de-
scribe galls and present biological infor-
mation.

Walsh’s collection was lost in the Chi-

Nematus, Pristiphora, Euura, Eupontania, sawflies, galls, willow, Salix,

cago fire of 1871 (Mallis 1971). However,
one specimen of Pontania, labeled as the
type of ‘““Nematus s-pomum Walsh” is in
the Museum of Comparative Zoology at
Harvard University, and more Walsh ma-
terial is in the Academy of Natural Sciences
of Philadelphia. None of these specimens
have ever been treated as syntypes and none
bear Walsh’s original identification label but
the specimens deposited in Philadelphia fit
the descriptions and labeling of the speci-
mens collected and described by Walsh. It
is improbable that this material had been
collected by anybody else. Walsh wrote that
he had sent a part of his material to E. Nor-
ton (at Philadelphia): “‘I sent a normal fe-
male of E. perturbans, and a female of E.
s. ovum to Mr. Norton, along with many
male and female varieties of Nematus s. po-
mum, each specimen numbered, but none of
them named’? (Walsh 1866b: 254). All
specimens are glued on triangular or pen-

VOLUME 102, NUMBER 4

tagonal (in case of Nematus salicispomum)
cardboard points and each bear a small
square label with printed letters “‘Ill.”’ (rare
in handwriting). None of them have iden-
tification labels written by Walsh himself.
The labeling varies and identification labels
(if any) were added by Norton or, for Pon-
tania species, also by Marlatt. Most speci-
mens deposited in Philadelphia have small
squares with numbers, and we assume that
these are the specimens sent by Walsh to
Norton.

In this paper we try to clarify the prob-
lems concerning sawfly species described
by Walsh. Lectotypes are selected for six
species: Euura salicisovum, E. salicisnodus,
Nematus salicispomum, N. salicisdesmo-
dioides, N. salicispisum, and N. quercicola;
a neotype is selected for Pristiphora gros-
sulariae, and lectotypes of E. salicisovum,
N. salicispomum, and N. salicisdesmodioi-
des are designated also as neotypes of E.
perturbans, N. hospes, and N. inquilinus
Walsh, respectively. Type material is still
absent for Euura salicisgemma, E. salici-
sovulum, N. mendicus, N. fur, and Pristi-
phora sycophanta.

Of the 14 names proposed by Walsh, sev-
en of them can be considered valid: Euura
salicisovum (= E. perturbans), E. saliciso-
vulum, E. salicisnodus, Eupontania salicis-
pomum (= E. hospes), E. salicisdesmodioi-
des (= N. inquilinus), E. salicispisum (= N.
quercicola), and Pristiphora sycophanta.
Pristiphora grossulariae, N. mendicus, and
N. fur are currently treated as synonyms of
Pristiphora rufipes Serville, 1823, Nematus
oligospilus Foerster, and Amauronematus
histrio Serville, 1823, respectively. How-
ever, placement of the latter two needs con-
firmation.

Most of the gall-making sawflies (and
midges) described by Walsh have abbrevi-
ated names. He named many of his species
like “‘Nematus s. pomum’” or “Euura s. no-
dus.” Some authors considered them as un-
available trinomens or used different spell-
ings for the same species (e.g., Marlatt
1896, Rohwer 1909, Ross 1951, Smith

975

1979). For example, one of the most com-
mon species creating apple-like galls in
eastern North America has been placed ei-
ther in Nematus or Pontania under the
names *“‘s. pomum Walsh,” “‘salicis-pomum
Walsh,” ‘“‘pomum Walsh,” ‘‘s-pomum
Walsh,” or “hospes Walsh (= s. pomum
Walsh).”’

Undoubtedly, “‘s.”” is only an abbrevia-
tion Walsh used. For the first species in any
genus, either of midges or sawflies, Walsh
clearly showed that ‘‘s.”” means “‘salicis.
For instance: ‘“‘Gall Salicis gemma, n. sp.”
and then “Euura s. gemma, n. sp.”’ (Walsh
1866b: 250, No. 16). In fact, these species
were described as trinomens like ‘“‘Euura
salicis gemma.’ According to the Interna-
tional Code of Zoological Nomenclature
(1985), these are valid names and we
should accept them, expanding the abbre-
viation and fusing “‘salicis’”’ with the third
word. Within sawflies, these names are: Eu-
ura salicisgemma, E. salicisovum, E. sali-
cisovulum, E. salicisnodus, Nematus sali-
cispomum, N. salicisdesmodioides, and N.
salicispisum.

Acronyms used for museums are: USNM
= National Museum of Natural History,
Smithsonian Institution, Washington, DC;
MCZ = Museum of Comparative Zoology,
Harvard University, Cambridge, MA;
ANSP = Academy of Natural Sciences of
Philadelphia, PA. Species headings are the
Walsh species in their original combina-
tions.

9

SPECIES DESCRIBED BY WALSH

Pristiphora grossulariae Walsh was de-
scribed from Davenport, Iowa (1866a). All
other sawfly species described by Walsh
were published in a paper on insects reared
from willow galls (1866b) “‘found on sev-
eral species of Willow in the neighborhood
of Rock Island, [linois’’ (Walsh 1864).

Pristiphora grossulariae Walsh
1866a: 123
(Figs. 1—2)
Type material—Described from ‘‘four
males and forty-nine females” and “forty

976

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-2.

3

larvae of various sizes.’’ One female at
ANSP probably belongs to the type series
of this species. It is mounted similar to oth-
er specimens from Walsh’s collection (Figs.
1—2), and bears Norton’s identification la-
bel. We hereby select it as neotype; it is

Sas
i don ns
SET isc A

Pristiphora grossulariae, neotype. 1, Dorsal view. 2, Lateral view.

labeled ‘‘Pristiphora grossulariae W. fe-
male.”

Valid name.—Pristiphora rufipes (Ser-
ville 1823) (= Pristiphora grossulariae
Walsh). Pristiphora rufipes was previously
attributed to Lepeletier; see Blank and Tae-

VOLUME 102, NUMBER 4

ger (1998) for correct authorship of species
previously attributed to Lepeletier.

Host.—Ribes spp.; larvae feed on differ-
ent currant species, but particularly on Ri-
bes grossularia L.

Notes.—This is a well-known currant
pest introduced from Europe. Larvae are
external leaf feeders.

Euura salicisgemma Walsh 1866b: 248,
250 (No. 16)
(Figs. 3—6)

Type material.—Described from 34 galls,
2 larvae, 1 male (reared May 5), and 1 fe-
male (reared May 2). Specimens possibly
lost.

There are no specimens under the name
“salicisgemma”’ in ANSP, but there are six
specimens under the name Euura orbitalis
Norton. All of them are from Illinois, each
bearing a small square “‘Ill.”” These six
specimens include one male of Euura,
which we cannot identify to species, one
female of Euura salicisovum Walsh, one
Phyllocolpa sp., and three females of E. or-
bitalis Norton (= salicisgemma Walsh).
One of the E. orbitalis specimens is labeled
we. Am Ent. Soc. Collection’’;
“TYPE”; “Euura orbitalis’”; “‘Euura orbi-
talis Norton [Norton’s label]’’; and a label
“Cannot be Type. Type came from
“Conn.” SAR” [S.A.Rohwer] (Figs. 3—6).
This specimen cannot be treated as the type
of Norton’s Euura orbitalis because Norton
described it from N.Y. (Brooklyn) and
Conn., and we cannot treat it as a type of
Walsh’s species. We were unable to find the
type of Euura orbitalis Norton elsewhere.
We doubt if any of these specimens belong
to Walsh’s material because they are pinned
and all other Walsh specimens that we have
seen are glued to points.

It would be desirable to select neotypes
(preferably from reared material) for Euura
orbitalis Norton and Euura_ salicisgemma
Walsh, to fix the usage of these names, but
we do not have reared material at present.

Valid name.—Euura orbitalis Norton

977

1862 (= Euura salicisgemma Walsh, syn-
onymized by Norton 1867).

Host.—According to Walsh (1866b: 248,
250), this species creates bud galls on Salix
humilis Marsh.

Notes.—This species is related to the
Holarctic Euura mucronata (Hartig), but E.
orbitalis is much paler colored (“‘head pale
luteous’’) than any of the European species
of Euura. Both species belong to the sub-
genus Gemmura E. L. Smith (1968), which
is characterized by a short ovipositor and
the habit of making bud galls. Euura orbi-
talis is separated from E. mucronata and E.
nigrella Rohwer by a short sawsheath in
dorsal view strongly narrowed at center
with very short medial flange (Fig. 4).

We studied specimens from Canada
(Quebec and Ontario) and U.S.A. (Illinois,
Missouri, New Hampshire, and New York).

Euura salicisovum Walsh 1866b: 248,
251-252 (No. 17)
(Figs. 7-10)

Type material.—Described from numer-
ous galls, 7 larvae, 10 males, and 5 females,
bred 16—27.

The lectotype female, here designated, is
glued on a whitish triangle and labeled
“Til.”?; “18°. Two paralectotype males are
labeled 19”; ‘‘Euura s. ovum male IIl.
Walsh” [on the underside ‘‘gall maker’’].
Another paralectotype male is labeled only
“TH.” (without a number). Deposited in
ANSP.

The female lectotype fits Walsh’s de-
scription. It is in rather good condition, but
the flagellum of both antennae, the entire
left hindtarsus, the apical parts of the right
hindtarsus, the apex of the left midtarsus,
and the apex of the left forewing are miss-
ing (Fig. 7). The lower part of the thorax is
not visible. The identification labels at-
tached to the male paralectotypes are in
Norton’s handwriting. There is little doubt
that the selected lectotype belongs to the
type series of Euura salicisovum. The only
other possibility is that it might be a syn-
type of Euura perturbans Walsh (a syno-

978 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 3-5. Euura orbitalis, specimen labeled “‘Ill.”’ 3, Dorsal view. 4, Abdomen, dorsal view. 5, Head, dorsal
view.

VOLUME 102, NUMBER 4

Fig. 6.

Euura orbitalis, head, front view.

nym of E. salicisovum), but, according to
Walsh, E. perturbans should be generally
darker than this female.

Valid name.—Euura salicisovum Walsh.

Host.—According to Walsh on “S. cor-
data.”’ Its currently accepted name is Salix
eriocephala Michaux (= cordata Muhlen-
berg not Michaux) (Argus 1997). The galls
are ‘““‘monothalomous, spongy, growing
from the side of the twig.’’ Galls obviously
belonging to this species were common on
S. eriocephala around Ithaca, NY, in 1997
(Fig. 10).

Notes.—The species is pale colored and
usually larger than Euura orbitalis. It is
characterized by its longer ovipositor, saw-
sheath in dorsal view with a more or less
gradually attenuate medial flange, and the
entire sawsheath triangular in outline with
the apical hairs distinctly bent apically (com-
pare Figs. 4, 9). In both sexes, it is separated
from E. salicisnodus (together with most of
other Euura species) by its distinctly con-
vex, glabrous and strongly shining inner or-
bits (Fig. 8) and, in dorsal view, the distinct
inner orbit margins above the eye.

For further notes, see also Euura salici-
sovulum Walsh.

Euura perturbans Walsh 1866b: 254

Type material.—Described from two
males and five females bred from cecido-

O72

mylidous galls (“‘S. strobiloides, O. S.; “‘S.
batatus Walsh,” S. rhodoides Walsh (galls
of the preceding years); and one from a bud
gall on “Vitis cordifolia.”

We did not find appropriate specimens in
ANSP to select a lectotype; however, to fix
the usage of this name, we hereby select the
lectotype of Euura salicisovum as neotype
for E. perturbans. See E. salicisovum for
labels on the specimen and Figs. 7-9.

Valid name.—Euura salicisovum Walsh
(= Euura perturbans Walsh).

Notes.—Euura perturbans was synony-
mized by Marlatt (1896). According to
Walsh, the specimens of E. perturbans
should be distinguished from E. saliciso-
vum only by “‘the dorsum of the abdomen
varying from honey-yellow, including the
lateral plates, through obfuscated [sic], to
deep black with the lateral plates also
black.”’ We consider this a color variation
and accept the previously proposed synon-
ymy under E. salicisovum.

Euura salicisovulum Walsh 1866b: 248,
253 (No. 18)

Type material.—Described from 30 galls
on Salix humilis and 6 larvae. Probably all
lost.

Valid name.—Euura
Walsh.

Notes.—The galls are the same as those
of Euura salicisovum; the larvae, according
to Walsh, are distinguished from those mak-
ing galls on Salix eriocephala being “all
decidedly pale greenish,” while those of E.
salicisovum were “all decidedly yellow-
ish.”” No galls were found by Walsh on Sa-
lix discolor which is closely related to S.
humilis.

The taxonomic status of this species and
its host plant specificity are not quite clear.
Galls of this type occur not only on Salix
eriocephala and S. humilis (including var.
tristis), as stated by Walsh, but also on S.
discolor and S. petiolaris. All such galls
seem to be associated with a bud, which is
however left unmodified, the swelling al-
ways situated just below the bud.

Specimens that seem to be conspecific

salicisovulum

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 7-9. Euura salicisovum, lectotype. 7, Dorsal view. 8, Head, front view; arrow indicates inner orbit. 9,
Apex of abdomen, dorsal view.

with E. salicisovum are known from Illinois Euura salicisnodus Walsh 1866b: 248,
and Virginia, but all of them are either cap- 253-254 (No. 19)

tured or reared from undetermined willows. (Figs. 11-12)

We are not sure if they belong to E. sali-

cisovum or to the complex of the sibling Type material.—Described from galls on
species involving E. saliciovulum. Thus, we ‘‘31 affected twigs,’ undetermined number
cannot designate a neotype yet. of larvae, and two males.

VOLUME 102, NUMBER 4

Fig. 10. Galls of Euura salicisovum on stems of Salix eriocephala, from near Ithaca, NY.

The lectotype male, here designated is at-
tached to a whitish elongated triangle and
labeled “Ill” [in handwriting, on the un-
derside ““‘B D W #3’’]; “‘Euura salicis nodus
Walsh.”’ (Fig. 3). Deposited in ANSP. It is
in rather good condition with only the apex
of the left flagellum and the left hindtarsus
missing.

Valid name.—Euura salicisnodus Walsh.

Host.—According to Walsh “Salix lon-
gifolia.”’ Its currently used name is either
Salix interior Rowlee (= longifolia Muhl-
enberg) or Salix exigua ssp. interior (Row-
lee) Cronquist (see Argus 1997). The galls
are “‘a mere enlargement of the twig, po-
lythalamous, pitchy inside, with its cells all
internal” (Fig. 12).

Notes.—The male lectotype fits the de-
scription well. It is separated from other
males of Euura described by Walsh, by
“black spot enclosing the ocelli is larger,
and is confluent with the eye for its entire
length, leaving no orbit between them.”’

This species is probably related to E. atra

Jurine, a species introduced from Europe
together with its host plants Salix alba L.
and S. fragilis L. and their hybrids. Both
species are characterized by the rather flat
inner orbits and by the narrow sawsheath
gradually tapering to its apex in dorsal
view. The longest lateral hairs are distrib-
uted throughout the entire apical and ven-
tral part of the sheath (in lateral view) and
a glabrous medial flange is lacking. Euura
salicisnodus is separated from E. atra by its
extensive paler coloration and the galls
which form an enlargement of the twig. Eu-
ura atra does not produce enlargements of
the twig; its galls are practically unnotice-
able.

Examined specimens are from Illinois,
New York, Michigan (?), Ohio, and Canada
(London, Ontario). Two species described
from the western United States, Euura mac-
gillivrayi Rohwer and Euura exigua E. L.
Smith, are associated with the same willow
species (or very closely related ones) and

982 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 11-12. Euura salicisnodis, \ectotype. 11, Dorsal view. 12, Gall on stems of Salix interior, from near
Ithaca, NY.

VOLUME 102, NUMBER 4

they might prove to be only geographical
forms of E. salicisnodus.

Nematus salicispomum Walsh 1866b: 248,
255-256 (No. 20)
(Figs. 13-16)

Type material.—Described from very nu-
merous specimens of galls, an undeter-
mined amount of larvae, 26 males and 46
females “‘bred April 16—25 and one female
bred many years ago.”

The lectotype female (Figs. 13—16), here
designated, is on a pentagonal piece of
brown cardboard, labeled ‘“‘Ill’’; ‘‘12’’;
“Return to Am. Ent. Soc.”’; “‘Pontania po-
mum Walsh’’; “‘N. s. pomum Walsh.” De-
posited in ANSP.

In the ANSP, there are 11 paralectotypes,
6 females and 5 males, all mounted on sim-
ilar pentagonal cardboard points: 2 females
and 3 males are without a number, and 1
female is labeled “Hl:; 1527; “Return to
Am. Ent. Soc.” The heads of one male and
of one female are missing, otherwise they
are in good condition. One female labeled
eile EYPE 14013": “Cresson”; “Ne-
matus s-pomum Walsh”’; ““MCZ // Museum
of Comparative Zoology” is deposited in
the MCZ. Two females and 2 males, all on
similar cardboard points and similarly la-
beled “‘Ill.’’; “‘Return to Am. Ent. Soc.”
with two of them bearing C. L. Marlatt’s
identification labels ‘‘Pontania pomum
Walsh” are deposited in the USNM.

Valid name.—Eupontania salicispomum
(Walsh), new combination. See Zinovjev
(1993) for the definition of Eupontania.

Host.—The galls are on Salix eriocepha-
la Michaux (= S. cordata). A subsidiary
host plant might be Salix discolor Muhl-
enberg. Walsh wrote about it as follows:
“On S. cordata, (and very rarely on Salix
discolor)”; he found galls of Nematus sal-
icispisum and galls “‘so identical in appear-
ance with S. pomum’’ on the same bushes.”
“In both the above two cases a few discolor
bushes were growing in the midst of very
large numbers of S. cordata, the species on
which S. pomum is normally found.’’ These

983

data on its host plant specificity seem to be
reliable, but confirmation by rearing adults
is needed.

The galls are roundish (apple-like), tran-
sected by the leaf blade with a large part of
the gall visible from the upper side of the
leaf, but the larger part of the gall is situated
below leaf surface (illustrated by Zinovjev
and Smith 1999: 361, fig. 2).

Notes.—In this species, the hind tibial
spur is shorter than the apical tibial breadth,
the frontal area lacks hairs on its anterior
part, the inner orbits are glabrous and shiny
(Fig. 15) with hairs only along the eye mar-
gin, and the sawsheath in dorsal view is a
narrow triangle with rounded margins (Fig.
16). The frontal wall of the frontal area is
usually considerably protruding (e.g., in
reared specimens from New York), but in
specimens of the type series, the whole up-
per head in lateral view is rather flat.

Nematus salicisdesmodioides Walsh
1866b: 248, 257-258 (No. 20)
(Fig. 17)

Type material—Described from an un-
determined number of galls, “three or
four” larvae, 2 males and 8 females, bred
April 2-15.

The lectotype female (Fig. 17), here des-
ignated, is attached to an elongated triangle
and labeled ‘“‘Ill.’’; “13’’; ““Return to Am.
Ent. Soc.”’; ““‘Pontania desmodioides female
Wish.” [identification label of Marlatt]; “‘N.
desmodioides Walsh [identification label of
Norton]. Deposited in ANSP.

A paralectotype male is deposited in
ANSP. It is mounted similar to the female,
and it is labeled the same (with number 13),
but without an identification label.

Valid name.—Eupontania salicisdesmo-
dioides (Walsh), new combination.

Host.—Salix humilis Marshall.

Notes.—The galls are ‘“‘semicircular in
outline, sessile’? (transected by the leaf
blade and equally developed on both sides
of the leaf).

The female can be separated from other

984

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 13-16.

Eupontania by the following combination
of characters: sawsheath in dorsal view as
a long triangle with an acute apex and
straight sides and with hairs directed more
posteriorly than laterally (with an angle of
less than 90°); upper part of head in lateral

Eupontania salicispomum, \ectotype. 13, Dorsal view. 14, Head, dorsal view. 15, Head, front
view; arrow indicated inner orbit. 16, Apex of abdomen, dorsal view.

view densely covered by hairs thoughout,
hind tibial spurs shorter than apical tibial
breadth; and inner orbits with dense hairs.

An additional female was examined from
Massachusetts (deposited in ANSP) [seen
by Marlatt because of his label].

VOLUME 102, NUMBER 4

985

Fig. 17.

Nematus salicispisum Walsh 1866b: 248,
258-260 (No. 21bis)
(Figs. 18—20)

Type material.—Described from many
galls, larvae, two males, and three females.

The lectotype female, here designated
(Figs. 18-20), is attached to a long trian-
gular cardboard point, labeled “‘Ill.’’; “14”;
“Return to Am. Ent. Soc.’’ Two paralec-
totypes males are similarly labeled as the
lectotype; one is labeled number 14 as is
the lectotype, and the other is labeled num-
ber 15.

Valid name.—Eupontania salicispisum
(Walsh), new combination.

Host.—Salix discolor: The gall is illus-
trated by Zinovjev and Smith (1999: fig. 1).
They are small, rounded, and attached to
the sideveins, with a very small scar visible
from the upperside and concave below the
leaf surface.

Notes.—In the key to Palearctic species
(Zinovjev 1993) this species runs to the vi-

Eupontania salicisdesmodiodes, \ectotype, dorsal view.

minalis group (length of inner hind tibial
spur subequal to maximum width of hind
tibial apex and rather thin). It is character-
ized also by its small size; short ovipositor
(Fig. 20); sawsheath in dorsal view as a
short triangle with rounded sides (Fig. 19)
with the longest hairs directed more later-
ally than posteriorly and more strongly bent
near their apices; upper part of head in lat-
eral view with dense erect hairs, but front
wall of frontal area glabrous; antennal hol-
low glabrous and shining; and inner orbits
with hairs well developed.

Nematus quercicola Walsh 1866b: 260
(Figs. 21—24)

Type material.—Described from 2 males,
7 females reared from ‘‘an undescribed,
cabbage-like, polythalamous, Cecidomyi-
idous gall on the White Oak ...”

The lectotype (Figs. 21—24) is a female
labeled “‘IIl’’; ““N. quercicola Walsh” [Nor-
ton’s label]; ‘“‘Return to Am. Ent. Soc.”’;

986

Figs. 18—20.
Apex of abdomen, lateral view.

‘‘Pontania pisum Wlsh female” [Marlatt’s
identification label]. Deposited in ANSP.
Valid name.—Eupontania salicispisum
(Walsh) (= Nematus quercicola Walsh).
Notes.—Nematus quercicola was treated
as a synonym of the preceding species by

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Eupontania salicispisum, lectotype. 18, Dorsal view. 19, Apex of abdomen, dorsal view. 20,

Marlatt (1896), or by others (e.g., Smith
1979) as a nomen nudum because the only
character mentioned by Walsh to distin-
guish it from the preceding species was that
‘tall N. s. pisum went underground to pup-

>

ize’? and ‘‘N. quercicola pupized in the

VOLUME 102, NUMBER 4 987

22

Figs. 21-24. Nematus quercicola, lectotype. 21, Dorsal view. 22, Apex of abdomen, dorsal view. 23, Head
and thorax, lateral view. 24, Abdomen, lateral view.

988

gall.”” A biological feature such as this, by
formal reasons, could be treated as a “‘char-
acter,” and we should accept this name as
an available one. According to Walsh, this
species ‘“‘cannot be distinguished from the
gallmaking N. s. pisum male, female.”

Nematus inquilinus Walsh 1866b:
260-261

Type material.—Described from one
male, three females, bred from the cecido-
myiidous gall “‘S. rhodoides Walsh. No ap-
propriate material was found in ANSP. The
neotype female, here designated is the lec-
totype of Nematus salicisdesmodiodes
Walsh (Fig. 17).

Valid name.—Eupontania salicisdesmo-
diodes (Walsh) (= Nematus inquilinus
Walsh).

Notes.—This species was placed in syn-
onymy with Nematus salicisdesmodioides
Walsh (Marlatt 1896) and was listed as
‘“‘unplaced”’ by Smith (1979). The type ma-
terial was reared from galls on Salix hu-
milis, the host plant of Eupontania salicis-
desmodioides. To fix the usage of this name
we designate the lectotype of Nematus sal-
icisdesmodioides as neotype of Nematus in-
quilinus Walsh.

Nematus hospes Walsh 1866b: 261

Type material.—Described from one
male and two females from the gall of s.
strobiloides Osten-Sacken. No type mate-
rial was located, but there is no strict evi-
dence that absolutely all specimens under
the name WN. salicispomum belong to the
original type series. To fix the usage of this
name we select the lectotype of N. salicis-
pomum Walsh (Figs. 13—16) as the neotype
of N. hospes Walsh.

Valid name.—Eupontania salicispomum
(Walsh) (= Nematus hospes Walsh).

Notes.—According to Walsh ‘‘absolutely
undistinguishable from the normal type of
the gall-making N. s. pomum.” Marlatt
(1896) synonymized N. hospes, and these
species have always been treated as con-
specific.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Nematus mendicus Walsh 1866: 261

Type material.—Described from one
male, three females: ‘“‘one female bred May
2 from the Tenthredinous gall S. pomum n.
sp. of the preceding year’s growth, and an-
other female, August 5, from the Cecido-
myidous gall S. brassicoides Walsh of the
same year growth; the other female and the
male captured at large.’’ No type material
was located.

Valid name.—Nematus oligospilus Foers-
ter 1854 (= Nematus mendicus Walsh).

Host.—Willows, Salix spp.

Notes.—Benson (1962) synonymized
Nematus mendicus, thus treating N. oligos-
pilus as a Holarctic species. It is in a com-
plex of green Nematus species with external
leaf-feeding larvae. Further studies may re-
veal several species in this complex. Cur-
rently, we accept Benson’s synonymy.

Nematus fur Walsh 1866: 263

Type material.—Described from a single
male ‘“‘bred March 29 from an old bored
subpeduncled spherical gall, .57 inch in di-
ameter, made by Cecidomyia s. batataus
Walsh on S. humilis.” The type was not lo-
cated.

Valid name.—Currently, Nematus fur is
treated as a synonym of Amauronematus
histrio Serville 1823. However, its taxo-
nomic position is not quite clear. It is pos-
sibly a valid species of Amauronematus.

Notes.—This species was treated by
Ross (1951) as a synonym of Amaurone-
matus luteotergum Norton 1861, and by
Smith (1979) as synonym of A. histrio Le-
peletier 1823 (= luteotergum Norton).
However, these treatments do not fit exactly
the description of N. fur. According to
Walsh, Nematus fur has “‘black legs” and
‘“‘wings subhyaline, slightly tinged with fu-
liginous.”’ This dark coloration of the wings
and legs is rare in the Nematinae, and we
could not find appropriate specimens in col-
lections that fit the description of N. fur.
Otherwise, according to its morphology
(particularly microsculpture) and its ability

VOLUME 102, NUMBER 4

to bore for pupation into plant tissue, it al-
most certainly belongs to Amauronematus.

Pristiphora sycophanta Walsh 1866b: 263

Type material.—Described from a single
male “‘bred August 9 from a cocoon found,
July 27, inside the Cecidomyidous gall S.
brassicoides Walsh of the same year
growth.” The type was not found.

Valid name.—Pristiphora sycophanta
Walsh.

Host plants.—This is a common Pristi-
phora on willows. The cecidomyidous gall
from which the type was reared was on Sa-
lix interior.

Notes.—Numerous specimens were tak-
en in Clarke Co., VA, where the predomi-
nant willows around a pond were Salix ni-
gra Marsh. Malaise trap collections at this
locality included P. sycophanta specimens
throughout the spring and summer, indicat-
ing that there are several generation a year.
This supports our interpretation of P. sy-
cophanta, which Walsh stated was reared
from the same years growth.

ACKNOWLEDGMENTS

We thank D. Azuma (Academy of Nat-
ural Sciences of Philadelphia, PA) and S.
Cover (Museum of Comparative Zoology,
Harvard University, Cambridge, MA) for
allowing study of important material in
their collection, and I. M. Kerzhner (Zoo-
logical Institute, Russian Academy of Sci-
ences, St. Petersburg) for advice on usage
of the Walsh names. This study was sup-
ported in part by Smithsonian Instituion
Short-Term Visitors Grants, 1997 and1999,
to A.G. Zinovjev. We thank Terry Nuhn
(Systematic Entomology Laboratory, USDA)
for taking the photographs and Cathy An-
derson (Systematic Entomology Laborato-
ry, USDA) for arranging the plates. We ap-
preciate the comments of the following re-
viewers: Henri Goulet (Agriculture and
Agri-Food Canada, Ottawa) and T, J. Henry
(Systematic Entomology Laboratory,
USDA).

989

LITERATURE CITED

Argus, G. W. 1997. Infrageneric classification of Salix
(Salicaceae) in the New World. Systematic Botany
Monographs 52: 1-121.

Benson, R. B. 1962. Holarctic sawflies (Hymenoptera:
Symphyta). Bulletin of the British Museum (Nat-
ural History) Entomology 12: 381—409.

Blank S. M. and A. Taeger. 1998. Comments on the
taxonomy of Symphyta (Hymenoptera) (Prelimi-
nary studies for a catalogue of Symphyta, part. 4),
pp.141—174. In Taeger, A. and S. M. Blank, eds.,
Pflanzenwespen Deutschlands (Hymenoptera,
Symphyta). Kommentierte Bestandsaufnahme.
Verlag Goecke & Evers, Keltern, 368 pp., 4/5 ta.

International Code of Zoological Nomencalture. 1985.
University of California Press, Berkeley and Los
Angeles, 338 pp.

Marlatt, C. L. 1896. Revision of the Nematinae of
North America, a subfamily of leaf-feeding Hy-
menoptera of the family Tenthredinidae. United
States Department of Agriculture, Division of En-
tomology, Technical Series No. 3, 135 pp.

Mallis, A. 1971. American Entomologists. Rutgers
University Press, New Brunswick, New Jersey,
549 pp.

Norton, E. 1862. Notice of several new species of Ten-
thredinidae. Proceedings of the Entomological So-
ciety of Philadelphia 1: 143-144.

. 1867. Catalogue of the described Tenthredi-
nidae and Uroceridae of North America. Trans-
actions of the American Entomological Society 1:
193-280.

Rohwer, S. A. 1909. The sawfly genus Cryptocampus
in boreal North America. Journal of the New York
Entomological Society 17: 7—25.

Ross, H. H. 1951. Symphyta, pp. 4-89. Jn Muesebeck,
C. E W., K. V. Krombein, and H. K. Townes, eds.,
Hymenoptera of America North of Mexico, Syn-
optic Catalog. United States Department of Agri-
culture, Agriculture Monograph No. 2, 1420 pp.

Smith, D. R. 1979. Symphyta, pp. 3-137. In Krom-
bein, D. R., P. D. Hurd, Jr, D. R. Smith, and B.
D. Burks, eds., Catalog of Hymenoptera in Amer-
ica North of Mexico, Vol. 1. Smithsonian Insti-
tution Press, Washington, D.C.

Smith, E. L. 1968. Biosystematics and morphology of
Symphyta. I. Stem-galling Euura of the California
region, and a new female genitalic nomenclature.
Annals of the Entomological Society of America
61: 1389-1407.

Walsh, B. D. 1864. On the insects, coleopterous, hy-
menopterous and dipterous, inhabiting the galls of
certain species of willow. Part 1st.—Diptera. Pro-
ceedings of the Entomological Society of Phila-
delphia 3: 543-644.

1866a. Imported insects;—The Gooseberry

sawfly. The Practical Entomologist. A Monthly

Bulletin 1(12): 117-125.

990 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

. 1866b. On the insects, coleopterous, hyme- viminalis-group (Hymenoptera: Tenthredinidae).
nopterous and dipterous, inhabiting the galls of Zoosystematica Rossica 2: 145—154.
certain species of willow.—Part 2d and last. Pro- _Zinovjev, A. G. and D. R. Smith. 1999. Types and
ceedings of the Entomological Society of Phila- biological notes of the eastern North American
delphia 6: 223-288. sawflies of Pontania Costa and Phyllocolpa Ben-
Zinovjev, A. G. 1993. Subgenera and Palaearctic spe- son (Hymenoptera: Tenthredinidae) described by
cies groups of the genus Pontania, with notes on Marlatt, Dyar, and Rohwer. Proceedings of the En-

the taxonomy of some European species of the tomological Society of Washington 101: 359-371.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 991-1002

A NEW SPECIES OF THE SUBGENUS SABETHOIDES OF SABETHES
(DIPTERA: CULICIDAE) FROM VENEZUELA AND BRAZIL

Dusty A. MOosgEs, THERESA M. HOWARD, AND RALPH E. HARBACH

(DAM) Department of Biology, Cumberland College, 7000 College Station Drive, Wil-
liamsburg, KY 40769, U.S.A.; (TMH, REH) Department of Entomology, The Natural
History Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: t-howard@nhm.ac.

uk; r.harbach@nhm.ac.uk)

Abstract.—The adult male, pupa and larva of Sabethes (Sabethoides) conditus, n. sp.,
are described from localities in western Brazil and northern Venezuela. The species is
distinguished from Sabethes chloropterus (von Humboldt), which it closely resembles in

all life stages.

Key Words: Diptera, Culicidae, Sabethes,

Venezuela

Mosquitoes of subgenus Sabethoides
Theobald of genus Sabethes Robineau-Des-
voidy are some of the more common insects
in Neotropical forests, yet they are poorly
known and rarely studied. No significant
taxonomic research has been done on the
subgenus since Lane (1953). This is sur-
prising since one of the species, Sa. chlo-
ropterus (von Humboldt), has repeatedly
been found infected with St. Louis enceph-
alitis virus (Galindo et al. 1959) and is
known to transmit yellow fever virus to hu-
mans (Galindo et al. 1956). In addition to
Sa. chloropterus, the subgenus includes Sa.
glaucodaemon (Dyar and Shannon), Sa. tri-
dentatus Cerqueira, and at least 2 undescri-
bed species. With the addition of the new
species described in this paper, Sabethoides
now includes 4 formally recognized spe-
cies.

The new species described here very
closely resembles Sa. chloropterus in all
life stages, and undoubtedly has been con-
fused as this species in the past. For this

Sabethoides, new species, mosquito, Brazil,

reason, future study may indicate that it is
also a vector of arboviruses.

MATERIALS AND METHODS

This study is based on specimens bor-
rowed from the National Museum of Nat-
ural History, Smithsonian Institution,
Washington, DC. Observations of the adults
were made under simulated natural light.
Larval and pupal chaetotaxy were studied
using a combination of bright field and dif-
ferential interference contrast microscopy.
Measurements and counts were taken from
all specimens of the type series where the
structures in question were present. Num-
bers in parentheses represent modes of the
reported ranges unless indicated otherwise.
The form of presentation, descriptive ter-
minology and abbreviations used in the spe-
cies description follow Harbach and Knight
(1980, 1982) and recent papers published as
part of an ongoing revision of the genus
Sabethes (Harbach and Peyton 1991; Har-

992

A B C D

Fig. 1. Tibia and tarsus of midleg of Sabethes (Sa-
bethoides) conditus (A, B) and Sa. chloropterus (C,
D). A, C, Anterior; B, C, posterior. Scale = 2.0 mm.

bach 1991la, 1994, 1995a, b; Harbach and
Petersen 1992; Hall et al. 1999).

Sabethes (Sabethoides) conditus, Moses,
Howard, and Harbach, new species
(Figs. 1—4)

Sabethes (Sabethoides) sp 4 of Heinemann
and Belkin 1978: 373.

This species exhibits the diagnostic char-
acteristics of the subgenus noted by Har-
bach (1991a): midtarsus marked with white
scaling; prespiracular, upper proepisternal
and lower mesokatepisternal setae present;
prealar setae absent; and legs without pad-
dles.

Male.—Medium-sized mosquito with
brilliant metallic-coloured scaling; scales of
head capsule, thorax and abdomen very
broad and flat; scales of vertex with differ-
ent combinations of metallic blue, violet
and green reflections depending on angle of
light; scales of postgena, thoracic pleura
and coxae silvery white; antepronotum
ranging from dark gold to bright cerulean

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

blue depending on angle of light; scaling of
postpronotum similar to pleura but with a
slight golden hue; scutal and scutellar scales
metallic gold with cerulean reflections; me-
sopostnotum without scales; proboscis and
legs predominantly metallic blue and violet,
proboscis darker; wing entirely dark-scaled
with blue, gold and violet iridescence when
viewed from certain angles; abdominal ter-
ga primarily metallic gold with narrow pale
basal bands, from dorsal angles golden ar-
eas appear cerulean to green and basal
bands mauve to violet, basal bands broader
laterally and silvery-white in lateral view.
Head: Eyes joined above and below. Oc-
ciput with transverse row of short semi-
erect scales at back of head. Ocular setae
short, dark, close to margin of eye; 2 long,
bronzy, approximated interocular setae pre-
sent. Antenna: Dark; length 1.29—1.36 mm
(x = 1.34 mm), significantly shorter than
proboscis; pedicel large, surface silvery pu-
bescent; flagellum rather strongly verticil-
late, proximal whorls with 9 or 10 setae,
longest setae about 0.33 length of antenna.
Clypeus and frons without setae and scales,
dark. Proboscis long, slender, straight;
length 2.00—2.10 mm (* = 2.05 mm); distal
0.3 gradually expanded laterally, becoming
twice as broad as proximal part; dark-scaled
with ventral patch of yellowish white scal-
ing extending 0.5 to 0.8 from base. Maxil-
lary palpus short, about 0.08 length of pro-
boscis; silvery-scaled dorsally, bare ven-
trally. Thorax: Integument brown. Dorsum
with dark setae on anterior promontory (10,
11), antepronotum (11-13), supraalar area
(10-15), scutellum (3,4 long setae on lat-
eral lobes; 2 long setae projecting down-
ward from midlobe) and mesopostnotum
(4). Pleura with prespiracular (2), upper
proepisternal (1), lower mesokatepisternal
(1,2) and upper mesepimeral setae (9-11);
prespiracular setae dark, others yellow or
golden. Lower part of proepisternum with-
out scales, scales on upper part contiguous
with scales on anteprocoxal membrane;
scales present on upper portion of postpro-
coxal membrane; mesopleuron with scales

VOLUME 102, NUMBER 4

covering all but lower anterior margin of
mesokatepisternum, extreme dorsal margin
of postspiracular area, narrow ventral and
upper posterior margins of mesepimeron
and mesomeron; scales absent from meta-
pleuron, metameron and postmetacoxal
membrane. Wing: Length 3.2 mm; dorsal
scales broader and slightly asymmetrical on
anterior and distal veins; alula with fine pil-
iform scales on margin distally; calypters
without setae. Halter: Scabellum without
scales, integument pale; pedicel and capi-
tellum dark-scaled. Legs: Coxae and tro-
chanters with silvery-white scales, trochan-
ters with some dark scales dorsally at apex;
femora dark above and golden below; all
tibiae and fore- and hindtarsi entirely dark-
scaled; anterior surface of midtarsus (Fig.
1) white-scaled from middle of tarsomere 2
to middle of tarsomere 4 (paratype from
Brazil with white scaling encircling tarso-
mere 3 and part of 4). Forefemur about 1.2
length of proboscis, same length as midfe-
mur, about 1.3 length of hindfemur; hind-
tibia about as long as hindfemur, hindtar-
somere 1 about 1.2 length of hindfemur.
Ungues small, simple, black. Abdomen:
Coloration as noted above, sternal scales
distinctly larger and less recumbent than
tergal scales. Genitalia (Fig. 2): Tergum
VIif (ventral in position; not illustrated)
with deep V-shaped emargination at middle
of posterior margin, posterior border on ei-
ther side of emargination with 3 or 4 irreg-
ular rows of long close-set setae, posterior
half of surface before setae covered with
recumbent spatulate scales that rather
abruptly grade into a cluster of much larger
scales on posterolateral corners, scales of
cluster about half length of marginal setae.
Tergum and sternum IX fused laterally,
forming a complete ring of sclerotization;
ninth tergal lobes small, not produced, each
bearing 3 flattened setae with apices bent
laterad; interlobular bridge moderately
broad. Gonocoxite stout, width more than
half length in lateral view, tapered in distal
third, tergomesal surface membranous, dis-
tal sternal area covered with scales and

993

short setae, bearing 3 long tergomesal setae
below basal mesal lobe; basal mesal lobe of
irregular shape, roughly trapezoidal in ven-
tral (tergal) view, partly covered with small
slender setae and bearing 2 large setae at
caudolateral angle. Gonostylus (side view)
large, about two-thirds length of gonocox-
ite; stem stout, short, less than half length
of head; head as illustrated, bearing 5 well
developed lobes (A, C, E, M, M’), a slight-
ly produced lobe B and an elongate tergal
fimbriate process; lobe A, elongate tapered
process arising tergomesal of base of lobe
M, bearing several short stout setae at apex;
lobe E, rather short conical lobe borne mes-
ally at bases of lobes A and M, with large
blade-like seta and 2 smaller needle-like se-
tae at apex; lobe B, lateral minutely spicu-
late swelling associated with sternomesal
margin of lobe E, bearing 3 large sternally
directed setae with distal ends flattened and
expanded subapically; lobe M, irregular
partially subdivided median apical lobe
bearing broadly V-shaped tergal process
with short striated arms and more proximal
long slender tergally directed fimbriate pro-
cess comprised of partially fused and coa-
lesced filaments with free ends bending to-
ward base of gonostylus; lobe M’, elongate
irregular process arising from sternolateral
margin at base of lobe M, subapical tergo-
lateral margin with several short thickened
setae and more proximal small flattened
sigmoidal seta, sternal margin with laterally
compressed projection bearing a narrowed
retrorse apex; lobe C, large mesally slanted
process arising from sternal margin at base
of lobe M’, bearing pattern of decumbent
spine-like spicules on mesal side of apical
margin and cluster of long sternally direct-
ed filaments on lateral side at proximal end.
Aedeagus longer than wide, widest in basal
half; submedian tergal arms fused to form
broad median tergal bridge; apical tergal
arms fused to form narrow apical bridge;
median sternal plate rather membranous,
apex not markedly flared, hood-like. Proc-
tiger (in lateral view) with broad basal
sclerotization (tergum X); paraproct slender,

994 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

lateral
tergal

lateral

tergal

Fig. 2. Male genitalia of Sabethes (Sabethoides) conditus. Aspects as indicated for (A) gonocoxopodite; (B)

tergum IX; (C) gonostylus; (D) proctiger; (E) aedeagus, with parameres attached. A, B, C, E, M and M’ =
gonostylar lobes; BML = basal mesal lobe; fp = fimbriate process; tms = tergomesal setae. Scale in mm.

VOLUME 102, NUMBER 4

N

LE

\

Yigg Teilard.

Fig. 3. Fourth-instar larva of Sabethes (Sabethoides) conditus. A, Head, dorsal (left) and ventral (right)
aspects of left side. B, Dorsomentum. C, Thorax and abdominal segments I-VI, dorsal (left) and ventral (right)

aspects of left side. D, Abdominal segments VII-X, left side. E, Comb scale. A = antenna; C = cranium; CS
= comb scale; Dm =

dorsomentum; P = prothorax; M = mesothorax; S = siphon; T = metathorax; I-X =
abdominal segments; 1-15 = setal numbers for specified areas, e.g., seta 5-C. Scales in mm.

995

996

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Numbers of branches for setae of fourth-instar larvae of Sabethes (Sabethoides) conditus.
Head Thorax Abdominal segments
Seta oc re, ae P M T I il
0) 1 10-13 — -- a i
1 1 5,6(5,6) 3—5(4) 3 2,3(3) Dy
2 — | 1 1 DES (2-3)) 3,4(3)
3 1 2 l 2,3(2) 1 1
4 1 6—9(7) i 2-4(2) 2—4(3) 4—6(4)
5 D, 1,2(1) ] 3—5(3) 3 5,6(6)
6 1 2, 1 1—3(2) 5,6(6) 5—8(5)
7 3—5(3) 7,10(10) i 10—12(10) 5-7(5) 46(6)
8 DES (23) 9-11(10) 5,6(6) 10—12(10) — 1,21)
9 y 2,3(2) 1,3(1) 7,8(7,8) 1 2
10 1e2@) 6—8(7) 5,6(5) 1 ] 1
11 6—-9(9) 1,2(2) 1,2(2) 1,2(1) 6—-8(7) 4
12 3,4(3) 1 1 1 — ]
13 1 — 13-16 5,6(6) 6,7(6) 5—7(6)
14 4,5(5) 2—4(4) 12—-19(19) — — -——
15 1—3(1) = = = = =

apex with 2 or 3 appressed teeth, rather
large subapical area distinctly sclerotized,
bearing 6—23 cercal setae.

Female.—Not definitely associated with
the male (see Systematics, below); 2 pre-
sumed females from Brazil resemble the
males from Venezuela except for the ab-
sence of a pale patch on the ventral surface
of the proboscis and in having slightly more
extensive pale scaling on the anterior sur-
face of the midtarsus, extending from prox-
imal 0.2 of tarsomere 2 to apex of 4.

Egg.—Unknown.

Larva, fourth instar (Fig. 3).—Exhibiting
the subgeneric characters noted by Harbach
1991a; similar to Sa. chloropterus (cf. Fig.
4 in Harbach 1991b); character and place-
ment of setae as figured, numbers of
branches in Table 1. Head: Slightly wider
than long, widest in posterior half; length
about 1.0 mm; width about 1.3 mm. Occip-
ital foramen widely V-shaped with arms ex-
tending dorsolaterally to point laterad of
level of seta 9-C, margins heavily tanned,
ventrocaudal margin with collar-like edge.
Anterior margin of labiogula weakly den-
ticulate; hypostomal suture complete, gent-
ly curved. Dorsomentum (Fig. 3B) short,
with 6,7 teeth on either side of median

tooth, median tooth and most lateral tooth
larger than others. Maxilla, including apical
tooth, about half length of head, with 8-10
lateral teeth, first lateral tooth larger than
the others which become progressively
smaller. Setae 4,6-C single, simple; 7-C
usually triple; 9-C inserted at level distinct-
ly posterior to 10-C; 10-C single or double;
14-C without thickened branches; 15-C in-
serted cephalad of 14-C near anterior mar-
gin of labiogula, long with 1—3(1) branches.
Antenna: Short, cylindrical; slightly ta-
pered distally, length about 0.28 mm. Seta
1-A single, simple, borne dorsally about 0.8
from base, length about 2 times width of
antenna at point of insertion. Thorax: In-
tegument hyaline, smooth. Setae 0,1,8,14-P,
1,13,14-M and 5,8-T with multiple short
flexible branches; 11-P,.M,T single or dou-
ble, 11-R.M with slender flexible branches,
11-T with stiff thickened branches; 13-T
with multiple branches, about length of tho-
rax. Abdomen: Integument hyaline, smooth
except for rows of minute spicules before
comb. Seta 1-I,II with thickened branches,
1-III-V long, double with one branch mark-
edly shorter; 2-I laterad of seta 1, 2-II-VII
well mesad and anterior to seta 1; 6,7-I, I
similar, strongly developed with multiple

VOLUME 102, NUMBER 4 997
Table 1. Continued.
Abdominal segments
= we) vy hh. vw. = ww wt hoo
1 1 1 1 1 1 =
2 2 2 2 223 (2) 5—8(5) 2
3,4(3,4) 3,4(4) 3)9\G)5) 3,4(3) 23D) 1 3,4(3,4)
1 ] 1 1 1 6,7(6,7) 2
] 1 8—11(8) HED En) 1 1 2
4,5(4,5) 3,4(4) 3,4(3) D2) 2—5(3) 2 —
1 ] 1 1 10—-14(13) —
9-12 11-16 11—13(12) 5—8(7) ] a —
2,3(3) 2 1,2(2) 14-17 19-25 1-S, 1
23 (2-5) 23) 2 SPX) 2) la-S, 1
1 1 1,2(1) i 1,2(2) 2a-S, 1
4,5(4,5) 3,4(3) 2-6 15-21 1 a —
1 1 | 1 1 oe
5-7(7) 5—6(5,6) 5,6(5) 29—34(29) 3,4(3) = =

aciculate branches, 6-III-VI long, single and
progressively shorter; 5-II-VI and 13-I-V
strongly developed, stellate, with stiff acic-
ulate branches; 13-III-V apparently with
more branches (x = 17) than Sa. chlorop-
terus (x 12); punctures present on seg-
ments III-V. Segment VIII: Comb a single
row of 21—26 long slender spine-like scales
(Fig. 3E) without fringe of minute spicules,
scales close-set with flared bases sometimes
partially joined. Siphon: Relatively long
and slender, gradually tapered from base to
apex; moderately tanned, surface more or
less evenly covered with short rows of mi-
nute spicules; length about 1.3 mm, width
at mid-length about 0.1 mm, index about
13. Pecten of about 30 fine filaments ex-
tending from below level of insertion of
seta 1-S to point distal to seta la-S. Seta 1-
S inserted 0.25 from base of siphon, la-S
includes 1 or 2, usually 2, and 2a-S includes
2 or 3, usually 3, single setae; seta 2-S
stout, slightly sinuous and minutely forked
at tip (not apparent in Fig. 3D). Segment X:
Saddle borne on posterodorsal quarter of
segment, surface more or less evenly cov-
ered with rows of minute spicules; length
about 0.2 mm; siphon/saddle index about

6.5. Setae 1—4-X equally well developed;
1,3,4-X double, 2-X usually triple (3,4).
Pupa (Fig. 4).—Exhibiting the subgener-
ic characters noted by Harbach 1991a; sim-
ilar to Sa. chloropterus; character and po-
sitions of setae as illustrated, numbers of
branches in Table 2. Cephalothorax: Light-
ly tanned. Seta 1-CT strongly developed,
double, branches sigmoidally curved with
hooked tips; 5-CT also well developed,
double or triple, more often triple. Trumpet:
Moderately tanned, slightly flattened but lit-
tle if at all expanded laterally; length 0.32—
0.39 mm (x 0.35 mm), width at mid-
length 0.11—0.14 mm, index 2.78—-2.91 (x
= 2.81); pinna short, length about 0.08 mm.
Abdomen: Lightly tanned, terga and sterna
darker anteriorly; length about 3.7 mm.
Seta 1-I well developed, with about 80 ter-
minal branches; 6,7-I long, 6-I single, lon-
ger than 7-I; 2-II-VII lateral to seta 1; 5-
II, with multiple branches, much shorter
than 5-IV-VI, 5-IV-VI very long, more than
twice length of following tergum; seta 4-IV
short, with multiple branches, generally
with more branches than in Sa. chloropte-
rus which usually has 1 or 2; 10-II and 13-
VI present on both sides in 2 available

998

Table 2.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Numbers of branches for setae of pupae of Sabethes (Sabethoides) conditus.

Cephalothorax

Abdominal segments

Seta (3h I I Il IV
0 — — 1 1 1
1 2 10-14(10)* 5,6(6) 2,3(3) 2,3(2)
2 2,3(3) 1 I 1 1
3 23323) 1,2(1) l 1 2-4(3)
4 2-4(3) 5—7(6) 5,6(5) 3-5(4) 2-6
5 3 1 6-11 5—8(5) i
6 1-3(2) 1 1 1 1
7 2-4(3) 3 3,4(3,4) 4,6(4,6) 3-8
8 3,4(3) = a 4-7 2
9 1 1 1 1 1
10 1,2(1) at 1,2(1) 120) 2
11 3,4(3) 1 2;3(2;3) 3 1,2(2)
12 12 @,2) = = = aa
13 — _ _ — =
14 -- — = == —

* Primary branches.
+ Presumed (see legend for Fig. 4).
+ Alveolus only.

specimens, 13-VI quite variable in size and
number of branches (3—12), apparently ab-
sent in Sa. chloropterus. Genital lobe:
Lightly tanned, length (male only) about
0.36 mm. Paddle: Lightly tanned, asym-
metrical, broadest at base, tapered distally,
inner part narrow, less than 0.5 width of
outer part, and minutely spiculate at apex;
length about 0.7 mm, width at widest point
about 0.45 mm, index 1.55.
Systematics.—Sabethes conditus resem-
bles Sa. tridentatus and Sa. glaucodaemon,
but differs in having distinctly shorter upper
meseperimeral setae. These setae reach the
lateral area of the mesopostnotum in Sa.
conditus and extend to near the middle of
the mesopostnotum in the other two spe-
cies. The male of Sa. conditus also differs
in the presence of ventral pale scaling on
the proboscis, the pattern of pale scaling on
the mid-tarsi, and the structure of the gen-
italia. Sabethes conditus has undoubtedly
been misidentified as Sa. chloropterus in
the past, mainly because most studies have
focused on females that appear to be essen-
tially isomorphic for the two species.
Knight and Stone (1977) list four junior
synonyms for Sa. chloropterus: Sabethes

nitidus (Theobald), Sabethoides confusus
(Theobald), Sabethoides rangeli (Surcouf
and Gonzales-Rincones), and Sabethoides
imperfectus (Bonne-Webster and Bonne).
Sabethes nitidus was included in error be-
cause the lectotype designation by Belkin
(1968) validated this nominal taxon as a
distinct species of subgenus Sabethes, and
the paralectotype male that Knight and
Stone used as a basis for synonymy with
Sa. chloropterus has no taxonomic status.
We know from concurrent studies of Sa-
bethoides that the holotype female of Sa-
bethoides imperfectus is not conspecific
with Sa. chloropterus and will be formally
synonymized with another species in a fu-
ture revision of the subgenus (by R.E.H.).
The holotype females of Sabethoides con-
fusus and Sabethoides rangeli are both in
very poor condition, and even though Sa-
bethoides confusus has no midlegs, both
specimens clearly belong to a species of Sa-
bethoides other than Sa. glaucodaemon or
Sa. tridentatus. The holotype of Sabethoi-
des rangeli has faint white scaling on the
ventral surface of the proboscis (0.5 to 0.8
from base), a feature that is not known in
females of Sa. chloropterus or the pre-

VOLUME 102, NUMBER 4 999
Table 2. Continued.
Abdominal segments Paddle

Vv VI VII VII IX le
1 1 1 1 — =>
2 2,3(2) 23 (6) —- — a
1 1 1 — — —
DS (2) 2,3(2) 3,4(3) a = =
3—6(6) 2 1 1,2(1,2) = =
1 1 3—5(4) — _ —
1E2(2) 1252) 5,6(5,6) = wer
5-8 1272) 1,2(1,2) — — —
2,3(2) 6—9(7) 10—15(14) — — —
1 1 13—15(14) 23—27(26) = =
1 1 Asi) = = ==
1,2(2) 4-6(4) 1 — — ae
el 12 at we ie he

sumed females of Sa. conditus examined in
this study. In the absence of any evidence
to suggest that the holotype of either Sa-
bethoides confusus or Sabethoides rangeli
may be conspecific with Sa. conditus rather
than Sa. chloropterus, these nominal forms
are retained in synonymy with the latter
species.

The male of Sa. conditus is easily distin-
guished from the male of Sa. chloropterus
in having a lesser amount of pale scaling
on the proboscis and midtarsus, and a very
differently constructed gonostylus. The pro-
boscis of Sa. conditus has ventral pale scal-
ing extending from 0.5 to 0.8 beyond the
base whereas in Sa. chloropterus it extends
from 0.3 to 0.8 beyond the base. Similarly,
the midtarsus has pale scaling on the ante-
rior surfaces of tarsomeres 2—4 in Sa. con-
ditus (also on the posterior surface of tar-
someres 3 and 4 in the paratype from Bra-
zil) whereas it occurs on the anterior sur-
faces of tarsomeres 2 and 3 and the
posterior surfaces of tarsomeres 1—4 in Sa.
chloropterus.

The two females of Sa. conditus from
Brazil are presumed to belong to this spe-
cies because specimen numbers on the la-
bels indicate that they were collected and
reared from pupae along with the paratype

male. Apparently no collection records, ad-
ditional data or associated pupal exuviae
exist in the Smithsonian Institution to con-
firm this (J. Pecor, personal communica-
tion). The two females have the same pat-
tern of pale scaling on the midlegs and ap-
pear to resemble verified females of Sa.
chloropterus in overall habitus (the midlegs
of Sa. chloropterus sometimes also have a
variable amount of pale scaling on the pos-
terior surfaces of tarsomeres 2—4). If these
two specimens are females of Sa. conditus,
then the female of this species is indistin-
guishable from the female of Sa. chlorop-
terus.

No diagnostic characters were found to
distinguish the larva and pupa of Sa. con-
ditus from those of Sa. chloropterus, but
some partially differential characteristics
have been noted that may prove to be useful
once additional material becomes available
for further study. In the pupa, seta 4-IV has
2—6 branches whereas specimens of Sa.
chloropterus only have 1 or 2 (usually 2),
and seta 13-VII, a unique feature among
mosquitoes, is present on both sides in the
two available specimens of Sa. conditus. In
the larva, the mean sum of branches of seta
13-III-V on one side is 17 compared to 12
in Sa. chloropterus, comb scales apparently

1000 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

eee Hoey

Fig. 4. A, B, Pupa of Sabethes (Sabethoides) conditus. A, Left side of cephalothorax, dorsal to night. B,
Dorsal (left) and ventral (right) aspects of metathorax and abdomen. C, Localities in Venezuela and Brazil where
type specimens of Sa. conditus were collected (see Material examined). Seta 5-[V was absent in available
specimens and is drawn similar to 5-V,VI based on this development in Sabethoides. CT = cephalothorax; P =
paddle; I-VIII = abdominal segments; 1—14 = setal numbers for specified areas, e.g., seta 3-I. Scale in mm.

VOLUME 102, NUMBER 4

lack the fringes of minute spicules that are
present in Sa. chloropterus, and seta 4-X is
double whereas it has 3 or 4 branches in
Sa. chloropterus.

Etymology.—The specific name of con-
ditus is a Latin adjective (masculine) mean-
ing unseen or hidden. The name refers to
the fact that the species undoubtedly has
been mistaken for Sa. chloropterus in the
past.

Bionomics.—The type specimens from
Venezuela were collected as larvae found in
brownish water contained in a small hole
located 0.6 m above ground level in a le-
guminous tree. The treehole also contained
larvae of Culex (Anoedioporpa) conserva-
tor Dyar and Knab. Nothing else is in def-
initely known about the bionomics of Sa.
conditus, but it is likely that some of the
information published about Sa. chloropte-
rus actually applies to this species.

Distribution.—The type specimens were
collected at the two localities indicated in
Fig. 4C, one in northern Venezuela and the
other in western Brazil. Sabethes conditus
is undoubtedly widely distributed between
and around these localities. Reports of Sa.
chloropterus from this region may apply to
Sa. conditus or a mixture of the two spe-
cies.

Material examined.—Eleven specimens
(3 6,2 6 genitalia, 2 2, 2 larval exuviae,
2 pupal exuviae), including 2 larval rear-
ings. VENEZUELA: Aragua, Ocumare de
la’ Costa, Puerto Ocumare, just E of
(19PFM3459), near sea level, 10 Jul 1969,
J. A. Bergland and J. Valencia, 1 ¢ with
associated larval and pupal exuviae and dis-
sected genitalia on separate slides (VZ133-
11), associated larval and pupal exuviae of
lost 6 (VZ133—22) and 1 6 (VZ133-1)
(small treehole in partial shade 4 m above
ground in leguminous tree in coconut plan-
tation; water temporary, brown, turbid).
BRAZIL: Rond6nia, Costa Marques, 31 Jan
1992, USAMRU-B personnel, 1 d6 (BR513
(65)-102) with dissected genitalia on micro-
scope slide and 2 2 (BR513 (65)-100 and
-102).

1001

The males above comprise the type series
of Sa. conditus. The holotype (VZ133-11),
with associated larval and pupal exuviae
and dissected genitalia on separate micro-
scope slides, and 3 paratypes (VZ133-1,
VZ133-22, BR513 (65)-102) are deposited
in the National Museum of Natural History,
Smithsonian Institution. The two females
are excluded from the type series because
they are not definitely associated with the
males, and may not be conspecific with
them.

ACKNOWLEDGMENTS

We thank personnel of the Walter Reed
Biosystematics Unit, Smithsonian Institu-
tion, Washington, DC, for the loan of spec-
imens. We are grateful to Robert Hancock,
Cumberland College, Williamsburg, Ken-
tucky, for assistance and advice to Dusty
Moses. Jane Carter of the same college is
acknowledged for co-ordinating the Inter-
national Enrichment Program that provided
the Biological Internship to Dusty Moses
that made this study possible.

LITERATURE CITED

Belkin, J. N. 1968. Mosquito studies (Diptera, Culic-
idae) IX. The type specimens of New World mos-
quitoes in European museums. Contributions of
the American Entomological Institute (Ann Ar-
bor) 3(4): 1-69.

Galindo, P., E. de Rodaniche, and H. Trapido. 1956.
Experimental transmission of yellow fever by
Central American species of Haemagogus and Sa-
bethes chloropterus. American Journal of Tropical
Medicine and Hygiene 5: 1022-1031.

Galindo, P., E. de Rodaniche, and C. M. Johnson.
1959. St. Louis encephalitis in Panama. 1. Isola-
tion of the virus from forest mosquitoes and hu-
man blood. American Journal of Tropical Medi-
cine and Hygiene 8: 557-560.

Hall, C. R., T. M. Howard, and R. E. Harbach. 1999.
Sabethes (Peytonulus) luxodens, a new species of
Sabethini (Diptera: Culicidae) from Ecuador. Me-
morias do Instituto Oswaldo Cruz 94: 329-338.

Harbach, R. E. 199la. A new subgenus of the genus
Sabethes (Diptera: Culicidae). Mosquito System-
atics 23: 1-9.

. 1991b. Ontogeny of the larval stage of Sa-

bethes chloropterus, with special reference to setal

development and phylogenetic implications for

1002

the family Culicidae (Diptera). Mosquito System-

atics 23: 10-24.

. 1994. The subgenus Sabethinus of Sabethes

(Diptera: Culicidae). Systematic Entomology 19:

207-234.

. 1995a. A new Sabethes of the subgenus Pey-

tonulus (Diptera: Culicidae) with an unusual

fourth-instar larva. Entomologica Scandinavia 26:

87-96.

. 1995b. Two new species of the subgenus Pey-
tonulus of Sabethes (Diptera: Culicidae) from Co-
lombia. Memorias do Instituto Oswaldo Cruz. 90:
583-587.

Harbach, R. E. and K. L. Knight. 1980. Taxonomists’
Glossary of Mosquito Anatomy. Plexus Publish-
ing, Inc., Marlton, New Jersey, xi + 415 pp.

. 1982. Corrections and additions to Taxono-

mists’ Glossary of Mosquito Anatomy. Mosquito

Systematics 13: 201—217.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Harbach, R. E. and J. L. Petersen. 1992. Two species
previously confused under the concept of Sabeth-
es tarsopus in Central America (Diptera: Culici-
dae). Mosquito Systematics 24: 102-124.

Harbach, R. E. and E. L. Peyton. 1991. Transfer of the
subgenus Davismyia from Wyeomyia to Sabethes
and description of the type species, Miamyia pe-
trocchiae (Diptera: Culicidae). Mosquito System-
atics 22: 149-159.

Heinemann, S. J. and J. N. Belkin. 1978. Collection
records of the project ““Mosquitoes of Middle
America” 11. Venezuela (VZ); Guianas: French
Guiana (FG, FGC), Guyana (GUY), Surinam
(SUR). Mosquito Systematics 10: 365—459.

Knight, K. L. and A. Stone. 1977. A Catalog of the
Mosquitoes of the World (Diptera: Culicidae). 2nd
edition. Thomas Say Foundation 6: ix + 1—611.

Lane, J. 1953. Neotropical Culicidae, Vol. Hl. Univer-
sity of SAo Paulo, Sao Paulo. pp. 1054-1098.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1003-1009

CORRECTIONS AND ADDITIONS TO THE “CATALOG OF
THE STILT BUGS, OR BERYTIDAE, OF THE WORLD
(INSECTA: HEMIPTERA: HETEROPTERA)”

THOMAS J. HENRY AND RICHARD C. FROESCHNER

(TJH) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S.
Department of Agriculture % National Museum of Natural History, Washington, DC
20560-0168, U.S.A. (e-mail: thenry @sel.barc.usda.gov); (RCF) Department of Entomol-
ogy, National Museum of Natural History, Smithsonian Institution, Washington, DC
20560-0105, U.S.A.

Abstract.—The catalog of the stilt bugs of the world, containing entries for 36 valid
genera and 168 valid species, appeared in early 1998. Since then, we have discovered a
number of errors thanks to the help of several colleagues. Corrections and additions range
from minor misspellings to the omission of taxa. Among the more important entries
concern the date of description of Metacanthus Costa and its type species and the addition
of the two overlooked species, Neides propinquus Horvath and Gampsocoris gatai Giin-

ther.

Key Words:
misspellings, literature cited

The world stilt bug catalog (Henry and
Froeschner 1998. Catalog of the stilt bugs,
or Berytidae, of the world (Insecta: Hemip-
tera: Heteroptera). Contributions of the
American Entomological Institute 30(4): 1—
72.) appeared in early 1998. Since then, we
have discovered a number of mistakes, in-
cluding those brought to our attention by
Drs. I. M. Kerzhner, Jean Péricart, Wolf-
gang Rabitsch, and J. L. Stehlik. These er-
rors, most of which involve Old World taxa,
have prompted us to provide this paper in
the interest of making the corrections avail-
able for inclusion in Dr. Péricart’s forth-
coming catalog of the Palearctic Berytidae.

The corrections, additions, and other
changes in this paper are arranged by page
and follow the general format of our Cata-
log (Henry and Froeschner 1998). The one
exception to the format is that we give only
“author and date’’ for text references doc-

Heteroptera, Berytidae, catalog, omitted taxa, gender, type fixations, dating,

umenting the country distribution. The user
should consult the bibliography for the full
citation of these references.

CORRECTIONS AND ADDITIONS

Page 6. To the distribution of Apoplymus
pectoralis Fieber add Albania (Josifov,
1986) and Cyprus (Lindberg, 1948).

Page 8. Under the genus Neides, the type
designation for junior synonym Podi-
cerus Duméril is not ‘Monotypic.”’
Correct type of designation to “‘Desig-
nated by Kirkaldy, 1900, Entomologist,
33(449): 265.”

Page 8. Under entry for Neides aduncus
Fieber, 1859, correct spelling of type lo-
cality “‘Corisca”’ to ‘‘Corsica.”’

Page 8. Under distribution for Neides af-
ghanus Seidenstiicker correct the distri-
bution of “‘Russia’”’ to ‘Tajikistan and
Turkmenistan.”

1004

Page 9. Add following species after Neides

gomeranus Heiss:
Neides propinquus Horvath, 1901

1901 Neides propinquus Horvath,
Drit. Asia. Fors:, Grafs Bug, Zich> 2:
259. [Siberia].

Distribution: Mongolia, Russia (Sibe-
rian, Far Eastern) (Hoberlandt, 1968,
Acta Faun. Entomol. Mus. Natl. Pragae,
13: 51; Kanyukova, 1988, In: Key In-
sects Far-Eastern USSR, Hemiptera, 2:
882).

Host: Unknown.

Page 9. Under 1865 Neides depressus entry,

correct date given for Reuter from 1971
to 1871.

Page 9. Under synonymy of Neides tipu-

larius (Linnaeus), the junior synonym
Cimex araneoides Goeze was synony-
mized by Harrer (G. A.), not Schaeffer.
From distribution, transfer “‘Far East-
ern’ records to Neides propinquus (I.
M. Kerzhner, personal communication,
following Kanyukova, 1988, Jn: Key In-
sects Far-Eastern USSR, Hemiptera 2:
882), and also drop ending “k’’ from
‘““‘Derzhaskyk“‘and change 2™ “u’’ in
‘“‘Lukushuk”’ to an “a.” To the distri-
bution of N. tipularius add Greece (Jos-
ifov 1986), Hungary (Horvath, 1898),
Luxembourg (Reichling and Gerend,
1994), and Slovakia (Stehlik and Vavii-
nova 1995); and to notes of N. tipular-
ius add ‘and Stehlik and Vavrinova
(1995, Acta Mus. Moraviae, 79: 159)
discussed the habitats, host, and life his-
tory.”

Page 11. Under host of Paleologus feanus

and distribution of Paraberytus similis,
correct spelling of “‘Linnavuouri’’ to
*““Linnavuori.”

Page 12. Authorship for the tribe Berytini-

ni, derived from the nominate family
name Berytinidae (based on the type ge-
nus Berytinus Kirkaldy), should be
credited to Southwood and Leston,
1959, not Henry, 1997a.

Page 12. To the distribution of Berytinus

clavipes (Fabricius) add Bulgaria (Josi-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

fov, 1986), Hungary (Horvath, 1898),
Luxembourg (Reichling and Gerend,
1994), Macedonia (Géllner-Scheiding,
1978), Romania (Sienkiewiz, 1964),
Slovakia (Stehlik and Vavrinova, 1995),
Slovenia (Gogala and Gogala, 1991),
Switzerland (Rampazzi and Dethier,
1997), and Yugoslovia (Horvath, 1903);
and to notes of B. clavipes add ‘‘and
Stehlik and Vavrinova (1995, Acta Mus.
Moraviae, 79: 154) discussed the habi-
tats, life cycle, and hosts.”

Page 13. To the distribution of Berytinus
consimilis (Horvath) add Slovakia
(Stehlik and Vavirinova, 1995); and to
notes of B. consimilis add ‘‘and Stehlik
and Vavrinova (1995, Acta Mus. Mo-
raviae, 79: 159) discussed the habitats
and the host.”

Page 13. To the distribution of Berytinus
crassipes (Herrich-Schaeffer) add Slo-
vakia (Stehlik and Vavfrinova, 1995,
Acta Mus. Moraviae, 79: 159); and to
notes of B. crassipes add ‘‘and Stehlik
and Vavrinova (1995, Acta Mus. Mo-
raviae, 79: 159) discussed the habitats,
host, and wing polymorphism.”

Page 14. To the distribution of Berytinus
geniculatus (Horvath) add Slovakia
(Stehlik and Vavrinova, 1995); and to
notes of B. geniculatus add ‘and Steh-
lik and Vavrinova (1995, Acta Mus.
Moraviae, 79: 160) discussed the habi-
tats, hosts, and the life cycle.”

Page 14. To the distribution of Berytinus
distinguendus (Ferrari) add Egypt (Lin-
navuori, 1964) and Hungary (Horvath,
1898).

Page 15. Change date of publication of Ber-
ytinus hiriticornis (Brullé) and subspe-
cies B. hirticornis hirticornis (Brullé)
from 1835 to 1836, following dating
used in Palearctic Catalog (Aukema and
Rieger, 1996); to the distribution of B.
hirticornis add Luxembourg (Reichling
and Gerend, 1994) and Slovakia (Steh-
lik and Vaviinova, 1995); and add to
notes of B. clavipes ‘‘and Stehlik and
Vavrinova (1995, Acta Mus. Moraviae,

VOLUME 102, NUMBER 4

79: 157) discussed the habitats and
hosts.”

Page 15. To the distribution of Berytinus
hirticornis nigrolineatus (Jakovlev) add
Bulgaria (Josifov, 1986) and Greece
(Rieger, 1995).

Page 15. To the distribution of Berytinus
hirticornis pilipes (Puton) add Greece
(Heiss, 1984) and Spain (Wagner,
1960).

Page 16. To the distribution of Berytinus
minor minor (Herrich-Schaeffer) add
Albania and Bulgaria (Josifov, 1986),
Austria, Romania, and Switzerland
(Sienkiewicz, 1964), Hungary (Hor-
vath, 1898), Luxembourg (Reichling
and Gerend, 1994), and Slovenia (Go-
gala and Gogala, 1991).

Page 17. Under distribution of Berytinus
montivagus (Meyer-Diir), drop second
‘“*k” in Derzhansky, 1997; to the distri-
bution of B. montivagus add Slovakia
(Stehlik and Vavirinova, 1995); and to
notes of B. montivagus add ‘“‘and Steh-
lik and Vavirinova (1995, Acta Mus.
Moraviae, 79: 161) discussed the habi-
tats, hosts, and the life cycle.”

Page 17. To the distribution of B. signoreti
(Fieber) add Luxembourg (Reichling
and Gerend, 1994), Slovakia (Stehlik
and Vaviinova, 1995), and Tunisia (Lin-
navuori, 1965), and to the notes of B.
signoreti add the reference “‘Stehlik and
Vaviinova, 1995, Acta Mus. Moraviae,
TI 2NO2

Page 18. Under heading of Berytinus stran-
gulatus (Rey), correct date from 1887
to 1888. Also, correct spelling of
‘“LExchange”’ to ““L’Echange”’ and the
type locality from ‘‘Europe’’ to
wEtance, ”’

Page 18. To the distribution of B. striola
(Ferrari) add Slovakia (Stehlik and Va-
vrinova, 1995) and to notes of B. striola
add ‘‘and Stehlik and Vavrinova (1995,
Acta Mus. Moraviae, 79: 163) discussed
the habitats, hosts, and life cycle.”

Page 20. To distribution of Gampsocoris
culicinus culicinus Seidenstiicker add

1005

Slovakia (Stehlik and Vavrinova, 1995);
and to notes of G. culicinus add ‘and
Stehlik and Vavrinova (1995, Acta Mus.
Moraviae, 79: 165) discussed the habi-
tats, host, and life history.”

Page 21. To the distribution of Gampsocor-
is enslini Seidenstiicker add Albania,
Armenia, Azerbaidzhan, Bulgaria,
Georgia, Greece, Iran, Yugoslavia (Per-
icart, 1984).

Page 21. Add following species entry:

Gampsocoris gatai Giinther, 1996

1996 Gampsocoris gatai Gitnther,
Mitt. Int. Entomol. Ver., 21: 125.
[Spain].

Distribution: Spain.

Host: Teucrium charidemi Sandwith
[Lamiaceae] (Giinther, 1998, Russian
Entomol. Soc. 1: 106); the host was giv-
en incorrectly as Ononis natris ssp. ra-
mosissima (Desf.) Batt. in original de-
scription (Giinther, 1996, Mitt. Int. En-
tomol. Ver., 21: 128).

Page 23. Under Gampsocoris punctipes
punctipes (Germar) entry, correct Bur-
meister, 1835, reference from ““Handbk”
to “Handbch.”” Under Wagner, change
date 1955 to 1954 and spelling of “‘sei-
denstuckeri”’ to “‘seidenstueckeri.”’ To
the distribution add Luxembourg
(Reichling and Gerend, 1994), Saudi
Arabia (Linnavuori, 1986), and Slovak-
ia (Stehlik and Vavrinova, 1995); and to
notes of G. punctipes add ‘‘and Stehlik
and Vavrinova (1995, Acta Mus. Mo-
raviae, 79: 166) discussed the habitats,
host, and life history.”

Page 30. Close parentheses and eliminate
one period at end of ‘Distribution’ for
Capyella horni Breddin.

Page 29. Under genus Capyella and in Lit-
erature Cited (p. 53), correct date for
Capytum Strand from 1926 to 1928.

Page 32. Under Jalysus albidus Stusak, cor-
rect spelling of host genus from ‘‘Ad-
ensotemma”’ to ‘‘Adenostemma.”’ Also
modify spelling in host index, p. 56, un-
der both the generic and species entries.

1006

Page 35. We incorrectly listed Berytus ele-
gans Costa, 1847 (nec Curtis) as a ju-
nior synonym of Berytus meridionalis
Costa, 1843 and as the type species of
Metacanthus Costa, 1847. This interpre-
tation is in error and should have re-
ferred to Neides elegans Curtis, as noted
on page 23 of our catalog, where we
correctly list Neides elegans Curtis as a
junior synonym of Berytus punctipes
(the type species of Gampsocoris Fuss).
The problem leading to this situation
began when Costa (1843) described his
species Berytus meridionalis and sug-
gested that it should be placed in a new
genus, which he tentatively (but not for-
mally) named Metacanthus. Later, he
(Costa, 1847) described and more for-
mally proposed the name Metacanthus,
but used Berytus elegans Curtis (based
on his figure) as the type species, in-
stead of B. meridionalis. If Costa’s
(1847) later description is followed,
with B. elegans as the type species, then
the genus Metacanthus becomes a ju-
nior synonym of Gampsocoris; thus,
disrupting the stability of the largest stilt
bug genus. Because of this stituation,
we fully agree with Dr. J. Péricart’s
(personal communication) suggestion
that Costa’s (1843) first description of
Metacanthus should be accepted, with
Berytus meridionalis as the type spe-
cies, a course of action followed by him
(Péricart, 1984) in his well-known book
on the Berytidae of the Mediterranean
Region.

Therefore, modify the genus Metacan-
thus as follows:

Genus Metacanthus Costa, 1843

1843 Metacanthus Costa, Cimic. Neo-
pol., p. 26. Type species: Berytus mer-
idionalis Costa, 1843. Original desig-
nation.

Note: We follow Péricart (1984), who
considered that Costa (1843) sufficient-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ly described and validated the name Me-
tacanthus and designated as its type, B.
meridionalis. To accept otherwise is to
seriously jeopardize stability and
change the name of the largest stilt bug
genus, Metacanthus Costa.

Page 35. To the distribution of Metacanthus
(Cardopostethus) annulosus (Fieber)
add Slovakia (Stehlik and Vavrinova,
1995):

Page 36. Modify the subgenus Metacanthus
as follows (following generic modifi-
cation above, p. 35):

Subgenus Metacanthus Costa, 1843

1843 Metacanthus Costa, Cimic. Neo-
pol., p. 26. Type species: Berytus mer-
idionalis Costa, 1843. Original desig-
nation.

Page 36. Under Metacanthus acintus, cor-
rect spelling of “‘Nannaizab” to ““Non-
naizab.”’

Page 36. Correct spelling of species names
Metacanthus ‘‘ataoensis” to “‘antaoen-
sis,” three times.

Page 36. The first combination of Gamp-
socoris delhiensis Menon and Ghai in
Metacanthus was by Josifov (1965, Rei-
chenbachia 5: 285), not Wheeler and
Schaefer (1982, Ann. Entomol. Soc.
Am., 75: 502).

Page 37. In note under Metacanthus linea-
tus (Jakovlev), correct spelling of M.
‘“‘maghrebrinus” to “‘maghrebinus.”

Page 37. Change date for description of
Metacanthus meridionalis (Costa) from
1844 to 1843; delete entry for “1847
Metacanthus elegans Costa, Cimic.
Neapol. Cent., p. 259. [Italy]. Synony-
mized by Bergroth, 1913, Mém. Soc.
Entomol. Belg., 22: 177.” as explained
under genus above (p. 35); correct spell-
ing of “‘Corisca”’ to “‘Corsica” under
Megalomerium pallidum entry; and cor-
rect spelling of “‘Linnavuouri” to “Lin-
navuori.”’

Page 38. In distribution under Metacanthus
mollis Stusak, correct spelling of Lin-
navuouri to Linnavuori; and to the dis-

VOLUME 102, NUMBER 4

tribution of M. mollis add Saudi Arabia,
Somalia, and Yemen (Linnavuori,
1986).

Page 39. In distribution under Metacanthus
nitidus Stusak, correct spelling of ‘‘Lin-
navuouri”’ to ““Linnavuori.”’

Page 39. Change spelling of Metacanthus
“pertenerus’’ (Breddin) to M. “‘perte-
nerum”’ three times at species and both
subspecies entries. Also add to index
entries for ‘““Metacanthus pertenerum”
(p. 68) and ‘‘Metacanthus, pertenerum
vittatus’’ (p. 72).

Page 40. For the type species of the genus
Neometacanthus and under the species
Neometacanthus picticornis, correct
spelling of ‘‘Noualheir’” to ‘‘Noual-
hier.”

Page 42. To the distribution of Yemma
gracilis Linnavuori add Yemen (Linna-
vuori and van Harten, 1997).

Page 43. The genus Yemma is feminine;
therefore correct spelling of the specific
epithet ““Yemma signatus’’ to “‘Yemma
signata’”’ in first line and add “‘[sic]”’ to
Hsiao, 1977 entry.

Page 44. Under Metatropis rufescens, cor-
rect Burmeister, 1835, reference from
*“Handbk.”’ to “‘Handbch.” To the dis-
tribution of M. rufescens add Luxem-
bourg (Reichling and Gerend, 1994)
and Slovakia (Stehlik and Vaviinova,
1995); and to notes of M. rufescens add
“and Stehlik and Vavrinova (1995, Acta
Mus. Moraviae, 79: 159) discussed the
habitats, host, and life history.”

Page 45. The genus Metatropis is feminine;
therefore correct spelling of the specific
epithet “Metatropis tesongsanicus’’ to
*““Metatropis tesongsanica’’ in first line
and add “‘[sic]” after original combi-
nation.

Page 45. Correct spelling of “Reiger” to
“Rieger” under Aukema and Rieger,
1995 and 1996.

Page 46. Change date of Brullé 1835 ref-
erence to 1836, following the Aukema
and Rieger (1996).

Page 46. Correct dates of Costa, A. from

1007

1844-1862 to 1843-1862, including
first part “‘1843 (separate, part 1, 7:143-
405 ...,” following Kerzhner (1983,
Mitt. Zool. Mus. Berlin 59: 191).

Page 46. Under Curtis (1824—40) reference
correct spelling of “‘Literatureae”’ [sec-
ond to last line] to ‘“‘Literaturae.”’

Page 48. Under Germar, 1817—1847, refer-
ence, switch order of dates from ‘‘16—
17 (1837); 18-19 (1936)” to 1936 and
1937; and under Gmelin, 1790, refer-
ence, correct spelling of ‘“‘editre’’ to
“‘edite’’; under Goeze, 1778, reference,
capitalize “‘Natursystems.”’

Page 49. Under Herrich-Schaeffer refer-
ence, change leading date 1851 to
1836-1853 and title from ‘‘Wazenarti-
gen Insecten” to “Die Wanzenartigen
Insecten’’; under Horvath, 1885, delete
“de” in Revue d’Entomologie; under
Horvath, 1901, change ‘“‘Forsehungsre-
ise’ to “‘Forschungsreise’’; and under
Horvath, 1912, change “‘Tijschrift’’ to
**Tijdschrift.”’

Page 50. Under Josifov, 1959, capitalize
‘“‘art’’ in Heteropteren-Art; under Josi-
fov, 1965, capitalize “‘systematik”’; un-
der Josifov, 1974, change “In Russian”
to “In Bulgarian”; under Lindberg,
1934, change “‘F’’ to lower case in
‘“*For’’; and under Lindberg, 1958, cor-
rect journal spelling to Societas ““Scien-
tiarum Fennicae.”’

Page 51. Under A. Puton, 1876, add to be-
ginning of title ““Notes pour.”

Page 52. Under Putshkov, 1974, add *‘[In
Ukranian]’’; under Rey, 1888, change
““L-Exchange”’ to “‘L_Echange’’; change
authorship of J. C. Schaeffer, 1784 to
G. A. Harrer, 1784 and move forward
alphabetically; and under Scholtz, 1847,
change “‘de’’ to “‘der.”’

Page 53. Under Stal, 1870-1876, correct
*“Svenskaps”’ to ““Svenska’’; and under
Strand: = 1926; cormece, the date to
**1928,”’ change “‘nomenclatoric’’ to
**nomenclatorica,’’ and change “‘Arkiv”’
to “Archiv” and insert “‘fiir.”’

1008

Page 54. Under Stusak, 1976 a 1976c, spell
out “Bulgarian Academy of Sciences.”

Page 55. Delete Wagner 1955 “Ergebnisse,
einer ....’’; under Wagner, 1965, cor-
rect spelling of ‘‘Commentiones”’ to
‘**Commentationes.”’

Add the following papers to the bibli-
ography:

Gogala, A. and M. Gogala. 1991. True bugs of Slo-
venia (Insecta, Heteroptera). Biologiski Vestnik
37: 11-44.

GGllner-Scheiding, U. 1978. Beitrag zur Kenntnis
der Heteropterenfauna Mazedoniens. Acta Musei
Macedonici Scientiarum Naturalium 15(6): 145—
149.

Giinther, H. 1996. Gampsocoris gatai n. sp., eine
neue Keulenwanzenart aus Stidspanien (Heterop-
tera: Berytidae). Mitteilungen der Internationalen
Entomologischen Vereins 21: 125—129.

Giinther, H. 1998. New and rare true bugs (Heter-
optera) from southern Spain. /n: The problems of
Entomology in Russia. Russian Academy of Sci-
ences, Russian Entomological Society 1: 106—
108.

Heiss, E. 1984. Heteropteren aus Kreta II (Insecta:
Heteroptera). Bericht des Naturwissenschaftlich-
Medizinischen Vereins in Innsbruck 71: 141—155.

Hoberlandt, L. 1968. Results of 1st Mongolian-Cze-
choslovak entomological and botanical expedition
to Mongolia. No. 17: Heteroptera (Berytidae, Pyr-
rhocoridae, Coreidae, Rhopalidae, Acanthosoma-
tidae, Cydnidae, Scutelleridae and Pentatomidae).
Acta Faunistica Entomologica Musei Nationalis
Pragae 13: 51—60.

Horvath, G. 1898. Ordo Hemiptera. In: Fauna Regni
Hungarieae: 1—64. Budapest.

Horvath, G. 1903. Szerbia Hemiptera Faunaja. (Fau-
na Hemipterorum Serbiae). Annales Musei Na-
tionalis Hungarici 1: 1-8.

Josifov, M. 1986. Verzeichnis der von der Balkan-
halbinsel bekannten Heteropterenarten (Insecta,
Heteroptera). Faunistische Abhandlungen Staa-
tliches Museum fiir Tierkunde Dresden 14: 61—93.

Kerzhner, I. M. 1983. Nachtrag zu den Publikations-
daten der Separatausgabe de Werkes “‘Cimicum
Regni Neapolitani. Centuriae’’ von A. Costa. Mit-
teilungen aus dem Zoologischen Museum in Ber-
lin 597 L911"

Kirkaldy, G. W. 1900. On the nomenclature of the
genera of the Rhynchota, Heteroptera and auchen-
orrhynchous Homoptera. The Entomologist
33(449): 262-265.

Lindberg, H. 1948. On the insect fauna of Cyprus.
Results of the expedition of 1939 by Harald Hak-
an and P. H. Lindberg, I-II. Commentationes Biol-
ogicae 8 (8): 1-32.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Linnavuori, R. 1964. Hemiptera of Egypt, with re-
marks on some species of the adjacent Eremian
region. Annales Zoologici Fennici 1: 306—356.

Linnavuori, R. 1965. Studies on the South-and East-
mediterranean hemipterous fauna. Acta Entomo-
logica Fennica 21: 1—69.

Linnavuori, R. 1986. Heteroptera of Saudi Arabia.
Fauna of Saudi Arabia. 8: 31—197.

Linnavuori, R. and A. van Harten. 1997. Note on
Heteroptera (Insecta: Hemiptera) of Yemen. Fauna
of Saudi Arabia. 16: 169-236.

Rampazzi, E and M. Dethier. 1997. Gli eterotteri
(Insecta: Heteroptera) delle torbiere a sfagni del
Cantone Ticino e de Moesano (Val Calanca e Val
Mesolcina—GR), Svizzera. Mitteilungen der
Schweizerischen Entomologischen Gesellschaft
70: 419-439.

Reichling, L. and R. Gerend. 1994. Liste des Hét-
éropteres du Grand-Duché de Luxembourg. Bul-
letin de la Société des Naturalistes Luxembour-
geois 95: 273-286.

Rieger, C. 1995. Die Fauna der Agiis-Insel Santorin.
Teil 9. Heteroptera. Stuttgarter Beitraege zur Na-
turkunde, Ser. A (Biologie) 520: 1—26.

Sienkiewicz, I. 1964. The catalogue of the “A. L.
Montandon collection” of Palaearctic Heteroptera
preserved in the “Grigore Antipa’”’ Museum of
Natural History, Bucharest. Bucharest, 146 pp.

Stehlik, J. L. and I. Vaviinova. 1995. Results of the
investigations on Heteroptera in Slovakia made by
the Moravian Museum. Acta Musei Moraviae,
Sci. Nat. 79: 149-168 (1994).

Wagner, E. 1954. Gampsocoris seidenstuekeri nov.
spec. Nachrichten des Naturwissenschaftliche
Museums der Stadt Aschaffenburg 45: 37—40.

Wagner, E. 1960. Beitrag zur Heteropteren-Fauna
Nordost-Spaniens. Trabajos del Museo de Zoolo-
gia, Nueva Serie Zoologica 2(3): 3—26.

ACKNOWLEDGMENTS

We are grateful to I. M. Kerzhner (Zoo-
logical Institute, Russian Academy of Sci-
ences, St. Petersburg) for his careful review
of the catalog and for noting the inadvertent
omission of Neides propinquus; J. Péricart
(Montereau, France) for his comments on
the serious type-species problem involving
the genus Metacanthus; J. L. Stehlik (Dept.
Entomology, Moravian Museum, Preslova,
Czech Republic) for pointing out an impor-
tant paper on the stilt bugs of Slovakia; and
to W. Rabitsch (Dept. of Ecophysiology, In-
stitute of Zoology, Vienna, Austria) for not-

VOLUME 102, NUMBER 4 1009

ing numerous omitted country records and Vandenberg, Systematic Entomology Lab-
for bringing to our attention the description oratory, U. S. Department of Agriculture,
of the recently described Gampsocoris ga- Washington, DC, for reviewing the manu-
tai. We also thank D. R. Smith and N. J. script and offering other useful suggestions.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1010-1013

THREE NEW SPECIES OF DIPOGON FOX (SUBGENUS DIPOGON)
(HYMENOPTERA: POMPILIDAE) FROM CENTRAL AND WESTERN
NORTH AMERICA

HOWARD E. EVANS

Department of Bioagricultural Science and Pest Management, Colorado State Univer-

sity, Fort Collins, CO 80523-1177, U.S.A.

Abstract.—Three new species are described in the genus Dipogon, subgenus Dipogon
(Pompilidae). They are: D. kiowa (Prowers and Delta counties, Colorado), D. kenza
(Douglas Co., Kansas), and D. anasazi (Montezuma Co., Colorado).

Key Words:

Species of Dipogon Fox are not often
taken by conventional collecting methods,
as they are rarely attracted to flowers or
honeydew. They occur principally in wood-
ed areas, where most nest in cavities in
wood. Males are rarely taken except when
reared from trap nests (Medler and Koerber
1957, Krombein 1967). One species of sub-
genus Deuteragenia Sustera has been
reared from stems of weeds and garden
plants (Williams 1966) and a species of
subgenus Dipogon has been reared from
stems of Sambucus (Wasbauer 1966), but
most specimens have been taken in asso-
ciation with trees. Townes (1957) recog-
nized five species of subgenus Dipogon
north of Mexico, but four more have since
been added (Wasbauer 1960, 1966; Evans
1987). Three more species are added here,
all thus far known only from females. All
have the frons and thoracic dorsum pol-
ished and largely devoid of recumbent pu-
bescence and the punctures of the frons dis-
tinct but minute and widely spaced. Thus
they belong to the graenicheri group of
Townes (1957) provided the limits of that
group are expanded to include species in
which the microtrichiae of the fore wing are
not necessarily much larger and more

Hymenoptera, Pompilidae, Dipogon, new species, North America

crowded in the fuscous bands than else-
where.

Dipogon (Dipogon) kiowa Evans,
new species

Holotype.—?, COLORADO: Prowers
Co., Lamar, tree trunks, 4—7 Sept. 1998 (D.
Leatherman) [National Museum of Natural
History].

Paratypes.—, same data; 2, same lo-
cality and collector, Willow Creek Park,
15-16 Aug. 1998; 6 2, same locality and
collector as type, 26-31 July 1999; 9,
COLORADO: Delta Co., Crawford State
Park, ex Utah juniper, 5 Aug. 1999 (D.
Leatherman) [Paratypes in National Muse-
um of Natural History and Colorado State
University].

Etymology.—Named for the Kiowa In-
dians that formerly roamed the Arkansas
Valley. On July 27, 1820, members of the
Long Expedition visited an encampment of
Kiowas not far from the type locality.

Description of type.—Length 5.5 mm,
fore wing length 4.2 mm. Head ferruginous,
including mouthparts and antenna, except
first two and last antennal segments par-
tially infuscated, ocellar triangle and paired
streaks just below them weakly infuscated.

VOLUME 102, NUMBER 4

Pronotum ferruginous, with a pair of fus-
cous spots anterodorsally; remainder of tho-
racic dorsum and central part of propodeum
black; meso- and metapleura and lateral
parts of propodeum ferruginous; thoracic
venter black. Metasoma black except sides
of first tergite stained with reddish. Legs
ferruginous except suffused with black as
follows: upper surface of mid and hind cox-
ae and trochanters, basal and apical extrem-
ities of fore and mid femora and much of
mesal surface of hind femur, outer surface
of fore and mid tibiae and all of hind tibia,
basal segment of mid and hind tarsi. Wings
hyaline, with a narrow brown band at the
basal vein and a large brown area extending
from the stigma through the third discoidal
cell; microtrichiae slightly denser in the
dark bands than elsewhere, especially dense
at the basal vein.

Surface of head polished, only very
faintly alutaceous; clypeus with several
strong setae; frons with small punctures
separated by 5—10 times their own diame-
ters, each giving rise to a very short, erect
seta. Thorax with integument shining, me-
soscutum slightly more alutaceous and
more close punctate than frons; scutellum
and metanotum with dense, recumbent sil-
very to golden pubescence; meso- and me-
tapleura polished, very weakly alutaceous;
mid and hind coxae strongly silvery-seri-
ceous; propodeum polished, with sparse,
small punctures; metasoma sparsely sil-
very-sericeous.

Width of head 1.05 times length of head
to margin of clypeus; middle interocular
distance 0.60 times width of head; eyes
convergent above, upper interocular dis-
tance 0.75 times lower. Postocellar and
ocello-ocular distances subequal. First four
antennal segments in a ratio of 17:9:29:22.
Posterior margin of pronotum arcuate; slope
of propodeum low and even, midline not
impressed. Second submarginal cell maxi-
mum length 3 times its maximum height
and 1.6 times maximum length of third sub-
marginal cell.

Variation—Two of the paratypes are

1011

considerably smaller than the type, with
fore wing length 3.2 mm. Otherwise, there
is little variation in size or morphology
within the series. Two of the Lamar para-
types have the frons wholly ferruginous,
and one Lamar paratype as well as the one
from Delta Co. have the frons as well as
the vertex and occiput black. These darker
specimens also have the thorax and legs
more heavily infuscated than in the other
paratypes.

Remarks.—The polished and nearly bare
integument, combined with the contrasting-
ly dense pubescence on the scutellum and
metanotum, plus the unusual color pattern,
distinguish this species. In contrast to
graenicheri Banks (1939), the body is more
extensively marked with black and the me-
tasoma wholly black; also the fasciae of the
fore wing are less intense and the microtri-
chiae only slightly more dense within the
wing bands. Dipogon parkeri Wasbauer
(1966), described from Nevada, has a color
pattern not unlike kiowa, but the head, tho-
racic dorsum, and propodeum are duller,
with stronger surface sculpturing and an
overlay of appressed pubescence. In parkeri
the eyes are less convergent above, the up-
per interocular distance measuring 0.89
times the lower.

Dipogon (Dipogon) konza Evans,
new species

Holotype-—?, KANSAS: Douglas Co.,
Lawrence, 25 Aug. 1986, Douglas Yanega,
on Gonolobus vine [Snow Entomological
Museum, University of Kansas].

Paratypes.—3 2, same data except 26
Aug., 4 and 27 Sept. 1896; 2, Lakeview,
near Lawrence, 24 Sept. 1966, G. C. Eick-
wort; 2, Breidenthal Reserve, 2 mi. N
Baldwin, Douglas Co., 22 Aug.—8 Sept.
1982, malaise trap, D. B. Wahl [paratypes
at University of Kansas, Colorado State
University, and National Museum of Nat-
ural History].

Etymology.—Named for the Konza In-
dians, using the spelling employed by early

1012

explorers in what is now the state of Kan-
sas.

Description of type.—Length 4.2 mm;
fore wing length 3.4 mm. Body entirely
glossy black except clypeus and sides of
pronotum brownish; mandible dark brown
basally, yellow-brown apically; antenna
wholly light ferruginous, legs also of this
color except coxae suffused with brown.
Wings hyaline, somewhat luteous, veins on
outer part translucent; fore wing unbanded
but with a small brown cloud just distad of
stigma; microtrichiae of uniform size and
distribution except somewhat more crowd-
ed along basal vein.

Clypeus with several strong setae. Frons
and vertex strongly polished, without sur-
face sculpturing, with minute punctures
separated by many times their own diame-
ters, each giving rise to a short, erect seta.
Thorax and propodeum similarly polished
and sparsely punctate except scutellum fine-
ly, closely punctate and mesosternum with
fine, appressed, silvery pubescence. Meta-
soma polished, very weakly alutaceous,
with the usual strong setae ventrally and
apically; first 3 tergites with only scattered,
short setae.

Width of head 1.06 times length of head;
middle interocular distance 0.61 times head
width; eyes weakly convergent above, up-
per interocular distance 0.9 times lower.
Vertex weakly elevated above eye tops,
postocellar and ocello-ocular distances sub-
equal. First 4 antennal segments in a ratio
of 17:9:20:17. Posterior margin of prono-
tum arcuate; slope of propodeum low and
even, midline not impressed. Second sub-
marginal cell with its maximum length 3
times the maximum height, 1.5 times max-
imum length of third submarginal.

Variation.—Individuals in this series
vary in fore wing length from 3.0 to 3.8
mm. All have the deep brown to black
glossy integument and the light ferruginous
antennae and legs (at least beyond the cox-
ae), but the largest female is streaked with
ferruginous along the sides of the thorax
and propodeum. Although there are no dis-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tinct wing fasciae in any of the specimens,
in two there is weak clouding along the bas-
al vein.

Remarks.—The clear wings, with only
faint localized clouding in some specimens,
set this species apart from other species of
Dipogon except for fulleri Krombein
(1962), a member of the subgenus Winne-
manella. The glossy black integument con-
trasting to the pale appendages also char-
acterizes this species.

Dipogon (Dipogon) anasazi Evans,
new species

Holotype.—?, COLORADO: Montezu-
ma Co., Soda Canyon, Mesa Verde Nation-
al Park, 6 Aug. 1999, B. Kondratieff, S.
Wells, W. Cranshaw, P. Pineda, & W. Paint-
er [National Museum of Natural History].

Etymology.—This species is named for
the Anasazi, the remains of whose homes
provide the major treasures of Mesa Verde
National Park.

Description of type.—Length 4.8 mm;
fore wing length 4.4 mm. Entire body and
appendages light ferruginous, eyes and
ocelli contrastingly nearly black. Wings hy-
aline, slightly luteous basally; fore wing
with a weak brown band at basal vein and
a broad brown cloud below the stigma and
basal half of marginal cell. Microtrichiae of
fore wing distinctly darker and more
crowded within the two dark bands than
elsewhere.

Integument of head, thorax, and propo-
deum strongly polished and without surface
sculpturing, also without appressed pubes-
cence. Clypeus with several strong setae,
but frons with only short, erect hairs arising
from small punctures separated by many
times their own diameters. Thorax and pro-
podeum similarly with sparse, small punc-
tures except posterior fifth of mesoscutum
and all of scutellum and metanotum with
dense, fine punctures. Metasoma also pol-
ished but with rather dense, fine punctures,
posterior segments with usual stiff bristles.

Head 1.04 times as wide as high, vertex
roundly elevated above eye tops. Middle in-

VOLUME 102, NUMBER 4

terocular distance 0.62 times head width;
upper interocular distance 0.84 times lower.
Ocelli in a compact triangle, ocello-ocular
distance 1.2 times postocellar distance. First
4 antennal segments in a ratio of 19:10:25:
21. Second submarginal cell with its max-
imum width 2.8 times maximum height, 1.3
times maximum width of third submarginal
cell.

Remarks.—This species has many fea-
tures in common with graenicheri Banks
(1939). It differs in the wholly pale color-
ation and weaker wing bands; also the body
is even more highly polished, the vertex
more elevated above the eye tops, and the
punctures of the frons and thorax are more
sparse.

The type and only known specimen was
collected by Samuel Wells by beating ju-
nipers in a relatively dry canyon that had
been burned over a few years earlier.

DISCUSSION

Dipogon diablo Wasbauer (1960) was as-
signed by the describer to the graenicheri
group, but the head and thoracic dorsum
and dull and minutely granulo-reticulate as
well as sparsely covered with appressed pu-
bescence. Thus it does not appear closely
related to the species considered here. The
following couplets may serve to separate
females of the graenicheri group as defined
here.

KEY

1. Fore wing not fasciate (may have a small
brown spot distad of stigma); body glossy
black, legs and antenna light ferruginous (body
may be streaked with ferruginous along sides
@laineSOSOMaA) wreak sehek el eek okene konza, 0.sp.

— Fore wing bifasciate; color not as above .... 2

2. Entire body pale ferruginous, with contrasting
dark eyes and ocelli; integument highly pol-
ished; vertex roundly elevated above eye tops
RSE ese. dks ao e4 anasazi, N. sp.

— Not entirely ferruginous nor integument as
highly polished; vertex more weakly elevated
ADOWUS CVS LO) SSeS Ske now closes oe oeaeeee 3

3. Mesosoma and at least basal third of metasoma
ferruginous; legs ferruginous

g. graenicheri Banks

1013

— Mesosoma fuscous at least dorsally, metasoma
wholly black; legs in large part fuscous .... 4
4. Body black, antenna, fore leg, and tarsi except
basally, tinged with fulvous; microtrichiae of
fore wing much darker and more crowded in
fasciae than elsewhere; scutellum and metan-
otum not more densely pubescent than meson-
(ORDtCH Ss wah oo Gd edict graenicheri atratus Townes
— Mesosoma black, pleura and lateral parts of
propodeum ferruginous; microtrichiae of fore
wing barely darker and more crowded in fas-
ciae than elsewhere; scutellum and metanotum
Genselyapubescentn sas aaa Kiowa, nN. sp.
ACKNOWLEDGMENTS

For the loan of types and other speci-
mens, I am indebted to Robert W. Brooks,
University of Kansas; Michael S. Kelley,
Museum of Comparative Zoology; Steve
Heydon, University of California at Davis;
and Wojciech Pulawski and Bob Zuparko
of the California Academy of Sciences.

LITERATURE CITED

Banks, N. 1939. Notes and descriptions of native
Psammocharidae. Canadian Entomologist 71:
225-231.

Evans, H. E. 1987. The genus Dipogon (Hymenoptera:
Pompilidae) in the Rocky Mountains. Entomolog-
ical News 98: 41—45.

Krombein, K. V. 1962. Natural history of Plummers
Island, Maryland. XIII. Descriptions of new wasps
from Plummers Island, Maryland, (Hymenoptera:
Aculeata). Proceedings of the Biological Society
of Washington 75: 1-18.

. 1967. Trap-nesting wasps and bees: Life his-
tories, nests, and associates. Smithsonian Press,
Washington, DC, 570 pp.

Medler, J. T. and T. W. Koerber. 1957. Biology of Di-
pogon sayi Banks (Hymenoptera, Pompilidae) in
trap-nests in Wisconsin. Annals of the Entomo-
logical Society of America 50: 621—625.

Townes, H. 1957. Nearctic Wasps of the Subfamilies
Pepsinae and Ceropalinae. Bulletin of the United
States National Museum 209. 286 pp.

Wasbauer, M. S. 1960. Taxonomic and distributional
notes on some western spider wasps (Hymenop-
tera: Pompilidae). Pan-Pacific Entomologist 36:
171-177.

. 1966. A new spider hunting wasp of the sub-
genus Dipogon from western Nevada (Hymenop-
tera: Pompilidae). Proceedings of the Biological
Society of Washington 79: 17—20.

Williams, E X. 1966. A spider-hunting wasp found
nesting in hollow plant stems (Hymenoptera:
Pompilidae, Pepsinae). Wasmann Journal of Bi-
ology 24: 33-47.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1014-1069

CATALOGUE OF THE TYPE SPECIMENS OF TORTRICIDAE
(LEPIDOPTERA) IN THE COLLECTION OF THE NATIONAL MUSEUM OF
NATURAL HISTORY, SMITHSONIAN INSTITUTION, WASHINGTON, D.C.

JOHN W. BROWN AND JON LEwIs

Systematic Entomology Laboratory, USDA, PSI, Agricultural Research Service, %
National Museum of Natural History, Washington, DC 20560-0168, U.S.A. (e-mail:
jbrown @sel.barc.usda.gov)

Abstract.—Type specimens of the family Tortricidae deposited in the collection of the
National Museum of Natural History, Smithsonian Institution, are listed alphabetically by
species (or subspecies), along with an abbreviated literature citation to the original de-
scription and collection data for the specimen. The type collection includes 1,001 name-
bearing types of Tortricidae: 851 holotypes, 50 lectotypes (6 of which are designated
herein), 60 species represented by one or more syntypes, 38 species represented by the
only known extant specimen from the original series, 1 neotype, and 1 “‘pseudo-type.”
The collection includes the vast majority of the tortricid species described by C. Fernald,
A. Busck, C. Heinrich, J. Clarke, A. Blanchard, and A. Kawabe. It also includes the types
of numerous species described by Lord Walsingham, W. Kearfott, H. Dyar, N. Obraztsov,

J. Razowski, J. Powell, W. Miller, R. Brown, and J. Brown.

Key Words:

The type collection of Tortricidae in the
National Museum of Natural History
(USNM), Smithsonian Institution, repre-
sents a significant worldwide resource for
researchers interested in tortricid moths
(i.e., the leafrollers). The strengths of the
collection lie in the faunas of North and
South America, the Orient (Japan, Taiwan,
Philippine Islands), and Oceania. The C. H.
Fernald Collection, purchased in 1924
(Clarke 1974), formed the foundation, con-
taining types (primarily “‘cotypes’’) of nu-
merous species described by Fernald, Lord
Walsingham, and W. D. Kearfott, and a few
described by A. Grote, B. Clemens, and A.
Packard. Walsingham sent Fernald “‘coty-
pes” of most of the species he described
from California and Oregon (Walsingham
1879) from material in The Natural History
Museum (formerly British Museum of Nat-

leafrollers, lectotype, holotype, syntype, original description, host plants

ural History). Kearfott also gave Fernald
examples from his type series; the majority
of Kearfott’s material eventually was de-
posited in the American Museum of Natural
History, New York. Klots (1942) resolved
most of the nomenclatorial difficulties as-
sociated with the Kearfott “‘cotypes,”’ des-
ignating numerous lectotypes, primarily in
the collection of the American Museum of
Natural History. Miller (1970) resolved
most of nomenclatorial problems associated
with the Fernald ‘‘cotypes,” designating
lectotypes for the vast majority of his spe-
cies of North American Olethreutinae. Few
lectotypes have been designated for the
Walsingham (1879) species from California
and Oregon.

From 1905-1930 August Busck and Carl
Heinrich, both U.S. Department of Agri-
culture employees at the museum, added

VOLUME 102, NUMBER 4

numerous tortricid types, primarily from
North and South America, through their
prolific taxonomic research on microlepi-
doptera. In particular, over 150 species were
described by Heinrich (1923, 1926) in his
revisions of the Olethreutinae of North
America. Also during the early part of the
century, Walsingham’s (1914) contribution
to the “‘Biologia Centrali-Americana”’ ap-
peared; it included the descriptions of many
new species, with most of the material split
between The Natural History Museum and
the USNM. By this time, Walsingham was
designating a type for each new species, so
there is little ambiguity regarding these pri-
mary types.

In the 1960s, revisions of New World
genera by Obraztsov (1961, 1962, 1963,
1964, 1966a, b, c) resulted in the addition
of many tortricid types to the USNM col-
lection. From 1968—1982 specific large re-
search projects such as Alexis Diakonoff’s
studies of the microlepidoptera of the Phil-
ippine Islands (Diakonoff 1968) and of Sri
Lanka (Diakonoff 1982), and J. E G.
Clarke’s work on Neotropical ‘‘Phaloni-
idae”’ (Clarke 1968) and the microlepidop-
tera of Rapa Island and Micronesia (Clarke
1971, 1976) added many types of Tortrici-
dae. The Blanchard Collection, donated in
1985 (Davis 1985), also added many tortri-
cid types, nearly all from Texas. The At-
sushi Kawabe collection, composed almost
exclusively of Oriental species and donated
in 1990 (Davis 1996), added significant
numbers to the tortricid type collection as
well, broadening greatly the geographic
coverage. Use of the tortricid holdings of
the USNM by specialists worldwide (e.g.,
Richard L. Brown, John W. Brown, John B.
Heppner, William E. Miller, Jerry A. Pow-
ell, Josef Razowski, and others) has contin-
ued to increase type holdings.

Through recent curatorial efforts, the
type specimens of the family Tortricidae
have been organized into a single collec-
tion. The specimens are arranged in alpha-
betic sequence by species (or subspecies) in
individual, labeled unit trays and stored in

1015

13 insect drawers in the USNM. The pur-
pose of this paper is to present a list of the
type specimens. The list is arranged alpha-
betically by species (or subspecies), fol-
lowed by the author, year of publication,
and genus (or genus and species for sub-
species) in which the taxon was originally
described. This information is followed by
an abbreviated citation of the publication in
which the description appears. The sex,
type designation (i.e., holotype, lectotype,
syntype, “‘type’’, neotype, ““‘pseudo-type’’),
and collection data also are included. We
attempt to provide the maximum collection
data, relying on both the text of the original
description and information on specimen la-
bels. For example, country, county, prov-
ince, etc. frequently are not present on the
specimen labels; this information is provid-
ed where retrievable accurately. Abbrevia-
tions used in the list include the following:
AMNH = American Museum of Natural
History, New York; BMNH = The Natural
History Museum, London; NHMW = Na-
turhistorisches Museum, Wien; r.f. = reared
from; em: = emerged; mi = mile(s); ' =
feet (elevation). Where the collecting date
can have more than one interpretation, it is
presented in quotes. For example, “‘Aug. 8/
15”’ could be interpreted as 8 August 1915
or 8—15 August [no year].

The modern practice of designating a ho-
lotype was not followed by most tortricid
workers before 1900 (e.g., Riley, Grote,
Zeller), and some early authors (e.g., Wal-
singham, Dyar) labeled all specimens of the
original series as “‘type.’’ Where the origi-
nal type series consists of a single speci-
men, as clearly stated by the author, we re-
fer to it as the holotype even though not
specifically designated as such in the orig-
inal description. Where only a single spec-
imen is presumed still to be extant from the
original series (and a type was not specifi-
cally designated), we refer to this specimen
as the “‘type’’ (in quotes). In this case, the
specimen is a potential lectotype because it
may be the only remaining representative
from the original series. Where two or more

1016

specimens of the original series are present
and no type was specifically designated in
the original description, we refer to these
specimens as syntypes. We defer the latter
two cases to the future work of specialists
to designate appropriate lectotypes for
these, as needed. Six specimens are unpub-
lished lectotypes unambiguously selected
and labeled by Obraztsov (n = 3), Powell
(n = 2), and Clarke (n = 1); these selec-
tions are designated in this paper.

The USNM tortricid type collection in-
cludes 851 holotypes, 50 lectotypes (6 of
which are designated herein), 60 species
represented by one or more syntypes, 38
“types” (1.e., the only known extant spec-
imen from the original series), 1 neotype,
and 1 “‘pseudo-type.”’

LIST OF SPECIES

abiephaga Issiki, 1962 (Ariola); in Issiki
and Mutuura, Publ. Entomol. Lab. Univ.
Osaka Pref. 7: 3. Holotype ¢, Japan,
Honsyu, Koozuke, Manza, 27 July
1958, S. Issiki & T. Yasuda.

abietana Fernald, 1908 (Argyroploce);
Canad. Entomol. 40: 349. Holotype <6,
USA, Massachusetts, Hampshire Co.,
Amherst. According to Fernald this spe-
cies was “Described from three male
specimens taken in Amherst, Mass., one
of which I make the type .. .”’

abievora Issiki, 1961 (Epagoge); in Issiki
and Mutuura, Microlepidoptera injuri-
ous to coniferous plants in Japan: 34.
Holotype 6, Japan, Osaka, Ikeda, 20
November 1958, T. Kodama.

abornana Busck, 1939 (Lorita); Bull.
South. Calif. Acad. Sci. 38: 101. Holo-
type 2, USA, California, Los Angeles
Co., El Segundo, rf. Cuscuta californi-
ca, em: 11 July 1938, W. Pierce.

abbreviatana Walsingham, 1879 (Paedis-
ca); Ill. Lepid. Heter. British Museum
4: 54. Syntype ¢, USA, Pennsylvania.
According to the original description,
this species was described from two
males from Washington [D.C. according
to Heinrich 1923], August 1872. The

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

other specimen is presumed to be in the
BMNH. The USNM specimen bears the
typical Walsingham “Type”’ label, but
the collecting locality is not consistent
with the original description.

abundantia Clarke, 1976 (Heleanna phys-
alodes); Insects of Micronesia 9: 15.
Holotype 3, Micronesia, Kusaie, Pu-
kusrik, | m, 3 April 1953, J. Clarke.

accessa Heinrich, 1931 (Epinotia); Proc.
U.S. Natl. Mus. 79: 9. Holotype 6, Pan-
ama, Trinidad River, March 1912, A.
Busck.

achlyoptera Clarke, 1976 (Cryptaspasma);
Insects of Micronesia 9: 58. Holotype
?, Micronesia, Guam, Ritidian, 7 Au-
gust 1945, J. Gressitt.

acrocroca Diakonoff, 1982 (Acanthoclita);
Zool. Verhandel. (Leiden) 193: 28. Ho-
lotype 6, Sri Lanka, Ratnapura District,
Uggalkaltota, 350’, Irrigation Bunga-
low, 31 January—8 February 1970, D.
Davis & B. Rowe.

acrospodia Diakonoff, 1982 (Spanistoneu-
ra); Zool. Verhandel. (Leiden) 193: 9.
Holotype 6, Sri Lanka, Kandy District,
1,800’, Peak View Motel, 15—24 Janu-
ary 1970, D. Davis & B. Rowe.

adana Heinrich, 1923 (Rhyacionia); Bull.
U.S. Natl. Mus. 123: 18. Holotype 6,
USA, Massachusetts, Essex Co., Forest
Hill, 5 April 1910, W. Raff.

ademonia Clarke, 1968 (razona); Proc.
U.S. Natl. Mus. 125: 42. Holotype ?,
Costa Rica, La Florida, 500’, W.
Schaus.

adjuncta Heinrich, 1924 (Gypsonoma), J.
Wash. Acad. Sci. 14: 389. Holotype 6,
Canada, Ontario, Toronto, June 1913,
Parish.

adobe Brown, 1992 (Catastega); J. New
York Entomol. Soc. 100: 224. Holotype
6, USA, Arizona, Cochise Co., Chiri-
cahua Mountains, 4 mi W Portal,
5,300’, 3-6 August 1964, D. Davis.

adoceta Diakonoff, 1964 (Bactra); Zool.
Verhandel. (Leiden) 70: 11. Holotype
6, Brazil, Parana, Castro, W. Schaus.

aechnemorpha Diakonoff, 1982 (Gypsono-

~VOLUME 102, NUMBER 4

ma); Zool. Verhandel. (Leiden) 193: 48.
Holotype 6, Sri Lanka, Ratnapura Dis-
trict, Uggalkaltota, 350’, Irrigation Bun-
galow, 31 January—8 February 1970, D.
Davis & B. Rowe.

aelina Diakonoff, 1982 (Cydia); Zool. Ver-
handel. (Leiden) 193: 21. Holotype <4,
Sri Lanka, Ratnapura District, Uggal-
kaltota, 350’, Irrigation Bungalow, 31
January—8 February 1970, D. Davis &
B. Rowe.

aenigmana Powell, 1964 (Acleris); Univ.
Calif. Publ. Entomol. 32: 99. Holotype
36, USA, California, Nevada Co., Truck-
ce Och 8-15.”

aequilibra Diakonoff, 1982 (Aemulatrix);
Zool. Verhandel. (Leiden) 193: 3. Ho-
lotype 6, Sri Lanka, Ratnapura District,
Uggalkaltota, 350’, Irrigation Bunga-
low, 31 January—8 February 1970, D.
Davis & B. Rowe.

aesculana Riley, 1881 (Proteopteryx);
Trans. St. Louis Acad. Sci. 4: 321.
“Type”? 6, USA, Missouri, “boring the
tender terminal twigs of Buckeye and of
Maple ...’’ Riley described this species
from eight specimens; we have been
able to locate only one. The specimen
is clearly labeled “‘Type”’ in Riley’s
hand.

aethalea Obraztsov, 1964 (Proeulia); Proc.
U.S. Natl. Mus. 116: 188. Holotype ¢,
Chile, Santiago, La Obra, October 1952,
Pena.

aglaia Clarke, 1955 (Orthocomotis); Trans.
Royal Entomol. Soc. London 107: 149.
Holotype 2, Brazil, Rio de Janeiro, Ita-
tiaya, 700 m, 12 October 1926, J. Zikan.

agricolana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. British Museum 4: 42.
Two syntypes (2 66), USA, California.
This species was described from five
males from California and Oregon, col-
lected in May and June. As is typical of
Walsingham specimens from this era,
both of the syntypes cited above are la-
beled ‘“‘Type.’’ Neither has an abdomen;
although there is a slide label attached

1017

to one of the specimens, the slide has
not been found.

ainslieana Obraztsov, 1962 (Anopina); Am.
Mus. Novit. 2082: 34. Holotype ¢,
USA, New Mexico, Dona Ana Co.,
Mesilla, C. Ainslie.

aktita Miller, 1978 (Rhyacionia); in Powell
and Miller, U.S. Dept. Agric., Agric.
Handb. 514: 24. Holotype ¢, USA,
New Jersey, Ocean Co., Lakehurst, 5
May 1962, R. Hodges.

alaskae Heinrich, 1923 (Epinotia cruci-
ana): ] Bull. WW:S. Natl Muse 1239229.
Holotype 6, USA, Alaska, Yukon, 3
August 1916, G. Harrington [cited as
*‘Huntington” by Heinrich]. The col-
lecting locality, probably referring to
Yukon River or some other locality in
Alaska, is somewhat ambiguous _ be-
cause Yukon Territory is in Canada, not
Alaska.

albafascia Heinrich, 1929 (Ancylis); Proc.
U.S. Natl. Mus. 75: 19. Holotype ¢,
USA, California, Tulare Co., Mineral
King, “June 24-30.”

albicapitana Busck, 1914 (Evetria); Proc.
Entomol. Soc. Wash. 16: 147. Holotype
6, Canada, Saskatchewan, Prince Al-
bert, rf. Pinus divaricata, em: 12 March
1914, J. Blumer. Although the type la-
bel indicates ‘“‘Evetria capitana”’ rather
than albicapitana, the USNM type
number (Cat. No. 18444) and the col-
lection data on the specimen match that
given in the original description.

albicaudana Busck, 1915 (Sparganothis);
Proc. Entomol. Soc. Wash. 17: 85. Ho-
lotype 36, USA, Pennsylvania, Notch,
rf. maple, em: 5 July 1913, A. Busck.

albiciliana Fernald, 1882 (Sericoris); Trans.
Am. Entomol. Soc. 10: 70. Lectotype
3d, USA, Maine, Penobscot Co., Orono.
Designated by Miller (1970).

albidula Heinrich, 1926 (Olethreutes buck-
ellana); Bull. U.S. Natl. Mus. 132: 189.
Holotype 6, USA, California, Inyo Co.,
15-30 June 1922, O. Poling.

albimaculana Fernald, 1879 (Grapholitha);
Canad. Entomol. 11: 157. Lectotype ¢,

1018

USA, Maine, Penobscot Co., Orono.
Designated by Miller (1970).

albipuncta Heinrich, 1926 (Bactra_ veru-
tana); Bull. U.S. Natl. Mus. 132: 84.
Holotype ¢, USA, Colorado, Adams
Co., Denver, Oslar.

albopunctana Brown, 1999 (Dimorphopal-
pa); Pan-Pac. Entomol. 75: 85. Holo-
type 6, Venezuela, Aragua, Rancho
Grande, 1,100 m, 16—19 January 1966,
S. & W. Duckworth.

aliana Kawabe, 1965 (Clepsis); Kontyt 34:
460. Holotype 6, Japan, Kurodake
(Hokkaido), Mt. Daisetsu, 22 July 1952,
A. Matsura.

alishana Kawabe, 1986 (Epiblema); Ento-
mol. Pap. Pres. Kurosawa, Tokyo: 82.
Holotype 6, Taiwan, Chiai Hsien, Al-
ishan, 2,300 m, 7 August 1971, Y. Shi-
bata.

alleniana Fernald, 1882 (Tortrix); Trans.
Am. Entomol. Soc. 10: 68. “Type” dé,
USA, Maine, Penobscot Co., Orono,
July. This species was described from
three males and three females; we have
been able to locate only one male. The
specimen is clearly labeled as “‘Type”’
in Fernald’s hand.

allochroma Diakonoff, 1968 (Costosa);
Bull. U.S. Natl. Mus. 257: 81. Holotype
3, Philippine Islands, Davao Province,
Mindanao, E slope Mt McKinley,
5,600’, 16 September 1946, CNHM
Philippine Zool. Exped. 1946—47, H.
Hoogstraal.

alphabetica Walsingham, 1914 (Eucosma);
Biol. Centr.-Am., Lepid., Het. 4: 236.
Holotype ¢, Mexico, Puebla, Popoca-
tepetl [Park], 8—11,000’, July 1906, W.
Schaus. Described from a single speci-
men identified as “‘Type.”’

alphitopa Clarke, 1968 (Hysterosia); Proc.
U.S. Natl. Mus. 125: 7. Holotype ¢,
Venezuela, Aragua, Rancho Grande,
16—23 October 1966, S. & W. Duck-
worth.

alterana Heinrich, 1923 (Thiodia); Bull.
U.S. Natl. Mus. 123: 46. Holotype 6,
USA, Maryland, Montgomery Co.,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Plummers Island, | August 1903, A.
Busck.

altissima Kawabe, 1978 (Eubrochoneura);
Tinea 10: 177. Holotype 3, Japan, Hon-
shu, Shizouka Pref., Odaru Spa, S Izu,
29 April 1968, A. Kawabe.

amatana Dyar, 1901 (Eulia); J. N. Y. En-
tomol. Soc. 9: 24. Three syntypes (3
22), USA, Florida, Palm Beach Co.,
Palm Beach, r.f. Nectandra [| = Ocotea|
(n = 1). Dyar described this species
from the three specimens cited above.

amblyopa Clarke, 1976 (Cryptophlebia);
Insects of Micronesia 9: 106. Holotype
3, Micronesia, Palau Islands, Koror Is-
land, Koror, January—May, light trap,
Beardsley.

ambodaidaleia Miller, 1983 (Phaneta);
Ann. Entomol. Soc. Am. 76: 101. Ho-
lotype 2, USA, South Carolina, Berke-
ley Co., Wedge Plantation, Mc-
Clellanville, 16 March 1968, D. Fergu-
son.

ambogonium Pogue, 1986 (Apolychrosis);
in Cibridn-Tovar et al., Cone and Seed
Insects of the Mexican Conifers: 21.
Holotype ¢, Mexico, Mexico, San Mi-
guel Atlautla, 2,500 m, rf. Pinus leio-
phylla, em: 14 September 1983, W. San-
chez.

americana Fernald, 1882 (Teras); Trans.
Am. Entomol. Soc. 10: 66. Lectotype
3, USA, Massachusetts, ‘“‘“Cambr. B.”’
[Cambridge?]. Designated by Obraztsov
(1963).

amoena Kawabe, 1986 (Phaecasiophora);
Entomol. Pap. Pres. Kurosawa, Tokyo:
77. Holotype 6, Taiwan, Hualien Hsien,
Hungyeh Spa, 200 m, 29-30 March
1984, A. Kawabe.

amorpha Clarke, 1986 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 123. Holo-
type 6, Marquesas Islands, Nuku Hiva,
Tapuaooa, 2,500’, 30 January 1968, J.
Clarke.

amphibola_ Diakonoff, 1982 (Diplosema-
phora); Zool. Verhandel. (Leiden) 193:
21. Holotype 6, Sn Lanka, Ratnapura
District, Uggalkaltota, 350’, Irrigation

VOLUME 102, NUMBER 4

Bungalow, 31 January—8 February
1970, D. Davis & B. Rowe.

anaranjada Miller, 1959 (Laspeyresia);
Florida Entomol. 42: 131. Holotype &,
USA, Georgia, Crisp Co., Cordele, rf.
Pinus ipalustris,,21 May %1950;) C.
Speers.

anastea Diakonoff, 1968 (Archidemis);
Bull. U.S. Natl. Mus. 257: 30. Holotype
36, Philippine Islands, Davao Province,
Mindanao, E slope Mt McKinley,
7,000’, 22 September 1946, CNHM
Philippine Zool. Exped. 1946—47, H.
Hoogstraal.

anaxia Clarke, 1968 (Amallectis); Proc.
U.S. Natl. Mus. 125: 32. Holotype °&,
Guatemala, Volcan Santa Maria, July,
Schaus and Barnes.

andromedana Barnes & McDunnough,
1917 (Olethreutes); Contrib. Nat. Hist.
Lepid. North-Am. 3: 223. “Type” ¢,
USA, Florida, Lee Co., Fort Meyers, r-f.
Andromeda, “Apr 24-30.’ According
to the original description, this species
was described from two males and four
females. Although a holotype was not
designated, the specimen cited above is
clearly labeled “Type.”

angusana Fernald, 1892 (Peronea); Canad.
Entomol. 24: 178. Two syntypes (1 <4,
1 2), USA, New York, West Farms, J.
Angus; and USA, Maine, Penobscot
Co., Orono, 1 August 1884. According
to the original description these two
specimens comprise the type series.

animosana Busck, 1907 (Tortrix); J. N. Y.
Entomol. Soc. 15: 235. Holotype °&,
Mexico, Jalapa, W. Schaus.

anisoneura Diakonoff, 1982 (Eupoecilia);
Zool. Verhandel. (Leiden) 193: 6. Ho-
lotype ¢, Sri Lanka, Ratnapura District,
Uggalkaltota, 350’, Irrigation Bunga-
low, 31 January—8 February 1970, D.
Davis & B. Rowe.

anisoptera Clarke, 1976 (Dudua); Insects
of Micronesia 9: 95. Holotype 6, Mi-
cronesia, Guam, Mt. Chachao, 16 May
1936, O. Swezey.

anthrocodelta Clarke, 1971 (Dichelopa);

1019

Smithsonian Contrib. Zool. 56: 104.
Holotype 3, Rapa Island, Maugaoa,
950’ (292 m), 23 November 1963, J. &
T. Clarke.

aperta Diakonoff, 1968 (Archips); Bull.
U.S. Natl. Mus. 257: 28. Holotype 2,
Philippine Islands, Luzon, Mountain
Province, Baguio, 7 May 1945, J. Fran-
clemont.

aporema Dognin, 1912 (Cnephasia); Het-
eroceres Nouveaux de LAmerique du
Sud 6: 49. Holotype ¢, Colombia, Fas-
sel.

aporrhegma Clarke, 1986 (Dichelopa phal-
aranthes); Smithsonian Contrib. Zool.
416: 136. Holotype 6, Marquesas Is-
lands, Nuku Hiva, Tapuaooa, 2,500’, 30
January 1968, J. & T. Clarke.

apospasta Obraztsov, 1964 (Proeulia);
Proc. U.S. Natl. Mus. 116: 191. Holo-
type @, Chile, Concepcion, October
1902, E. Reed.

approximana Heinrich, 1919 (Olethreutes);
Insect. Inscit. Menst. 7: 65. Holotype 3,
USA, New York, Rensselaer, rolling ter-
minal leaves of loosestrife [Lysima-
chiala12 June 19116, NYS Coll:

aprilana Grote, 1877 (Exentera); Canad.
Entomol. 9: 227. Two syntypes (1 d, 1
2), USA, New York, Albany Co., Al-
bany, April. According to the original
description, this species was described
from one male and one female collected
the ‘‘end of April” by Lintner and Hill.
The syntypes consist only of wings
glued to pieces of cardboard: two fore-
wings of the male, one forewing of the
female. The sexes of the specimens are
indicated on the specimen labels.

aquila Busck, 1914 (Homona); Proc. U.S.
Natl. Mus. 47: 53. Holotype 6, Panama,
Cabima, May 1911, A. Busck.

arammclaina Razowski, 1987 (Heppnero-
grapha); Bull. Polish Acad. Sci. 35: 64.
Holotype ¢, Costa Rica, Cartago Prov-
ince, Tuis, 28 May—4 June, W. Schaus.

argema Clarke, 1986 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 152. Holo-
type 2, Marquesas Islands, Fatu Hiva,

1020

Mt. Teoaiua, 2,000’, 22 March 1968, J.
& T. Clarke.

argentifasciata Heppner, 1989 (Choristo-
neura); Florida Entomol. 72: 104. Ho-
lotype 6, USA, Florida, Glades Co.,
Fisheating Creek, Palmdale, 7-10 May
1964, R. Hodges.

argentifurcatana Grote, 1876 (Conchylis);
Canad. Entomol. 8: 206. Syntype ¢é,
Canada, Ontario, Port Stanley, W. Saun-
ders. Although the specimen bears a la-
bel “‘London, Ont.,” according to the
original description the specimens were
taken at Port Stanley, by ““W. Saunders
from London, Ontario.”’ The disposition
of the other syntype is unknown to us.

argentina Brown, 1991 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 64. Holotype 2, Argen-
tina, Misiones, Puerto Rico, 4—8 April
LOW, Crs2O, Flint.

argillacea Clarke, 1976 (Ruthita); Insects
of Micronesia 9: 26. Holotype 2, Mi-
cronesia, Guam, May 1936, O. Swezey.

argodonta Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107g lS4 yHoletype 925) Brazile St
Catherines”’ [Santa Catarina], EK John-
son.

argutipunctana Blanchard & Knudson,
1983 (Phaneta); J. Lepid. Soc. 37: 143.
Holotype ¢d, USA, Texas, Hemphill
Co., Canadian, 15 August 1971, A. &
M. E. Blanchard.

argyraspis Razowski, 1984 (Saphenista);
Ann. Zool. Warsaw 38: 278. Holotype
2, Venezuela, Distrito Federal, 14 km
NE Tovar, 28—29 February 1976, C. &
O. Flint.

argyrospiloides Clarke, 1971 (Dichelopa);
Smithsonian Contrib. Zool. 56: 118.
Holotype 2, Rapa Island, Maurua, 600’
(184 m), 25 October 1963, J. Clarke.

arizonana Powell, 1975 (Epiblema); Pan-
Pac. Entomol. 51: 101. Holotype ¢,
USA, Arizona, Coconino Co., Oak
Creek Canyon, Flagstaff, rf. Rudbeckia
laciniata, em: 28 June 1936, G. Engle-
hardt.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

arizonensis Heinrich, 1914 (Evetria albi-
capitana); Proc. U.S. Natl. Mus. 57: 57.
Holotype 6, USA, Arizona, Santa Cat-
alina Mountains, r.f. Pinus cembroides,
23 June 1917, Hopkins no. 13977, G.
Hofer.

arrhostia Clarke, 1968 (Phtheochroa);
Proc. U.S. Natl. Mus. 125: 56. Holotype
2, Peru, Cusco, Machu Picchu, 2,385
m, 5 March 1959, J. Clarke.

arthuri Dang, 1984 (Cochylis); Canad. En-
tomol. 116: 253. Holotype ¢, Canada,
Saskatchewan, S.E. Saskatoon, r.f. He-
lianthus sp., August 1980.

asthenia Clarke, 1968 (Carolella); Proc.
U.S. Natl. Mus. 125: 55. Holotype 4,
Guatemala, Palin, July, Schaus &
Barnes.

asynthetes Diakonoff, 1968 (Gatesclark-
eana); Bull. U.S. Natl. Mus. 257: 42.
Holotype 6, Philippine Islands, Cadiz,
Occ. Negros, **3/8/29,”’ Batabas.

atascosana Blanchard, 1979 (Eucosma); J.
Lepid. Soc. 33: 212. Holotype ¢, USA,
Texas, Cameron Co., Laguna Atascosa,
22 November 1973, A. & M. Blan-
chard.

atayalicana Kawabe, 1989 (Acleris); Tinea
12: 195. Holotype 3, Taiwan, Hualien
Hsien, Tayuling, forest, 2,500 m, 9-18
June 1980, D. Davis.

atomosana Busck, 1907 (Phalonia); J. N.
Y. Entomol. Soc. 15: 22. Holotype 6,
USA, Pennsylvania, Allegheny Co.,
Pittsburgh, 17 August 1905, H. Engel.

atomosana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 42. Syn-
type ¢, USA, California [San Francisco
or Bear Valley according to the original
description], May—June, 1871. This spe-
cies was described from two males; the
other syntype is presumably in the
BMNH.

atrata Diakonoff, 1973 (Lobesia); Zool.
Monogr. Rijksmus. Nat. Hist. 1: 382.
Holotype ¢, Marianas Islands, Guam,
Piti, “‘7/9/36,”” O. Swezey.

atrilinea Clarke, 1976 (Cryptophlebia); In-
sects of Micronesia 9: 115. Holotype °,

VOLUME 102, NUMBER 4

Micronesia, Ponape, Colonia, 14 Janu-
ary 1953, J. Clarke.

atriplaga Clarke, 1976 (Duessa); Insects of
Micronesia 9: 28. Holotype 2, Micro-
nesia, Rota, nr. Sabana, 21 June 1946,
Townes 807, at light.

atristriga Clarke, 1953 (Epinotia); J. Wash.
Acad. Sci. 43: 228. Holotype 6, USA,
Illinois, Putnam Co., 24 March 1938,
M. Glenn. Although the collection date
of the holotype is cited as “‘March 17,
1945”’ in the original description, the
specimen listed above is unambiguously
labeled as the holotype.

atrodentana Fernald, 1882 (Eccopsis);
Trans. Am. Entomol. Soc. 10: 71. Lec-
totype ¢, Canada, Ontario, London, W.
Saunders. Designated by Miller (1970).

atsushii Bae, 1993 (Lobesia); Japan. J. En-
tomol. 61: 516. Holotype 6, Taiwan,
Hualien Hsien, Mt. Hohuanshan, 3,100
m, 30 July—1 August 1983, A. Kawabe.

audaculana Busck, 1907 (Archips); J. N. Y.
Entomol. Soc. 15: 235. Holotype °,
Mexico, Veracruz, Orizaba, R. Muller.

aurantia Clarke, 1976 (Adoxophyes); In-
sects of Micronesia 9: 127. Holotype 2,
Micronesia, Ponape, Colonia, 12—17
January 1953, J. Clarke.

aurantica Busck, 1920 (Epagoge); Insect.
Inscit. Menst. 8: 84. Holotype 2, Costa
Rica, Cartago Province, Juan Vinas, W.
Schaus.

auraria Clarke, 1949 (Eulia); Acta Zool.
Lilloana (Tucuman) 7: 583. Holotype
3, Chile, Santiago Province, Cajon de
Maypo, Cordillera, El Cencio, 12—20
January 1948, T. Ramirez.

aurata Diakonoff, 1968 (Adoxophyes);
Bull. U.S. Natl. Mus. 257: 11. Holotype
3, Philippine Islands, Luzon, Mt. Mak-
iling, Baker.

aureana Busck, 1907 (Phalonia); J. N. Y.
Entomol. Soc. 15: 26. Holotype 9°,
USA. Pennsylvania, Allegheny Co.,
Oak Station, 17 June 1906, FE Marloff.

aureola Diakonoff, 1977 (Nexosa); Zool.
Verhandel. (Leiden) 158: 15. Holotype
2, British New Guinea, Hydrographer

1021

Mountains, 2,500’, March 1918, Ei-
chhorn Bros.

auricaput Razowski, 1971 (Acleris); Acta
Zool. Cracov. 16: 550. Holotype °, Tai-
wan, Hassenzan, 6 June 1942, S. Issik1i.

aurichalceana Riley, 1881 (Melissopus);
Trans) StieouishAcad: «Seis 41323:
“Type” ¢, USA, Fernald, rf. acorns,
**1/6/76.”” Riley did not mention the
number of specimens he examined; this
is the only Riley specimen of aurichal-
ceana we have been able to locate. It is
labeled *““Type”’ in Riley’s hand.

auricomana Busck, 1907 (Tortrix); J. N. Y.
Entomol. Soc. 15: 236. Holotype 6,
Mexico, June 1906, R. Muller.

auriferana Busck, 1911 (Tortrix); Proc.
U.S. Natl. Mus. 40: 227. Holotype ¢,
Brazil, Parana, Castro, W. Schaus,
“6187 Wlsm. 1908.”

austrina Miller, 1985 (Eucosma); Ann. En-
tomol. Soc. Am. 78: 243. Holotype ¢,
USA, Texas, Hemphill Co., Canadian,
15 August 1971, A. & M. Blanchard.

azumina Yasuda & Kawabe, 1980 (Ar-
chips); Tinea 11: 13. Holotype @, Ja-
pan, Nagano Pref., Azumi V., Shimaji-
ma-Dan, 780 m, 15 April 1977, N. Hi-
bano.

baccharivora Pogue, 1988 (Lorita); Proc.
Entomol. Soc. Wash. 90: 449. Holotype
6, USA, Florida, Okeechobee Co.,
Lake Okeechobee, 27°N, 81°W, Septem-
ber 1984 (lab. reared), D. Green.

bactrana Heinrich, 1923 (Eucosma); Bull.
U.S. Natl. Mus. 123: 117. Holotype ¢,
USA, Colorado, San Juan Co., Silver-
ton, “July 16—23.”

baea Razowski, 1987 (Transtillaspis); Bull.
Polish Acad. Sci. 35: 79. Holotype <6,
Colombia, Narino, Volcan Galeras,
2,900 m, 13 January 1959, J. Clarke.

bakeri Diakonoff, 1968 (Homona); Bull.
U.S. Natl. Mus. 257: 20. Holotype ¢,
Philippine Islands, Luzon, Mt. Makil-
ing, Baker.

balia Diakonoff, 1982 (Acanthoclita); Zool.
Verhandel. (Leiden) 193: 30. Holotype

1022

6, Sri Lanka, Ratnapura District, Ug-
galkaltota, 350’, Irrigation Bungalow,
31 January—8 February 1970, D. Davis
& B. Rowe.

balioleuca Clarke, 1976 (Adoxophyes); In-
sects of Micronesia 9: 136. Holotype 6,
Micronesia, Ponape, N. slope Tamata-
mansakir, 19 January 1953, J. Clarke.

bana Kearfott, 1907 (Phalonia); Trans. Am.
Entomol: “Soci9332775) Lectotype: d,
USA, Illinois, Cook Co., Chicago, June,
J. Reading. Designated by Klots (1942).

banana Busck, 1906 (Lipoptycha); Proc.
Biol. Soc. Wash. 19: 182. Holotype ¢,
USA, Colorado, South Park, Oslar.

baracana Busck, 1907 (Aysterosia); J. N.
Y. Entomol. Soc. 15: 33. Holotype 6,
USA, Missouri, St. Louis Co., St. Louis,
11 July 1906, McElhose.

bascanion Razowski, 1987 (Transtillaspis);
Bull. Polish Acad. Sci. 35: 75. Holotype
6, Peru, Cusco, Machu Picchu, 2,700
m, 6 February 1959, J. Clarke.

basipunctana Walsingham, 1879 (Paedis-
ca); Ill. Lepid. Heter. Brit. Mus. 4: 40.
Syntype 2, USA, California, ‘“‘Lower
Lake,”’ 22 June 1871. Walsingham de-
scribed this species from two males and
three females; the other syntypes are
presumed to be in the BMNH.

batesi Heinrich, 1932 (Talponia); Proc. En-
tomol. Soc. Wash. 34: 20. Holotype 6,
Guatemala, Antiqua, 5,000’, rf. Annona
cherimoia, M. Bates.

batoidea Razowski, 1987 (Transtillaspis);
Bull. Polish Acad. Sci. 35: 75. Holotype
3, Peru, Cusco, Machu Picchu, 5 Feb-
ruary 1959, J. Clarke.

bauhiniae Busck, 1934 (Ancylis); Entomol.
Am. 13: 155. Holotype 6, Cuba, San-
tiago de Vegas, Hab., leaftier in Bauhi-
nia heterophylla, 4 December 1931, R.
Olera.

bebela Razowski, 1987 (Transtillaspis);
Bull. Polish Acad. Sci. 35: 82. Holotype
3d, Colombia, Bogota, Chico, 25 Janu-
ary 1959) ad#Clarke:

beckeri Clarke, 1973 (Eumarozia); J. Lepid.
Soc. 27: 269. Holotype 6, Costa Rica,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Cartago Province, Turrialba, rf. Jug-
lans, em: 29 August 1972, V. Becker.

benjamini Heinrich, 1923 (Thiodia); Bull.
U.S. Nat. Mus. 123: 66. Holotype ¢,
USA, Utah, Vineyard, 12 September
1912, T. Spalding.

bicolor Kawabe, 1963 (Acleris); Tyo to Ga
14: 70. Holotype 6, Japan, Nagano
Pref., Honshu, Naka-karuisawa, 23 Sep-
tember 1962, T. Maenami.

bicolor Kawabe, 1978 (Aterpia); Tinea 10:
174. Holotype ¢, Japan, Tsushima Is-
land, Taterayama, 2 November 1973, T.
Watanabe.

bicolor Powell, 1961 (Decodes); J. Lepid.
Soc. 14: 122. Holotype 2, USA, Cali-
fornia, Napa Co., Mt. St. Helena, 18
April 1939, E. Johnston.

bicolora Kawabe, 1976 (Zeiraphera); Tinea
10: 42. Holotype ¢, Japan, Gunma
Pref., Kumanotaira, 14 July 1952, M.
Hoshino.

bicornigera Razowski, 1984 (Histura);
Acta Zool. Cracov. 27: 212. Holotype
3, Colombia, Fassel.

bicornis Diakonoff, 1968 (Homona); Bull.
U.S. Natl. Mus. 257: 18. Holotype 6,
Philippine Islands, Luzon, Los Banos,
Baker.

biemina Kawabe, 1980 (Apotomis); Tinea
11: 18. Holotype ¢, Japan, Miyagi
Pref., Tokatta, 13 September 1971, W.
Watanabe.

bigaulae Brown, 1999 (Eubetia); J. New
York Entomol. Soc. 106: 181. Holotype
6, Venezuela, Aragua, Rancho Grande,
cloud forest, 1,100 m, 30—31 March
1978, blacklight, J. Heppner.

binotata Brown & Obraztsov, 1991 (Cu-
proxena); in Brown and Powell, Univ.
Calif. Publ. Entomol. 111: 61. Holotype
?, Brazil, Santa Catarina, 22 July 1935,
FE Hoffman.

biquadrana Walsingham, 1876 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 45. Syn-
type 6, USA, California, Shasta Co.,
Pitt River, “end of July” 1871. Wal-
singham described this species from

VOLUME 102, NUMBER 4

two males; the other syntype is pre-
sumed to be in the BMNH.

bira Kawabe, 1976 (Endothenia); Tinea 10:
43. Holotype 6, Japan, Ishikawa Pref.,
Anamizu, Bira, 18 August 1961, A. Ka-
wabe.

birdana Busck, 1907 (Hysterosia); J. N. Y.
Entomol. Soc 15: 32. Holotype 9°,
USA, New York, Rye, “‘bores Helian-
thus,’ H. Bird.

biserrata Brown, 1991 (Paraptila); J. Lep-
id. Soc. 44: 269. Holotype ¢, Costa
Rica, Cartago Province, Turrialba, 22—
28 February 1965, S. & W. Duckworth.

bittana Busck, 1906 (Hemimene); Proc.
Biol. Soc. Wash. 19: 179. Holotype ¢,
USA, Pennsylvania, Allegheny Co.,
Pittsburgh, 29 May 1905, H. Engel.

blackmorei Obraztsov, 1963 (Acleris emar-
gana); Proc. U.S. Natl. Mus. 114: 208.
Holotype ¢, Canada, British Columbia,
Goldstream, 7 August 1923, E. Black-
more.

blanchardi Miller, 1978 (Rhyacionia); in
Powell and Miller, U.S. Dept. Agric.,
Agric. Handb. 514: 19. Holotype ¢d,
USA, Texas, Montgomery Co., Conroe,
10 March 1968, A. & M. Blanchard.

blechra Razowski, 1987 (Transtillaspis);
Bull. Polish Acad. Sci. 35: 79. Holotype
3, Colombia, Narifio, Volcan Galeras,
2,900 m, 13 January 1959, J. Clarke.

bobana Kearfott, 1907 (Eucosma); Trans.
Am. Entomol. Soc. 33: 26. ‘“‘Type’’ 3,
USA, Colorado, Salida, 11 June 1888,
W. Dietz. This species was described
from three ‘“‘co-types”’ (Salida, Colora-
do; Southwestern Colorado; Harris Co.,
Texas). Although Heinrich (1923) indi-
cated that the “type” (from Salida, Col-
orado) was in the AMNH, Klots (1942)
correctly identified the USNM as the
place of disposition. Klots designated
the AMNH specimen (Southwestern
Colorado) as a “‘lectoparatype.”” The
disposition of the third syntype is un-
known.

bolanderana Walsingham, 1876 (Paedis-
ca); Ill. Lepid. Heter. Brit. Mus. 4: 42.

1023

Two syntypes (2 66), USA, California,
Siskiyou Co., Mount Shasta, August
1871. This species was described from
three males and two females; the other
syntypes are presumed to be in the
BMNH.

boliviae Razowski, 1988 (Proeulia); Acta
Zool. Cracov. 31: 407. Holotype 6, Bo-
livia, Cochabamba, Incachaca, tropical
cloud area, 2,100 m, 27 August—5 Sep-
tember 1956, L. Pefia.

boliviana Brown, 1991 (Bidorpitia); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 75. Holotype @, Bolivia,
Cochabamba, Incachaca, tropical cloud
area, 2,100 m, 27 August—5 September
1956, Lk. Pena:

boliviana Razowski, 1984 (Histura); Acta
Zool. Cracov. 27: 212. Holotype 3, Bo-
livia, Cochabamba, Incachaca, tropical
cloud area, 2,100 m, 27 August—5 Sep-
tember 1956, L. Pefia.

boscantica Dognin, 1912 (Tortrix); Heter-
oceres L-Amerique du Sud 6: 48. Ho-
lotype ¢, Colombia, Cali, San Antonio,
Fassel.

bourquini Clarke, 1949 (Eulia); Acta Zool.
Lilloana (Tucuman) 7: 581. Holotype
?, Brazil, Parana, Castro, W. Schaus.

brachistocera Razowski, 1987 (Transtillas-
pis); Bull. Polish Acad. Sci. 35: 81. Ho-
lotype 6, Colombia, Bogota, Chico, 25
January 1959, J. Clarke.

brachystigma Clarke, 1965 (Nesochoris);
Proc. U.S. Natl. Mus. 117: 76. Holotype
3d, Chile, Juan Fernandez Island, Mas-
atierra, E] Rabanal, 350 m, 27 February
195i 0P) Kuschel:

bracteatana Fernald, 1881 (Grapholitha);
in Comstock, Annu. Rept. Dept. Agric.
1881: 265. Lectotype 6, USA, Califor-
nia, Jolon, em: 13 September 1880.
Designated by Miller (1970).

bramiliana Brown, 1991 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 61. Holotype ¢, Brazil,
Rio de Janeiro, Petropolis, 600 m, 10
October 1985, S. Miller.

brandinojuxta Razowski, 1987 (Transtillas-

1024

pis); Bull. Polish Acad. Sci. 35: 77. Ho-
lotype ¢, Bolivia, Cochabamba, Inca-
chaca, tropical cloud area, 2,100 m, 27
August—5 September 1956, L. Pena.

brauni Heinrich, 1931 (Anchylopera); Proc.
U.S. Natl. Mus. 79: 11. Holotype 4,
USA, Ohio, Adams Co., Beaver Pond,
11 May 1927.

braziliana Brown, 1991 (Punctapinella);
Los Angeles Co. Mus. Contrib. Zool.
423: 4. Holotype °, Brazil, Santa Ca-
tarina, 20 November 1936, E Hoffman.

britana Busck, 1906 (Hemimene); Proc.
Biol. Soc. Wash. 19: 178. Lectotype ¢,
Canada, British Columbia, Kaslo, H.
Dyar. Designated by Miller (1983).

britannia Kearfott, 1904 (Acleris); Canad.
Entomol. 36: 138. Holotype ¢, Canada,
British Columbia, Kaslo, H. Dyar.

broui Knudson, 1986 (Dichrorampha); J.
Lepid. Soc. 40: 323. Holotype 6, USA,
Louisiana, St. Tammany Parish, 4.2 mi
NE Abita Springs, sec. 24, T6, SR 128,
uv [light], 12 April 1985,V. Brou.

brunana Brown, 1990 (Hynhamia); Ento-
mol. Scand. 21: 325. Holotype d, Peru,
Angasmarca.

brunneopurpuratum Heinrich, 1923 (Exar-
tema); Proc. Entomol. Soc. Wash. 25:
118. Holotype 2, USA, Virginia, Fair-
fax Co., Falls Church, rf. alder, em: 1
August 1913.

bumeliana Heinrich, 1926 (Goditha); Bull.
U.S. Natl. Mus. 132: 8. Holotype 6,
USA, Texas, Dallas Co., Dallas, “‘Di-
chrorampha boumelliana Boll mss.,
two generations, June and Oct. in the
rolled leaves of Boumellia languinosa.”
Although misspelled as ‘‘rumeliana”
immediately prior to its description, it is
spelled correctly as “‘bumeliana’’ pre-
viously on the same page, where it is
identified as the type species of Godi-
tha.

burgessiana Zeller, 1875 (Phoxopteris);
Verhandel. Zool.-Bot. Ges. Wein 25:
252. “Type” ¢°, USA, Massachusetts,
Essex Co., Beverly, 18 June 1869. This
species was described from a male and

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

two females; the specimen listed above
is the only Zeller specimen of burges-
siana we could find.

bununa Kawabe, 1989 (Acleris); Tinea 12:
196. Holotype 2, Taiwan, Hualien
Hsien, Tayuling, 2,560 m, 1 April 1984,
A. Kawabe.

busckana J. A. Comstock, 1939 (Pharma-
cis); Bull. South. Calif. Acad. Sci. 38:
112. Holotype ¢, USA, California, Los
Angeles Co., El Segundo, em: 16 No-
vember 1938, W. Pierce.

busckana Heinrich, 1923 (Rhyacionia);
Bull. U.S. Natl. Mus. 123: 17. Holotype
3d, USA, New York, Nassau Co., Long
Island, Bellmore, April 1913, G. Engle-
hardt.

busckana Keifer, 1933 (Clepsis); Calif.
Dept. Agric. Mon. Bull. 22: 351. Ho-
lotype 6, USA, California, San Francis-
co, 8 February 1933, rf. Scophularia
californica, em: 3 March 1933, H. Kei-
fer.

buscki Brown, 2000 (Accuminulia); J. Lep-
id. Soc. 53: 61. Holotype 3, Chile, San-
tiago Province, reared from grape
(fruit), em: 12 April 1954, M. Ramsay.

bushiensis Kawabe, 1980 (Epinotia); Tinea
11: 22. Holotype 6, Japan, Saitama
Pref., Bushi, Iruma, 20 October 1978,
H. Inoue.

bushnelli Busck, 1914 (Evetria); Proc. En-
tomol. Soc. Wash. 66: 144. Holotype @,
USA, New Mexico, Ft. Bayard, rf. Pi-
nus ponderosa, em: 19 March 1914, C.
Bushnell.

caeruleana Walsingham, 1879 (Grapholi-
tha); Ill. Lepid. Heter. Brit. Mus. 4: 66.
Syntype 6, USA, Oregon, Rouge River,
May 1872. Walsingham described this
species from one male and one female;
the female is presumed to be in the
BMNH.

caeruleumana Kawabe, 1980 (Zeiraphera);
Tinea 11: 24. Holotype 3, Japan, Akita
Pref., Tamagawa, 2 September 1970, T.
Watanabe.

caesiata Clarke, 1968 (Cochylis); Proc.

VOLUME 102, NUMBER 4

U.S. Natl. Mus. 125: 18. Holotype °&,
Venezuela, Aragua, Rancho Grande,
1,100 m, 16—23 October 1966, S. & W.
Duckworth.

californiae Heinrich, 1923 (Hystricophora
strygiana); Bull. U.S. Natl. Mus. 123:
256. Holotype ¢, USA, California,
Lake Tahoe, Deer Park Springs, “‘July
8—15.’’ Heinrich (1923) consistently
misspelled Hystrichophora in his revi-
sion.

callosoma Clarke, 1976 (Cryptophlebia);
Insects of Micronesia 9: 112. Holotype
2, Micronesia, Guam, Pt. Oca, near
Agana, 12 May 1945, Bohart & Gres-
sitt.

campestrana Zeller, 1875 (Sericoris); Ver-
handel. Zool.-Bot. Ges. Wein 25: 282.
“Type” 6, USA [Maine or Massachu-
setts according to the original descrip-
tion], Packard. Zeller did not mention
the number of specimens examined, al-
though it is clear that he had both sexes.
This is the only Zeller specimen of cam-
pestrana that we were able to find.

campicolana Walsingham, 1879 (Cochylis);
Ill. Lepid. Heter. Brit. Mus. 4: 29. Two
syntypes (6, 2), USA, California, Men-
docino Co., 10 June 1871. Walsingham
described this species from three males
and two females; the other syntypes are
presumed to be in the BMNH.

canariana Barnes & Busck, 1920 (Ayster-
osia); Contrib. Nat. Hist. Lepid. North
Am. 4: 218. Holotype 6, USA, Arizo-
na, White Mountains.

candidus Pogue, 1986 (Apolychrosis); in
Cibrian-Tovar et al., Cone and Seed In-
sects of the Mexican Conifers: 23. Ho-
lotype 6, Mexico, Puebla, Atotocoyan,
rf. Pinus strobus, em: 2 October 1981,
T. Mendez.

canitia Clarke, 1986 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 149. Holo-
type °, Marquesas Islands, Nuku Hiva,
Tapuaooa, 2,500’, 30 January 1968, J.
& T. Clarke.

canofascia Forbes, 1930 (Olethreutes); Sci.
Surv. Porto Rico and Virgin Islands 12:

1025

86. Holotype 3, Puerto Rico, Rio Pie-
dras, 10 July 1916, E. Smith.

canusana Wright, 1997 (Phaneta); in
Wright, Brown and Gibson, J. Lepid.
Soc. 51: 122. Holotype ¢, USA, Ohio,
Adams Co., Lynx Prairie Preserve, Sta-
tion 6, 17 March 1989, D. Wright.

capitana Busck, 1906 (Hemimene); Proc.
Biol. Soc. Wash. 19: 178. Holotype 3,
USA, Colorado, South Park, Oslar.

capizziana Obraztsov, 1963 (Acleris); Proc.
U.S. Natl. Mus. 114: 252. Holotype °,
USA, Oregon, Bendon, 24 September
1956, J. Capizzi.

capronata Razowski, 1988 (Ernocornutia);
Acta Zool. Cracov. 31: 398. Holotype
6, Colombia, Cauca, Paramo de Parace,
Lake San Rafael, 3,570 m, 27 January
195934).. Clarke:

carduana Busck, 1907 (Polychrosis); J. N.
Y. Entomol. Soc. 15: 134. Holotype &,
USA, Maryland, Montgomery Co., Hy-
attsville, rf. thistle, August 1906, A.
Busck.

carnana Barnes & Busck, 1920 (Tortrix);
Contrib. Nat. Hist. Lepid. North Am. 4:
214. Holotype 6, USA, California, San
Bernardino Co., San Bernardino Moun-
tains, Camp Baldy, “‘June 24—20.”

carpophagoides Clarke, 1951 (Cryptophle-
bia); J. Wash. Acad. Sci. 41: 299. Ho-
lotype 6, Argentina, Tucuman, rf. seeds
of pacara, K. Hayward.

caryana Fitch, 1856 (Grapholitha); New
York Agric. Rept. 16: 459. Two syn-
types (2 6d), USA, New York [no data
on specimen labels],“‘14,957,’’ Type
No. 394. Fitch did not indicate the num-
ber of specimens examined.

cataclasta Diakonoff, 1982 (Dicnecidia);
Zool. Verhandel. (Leiden) 193: 42. Ho-
lotype ¢, Sri Lanka, Anuradhapura Dis-
trict, Wildlife Sanctuary Bungalow, Hu-
nuwipagama, Wilpattu, 200’, 10-19
March 1970, D. Davis & B. Rowe.

cathedra Clarke, 1976 (Lobesia); Insects of
Micronesia 9: 105. Holotype 6, Micro-
nesia, Guam, Pt. Oca, 30 May 1945,
Bohart & Gressitt.

1026

catopta Razowski, 1988 (Ernocornutia);
Acta Zool. Cracov. 31: 397. Holotype
6, Colombia, Cauca, Paramo de Parace,
Lake San Rafael, 3,570 m, 27 January
1959, J: Clarke.

caulocatax Razowski, 1984 (Conchylis);
Ann. Zool. Cracov. 38: 278. Holotype
36, Venezuela, Bolivar, Morichal Tauca,
22 km E Rio Caura, 8—9 February 1976,
C. & O. Flint.

celiae Clarke, 1976 (Laspeyresia); Insects
of Micronesia 9: 120. Holotype 2, Mi-
cronesia, Yap Islands, Yap Island, Co-
lonia, 21 June 1957, C. Sabrosky.

celtisana Riley, 1881 (Paedisca); Trans. St.
Louis Acad. Sci 4: 319. Holotype ¢,
USA, Texas, rf. Celtis, Boll. Riley de-
scribed this species from one specimen.

cephalanthana Heinrich, 1921 (Phalonia);
J. Agric. Res. 20: 825. Holotype ¢,
USA, Texas, Charlotte, rf. Cephalan-
thus occidentalis, em: 16 September
1918, C. Heinrich.

cercocarpana Dyar, 1903 (Eucosma); Proc.
Entomol. Soc. Wash. 5: 297. Three syn-
types (3 66), USA, Colorado, Platte
Canyon, r.f. Cercocarpus parvifolius,
Dyar & Caudell. Dyar described this
species from three males and one fe-
male.

cerinus Kawabe, 1978 (Neoansthamna);
Tinea 10: 182. Holotype 6, Japan, Hon-
shu, Nippara, Tokyo, 20 June 1964, A.
Kawabe.

cervinana Fernald, 1882 (Teras); Trans.
Am. Entomol. Soc. 10: 65. Lectotype
3d, USA, Massachusetts, Cambridge (7).
Designated by Obraztsov (1963).

chalcana Packard, 1867 (Conchylis); Proc.
Boston Soc. Nat. Hist. 11: 56. ““Type”’
(sex unknown; only a forewing), Can-
ada, Labrador, Strawberry Harbor, near
Cape Webuc, 26 July. The description
does not indicate how many specimens
Packard examined, but the species is
listed as “‘uncommon.”’

chalybeana Fernald, 1882 (Peronea);
Trans. Am. Entomol. Soc. 10: 65. Two
syntypes (1 ¢6, 1 2), USA, New York,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

“L.I.”’ [Long Island]. Of the three spec-
imens referred to by Fernald (1.e., two
males and one female from Orono,
Maine and New York), only the two
specimens from New York have been
located by us.

changi Kawabe, 1989 (Capua); Tinea 12:
192. Holotype 6, Taiwan, Haulien
Hsien, Hohuanshan, 3,100 m, 30 July—
1 August 1983, A. Kawabe.

charma Clarke, 1968 (Phalonia); Proc.
U.S. Natl. Mus. 125: 16. Holotype 2°,
Argentina, Tucuman, Ciudad Universi-
taria, 800 m, 17 February 1959, J.
Clarke.

chica Brown, 1984 (Corticivora); Proc. En-
tomol. Soc. Wash. 86: 283. Holotype 3,
USA, Florida, Highlands Co., Lake
Placid, Archbold Biol. Sta., 1-7 May
1964, R. Hodges.

chiquitana Barnes & Busck, 1920 (Platy-
nota); Contrib. Nat. Hist. Lepid. North
Am. 4: 213. Holotype 6, USA, Califor-
nia, San Bernardino Co., Loma Linda.

chlamydata Dognin, 1912 (Polyortha); Het-
eroceres Nouveaux L Amerique du Sud
6: 50. Holotype ¢, Colombia, Cali, San
Antonio.

chloromonas Razowski, 1984 (Chlorortha);
Acta Zool. Cracov. 27: 215. Holotype
6, Venezuela, Aragua, Rancho Grande,
1,100 m, 16—23 October 1966, S. & W.
Duckworth.

chortodes Diakonoff, 1968 (Eudemis); Bull.
U.S. Natl. Mus. 257: 52. Holotype ¢,
Philippine Islands, Mountain Province,
Luzon, Baguio, 2 June 1945, J. Francle-
mont.

chrysea Heinrich, 1926 (Bactra verutana);
Bull. U.S. Natl. Mus. 132: 85. Holotype
6, USA, California, San Bernardino
Co., Loma Linda, ‘“‘March 24-30.”

cibdela Razowski, 1988 (Argyrotaenia);
Acta Zool. Cracov. 31: 409. Holotype
3d, Colombia, Cusco, Tambomachay, 3
February 1959, J. Clarke.

cibriani Miller, 1988 (Rhyacionia); J. Lep-
id. Soc. 42: 236. Holotype ¢, Mexico,

VOLUME 102, NUMBER 4

Mexico, Paso de Cortes, r.f. Pinus har-
twegii, 12 March 1984, D. Cibrian.
cinderella Riley, 1872 (Acleris); Board of
Agric., 4th Annu. Rept. Noxious, Ben-
eficial and Other Insects . . . of Missou-
ri: 46. “Type’”” 2, USA, Missouri [no
data on specimen label], “*928.P.”’ The
description does not indicate how many
specimens Riley examined; this is the
only Riley specimen of A. cinderella we
could find. Although there is no indi-
cation in Riley’s hand that is was among
the original type series, two subsequent
labels identify it as “‘type.”’

cinereolineana Heinrich, 1923 (Thiodia);
Bull. U.S. Natl. Mus. 123: 52. Holotype
6, USA, Utah, Juab Co., Eureka, 21
April 1910, T. Spalding.

cirrholepida Clarke, 1976 (Icelita); Insects
of Micronesia 9: 43. Holotype 3, Mi-
cronesia, Palau, Koror Island, Koror, 25
April 1957, C. Sabrosky.

citrana Fernald, 1889 (Tortrix); Entomol.
Am. 5: 18. ““Type”’ 2, USA, California,
“the larva had eaten into an orange
from Coquillett.”’ There is no indication
of how many specimens Fernald ex-
amined; this specimen is clearly labeled
; ype.

citrogramma Clarke, 1976 (Cryptophlebia);
Insects of Micronesia 9: 115. Holotype
?, Micronesia, Kusaie, Hill 541, 165 m,
18 April 1953, J. Clarke.

claduncus Razowski, 1988 (Chilips); Acta
Zool. Cracov. 31: 389. Holotype <4,
Chile, Centro-Austral, January—March
1898, V. Izquerdo.

clarkeana Razowski, 1984 (Polyortha);
Acta Zool. Cracov. 27: 224. Holotype
2, Argentina, Tucuman, Ciudad Univ-
ersitaria, 800 m, 17 February 1959, J.
Clarke.

clarkei Blanchard & Knudson, 1983 (Pha-
neta); Proc. Entomol. Soc. Wash. 85:
847. Holotype ¢6, USA, Texas, Hem-
phill Co., Canadian National Grassland,
Lake Marvin, 9 October 1982, E. Knud-
son.

clarkei Diakonoff, 1964 (Bactra); Zool.

1027

Verhandel. (Leiden) 70: 8. Holotype 6,
British Guiana, Vrijheidslust, larva in
stem of sedge (Cyperus), H. Moore.

clarkei Obraztsov, 1963 (Acleris); Proc.
U.S. Natl. Mus. 114: 251. Holotype 6,
USA, Washington, Kittitas Co., Cle
Elum, 9 April 1931, J. Clarke.

clarkei Obraztsov, 1966 (Pseudomeritastis);
Proc. U.S. Natl. Mus. 118: 226. Holo-
type 6, Colombia, Cauca, 17 km SE
Popayan, 2,000 m, 10 January 1959, J.
Clarke.

clarki Clarke, 1951 (Corticivora); J. Wash.
Acad. Sci. 41: 46. Holotype ¢, USA,
Connecticut, North Guilford, rf. red
pine (Pinus resinosa), G. Plumb.

clavana Fernald, 1882 (Semasia); Trans.
Am. Entomol. Soc. 10: 72. Lectotype
6, USA, Massachusetts, Truro, August
8. Designated by Miller (1970).

clavosa Diakonoff, 1973 (Lobesia); Zool.
Monogr. Rijksmus. Nat. Hist. 1: 381.
Holotype 36, Samoa, Fagatogo, Tutuila,
12 August 1940, ex-Trema(?), O. Swez-
ey.

claypoleana Riley, 1882 (Steganoptycha);
Am. Nat. 16: 914. “Type” @, USA,
Ohio, em: 2 June 1883. There are two
identically labeled Riley specimens (i.e.,
‘**360L, Iss. June 2 *83’’), one of which
has a USNM type label affixed. The
original description does not indicate
how many specimens Riley examined.

clemensiana Fernald, 1879 (Tortrix); Can-
ad. Entomol. 11: 155. Lectotype d (des-
ignated here), USA, Maine/Massachu-
setts/New York/Wisconsin [no locality
data on specimen label], 10 August
1880. Selected by Obraztsov in 1956.
Fernald described this species from 20
males and 17 females; he did not des-
ignate a type specimen or a type local-
ity, instead listing the states from which
he had specimens.

clenchi Clarke, 1980 (Proeulia); J. Lepid.
Soc. 34: 182. Holotype 6, Chile, Des-
venturadas Islands, San Ambrosio Is-
land, 450 m, 8 November 1960, G. Kus-
chel.

1028

cneca Obraztsov, 1964 (Proeulia); Proc.
U.S. Natl. Mus. 116: 193. Holotype ¢,
Chile, Santiago, Guayacan, 1,100 m,
October 1952, L. Pefia.

cockerellana Kearfott, 1907 (Tortrix);
Trans. Am. Entomol. Soc. 33: 71. Lec-
totype 6, USA, Colorado, Garfield Co.,
Glenwood Springs, August 1889. Des-
ignated by Rubinoff and Powell (1999).

cockleana Kearfott, 1904 (Enarmonia);
Canad. Entomol. 36: 137. Three syn-
types as follows: Canada, Alberta,
Banff, ““VII.9” (2 36) (one labeled
‘““cotype,”’ the other with a red USNM
type label); Canada, Manitoba, Awene,
20 July 1905, Criddle (1 @) (labeled
““cotype’’). Although Klots (1942) des-
ignated lectotypes for nearly all other
Kearfott Olethreutinae in the AMNH,
he refers to “two paralectotypes”’ of this
species.

collilonga Blanchard & Knudson, 1984
(Pelochrista); Proc. Entomol. Soc.
Wash. 86: 446. Holotype 6, USA, Tex-
as, Brown Co., Lake Brownwood State
Park, 21 April 1966, A. & M. Blan-
chard.

coloradanus Fernald, 1882 (Lophoderus);
Trans. Am. Entomol. Soc. 10: 67. Two
syntypes (1 6, 1 ¢), USA, Colorado.
According to the original description,
this species was described from one
male and one female from Colorado.

coloradensis Adamski, 1986 (Apotomis);
Canad. Entomol. 118: 662. Holotype ¢d,
USA, Colorado, Maysville, 17 August
1945, H. Ramstadt.

coloradensis Heinrich, 1920 (Evetria col-
jaxiana); Proc. U.S: Natl» Mus: 57: 55.
Holotype 6, USA, Colorado, Mount
Manitou, r.f. cones of Abies concolor, 6
October 1915, J. Pollock, W. Edmon-
ston, G. Hofer, and A. Champlain.

comandrana Fernald, 1892 (Teras); Canad.
Entomol. 24: 121. Lectotype 2, USA,
Massachusetts, Hampshire Co., Am-
herst, “‘Hatch Ex. Station.”’ Designated
by Obraztsov (1963).

comandranum Clarke, 1953 (Exartema); J.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Wash. Acad. Sci. 43: 230. Holotype 6,
USA, Illinois, Putnam Co., rf. Coman-
dra umbellata, 11 June 1942, M. Glenn.

comstocki Clarke, 1952 (Sonia); Bull.
South. Calif. Acad. Sci. 51: 62. Holo-
type ¢d, USA, California, San Diego
Co., San Felipe Wash, 11 September
1938, J. Comstock.

comstockiana Fernald, 1879 (Retinia); Can-
ad. Entomol. 11: 157. Lectotype’ d,
USA, New York, Tompkins Co., Ithaca.
Designated by Miller (1970).

concitatricana Heinrich, 1923 (Gwendoli-
na); Bull. U.S. Natl. Mus. 123: 189. Ho-
lotype 3d, USA, Texas, Kerr Co., Kerr-
ville, 1 June 1906, E Pratt.

concubitana Heinrich, 1923 (Gretchena);
Bull. U.S. Natl. Mus. 123: 181. Holo-
type 6, USA, Florida, Leon Co., Mon-
ticello; rf. “Hickoria’ *(\Caryale iS
March 1914, J. Gill.

conditana Walsingham, 1879 (Penthina);
Ill. Lepid. Heter. Brit. Mus. 4: 31. Syn-
type 6, USA, California, Mendocino
Co., 24 May 1871. Walsingham de-
scribed this species from two males; the
other syntype is presumed to be in the
BMNH.

confusa Obraztsov, 1962 (Anopina); Am.
Mus. Novit. 2082: 30. Holotype 6,
Mexico, Guerrero, Sierra de las Aguas
Escondidas, 9,500’, July, H. Smith.

conigerana Zeller, 1875 (Tortrix); Verhan-
del. Zool.-Bot. Ges. Wien 25: 227.
“Type”? 3d, USA, Maine, Packard. Zell-
er described this species from one male
from Maine, three females from Mas-
sachusetts, and one female from New
York. This is the only specimen of the
original series that we could find.

coniogramma Clarke, 1976 (Eucosma); In-
sects of Micronesia 9: 53. Holotype 4,
Micronesia, Kusaie, Pukusrik, 9-14
February 1953, J. Clarke.

consacculuana Brown, 1991 (Dorithia); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 47. Holotype 6, Mexico,
Chiapas, San Cristébal [de] las Casas,
17-21 July 1964, P. Spangler.

VOLUME 102, NUMBER 4

consobrina Busck, 1914 (Homona); Proc.
U.S. Natl. Mus. 46: 54. Holotype °&,
Panama, Porto Bello, March 1911, A.
Busck.

consobrinana Heinrich, 1923 (Eucosma);
Bull. U.S. Natl. Mus. 123: 128. Holo-
type ¢, USA, South Dakota, Union Co.,
Elk Point, August 1913, C. Ainslie.

consociana Heinrich, 1923 (Eucosma);
Bull. U.S. Natl. Mus. 123: 101. Holo-
type 6, USA, Utah, Juab Co., Eureka,
23 July 1911, T. Spalding.

constantia Clarke, 1968 (Cochylis); Proc.
U.S. Natl. Mus. 125: 22. Holotype 2°,
Peru, Cusco, Machu Picchu, 2,700 m, 6
February 1959, J. Clarke.

constellatana Zeller, 1875 (Sericoris); Ver-
handel. Zool.-Bot. Ges. Wein 25: 279.
milype + .d.USA, New «York, 2/71,”
Speyer. Zeller described this species
from six males and two females from
Ohio and New York. This is the only
Zeller specimen of S. constellatana we
could find.

contrasta Brown, 2000 (Lobogenesis);
Proc. Entomol. Soc. Wash. 102: 32. Ho-
lotype 6, Bolivia, Incachaca, Cocha-
bamba, tropical cloud forest area, 2,100
m, 27 August-5 September 1956, L.
Pena.

contrastana Kearfott, 1907 (Commophila);
Canad. Entomol. 39: 160. Lectotype <,
USA, Pennsylvania, Allegheny Co.,
Oak Station, 13 June 1907, E Marloff.
Designated by Klots (1942).

conversana Walsingham, 1879 (Grapholi-
tha); Ill. Lepid. Heter. Brit. Mus. 4: 66.
Syntype 6, USA, Oregon, Camp Wat-
son, “John Day’s river,” “‘beginning of
April 1872.’ This species was de-
scribed from three males and one fe-
male; the other syntypes presumably are
in the BMNH.

coppelia Clarke, 1976 (Icelita tatarana);
Insects of Micronesia 9: 36. Holotype
3d, Micronesia, Guam, Piti, 15 Septem-
ber 1936, rf. Intsia bijuga, O. Swezey.

cordiae Busck, 1934 (Ancylis); Entomol.
Am. 13: 154. Holotype 6, Cuba, Ha-

1029

vana, Santiago de Vegas, leaf tier on
Cordia globosa, 26 November 1931, A.
Otero.

cornifoliana Riley, 1881 (Phoxopteris);
Trans. St. Louis Acad. Sci. 4: 324.
“Type” 3, USA, Kansas, Riley Co.,
Manhattan, r.f. Cornus paniculata, em:
7 April 1873. Of two specimens re-
ferred to in the original description, this
is the only specimen we have found.

cornucopis Walsingham, 1914 (Enarmon-
ia); Biol. Centr.-Am., Lepid., Heter. 4:
240. Holotype 2, Mexico, Oaxaca, Sa-
lina Cruz, W. Schaus. Described from
the single female ‘“‘Type.”’

cornuta Brown & Obraztsov, 1990 (Cu-
proxena); in Brown and Powell, Univ.
Calif. Publ. Entomol. 111: 53. Holotype
36, Costa Rica, Cartago Province, Juan
Vinas, W. Schaus.

cornutana Dyar, 1903 (Epinotia); Proc. En-
tomol. Soc. Wash. 5: 231. Holotype °,
USA, Arizona, Coconino Co., Williams,
“22.7, H. Barber. According to the
original description, this species was de-
scribed from a single female.

cortesi Clarke, 1987 (Cryptophlebia); Acta
Entomol. Chileana 14: 8. Holotype 6,
Chile, Iquique Province, Region Tara-
paca, Valle de Azapa, 25 m, r.f. Acacia
macracantha, 10 January 1969, R.
Mendoza.

corylana Fernald, 1882 (Eccopsis); Trans.
Am. Entomol. Soc. 10: 71. Lectotype
3d, USA, New Hampshire, White Moun-
tains. Designated by Miller (1970).

corynetes Diakonoff, 1982 (Epinotia);
Zool. Verhandel. (Leiden) 193: 59. Ho-
lotype 6, Sri Lanka, Ratnapura District,
Uggalkaltota, 350’, Irrigation Bunga-
low, 31 January—8 February 1970, D.
Davis & B. Rowe.

cosmocosta Razowski 1987 (Coryssoval-
va); Tinea 12(suppl.): 130. Holotype ¢,
Colombia, Cauca, Paramo de Parace,
Lake San Rafael, 3,570 m, 29 January
1959; Je Clarke:

costaricana Razowski 1984 (Histurodes);
Acta Zool. Cracov. 27: 213. Holotype

1030

36, Costa Rica, San Pedro de Montes de
Oca.

costimaculana Fernald, 1882 (Penthina);
Trans. Am. Entomol. Soc. 10: 70. Lec-
totype 2, USA, Maine, Penobscot Co.,
Orono, 17 June 1881. Designated by
Miller (1970).

costinotana Franclemont, 1986 (Lozotaen-
ia); Proc. Entomol. Soc. Wash. 88: 57.
Holotype ¢, USA, Maine, Penobscot
Co., Passadumkeag, 30 June 1956, J.
Franclemont.

cracens Diakonoff, 1982 (Eupoecilia);
Zool. Verhandel. (Leiden) 193: 6. Ho-
lotype ¢, Sri Lanka, NE District, Kan-
da-ela Reservoir, 5.6 mi SW Nuwara
Eliya, 6,200’, 10—21 February 1970, D.
Davis & B. Rowe.

crambitana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 43. Syn-
type 6, USA, California, Mount Shasta,
August 1871. This species apparently
was described from one male and one
female; the female presumably is in the
BMNH.

cristata Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 155. Holotype 6, Costa Rica, Ca-
chi, W. Schaus.

crocoptila Diakonoff, 1968 (Peridaedala);
Bull. U.S. Natl. Mus. 257: 79. Holotype
?, Philippine Islands, Davao Province,
Mindanao, E slope Mt. McKinley,
7,000’, 22 September 1946, CNHM
Philippine Zool. Exped. 1946—47, H.
Hoogstraal.

cruentana Blanchard & Knudson, 1981
(Phaneta); J. Lepid. Soc. 35: 169. Ho-
lotype 6, USA, Texas, Anderson Co.,
Engeling Wildlife Management Area,
near Tennessee Colony, 28 June 1978,
A. & M. Blanchard.

cryptica Brown, 1991 (Bidorpitia); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 75. Holotype 2°, Vene-
zuela, Aragua, Rancho Grande, 1,100
m, 1—5 October 1966, S. & W. Duck-
worth.

cupressi Heinrich, 1923 (Epinotia hopkin-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

sana); Bull. U.S. Natl. Mus. 123: 207.
Holotype 6, USA, California, Cypress
Point, rf. cones of Cupressus macro-
carpus, 4 November 1915, J. Miller.

cyanosticha Clarke, 1976 (Cymolomia); In-
sects of Micronesia 9: 55. Holotype 6,
Micronesia, Kusaie, Hill 1010, 300 m,
13 April 1953, J. Clarke.

cyclopiana Heinrich, 1926 (Polychrosis);
Bull. U.S. Natl. Mus. 132: 97. Holotype
2, USA, New Jersey, Burnt Hills, rf.
seed pods of swamp magnolia, July
1920, H. Weiss.

cydna Razowski, 1993 (Apotoforma); Acta
Zool. Cracov. 36: 185. Holotype 2,
Venezuela, Rancho Grande, 1,100 m,
16-23 October 1966, S. & W. Duck-
worth.

cylichna Razowski, 1994 (Cylichneulia);
SHILAP Revta. Lepid. 22: 68. Holotype
?, Venezuela, Aragua, Rancho Grande,
15-21 June 1967, 1,100 m, R. Poole.

daemonicana Heinrich, 1923 (Eucosma);
Bull. U.S. Natl. Mus. 123: 111. Holo-
type 6, USA, New Mexico, Manzano
National Forest, Hell Cafion, “‘flying,”’
14 September 1916, C. Heinrich.

dapsilis Heinrich, 1929 (Eucosma); Proc.
U.S. Natl. Mus. 75: 5. Holetypecé:
USA, Wyoming, Park Co., Yellowstone
National Park, North Park.

dativa Heinrich, 1928 (Rhyacionia); Proc.
Entomol. Soc. Wash. 30: 61. Holotype
36, Japan, Yokahama, r.f. Pinus thunber-
gii, em: 1 July 1925, A. Kariya.

davisi Kawabe, 1989 (Archips); Tinea 12:
191. Holotype 6, Taiwan, Hualien
Hsien, Tayulin, 2,500 m, 9-18 June
1980, D. Davis.

deceptana Busck, 1907 (Pharmacis); J. N.
Y. Entomol. Soc. 15: 29. Holotype 2,
USA, Texas, Kerr Co., Kerrville, W.
Barnes.

deceptiva Clarke, 1949 (Eulia); Acta Zool.
Lilloana (Tucuman) 7: 581. Holotype
?, Brazil, Santa Catarina, Nova Teuton-
ia, K Plaumann.

decor Kawabe, 1978 (Enarmonia); Tinea

VOLUME 102, NUMBER 4

10: 188. Holotype ¢6, Japan, Honshu,
Chiba Pref., Tokiwadaira, 5 June 1977,
A. Kawabe.

decora Obraztsov, 1966 (Pseudomeritastis);
Proc. U.S. Natl. Mus. 118: 230. Holo-
type 6, Bolivia, Cochabamba, Incacha-
ca, tropical cloud area, 2,100 m, 27 Au-
gust—5 September 1956, L. Pena.

decorosa Heinrich, 1929 (Hystricophora);
Proc. U.S. Natl. Mus. 75: 20. Holotype
3, USA, Florida, Enterprise, “*4.16.”’
Heinrich (1923) consistently misspelled
Hystrichophora in his revision.

definitivana Heinrich, 1923 (Anchylopera),;
Bull. U.S. Natl. Mus. 123: 270. Holo-
type ¢, USA, Nevada, “July 16—23.”’

deflexana Heinrich, 1923 (Epiblema), Bull.
U.S. Natl. Mus. 123: 144. Holotype ¢,
USA, Texas, Cameron Co., Browns-
ville, 27 May 1917, A. Busck.

delicata Yasuda & Kawabe, 1980 (Croe-
sia); Tinea 11: 13. Holotype ¢, Japan,
Nagano Pref., Okutatesina, 23 August
1963, A. Kawabe.

delicatana Heinrich, 1923 (Gretchena);
Bull. U.S. Natl. Mus. 123: 185. Holo-
type 6, USA, Pennsylvania, Allegheny
Co., Oak Station, ““May 1-10,” E Mar-
loff.

delphinoides Heinrich, 1923 (Thiodia),
Bull. U.S. Natl. Mus. 123:59. Holotype
3, USA, Utah, Juab Co., Eureka, 16
July 1911, T. Spalding.

delphinus Heinrich, 1923 (Thiodia); Bull.
U.S. Natl. Mus. 123: 45. Holotype ¢,
USA, California, Lake Tahoe, Deer
Park Springs, ‘“‘July 1-7.”

dendrophila Clarke, 1971 (Dichelopa),
Smithsonian Contrib. Zool. 56: 105.
Holotype 6, Rapa Island, Maugaoa,
950’ (292 m), r.f. Cyathea rapensis, em:
3 November 1963, J. & T. Clarke.

deprecatoria Heinrich, 1926 (Olethreutes),
Bull. U.S. Natl. Mus. 132: 177. Holo-
type 6, Canada, British Columbia, Wel-
lington, July, G. Taylor.

derelicta Heinrich, 1929 (Eucosma); Proc.
U.S. Natl. Mus. 75: 13. Holotype ¢d,

1031

USA, North Carolina, Polk Co., Tryon,
13 August 1904, Fiske.

diabolana Blanchard, 1979 (Eucosma),; J.
Lepid. Soc. 33: 214. Holotype 6, USA,
Texas, Culberson, Co., Sierra Diablo
Wildlife Management Area, 6,000’, 31
March 1970, A. & M. Blanchard.

diamphidia Clarke, 1968 (Lasiothyris),
Proc. U.S. Natl. Mus. 125: 49. Holotype
3, Peru, Cusco, Machu Picchu, 5 Feb-
ruary 1959, J. Clarke.

dicaeus Diakonoff, 1968 (Archips); Bull.
U.S. Natl. Mus. 257: 26. Holotype 6,
Philippine Islands, Luzon, Mt. Makil-
ing, Baker.

digitana Heinrich, 1923 (Epinotia); Bull.
U.S. Natl. Mus. 123: 215. Holotype 6,
Canada, British Columbia, Kaslo Creek,
H. Dyar.

dilutifuscana Walsingham, 1879 (Sericor-
is); Ill. Lepid. Heter. Brit. Mus. 4: 33.
Syntype 6, USA, southern Oregon, 2
June 1872. Walsingham described this
species from two males; the other syn-
type is presumed to be in the BMNH.

dimorpha Clarke, 1949 (Eulia); Acta Zool.
Lilloana (Tucuman) 7: 585. Holotype
?, Brazil, Santa Catarina, 28 March
1936, E Hoffman.

dimorphana Barnes & Busck, 1920 (Tor-
trix); Contrib. Nat. Hist. Lepid. North
Am. 4: 215. Holotype 6, Canada, Brit-
ish Columbia, Victoria.

discobola Diakonoff, 1968 (Strepsicrates),;
Bull. U.S. Natl. Mus. 257: 85. Holotype
36, Philippine Islands, Davao Province,
Mindanao, E slope Mt. McKinley,
7,200’, 10 September 1946, mossy
stunted forest, CNHM Philippine Ex-
ped. 1946-47, H. Hoogstraal.

discretivana Heinrich, 1921 (Eucosma); J.
Agric. Res. 20: 823. Holotype ¢, USA,
Texas, Harris Co., Sheldon, 10 April
1919, Johnson.

dispersa Brown, 1990 (Auratonota), Flori-
da Entomol. 73: 154. Holotype 6, Pan-
ama, Cocle Province, Valle, 22 April
1965, S. & W. Duckworth.

disputabilis Obraztsov 1963 (Acleris); Proc.

1032

U.S. Natl. Mus. 114: 262. Holotype <6,
Canada, British Columbia, Goldstream,
16 October 1902.

dissitana Grote, 1879 (Ptycholoma); North
Am. Entomol. 1: 29. “Type”? 2, USA,
4 July 1878. This specimen, labeled
‘“‘type’’ by someone other than Grote, is
the only Grote specimen of P. dissitana
we could locate. The original descrip-
tion does not indicate how many spec-
imens were examined.

distincta Obraztsov, 1966 (Pseudomeritas-
tis); Proc. U.S. Natl. Mus. 118: 228.
Holotype 2, Costa Rica, Cartago Prov-
ince, La Florida, 500’, W. Schaus.

dognini Obraztsov, 1966 (Pseudatteria);
Proc. U.S. Natl. Mus. 118: 588. Holo-
type 6d, Ecuador, Environs of Loja,
1887.

dominica Brown, 1993 (Auratonota); Pan-
Pac. Entomol. 69: 314. Holotype 6,
West Indies, Dominica, 1.7 mi E Point
Casse, light trap, 24 March 1965, W.
Wirth.

domna Clarke, 1968 (Amallectis); Proc.
U.S. Natl. Mus. 125: 34. Holotype 6,
Colombia, Narifo, Volcan Galeras,
3,000 m, 14 January 1959, J. Clarke.

donaldana Kawabe, 1993 (Notocelia); Tin-
ea 13: 238. Holotype 6, Taiwan, Nan-
tou Hsien, Mei-feng, 30 km S Tayuling,
2,200 m, 1—8 July 1980, D. Davis.

doria Clarke, 1976 (Laspeyresia); Insects
of Micronesia 9: 123. Holotype 6, Mi-
cronesia, Palau, Babelthuap, Ngarhe-
long, 1 May 1957, C. Sabrosky.

dorsalana Dyar, 1903 (Tortrix); Proc. En-
tomol. Soc. Wash. 5: 231. Syntypes (4
3636, 2 22), USA, Arizona, Coconino
Co., Williams, rf. oak, em: 14 June
1901, A. Schwartz. According to the
original description, Dyar described this
species from four males and four fe-
males; all of the syntypes cited above
are labeled “Type No. 6736 USNM,”’
consistent with the original description.

dorsata Clarke, 1988 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 133. Holo-
type 2, Marquesas Islands, Hiva Oa,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Mt. Feani, 3,400’, 1 March 1968, J. &
T. Clarke.

druana Walsingham, 1914 (Tortrix); Biol.
Centr.-Am., Lepid., Heter., 4: 288. Ho-
lotype ¢, Mexico, Durango, r.f. cynipid
gall on Quercus, em: 11 February 1897.
This species was described from the sin-
gle male.

dryocremna Meyrick, 1932 (Polyortha);
Exotic Microlepidoptera 4: 343. Lecto-
type ¢ (designated here), Guatemala,
Palin, Schaus & Barnes. Selected by J.
Clarke. Meyrick’s description indicates
*“6 ex. [examples] (type U.S. Nat.
Muss):

duckworthorum Brown, 1991 (Cuproxena);
in Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 67. Holotype 3, Costa
Rica, Cartago Province, Turrialba, 17—
21 February 1965, S. & W. Duckworth.

dulciana Heinrich, 1923 (Gretchena); Bull.
U.S. Natl. Mus 123: 182. Holotype 6,
USA, New Jersey, Greenwood Lake,
“*V.30,”” W. Kearfott.

dysmorphia Clarke, 1968 (Lasiothyris);
Proc. U.S. Natl. Mus. 125: 47. Holotype
3, Bolivia, Cochabamba, Incachaca,
tropical cloud area, 2,100 m, 27 Au-
gust—5 September 1956, L. Pefia.

eburata Heinrich, 1929 (Eucosma); Proc.
U.S. Natl. Mus. 75: 6. Holotype ¢,
USA, Arizona, Mohave Co., “Aug. 24—
Sey

edemoidana Dyar, 1903 (Eucosma); Proc.
Entomol. Soc. Wash. 5: 229. Two syn-
types (2 22), USA, Arizona, Coconino
Co., Williams, ‘*19-7,’’ W. Barnes;
USA, New Mexico, Las Vegas HS,
18.8." According to the original de-
scription, this species was described
from the two females cited above.

egens Razowski, 1999 (Netechma); Polskie
Pismo Entomol. 68: 97. Holotype 6,
Colombia, Narifo, Volcan Galeras,
2,900 m, 13 January 1959, J. Clarke.
Razowski (1999) incorrect identifies the
disposition of the type as NHMW.

elaborata Kawabe, 1976 (Pseudohedya);

VOLUME 102, NUMBER 4

Tinea 10: 45. Holotype 6, Japan, Ao-
mori Pref., Tuta Spa, 20 July 1957, A.
Kawabe.

eleonora Obraztsov, 1962 (Anopina); Am.
Mus. Novit. 2082: 12. Holotype ¢,
USA, Arizona, Apache Co., Alpine, 6
June 1937, G. & J. Sperry.

elitha Clarke, 1976 (Heleanna physalodes);
Insects of Micronesia 9: 19. Holotype
36, Micronesia, Ponape Island, Colonia,
13 January 1953, J. Clarke.

elongana Brown, 1990 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 54. Holotype ¢, Vene-
zuela, Amazonas (territory), Cerro de
Neblina Camp VII, 1,850 m, 0°5S1’N,
65°58'W, 2—4 December 1984, Brown.

embaphion Razowski, 1984 (Saphenista),;
Ann. Zool. Warsaw 38: 276. Holotype
6, Venezuela, Mérida, 8 km SE Apar-
taderos, 22 February 1976, C. & O.
Flint.

embolina Razowski, 1984 (Saphenista);
Ann. Zool. Warsaw 38: 277. Holotype
2, Venezuela, Mérida, 4 km S Santo
Domingo, 19-23 February 1976, C. &
O. Flint.

emera Razowski, 1993 (Acleris); Acta
Zool. Cracov. 36: 191. Holotype 6, Bo-
livia, Cochabamba, Incachaca, 27 Au-
gust—5 September 1956, L. Pena.

emigratella Busck, 1909 (Amorbia), Proc.
Entomol. Soc. Wash. 11: 201. Syntype
6, Hawaiian Islands, Oahu, Tantulus,
O. Swezey. Busck did not indicate how
many specimens he examined.

enucleata Razowski, 1999 (Netechma);
Polskie Pismo Entomol. 68: 95. Holo-
type 6, Colombia, Narifio, Volcan Ga-
leras, 300 m, 14 January 1959, J.
Clarke. Razowski (1999) incorrect iden-
tifies the disposition of the type as
NHMW.

episticta Clarke, 1949 (Eulia); Acta Zool.
Lilloana (Tucuman) 7: 584. Holotype
6, Brazil, Santa Catarina, Nova Teuton-
ia, September 1948, E Plaumann.

equadora Brown, 1990 (Paraptila); J. Lep-

1033

id. Soc. 44: 271. Holotype 6, Ecuador,
Shell-Mera, 18 April 1958, R. Hodges.

erigeronana Riley, 1881 (Conchylis),;
Trans. St. Louis Acad. Sci. 4: 316. Ho-
lotype 6, USA, Texas, Columbia, rf.
‘‘cecidomyidous gall on Erigeron,”’ em:
24 February 1879, E. Schwarz. Riley
described this species from a single
specimen.

erotella Heinrich, 1923 (Carpocapsa),
Proc. Entomol. Soc. Wash. 25: 121. Ho-
lotype 2, USA, Maryland, Montgom-
ery Co., Hyattsville, rf. Pinus taeda,
em: 22 May 1915, A. Busck.

erubesca Kawabe, 1978 (Gypsonoma); Tin-
ea 10: 187. Holotype 6, Japan, Hok-
kaido, Sapporo, Matuyama Park, 30
July 1950, A. Mutsuura.

escharia Clarke, 1976 (Trymalitis); Insects
of Micronesia 9: 142. Holotype 3d, Mi-
cronesia, Guam, Ritidian, 2 August
1945, J. Gressitt.

essigana Busck, 1929 (Amorbia); Calif.
Dept. Agric. Mon. Bull. 18: 276. Ho-
lotype 6, USA, California, San Diego
Co., Chula Vista, rf. avocado leaves, 8
December 1928, R. McLean.

ethnica Heinrich, 1923 (Epinotia); Bull.
U.S. Natl. Mus. 123: 201. Holotype ¢,
USA, California, San Diego Co., San
Diego, “‘June 16—23.”

euchaldera Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 146. Holotype ¢, Colombia, June
1909, Fassel.

eupista Diakonoff, 1968 (Stenarchella);
Bull. U.S. Natl. Mus. 257: 99. Holotype
3, Philippine Islands, Davao Province,
Mindanao, La Lum Mt., Calian, 5,000’,
29-21 December 1920, Clagg, CM
Acc. 9163.

evestigana Razowski, 1984 (Polyortha),
Acta Zool. Cracov. 27: 221. Holotype
6, El Salvador, Cerro Miramundo,
2,300 m, 11 January 1969, S. Steinhau-
ser.

exacerbatricana Heinrich, 1923 (Epible-
ma); Bull. U.S. Natl. Mus 123: 146. Ho-

1034

lotype 6, USA, North Carolina, South-
ern Pines, ““Aug. 16—23.”

exaeresimum Heinrich, 1926 (Exartema);
Bull. U.S. Natl. Mus. 132: 160. Holo-
type 6, USA, Texas, Dallas Co., Dallas.

excerptionana Heinrich, 1923 (Eucosma);
Bull. U.S. Natl. Mus. 123: 117. Holo-
type 6, USA, Nevada, Washoe Co.,
Verdi, ‘“‘June 1 to 10,’’ A. Vachell.

exclusoriana Heinrich, 1923 (Eucosma);
Bull. U.S. Natl. Mus. 123: 110. Holo-
type d, USA, Texas, La Salle Co., Co-
tulla, 12 May 1906, Crawford & Pratt.

excusabilis Heinrich, 1923 (Eucosma);
Bull. U.S. Natl. Mus. 123: 123. Holo-
type 6, USA, California, Lake Tahoe,
Deer Park Springs.

exolivata Clarke, 1955 (Orthocomotis);
Trans. Entomol. Soc. London 107: 148.
Holotype @, Brazil, Santa Catarina,
Nova Teutonia, August 1936, H. Parish.

exomilana Franclemont, 1986 (Lozotaenia);
Proc. Entomol. Soc. Wash. 88: 58. Ho-
lotype ¢, USA, North Carolina, Macon
Co., Highlands, 3,865’, 11 July 1958, J.
Franclemont.

exoristus Razowski, 1988 (Exoletuncus);
Acta Zool. Cracov. 31: 390. Holotype
6, Colombia, Cauca, Paramo de Parce,
Lake San Rafael, 3,570 m, 29 January
1959, J. Clarke.

expolitana Heinrich, 1923 (Eucosma); Bull.
U.S. Natl. Mus. 123: 132. Holotype 6,
USA, Utah, Utah Co., Provo, 11 August
1908, T. Spalding.

exulis Issiki & Stringer, 1932 (Gnorismo-
neura),; Stylops 1: 125. Holotype 6,
Taiwan, Taihoku, 18 October 1923, S.
Issiki.

famula Zeller, 1875 (Teras tristana); Ver-
handel. Zool.-Bot. Ges. Wien 25: 214.
Lectotype 6, North America [no col-
lecting data]. Designated by Obraztsov
(1963). Zeller’s original description re-
fers to a male, consistent with the spec-
imen cited above. However, the associ-
ated Busck genitalia slide (Dec 6 1924;
USNM 25944) is of a female, and Ob-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

raztsov’s (1963: 236) lectotype desig-
nation also refers to the specimen as fe-
male.

fernaldana Grote, 1880 (Paedisca); N. Am.
Entomol.: 98. “‘Type’” 6, USA, Colo-
rado. This is the only Grote specimen
of fernaldana we could locate; it bears
a labeling identifying it as the type, but
not in Grote’s hand. The original de-
scription does not indicate how many
specimens were examined.

fernaldana Walsingham 1879 (Cochylis);
Ill. Lepid. Heter. Brit. Mus. 4: 27. Syn-
type 6, USA, northern California, Sep-
tember 1871. This species was de-
scribed from four males and one female;
all of the other syntypes are presumed
to be in the BMNH.

ferreana Busck, 1915 (Sparganothis); Proc.
Entomol. Soc. Wash. 17: 86. Holotype
2, USA, New York, Herkimer Co., II-
ion, 10 July 1912, H. McElhose.

ferruginana Fernald, 1882 (Semasia);
Trans. Am. Entomol. Soc. 10: 72. Lec-
totype 2, USA, Massachusetts, Good-
ell. Designated by Miller (1970).

ferrugineana Riley, 1881 (Exartema);
Trans. St. Louis Acad. Sci.. 4: 317.
“Type’’ 3, USA, Missouri, St. Louis
Co., St. Louis, rf. leaves of plum. Riley
described this species from two males,
only one of which could be located by
us.

ferruginiguttana Fernald, 1882 (Teras);
Trans. Am. Entomol. Soc. 10: 65.
“Type”? 2, USA, Colorado. Of two fe-
males that comprised the type series, we
could find only one. It is represented by
a portion of the thorax bearing a single
leg and a small piece of the left fore-
wing.

ferruginus Pogue, 1986 (Apolychrosis); in
Cibridn-Tovar et al., Cone and Seed In-
sects of the Mexican Conifers: 21. Ho-
lotype ¢, Mexico, Tlaxcala, Municipio
de Terrenate, Villareal, rf. Pseudotsuga
macrolepis, em: 18 August—6 Septem-
ber 1981, J. Mendez & R. Campos.

fertoriana Heinrich, 1923 (Thiodia); Bull.

VOLUME 102, NUMBER 4

U.S. Natl. Mus. 123: 264. Holotype 6,
Canada, British Colombia, Goldstream,
**10-5-03.”

filiana Busck, 1907 (Hendecaneura); J. N.
Y. Entomol. Soc. 15: 135. Holotype 6,
USA, California, Riverside Co., West
Riverside, 26 October 1905.

finitimana Heinrich, 1923 (Kundrya); Bull.
U.S. Natl. Mus. 123: 192. Holotype 6,
USA, New Hampshire, Rockingham
Co., Hampton, 11 June 1909, S. Shaw.

fishiana Fernald, 1882 (Peronea); Trans.
Am. Entomol. Soc. 10: 66. Lectotype
6, USA, Maine, Penobscot Co., Orono,
17 September 1879. Designated by Ob-
raztsov (1963).

flavana Fernald, 1905 (Eucosma pergan-
deana); Canad. Entomol. 37: 399. Lec-
totype 6, USA, Texas. Designated by
Miller (1970).

flavibasana Fernald, 1882 (Cenopis); Trans.
Am. Entomol. Soc. 10: 69. Lectotype 3
(designated here), USA, Illinois. Select-
ed by Powell in 1965. Although there is
little doubt that the lectotype is one of
2 specimens referred to in the original
description, Fernald misidentified the
sex, indicating two females, one from
Texas and one from Illinois.

flavifasciana Kawabe, 1976 (Olethreutes),;
Tinea 10: 47. Holotype 2, Japan, Gun-
ma Pref., Kuridaira, 20 July 1951, M.
Hoshino.

flavillana Dyar, 1903 (Epinotia); Proc. En-
tomol. Soc. Wash. 5: 230. Three syn-
types (1 6, 2 22), USA, Arizona, Co-
conino Co., Williams, bred from flower
stalk, June. According to the original
description, this species was described
from three specimens.

flexicostalis Dognin, 1908 (Tortrix); Ann.
Soc. Entomol. Belgium 52: 32. Holo-
type ¢, Peru (SE), Oconeque, Cara-
baya, ‘10.07,’ Warren.

flexura Clarke, 1986 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 124. Holo-
type 6, Marquesas Islands, Hiva Oa,
Feani, 3,800’, 20 February 1968, J. &
T. Clarke.

1035

flintana Brown, 1991 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 69. Holotype 2, Mexico,
Jalisco, Rt. 94, km 42, N Chapala [4 mi
toward Guadalajara from Chapala; top
of hill], 16-18 July 1966, O. Flint & A.
Ortiz.

footiana Fernald, 1882 (Eccopsis); Bull.
Buffalo Soc. Nat. Sci. 4: 53. Lectotype
2, USA, New York. Designated by
Miller (1970).

formosanus Kawabe, 1987 (Archippus);
Tinea 7: 122. Holotype ¢, Taiwan, Chi-
ayi, Alishan, 2,200 m, 9-11 July 1964,
H. Inoue.

foxcana Kearfott, 1907 (Phalonia); Trans.
Am. Entomol. Soc. 33: 84. Lectotype
36, USA, Ohio, Hamilton Co., Cincin-
nati, 18 August 1903, A. Braun. Kear-
fott described this species from three
specimens from Cincinnati, Ohio, and
Plummers Island, Maryland, with no in-
dication of which locality was the
source of two of the three. Klots (1942)
designated the above-mentioned lecto-
type, indicating that there were two lec-
toparatypes in the AMNH: a female
from Plummers Island and another
specimen without an abdomen (he
didn’t provide its collection data). Al-
though he correctly cited the date of
collection of the lectotype, he incorrect-
ly cited its collection locality as Plum-
mers Island.

fragariae Walsh & Riley, 1869 (Anchylo-
pera); Am. Entomol. 1: 89. Lectotype
2, USA, Illinois, Whiteside or Bureau
Co. According to the original descrip-
tion, this species was described from
nine specimens, only two of which
could be located by us. Designated by
Miller (1973).

fragariana Busck, 1919 (Tortricodes);
Proc. Entomol. Soc. Wash. 21: 52. Ho-
lotype 6, Canada, British Columbia,
Victoria, 17 August 1918.

frangula Clarke, 1968 (Amallectis); Proc.
U.S. Natl. Mus. 125: 30. Holotype °,
Venezuela, Aragua, Rancho Grande,

1036

1,100 m, 16—23 October 1966, S. & W.
Duckworth.

fraternana Busck 1907 (Hendecaneura); J.
N. Y. Entomol. Soc. 15: 134. Holotype
6, USA, California, Riverside Co.,
West Riverside, October.

fratruelis Heinrich, 1923 (Eucosma); Bull.
U.S. Natl. Mus. 123: 98. Holotype <4,
USA, North Carolina, Southern Pines,
8-15 August.

fraudabilis Heinrich, 1923 (Eucosma);
Bull. U.S. Natl. Mus. 123: 98. Holotype
3, USA, North Carolina, Southern
Pines. According to the original descrip-
tion, the type series was collected be-
tween 1 June and 23 July.

fritillana Blanchard & Knudson, 1981 (Eu-
cosma); J. Lepid. Soc. 35: 170. Holo-
type 6, USA, Texas, Anderson Co., En-
geling Wildlife Management Area, near
Tennessee Colony, 28 June 1978, A.
Blanchard.

frustrana W. Comstock, 1880 (Retinia);
Rept. U.S. Dept. Agric. 1879: 236. Lec-
totype 2, USA, Virginia, near Washing-
ton (D.C.), “Tortrix on P. inops,”’ em:
18 June 1879. Designated by Miller
(1967).

fucana Walsingham, 1879 (Lozotaenia); Il.
Lepid. Heter. Brit. Mus. 4: 12. Syntype
3, USA, Oregon (southern), May 1872.
This species was described from four
males; the other syntypes are presum-
ably in the BMNH.

fullerea Riley, 1869 (Penthina); J. Horti-
cult., Boston, Oct. 1868: 12. ““Type”’ 2,
USA, “seed pods of Antherrinum.”” An
associated label indicates that the spec-
imen is an unpublished lectotype se-
lected by Miller in 1982. Riley did not
indicate how many specimens he ex-
amined.

fulvomixtana Kawabe, 1974 (Zeiraphera);
Tyo to Ga 25: 98. Holotype d, Japan,
Gunma Pref., Doaiguchi, 12 July 1972,
S. Shimeki.

fumipennis Dognin, 1904 (Atteria); Ann.
Soc. Entomol. Belgium 48: 133. Holo-
type 6, Colombia, Micay, Aoiat, 1869.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

fumosana Powell, 1978 (Rhyacionia); in
Powell and Miller, U.S. Dept. Agric.,
Agric. Handb. 514: 21. Holotype 4,
USA, Colorado, El Paso Co., Colorado
Springs, Rock Creek Canyon, 2 May
1959, M. May.

fumoviridana Heinrich, 1923 (Epinotia);
Bull. U.S. Natl. Mus. 123: 208. Holo-
type 3d, USA, California, Siskiyou Co.,
Shasta Retreat, “Aug. 16-23.”

fuscalbana Zeller, 1876 (Sericoris); Verhan-
del. Zool.-Bot. Ges. Wein 25: 284.
“Type”? 6, USA, [Maine or Massachu-
setts according to the original descrip-
tion], Packard. This is the only Zeller
specimen of fuscalbana that we were
able to locate. The original description
does not indicate how many specimens
were examined, but it does indicate that
the series included only males.

fuscana Barnes & Busck, 1920 (Peronea);
Contrib. Nat. Hist. Lepid. North Am. 4:
216. Holotype ¢6, Canada, Manitoba,
Aweme, 18 April 1905, Criddle.

fuscodorsana Kearfott, 1904 (Commophi-
la); Canad. Entomol. 36: 141. Holotype
36, Canada, British Colombia, Kaslo, 7
June 1903, J. Cockle. Klots (1942) cor-
rectly stated that the holotype is in the
USNM, consistent with the original de-
scription.

fuscomaculatus Brown, 2000 (Odonthali-
tus); Proc. Entomol. Soc. Wash. 102:
42. Holotype 2, Mexico, Michoacan,
San Lorenzo, Rt. 15, km 206, 19 July
1966, O. Flint & A. Ortiz.

galena Clarke, 1971 (Tritopterna); Smith-
sonian Contrib. Zool. 56: 124. Holotype
3, Rapa Island, Teumukopuke, 500’, 7
October 1963, J. & T. Clarke.

galerasiana Razowski, 1988 (Uncicida);
Acta Zool. Cracov. 31: 396. Holotype
3, Colombia, Narifio, Volcan Galeras,
2,900 m, 13 January 1959, J. Clarke.

gallaesaliciana Riley, 1881 (Grapholitha);
Trans. St. Louis Acad. Sci. 4: 320.
“Type” 2, USA, Missouri, St. Louis
Co., St. Louis, ‘“‘Lepidopterous gall on

VOLUME 102, NUMBER 4

willow,” 17 June 1873. This is one of
two specimens cited by Riley in the
original description; the location of the
other is unknown.

gambra Razowski, 1988 (Ernocornutina);
Acta Zool. Cracov. 31: 399. Holotype
36, Argentina, Tucuman, Ciudad Univ-
ersitaria, 800 m, 17 February 1959, J.
Clarke.

gampsognathos Razowski, 1988 (Gaurun-
cus); Acta Zool. Cracov. 31: 405. Ho-
lotype 6, Bolivia, Cochabamba, Inca-
chaca, tropical cloud area, 2,100 m, 27
August—5 September 1956, L. Pefia.

garai Miller, 1987 (Gretchena); J. Lepid.
Soc. 41: 151. Holotype 2, Ecuador, 4
km S Loja, rf. Juglans neotropica, 28
June 1986, A. Samaniego.

gatesclarkei Kawabe, 1992 (Acleris); Tinea
13: 171. Holotype 6, Taiwan, Nantou
Hsien, Tsuifeng, 2,400 m, 29 December
1989, A. Kawabe.

gelastes Razowski, 1988 (Gauruncus); Acta
Zool. Cracov. 31: 404. Holotype d, Ar-
gentina, Tucuman, Ciudad Universitar-
ia, 800 m, 17 February 1959, J. Clarke.

gemellana Heinrich, 1923 (Eucosma cir-
culana); Bull. U.S. Natl. Mus. 123: 96.
Holotype 6, USA, Florida, Hillsbor-
ough Co., St. Petersburg, May.

gentilii Brown, 1998 (Argentulia); J. Lepid.
Soc. 52: 180. Holotype ¢, Argentina,
Neuquén, Paso Cordoba, 1,300 m, 20
February 1980, M. Gentili.

georgiella Hulst, 1887 (Myelois); Entomol.
Am. 3: 136. “Type” 6, USA, Colora-
do. This is the only Hulst specimen we
could find of M. georgiella; it is labeled
““type.”’ The original description does
not indicate how many specimens were
examined.

gerda Busck, 1911 (Olethreutes); Proc.
U.S. Natl. Mus. 40: 227. Holotype 2,
French Guiana, St. Jean, Maroni, W.
Schaus.

giganteana Riley, 1881 (Paedisca); Trans.
St. Louis Acad. Sci. 4: 318. “‘Pseudo-
type” ¢, USA, Missouri, “‘Barlon.”
According to Riley’s original descrip-

1037

tion, this species was described from 2
females, one from ‘‘Kansas (G. E Gau-
mer)’ and one from “Iowa, March,
1874 (Hoffmeister).’’ Although not in-
cluded in the original series, the speci-
men from Missouri bears a type label in
Riley’s hand and represents a potential
neotype if one is deemed necessary.

gigantica Busck, 1920 (Hysterosia); Insect.
Inscit. Menst. 8: 87. Holotype 2, Mex-
ico, Distrito Federal, Mexico City, R.
Muller.

gilletteana Dyar, 1903 (Eucosma); Proc.
Entomol. Soc. Wash. 5: 229. Syntype 6,
USA, Arizona, Coconino Co., Williams,
16.6; foun syntypes. (Gi Gc. 2),
USA, Colorado. According to Dyar, this
species was described from the series (n
= 5) cited above; all of the specimens
bear red labels “Type No: 6737
U.S.N.M.”

gloriola Heinrich, 1931 (Eucosma); Proc.
Entomol. Soc. Wash. 33: 196. Holotype
3, USA, Connecticut, Stamford, B.T.R.
Lab colony, rf. white pine tip, 6 May
193i"

gnoma Clarke, 1986 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 138. Holo-
type 6, Marquesas Islands, Fatu Hiva,
Hanavave, 10’, 12 April 1968, J. & T.
Clarke.

goodelliana Fernald, 1882 (Phoxopteris);
Trans. Am. Entomol. Soc. 10: 69. Lec-
totype 6, USA Maine, New Hampshire,
or Massachusetts [no locality data on
specimen labels]. Designated by Miller
(1970).

graceana Powell, 1960 (Argyrotaenia);
Pan-Pac. Entomol. 36: 93. Holotype <6,
USA, California, San Bernardino Co.,
San Bernardino Mts., Hathaway Creek,
2 August 1940, C. Henne.

grandicula Heinrich, 1926 (Laspeyresia);
Bull. U.S. Natl. Mus. 132: 54. Holotype
3, USA, Virginia, Giles Co., Mountain
Lake, 14—21 June 1907. A. Braun.

grandiflavana Walsingham, 1879 (Paedis-
ca); Ill. Lepid. Heter. Brit. Mus. 4: 50.
Syntype ¢, USA, California, Lake Co.,

1038

17-19 June 1871. This species was de-
scribed from one male and two females;
the other syntypes presumably are in the
BMNH.

grandis Busck, 1907 (Phalonia); J. N. Y.
Entomol. Soc. 15: 23. Holotype 2°,
USA, Colorado, Jefferson Co., Golden,
Chimney Gulch, Oslar, 1 July 1904.

gratiana Kawabe, 1974 (Hedya); Tyo to Ga
25: 101. Holotype ¢, Japan, Honmura,
Kuchinoerabujima Island, 14 August
1973, r.f. Glochidon obovatum, A. Ka-
wabe.

gratuitana Heinrich, 1923 (Epiblema);
Bull. U.S. Natl. Mus. 123: 268. Holo-
type 6, USA, Washington, East Sound,
11 July 1901.

graziella Blanchard, 1968 (Eucosma); J.
Lepid. Soc. 22: 143. Holotype ¢, USA,
Texas, Brewster Co., Big Bend National
Park, Green Gulch, 11 October 1966, A.
& M. Blanchard.

grindeliana Busck, 1906 (Cydia); Canad.
Entomol. 38: 211. Two syntypes (1 6,
1 2), USA, Texas, Clarendon, r.f. Grin-
delia squarrosa, 28 September 1905,
em: 4 October 1905, Hunter. In the orig-
inal description, Busck identified the
type as labeled “‘Type #9804”’; howev-
er, both of the specimens listed above
have identical red type labels with this
number.

griselda Blanchard & Knudson, 1981 (Eu-
cosma); J. Lepid. Soc. 35: 173. Holo-
type 6, USA, Texas, Brewster Co., Big
Bend National Park, Chisos Basin, 7
April 1967, A. & M. Blanchard.

groteana Fernald, 1882 (Cenopis); Trans.
Am. Entomol. Soc. 10: 69. Holotype 2,
USA, Ohio. This species was described
from a single specimen.

guerrerana Obraztsov, 1964 (Anopina);
Am. Mus. Novit. 2082: 15. Holotype ¢,
Mexico, Guerrero, Sierra de las Aguas
Escondidas, 9,500’, July, H. Smith.

guiana Busck, 1913 (Olethreutes); Insect.
Inscit. Menst. 1: 92. Holotype 6, British
Guiana [Guyana], Georgetown, “larvae

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

feed on a common weed in the cane
field.”

gunniana Busck, 1907 (Phalonia); J. N. Y.
Entomol. Soc. 15: 26. Holotype 2°,
USA, Maryland, Montgomery Co.,
Plummers Island, September 1903, A.
Busck.

guttulana Blanchard, 1979 (Eucosma); J.
Lepid. Soc. 33: 214. Holotype ¢, USA,
Texas, Kennedy Co., Padre Island Na-
tional Seashore, 19 July 1976, A. & M.
Blanchard.

gyraleus Diakonoff, 1982 (Archips); Zool.
Verhandel. (Leiden) 193: 87. Holotype
6, Sri Lanka, Ratnapura District, Ug-
galkaltota, 350’, Irrigation Bungalow,
31 January—8 February 1970, Davis &
Rowe.

habrosana Heinrich, 1923 (Exentera); Bull.
U.S. Natl. Mus. 123: 178. Holotype 6,
USA, California, San Diego Co., San
Diego, 17 March 1912, W. Wright.

hadrotes Clarke, 1986 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 126. Holo-
type 6, Marquesas Islands, Nuku Hiva,
Tunoa Ridge, 2,900’, 23 January 1968,
J. & T. Clarke.

hasseanthi Clarke, 1952 (Eucosma); Bull.
South. Calif. Acad. Sci. 52: 60. Holo-
type 6, USA, California, Orange Co,
‘“‘bred from Hasseanthus variegatus,”
23 August 1936, T. Hower.

heindelana Fernald, 1905 (Acleris); Am.
Nat. 39: 870. Lectotype ¢, Canada,
Manitoba, Winnipeg, A. W. Hanham.
Designated by Obraztsov (1963).

heliocausta Dognin, 1912 (Atteria); Heter-
oceres Nouveaux de L Amerique du Sud
6: 51. Holotype 6, Colombia, Cali, San
Antonio, 2,000 m, 22 September 1908,
Fassel.

helianthana Riley, 1881 (Grapholitha);
Trans. St.Louis’ Acad’ Scite=t-ws13:
“Type” 6, USA, Texas, “gall on sun-
flower,” 12 August 1873. Riley de-
scribed this species from two speci-
mens, only one of which has been lo-
cated by us.

VOLUME 102, NUMBER 4

hemeropis Dognin, 1912 (Olethreutes);
Heteroceres Nov. L Amerique du Sud 6:
49. Holotype 6, Colombia, Cali, San
Antonio, Fassel.

hemitephras Clarke, 1976 (Lasiothyris);
Proc. U.S. Natl. Mus. 125: 51. Holotype
2, Mexico, Puebla, Tehuacan, 10 Oc-
tober.

henicodes Razowski, 1988 (Bicavernaria);
Acta Zool. Cracov. 31: 400. Holotype
36, Peru, Cusco, Machu Picchu, 2,385
m, 6 February 1959, J. Clarke.

hennei Clarke, 1947 (Eucosma); Bull.
South. Calif. Acad. Sci. 46: 51. Holo-
type 6, USA, California, Los Angeles
Co., El Segundo sand dunes, em: 3 Oc-
tober 1940, C. Henne.

heos Razowski, 1988 (Helicteulia); Acta
Zool. Cracov. 31: 388. Holotype 6, Bo-
livia, Cochabamba, Incachaca, tropical
cloud area, 2,100 m, 27 August—5 Sep-
tember 1956; 1: "Pena:

herbacea Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 151. Holotype 6, Costa Rica, San
Pedro de Montes de Oca, r.f. avocado,
em: 15 January 1933, C. Ballou.

herbaria Busck, 1920 (Sociphora); Insect.
Inscit. Menst. 8: 85. Holotype 6, Gua-
temala, Cuyuga, W. Schaus.

heterophaea Clarke, 1968 (Phalonidia);
Proc. U.S. Natl. Mus. 125: 36. Holotype
6, Colombia, Antioquia, La Estrella,
1,730 m, 13 December 1959, FE Luis
Gallego M.

heucherana Heinrich, 1923 (Epinotia);
Bull. U.S. Natl. Mus. 123: 217. Holo-
type 6, USA, Virginia, Arlington Co.,
Rosslyn, larvae mining leaves of ‘“‘alum
root’? [Heuchera americana], C. Hein-

rich.
hieroglyphana Blanchard & Knudson, 1984
(Grapholita); Proc. Entomol. Soc.

Wash. 86: 448. Holotype ¢, USA, Tex-
as, Culberson Co., Guadalupe Moun-
tains, Nickel Creek, 10 July 1968, A. &
M. Blanchard.

hieroglypta Walsingham, 1914 (Olethreu-
tes); Biol. Centr.-Am. Lepid., Heter. 4:

1039

250. Holotype 6, Mexico, Veracruz,
Orizaba, W. Schaus. Walsingham clear-
ly designated a “‘Type”’ male deposited
in the USNM.

hipeana Grote, 1876 (Conchylis); Canad.
Entomol. 8: 207. Holotype ¢, Canada,
Ontario, Port Stanley, W. Saunders. Al-
though the specimen bears a label
‘“‘London, Ont.,”” according to the orig-
inal description it was taken at Port
Stanley, by ““W. Saunders from London,
Ontario.”” This species was described
from a single male.

hiranoi Kawabe, 1980 (Gypsonoma); Tinea
11: 27. Holotype 6, Japan, Nagano
Pref., Nakakaruizawa, 14 June 1964, A.
Kawabe.

hiroshii Kawabe, 1980 (Zeiraphera); Tinea
11: 26. Holotype 6, Japan, Gunma
Pref., Kumaneta’ra, 14 July 1952, M.
Hoshino.

hirsutana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 50. Syn-
type 6, USA, California, Sonoma Co.,
23 May 1871. This species was de-
scribed from four males; the other three
presumably are in the BMNH.

hodgesi Heppner, 1989 (Argyrotaenia);
Florida Entomol. 72: 102. Holotype ¢,
USA, Florida, Glade Co., Fisheating
Creek, 7-10 May 1964, R. Hodges.

hodgesi Razowski, 1993 (Apotoforma);
Acta Zool. Cracov. 36: 186. Holotype
?, Panama, Cerro Campana, 11—14 July
i967, O>S. Fling

hodsoni Miller, 1986 (Pseudexentera); J.
Lepid. Soc. 40: 223. Holotype 3, USA,
Pennsylvania, Allegheny Co., Oak Sta-
tion, 10 April 1910, E Marloff.

hoffmanana Brown, 1991 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 60. Holotype 2, Brazil,
Santa Catarina, Neu-Bremen, 12 June
1931, E Hoffman.

hohuanshana Kawabe, 1986 (Zeiraphera);
Entomol. Pap. Pres. Kurosawa, Tokyo:
79. Holotype ¢, Taiwan, Hualien Hsien,
Hohuanshan, 3,100 m, 30 July—1 Au-
gust 1983, A. Kwabe.

1040

hohuanshana Kawabe, 1989 (Acleris); Tin-
ea 12: 194. Holotype 3, Taiwan, Hu-
alien Hsien, Hohuanshan, 3,100 m, 30
July—1 August 1983, A. Kawabe.

hohuanshanensis Kawabe, 1985 (Clepsis);
Tinea 12: 5. Holotype ¢, Taiwan, Hu-
alien Hsien, Hohuanshan, 3,100 m, 30
July—1 August 1983, A. Kawabe.

holographa Clarke, 1965 (Nesochoris);
Proc. U.S. Natl. Mus. 117: 74. Holotype
6, Chile, Masatierra, Plazoleta del
Yunque; 200%m=s 9). February “1952; P.
Kuschel.

homolopa Diakonoff, 1968 (Eudemis);
Bull. U.S. Natl. Mus. 257: 54. Holotype
2, Philippine Islands, Davao Province,
Minandao, 5717.

horii Kawabe, 1987 (Cryptophlebia); Tinea
12: 141. Holotype 6, Japan, Okinawa
Island, Okubi, Kin, rf. Bruguiera gym-
norrhyza, 8 May 1983, S. Hori.

hoshinoi Kawabe, 1964 (Hastula); Tyo to
Ga 15: 2. Holotype 6, Japan, Honshu,
Tokyo, Setagaya, 16 June 1963, A. Ka-
wabe.

hostilis Diakonoff, 1956 (Bactra); Zool.
Verhandel. (Leiden) 29: 57. Holotype
3d, Japan, Honshiu, Funakoshi, Yoko-
suka, 11 September 1953.

houseri Miller, 1959 (Petrova); Ohio J. Sci.
59: 230. Holotype 6, USA, Ohio,
Washington Co., Veto, r.f., Pinus echin-
ata, 18 June 1954, W. Miller.

huachucensis Obraztsov, 1961 (Argyro-
taenia montezumae); Am. Mus. Novit.
2048: 7. Holotype ¢, USA, Arizona,
Cochise Co., Huachuca Mtns., August
1905, H. Skinner.

hubbardana Busck, 1907 (Phalonia); J. N.
Y. Entomol. Soc. 15: 27. Holotype 2,
USA, Arizona, Pima Co., Tucson, rf.
Koeberlinia spinosa, em: 12 May 1897,
H. Hubbard.

hypericana Ely, 1910 (Peronea); Proc. En-
tomol. Soc. Wash. 12: 68. Holotype 2,
USA, Virginia, Fairfax Co., Great Falls,
r.f., Hypericum prolificum, em: 17 June
1909, C. Ely.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

icelitodes Diakonoff, 1982 (Eucosma);
Zool. Verhandel. (Leiden) 193: 45. Ho-
lotype 6, Sri Lanka, Mannar District,
Olaithoduvai, 10 mi NW of Mannar, 0—
50’, 4—5 November 1976, G. Hevel, R.
Dietz, S. Karunaratne & D. Balasooriya.

icogramma Clarke, 1968 ([razona); Proc.
U.S. Natl. Mus. 125: 38. Holotype 2°,
Guatemala, Volcan Santa Maria, Octo-
ber, Schaus and Barnes.

illotana Walsingham, 1879 (Paedisca); Ill.
Lepid. Heter. Brit. Mus. 4: 48. Two syn-
types (1 6, 1 2), Oregon, Rogue River,
May 1872. Walsingham described this
species from two males and one female;
the other male syntype is presumed to
be in the BMNH.

imbrica Kawabe, 1978 (Hedya); Tinea 10:
181. Holotype ¢, Japan, Yakushima Is-
land, Shiratani, 14 September 1972, T.
Watanabe.

imbridana Fernald, 1905 (Cydia); Canad.
Entomol. 37: 400. Lectotype 6, USA,
Kansas, Pottawatomie Co., Onaga, F
Crevecoeur. Designated by Miller
(1970).

imitativa Heinrich, 1926 (Grapholitha);
Bull. U.S. Natl. Mus. 132: 34. Holotype
36, USA, California, San Francisco.

implicata Heinrich, 1924 (Proteoteras); J.
Wash. Acad. Sci. 14: 390. Holotype 6,
USA, Florida, Dade Co., Everglades,
ex-larva, “April 16—23.”

implicata Heinrich, 1931 (Thiodia); Proc.
U.S. Natl. Mus. 79: 7. Holotype ¢,
USA, Washington, Rochester, 13 June
1929, W. Baker.

improvisana Heinrich, 1923 (Epinotia);
Bull. U.S. Natl. Mus. 123: 269. Holo-
type 6, USA, California, Tulare Co.,
Mineral King, “‘June 16—23.”

incognita Obraztsov, 1963 (Acleris); Proc.
U.S. Natl. Mus. 114: 254. Holotype °,
USA, Idaho, Moscow Mountains, 7 Au-
gust 1933, J. Clarke.

inconspicua Obraztsov, 1964 (Proeulia);
Proc. U.S. Natl. Mus. 116: 190. Holo-
type 6, Chile, Santiago, La Obra, Oc-
tober 19525 L.jRena:

VOLUME 102, NUMBER 4

indagatricana Heinrich, 1923 (Thiodia);
Bull. U.S. Natl. Mus. 123: 56. Holotype
3, USA, Utah, Utah Co., Provo, 26 Au-
gust 1908, T. Spalding.

indentanus Dyar, 1903 (Phthinolophus);
Proc. Entomol. Soc. Wash. 5: 306.
Twenty-nine syntypes from various lo-
calities in the eastern United States. A
female from USA, Virginia, Fortress
Munroe, 24 June 1884, H. Dyar, was
placed in the type collection, presum-
ably by J. Clarke; the rest of the syn-
types are in the main collection. Ac-
cording to the original description, this
species was described from 17 males
and 21 females.

indigena Yasuda, 1978 (Eurydoxa); Trans.
Lepid. Soc. Jap. 29: 119. Holotype <6,
Taiwan, Rengwati, 29 March 1929, S.
Issiki.

infelix Heinrich, 1923 (Epiblema); Bull.
U.S. Natl. Mus. 123: 151. Holotype 6,
USA, North Carolina, Polk Co., Tryon,
25 May 1904, Fiske.

infernalis Heinrich, 1920 (Commophila);
Proc. U.S. Natl. Mus. 57: 61. Holotype
3, USA, New Mexico, Manzano Natl.
Forest, Hell Canyon, rf. berries of Jun-
iperus, 31 June 1917, C. Heinrich.

infida Heinrich, 1926 (Aphania); Bull. U.S.
Natl. Mus. 132: 121. Holotype 3, Can-
ada, Quebec, St. Johns Co., St. Therese
Island, 9 June 1915, W. Chagnon.

infimbriana Dyar, 1904 (Thiodia); Proc.
U.S. Natl. Mus. 27: 927. Holotype °,
Canada, British Columbia, Kaslo, H.
Dyar.

influana Heinrich, 1923 (Thiodia); Bull.
U.S. Natl. Mus. 123: 49. Holotype ¢,
USA, California, Siskiyou Co., Shasta
Retreat, “July 1-7.”

infuscata Heinrich, 1923 (Endothenia);
Proc. Entomol. Soc. Wash. 25: 109. Ho-
lotype ¢6, USA, Maryland, Forest Glen,
10 July 1914, O. Heidemann.

ingens Heinrich, 1926 (Laspeyresia); Bull.
WS: NatliMus: 132:63:) Holotype ¢;
USA, Florida, Hillsborough Co., St. Pe-
tersburg.

1041

ingrata Heinrich, 1926 (Laspeyresia); Bull.
U.S. Natl. Mus. 132: 50. Holotype ¢,
Canada, Manitoba, Aweme, 25 May
1905, Criddle.

injectiva Heinrich, 1926 (Hedulia); Bull.
U.S. Natl. Mus. 132: 65. Holotype ¢d,
USA, Nevada, Reno, rf. cones of Pinus,
em: 24 January 1911, J. Smith.

inopinata Heinrich, 1928 (Laspeyresia);
Proc. Entomol. Soc. Wash. 30: 91. Ho-
lotype ¢, China, South Manchuria, Kin-
shu, on apple, 4 August 1927, T. Kondo.

inopiosa Heinrich, 1926 (Laspeyresia);
Bull. U.S. Natl. Mus. 132: 46. Holotype
2, USA, Idaho, Kootenai Co., Coeur
d’Alene, rf. Pinus contorta, em: 11
May 1916, J. Evendon.

inoueit Kawabe, 1968 (Pandemis); Tinea 7:
121. Holotype 36, Taiwan, Chiayi Coun-
ty, Alishan, 2,200 m, 9-11 July 1964,
H. Inoue.

inouei Kawabe, 1972 (Eupoecilia); Tinea 9:
250. Holotype 6, Japan, Kagosaka-
toge, base of Mt. Fuji, 13 August 1969,
H. Inoue.

inouei Kawabe, 1987 (Metendothenia); Tin-
ea 12: 139. Holotype ¢, Japan, Mie
Pref., Hokusei-machi, Otsujishinden, 8
May 1986, T. Mano.

insidiosana Heinrich, 1923 (Epiblema);
Bull. U.S. Natl. Mus. 123: 145. Holo-
type 6, USA, North Carolina, Southern
Pines, “‘June 1-7.”

insignata Heinrich, 1924 (Thiodia); J.
Wash. Acad. Sci. 14: 386. Holotype ¢,
USA, Colorado, San Juan Co., Silver-
ton, “July 8-15” [genitalia slide only;
pinned specimen has not been found].

insignis Heinrich, 1928 (Petrova); Proc.
Entomol. Soc. Wash. 30: 63. Holotype
3, Japan, Yokohama, rf. Pinus thun-
bergii, em: 27 August 1924.

inspersa Heinrich, 1931 (Phaecasiophora);
Proc. U.S. Natl. Mus. 79: 13. Holotype
3, USA, Florida, Hillsborough Co., St.
Petersburg, 3 March 1915, R. Ludwig.

insulanus Kawabe, 1965 (Archippus); Tyo
to Ga 16: 23. Holotype 3, Japan, Oki-

1042
noerabu-jima Is., 8 April 1957, M.
Umebayashi.

interruptolineana Fernald, 1882 (Penthina);
Trans. Am. Entomol. Soc. 10: 70. Lec-
totype 6, USA, New Hampshire or
Massachusetts [no locality data on spec-
imen labels]. Designated by Miller
(1970).

invidana Barnes & Busck, 1920 (Tortrix);
Contrib. Nat. Hist. Lepid. North Am. 4:
215. Holotype 6, Canada, British Co-
lumbia, Vancouver Island, Duncans,
Hanham.

iodes Clarke, 1968 (Hysterosia); Proc. U.S.
Natl. Mus. 125: 4. Holotype 6, Guate-
mala, Volcan Santa Maria, Schaus &
Barnes.

iresinephora Razowski, 1988 (Eriotortrix);
Acta Zool. Cracov. 31: 403. Holotype
3d, Colombia, Bogota, Chico, 23 Janu-
ary 1959, J. Clarke.

iridana Barnes & Busck, 1920 (Platynota);
Contrib. Nat. Hist. Lepid. North Am. 4:
212. Holotype 2, USA, Florida, Hills-
borough Co., St. Petersburg, June.

irroratana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 48. Syn-
type 6, USA, California, Mendocino
Co., 7-10 June 1871. Walsingham de-
scribed this species from five males; all
but the USNM syntype are presumed to
be in the BMNH.

isipida Razowski, 1988 (Eriotortrix); Acta
Zool. Cracov. 31: 403. Holotype 3, Co-
lombia, Bogota, Chico, 23 January
1959, J. Clarke.

issikii Kawabe, 1980 (Aterpia); Tinea 11:
17. Holotype 3, Japan Honmura, Ku-
chinoarabu Island, 27—31 July 1973, A.
Kawabe.

issikii Razowski, 1977 (Hysterosia); Tyo to
Ga 28: 35. Holotype 2, Japan, Kuway-
ama, 23 June 1916.

ivana Fernald, 1901 (Tortrix); J. N. Y. En-
tomol. Soc. 9: 51. Holotype 6, USA,
Florida, rf. Iva imbricata, 10 March
1900. This species was described from
a single male.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

jJaponensis Kawabe, 1980 (Cydia); Tinea
11: 28. Holotype ¢, Japan, Akita Pref.,
Onuma Spa, Hachimantai, 9-10 August
1974, R. Sato.

Japonica Kawabe, 1978 (Eucoenogenes);
Tinea 10: 185. Holotype 3, Japan, Hon-
shu, Gunma Pref., Mt. Mikaboyama, 2
August 1968, S. Shimeki.

Jenningsi Powell, 1978 (Rhyacionia); in
Powell and Miller, U.S. Dept. Agric.,
Agric. Handb. 514: 17. Holotype 6,
USA, Arizona, Sitgreaves National For-
est, r.f. Pinus ponderosa, D. Jennings.

Jinboi Kawabe, 1965 (Clepsis); Kontyu 34:
459. Holotype 6, Japan, Honshu, South
Alps, Sanpuku-goya, | August 1964, K.
Jinbo [‘‘Jimbo”’ on label].

Jinboi Kawabe, 1976 (Cymolomia); Tinea
10: 44. Holotype ¢, Japan, Hokkaido,
Mt. Daisetsu (Kurodake), 16 August
1969, K. Jinbo.

jJordani Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 142. Holotype ¢, Brazil, New Bre-
men, 11 August 1936, EK Hoffman.

juglandana Fernald, 1879 (Tortrix); Canad.
Entomol. 11: 155. Two syntypes (1 6,
1 2), USA, Massachusetts/Ohio/Wis-
consin, and Canada (Ontario) [no local-
ity data on specimen labels]. Fernald
described this species from 11 males
and 15 females; he did not designate a
type specimen or a type locality, instead
listing the states from which he had
specimens.

juncticiliana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 75. Two
syntypes (2 6¢), USA, California,
Shasta Co., 16 July 1871. This species
was described from seven males; the
other five presumably are in the BMNH.

kana Busck, 1906 (Hemimene); Proc. Biol.
Soc. Wash. 19: 182. Lectotype ¢, Can-
ada, British Columbia, Kaslo, H. Dyar.
Designated by Miller (1983).

khasiensis Miller, 1977 (Petrova); J. Lepid.
Soc. 31: 135. Holotype 6, India, Assam

VOLUME 102, NUMBER 4

(Meghalaya), Upper Shillong, March
1963.

kincaidiana Fernald, 1900 (Phoxopteris); in
Dyar, Proc. Wash. Acad. Sci. 2: 500.
Holotype 6, USA, Alaska, Metlakahtla,
“6.4.99,” Harriman Expedition ‘99, T.
Kincaid.

kirishimensis Kawabe, 1974 (Eudemopsis);
Kontyu 42: 390. Holotype ¢, Japan,
Kagoshima Pref., Mt. Kirishima [Kari-
sima], 30 June 1958, A. Kawabe.

kiyosatoensis Kawabe, 1980 (Endothenia);
Tinea 11: 20. Holotype ¢, Japan, Ya-
mamshi Pref., Kiyosato, 1,300 m, 25—
28 August 1971, A. Kawabe.

knudsoni Miller, 1986 (Pseudexentera); J.
Lepid. Soc. 40: 224. Holotype 6, USA,
Texas, Kleberg Co., Riviera Beach site,
24 February 1984, E. Knudson.

koebelei Obraztsov, 1959 (Aphelia); Am.
Mus. Novit. 1964: 7. Holotype 6, USA,
Washington, Easton, Koebele.

komonana Kearfott, 1907 (Hysterosia);

Canad. Entomol. 37: 121. Lectotype 6,
USA, California, Santa Clara Co. Des-
ignated by Klots (1942).

kurosawai Kawabe, 1986 (Notocelia); En-
tomol. Pap. Pres. Kurosawa, Tokyo: 80.
Holotype 6, Taiwan, Hualien Hsien,
Hohuanshan, 3,100 m, 30 July—1 Au-
gust 1983, A. Kawabe.

kusaiensis Clarke, 1976 (Dudua aprobola);
Insects of Micronesia 9: 86. Holotype
6, Micronesia, Kusaie, Mutunlik, 14
February 1953, J. Clarke.

kuscheli Clarke, 1980 (Proeulia); J. Lepid.
Soc. 34: 184. Holotype ¢, Chile, Des-
venturadas Islands, San Ambrosia Is-
land, 450 m, on flowers of Thamnoseris
lacerata, 14 November 1960, G. Kus-
chel.

kusunokii Kawabe, 1993 (Apotomis); Tyo to
Ga 43: 257. Holotype 6, Japan, Keun-
shita, Mt. Daisetsu, 28 July 1983.

laciniana Zeller, 1875 (Phoxopteris); Ver-
handel. Zool.-Bot. Ges. Wein 25: 253.
“Type’’ 6, USA, Massachusetts, 1871.
Of the three males and one female re-

1043

ferred to in the original description, this
is the only specimen we have been able
to locate.

laetitia Clarke, 1968 (Cochylis); Proc. U.S.
Natl. Mus. 125: 25. Holotype ¢, Argen-
tina, Tucuman, Ciudad Universitaria,
800 m, 20 February 1959, J. Clarke.

lagopana Walsingham, 1879 (Steganopty-
cha); Ill. Lepid. Heter. Brit. Mus. 4: 71.
Syntype 2, USA, California, Colusa
Co., 28 June 1871. This species was de-
scribed from two males and three fe-
males; presumably all but the above
mentioned syntype are in BMNH.

lamberti Franclemont, 1986 (Sparganoth-
is); Proc. Entomol. Soc. Wash. 88: 56.
Holotype 3d, USA, South Carolina,
Oconee Co., Cherry Hill Recreation
Area, Route 107, 2,000’, 22 August
1958, J. Franclemont.

lambertiana Busck, 1915 (Tortrix); Proc.
Entomol. Soc. Wash. 17: 86. Holotype
3, USA, Oregon, Jackson Co., Ashland,
rf. Pinus lambertii, em: 27 June 1914,
P. Sergent. Although the original de-
scription gives “‘Oakland, Oreg.”’ as the
type locality, the label on the specimen
indicates Ashland.

lantana Busck, 1910 (Crocidosema); Proc.
Entomol. Soc. Wash. 12: 132. Holotype
36, Hawaii, Oahu, Tantalus, O. Swezey.

larana Brown, 2000 (Lobogenesis); Proc.
Entomol. Soc. Wash. 102: 29. Holotype
3d, Venezuela, Lara, Yacumba National
Park, 13 mk SE Sanare, 4,800’, cloud
forest, blacklight, 4-7 March 1978, J.
Heppner.

largo Heppner, 1981 (Cydia); J. Lepid. Soc.
35: 278. Holotype 36, USA, Florida,
Monroe Co., Key Largo, 15 mi NE Key
Largo City, 16 June 1974, J. Heppner.

laricana Busck, 1916 (Laspeyresia); Proc.
Entomol. Soc. Wash. 18: 152. Holotype
2, USA, Montana, Evaro, r.f., Larix oc-
cidentalis, May 1914, J. Brunner.

lariciana Kawabe, 1980 (Zeiraphera); Tin-
ea 11: 24. Holotype 6, Japan, Nagano
Pref., Shig-k6gen, 28 June 1964, Oza-
wa.

1044

latens Heinrich, 1929 (Thiodia); Proc. U.S.
Natl. Mus. 75: 2. Holotype 6, USA,
California, Tulare Co., Monachee
Meadows, 8,000’, “July 8-14.”

lathami Forbes, 1937 (Eucosma); J. N. Y.
Entomol. Soc. 43: 131. Holotype <4,
USA, New York, Long Island, Orient,
18 June 1935, R. Latham.

latiana Brown, 1991 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 52. Holotype ¢, Vene-
zuela, Aragua, Rancho Grande, 1,100
m, 10 May 1967, L. Rodriguez & C.
Andara.

laticurva Heinrich, 1929 (Eucosma); Proc.
U.S. Natl. Mus. 75: 4. Holotype ¢,
USA, California, Sierra Nevada.

latipunctana Walsingham, 1879 (Cochylis);
Ill. Lepid. Heter. Brit. Mus. 4: 29. Syn-
type 6, USA, California, Mendocino
Co., 31 May 1871. This species was de-
scribed from two males; the other syn-
type presumably is in the BMNH.

lautana Powell, 1960 (Argyrotaenia); Pan-
Pac. Entomol. 36: 90. Holotype <6,
USA, California, San Bernardino Co.,
San Bernardino Mtns., Camp Baldy,
“June 24-30.”

lavana Busck, 1907 (Phalonia); J. N. Y.
Entomol. Soc. 15: 27. Holotype ¢,
USA, Maryland, Montgomery Co., Hy-
attsville, A. Busck.

leguminana Busck, 1907 (Phalonia); J. N.
Y. Entomol. Soc. 15: 28. Holotype ¢,
USA, Washington, D.C., rf. Gleditchia
horrida, Feburary.

leguminis Heinrich, 1943 (Laspeyresia);
Proc. Entomol. Soc. Wash. 45: 71. Ho-
lotype 6, Peru, rf. beans, 19 August
1930, Willie.

leopardana Busck, 1906 (Hemimene); Proc.
Biol. Soc. Wash. 19: 181. Holotype 2,
USA, Maryland, Montgomery Co., Hy-
attsville, June 1906, A. Busck.

lepida Heinrich, 1924 (Epinotia cruciana);
J. Wash. Acad. Sci. 14: 391. Holotype
36, USA, New Hampshire, Coos Co.,
Mt. Washington, 4,000’, ‘“‘July 24-31.”

leucobasis Busck, 1916 (Laspeyresia);

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Proc. Entomol. Soc. Wash. 18: 152. Ho-
lotype 6, USA, Montana, Evaro, r.f. La-
rix occidentalis, May 1914, J. Brunner.

leucognoma Clarke, 1976 (Eumarissa); In-
sects of Micronesia 9: 32. Holotype 6,
Micronesia, Guam, Mt. Alifan, ex-um-
bellifer(?), 21 May 1936, O. Swezey.

leucothorax Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 150. Holotype 6, Brazil, New Bre-
men, 6 June 1936, E Hoffman.

lindana Fernald, 1892 (Steganoptycha);
Canad. Entomol. 24: 178. Lectotype d,
Canada, Ontario, Hamilton. Designated
by Miller (1970).

lindseyana Obraztsov, 1962 (Choristoneura
lambertiana); Am. Mus. Novit. 2101:
16. Holotype 6, USA, California, Mo-
doc Co., Warner Mtns., 3 mi E Davis
Creek, 15-23 July 1922, 5,500’, A.
Lindsey.

lineana Fernald, 1901 (Eucosma); J. N. Y.
Entomol. Soc. 9: 50. Holotype 2, USA,
Florida, Palm Beach Co., Palm Beach,
r.f. Anona laurifolia, H. Dyar. This spe-
cies was described from a single female.

linitipunctana Blanchard & Knudson, 1983
(Phaneta); J. Lepid. Soc. 37: 140. Ho-
lotype 6, USA, Texas, Nueces Co.,
North Padre Island, 9 September 1974,
A. & M. Blanchard.

listerana Kearfott, 1907 (Pharmacia);
Trans. Am. Entomol. Soc. 33: 80. Lec-
totype 6 (designated here), USA, Penn-
sylvania, Lackawanna Co., Nicholson, 6
July 1904, A. Lister. Selected by Obraz-
tsov in 1953.

lobata Razowski, 1988 (Argyrotaenia);
Acta Zool. Cracov. 31: 408. Holotype
3d, Bolivia, Cochabamba, Incachaca,
tropical cloud area, 2,100 m, 27 Au-
gust-5 September 1956, L. Pena.

loricana Grote, 1880 (Phoxopteris); Canad.
Entomol, 12: 218. ‘‘Type*> 62sAe
Ohio, Montgomery Co., Dayton, G. Pi-
late. According to the original descrip-
tion, the “‘type is in the collection of
Fernald,’’ which almost certainly refers
to the specimen cited above.

VOLUME 102, NUMBER 4

louisiana Busck, 1907 (Phalonia); J. N. Y.
Entomol. Soc. 15: 24. Holotype 6,
USA, Missouri, St. Louis Co., 2 mi W
St. Louis, August 1904, A. Busck.

luciferana Kawabe, 1980 (Zeiraphera);
Tinea 11: 26. Holotype 6, Japan, Mi-
yagi Pref., Sakunmi, 22 June 1969, T.
Watanabe.

luctuosana Blanchard, 1979 (Epiblema); J.
Lepid. Soc. 33: 184. Holotype ¢, USA,
Texas, Nueces Co., N. Padre Island, 6
April 1978, A. & M. Blanchard.

luculentana Heinrich, 1920 (Evetria); Proc.
U.S. Natl. Mus. 57: 56. Holotype ¢,
USA, Colorado, El Paso Co., r.f. Pinus
scopulorum, 5 May 1916, W. Edmon-
ston.

luoyingensis Kawabe, 1992 (Acleris); Tinea
13: 175. Holotype 6, Taiwan, Hualien
Hsien, Houhuanshan, Luoying Lodge,
2,800 m, 31 December 1988, A. Ka-
wabe.

lupicinia Clarke, 1971 (Dichelopa); Smith-
sonian Contrib. Zool. 56: 113. Holotype
36, Rapa Island, Pt. Maraia, rf. Rumex
crispus, em: 26 October 1963, J. & T.
Clarke.

macdunnoughi Obraztsov, 1963 (Acleris);
Proc. U.S. Natl. Mus. 114: 214. Holo-
type 2, USA, Massachusetts, Worcester
Co., Winchendon, 29 September 1902.

machimiana Barnes & Busck, 1920 (Spar-
ganothis); Contrib. Nat. Hist. Lepid.
North Am. 4: 211. Holotype °, USA,
Arizona, Cochise Co., Paradise, July.

macswaini Powell, 1980 (Decodes); Pacific
Insects 22: 89. Holotype 3, Mexico,
Nuevo Leon, 3 mi E Galeana, 5,000’,
7-9 August 1963, W. Duckworth & D.
Davis.

maculana Fernald, 1901 (Lipoptycha); J. N.
Y. Entomol. Soc. 9: 51. Holotype ¢,
USA, Florida, Palm Beach Co., Palm
Beach, r.f. Schoepfia arborescens, em:
24 February 1900, H. Dyar. There are
three specimens in the USNM collec-
tion with virtually identical collecting
data, two of which bear labels in Fer-

1045

nald’s hand indicating “‘type’’; the latter
two are undoubtedly the two males that
comprise the original series. Although
Miller (1970) was unable to find the ho-
lotype, one of the males bears a red
USNM type label (“‘Type 5413”’’) con-
sistent with the original description, im-
plicating it as the holotype.

maculatana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 48. Two
syntypes (2) 22), USA, California;
Lake Co., 17-19 June 1871. This spe-
cies was described from three males and
three females; the other syntypes pre-
sumably are in the BMNH.

maenamii Kawabe, 1974 (Olethreutes); Tyo
to Ga 25: 102. Holotype ¢, Japan, Izu
Island, Shikinejima, 16 June 1966, T.
Maenami.

magnifica Razowski & Becker, 1999 (Au-
ratonota); Revta. Bras. Zool. 16: 1174.
Holotype 6, Venezuela, Aragua, Ran-
cho Grande, 1,100 m, at light in mon-
tane tropical forest, 16 June 1973, J. C.
& K. G. Schaffer.

magnoliana Fernald, 1892 (Cacoecia);
Canad. Entomol. 24: 121. ‘“‘Type” 6,
USA, New York, Tompkins Co., Ithaca,
C.U. Exp. No. 292, rf. Magnolia acu-
minata, 22 June 1891, M. Slingerland.
Fernald did not indicate how many
specimens he examined.

maiana Kearfott, 1907 (Phalonia); Trans.
Am. Entomol. Soc. 33: 82. Lectotype
36, USA, New Jersey, Essex Co., Essex
County Park, 9-14 May 1910, W. Kear-
fott. Designated by Klots (1942).

maiorina Heinrich, 1923 (Bactra); Proc.
Entomol. Soc. Wash. 25: 105. Holotype
3, USA, Virginia, Arlington Co., Ar-
lington, rf. Scirpus fluviatilis, 1 July
1920.

malana Fernald, 1882 (Eccopsis); Trans.
Am. Entomol. Soc. 10: 72. Lectotype
36, USA, New York, Kings Co., Brook-
lyn, J. Smith. Designated by Miller
(1970).

manilkara Heppner, 1981 (Dichrorampha);
Florida Entomol. 64: 274. Holotype 6,

1046

USA, Florida, Monroe Co., Middle
Torch Key, 12 June 1974, r.f. Manilkara
emarginata, em: 6 July 1974, J. He-
ppner.

manoi Kawabe, 1987 (Hedya); Tinea 12:
140. Holotype 6, Japan, Mie Pref.,
Hokushei-machi, Otsujishenden, 8 May
1986, T. Mano.

mareda Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 142. Holotype 6, Brazil, Nova
Teutonia, May 1938, E Plaumann.

mariana Fernald, 1882 (Lophoderus);
Trans. Am. Entomol. Soc. 10: 67. Lec-
totype 6 (designated here), USA,
Maine, Penobscot Co., Orono. Selected
by Obraztsov 1956. According to the
original description, Fernald had four
males from Maine (Orono), Massachu-
setts, and New York.

maritima Dyar, 1904 (Ancylis); Proc. En-
tomol. Soc. Wash. 6: 221. Eight syn-
types, USA, Rhode Island, Washington
Co., Weekapaug, r.f. Lathyrus maritima,
H. Dyar. One of the syntypes was
placed in the type collection, presum-
ably by J. Clarke; the remainder are in
the main collection.

marloffiana Busck, 1907 (Phalonia); J. N.
Y. Entomol. Soc. 15: 26. Holotype °,
USA, Pennsylvania, Allegheny Co.,
Oak Station, 7 June 1906, E Marloff.

marmoreana Heinrich, 1923 (Epinotia);
Bull. U.S. Natl. Mus. 123: 222. Holo-
type d, USA, Utah, Tooele Co., Stock-
ton, 16 July 1913, T. Spalding.

marquesana Clarke, 1986 (Duessa); Smith-
sonian Contrib. Zool. 416: 155. Holo-
type 6, Marquesas Islands, Nuku Hiva,
Tunoa Ridge, 2,900’, 23 January 1968,
J. & T. Clarke.

maurodicha Clarke, 1976 (Herpystis); In-
sects of Micronesia 9: 74. Holotype 6,
Micronesia, Kusaie, Mutunlik, 22 m, 21
April 1953, J. Clarke.

maximana Barnes & Busck, 1920 (Pero-
nea), Contrib. Nat. Hist. Lepid. North
Am. 4: 216. Holotype 6, Canada, Brit-
ish Columbia, Victoria, A. Croker.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

mayelisana Blanchard, 1979 (Phaneta); J.
Lepid. Soc. 33: 209. Holotype 6, USA,
Texas, Cottle Co., Paducah, 17 April
1968, A. & M. Blanchard.

medioalbana Knudson, 1986 (Pammene); J.
Lepid. Soc. 40: 325. Holotype ¢, USA,
Texas, Gonzales Co., Palmetto State
Park, 23 March 1985, E. Knudson.

mediopartitum Heinrich, 1923 (Exartema);
Proc. Entomol. Soc. Wash. 25: 113. Ho-
lotype ¢, USA, Virginia, 1 June 1884.

megalorhis Diakonoff, 1982 (Sychnochlae-
na); Zool. Verhandel. (Leiden) 193:
117. Holotype 2, Sri Lanka, NE Dis-
trict, Kanda-ele Reservoir, 5.6 mi NW
Nuwara Eliya, 6,200’, 10—21 February
1970, D. Davis & B. Rowe.

megasaccula Brown, 1991 (Bidorpitia); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 73. Holotype ¢, Guate-
mala, Purulha, July, Schaus & Barnes.

melanantha Diakonoff, 1968 (Peridaeda-
la); Bull. U.S. Natl. Mus: 257: 77 sHe-
lotype 2, Philippine Islands, Mindanao,
Davao Prov., E slope Mt. McKinley,
mossy stunted forest, at light, 19 Sep-
tember 1946, CNHM Philippine Zool.,
Exped. 1946—47, H. Hoogstraal & D.
Heyneman.

melania Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 153. Holotype 6, Jamaica, St. Ann
Parish, 1,750’, E. Bell.

melanomesum Heinrich, 1923 (Exartema);
Proc. Entomol. Soc. Wash. 25: 119. Ho-
lotype 6, USA, Maine, Piscataquis Co.,
Sebec Lake, “July 16—23.”

melanoleuca Clarke, 1968 (Irazona); Proc.
U.S. Natl. Mus. 125: 44. Holotype ¢d,
Mexico, Puebla, rf. Pinus, January
1960, Guevara.

melasma Clarke, 1968 (Hysterosia); Proc.
U.S. Natl. Mus. 125: 9. Holotype 4,
Guatemala, Chejel, June, Schaus and
Barnes.

melia Clarke, 1976 (Adoxophyes); Insects
of Micronesia 9: 133. Holotype 6, Mi-
cronesia, Guam, Fadian, 19 August
1936, ex-Colubrina, O. Swezey.

VOLUME 102, NUMBER 4

melidora Razowski 1984 (Ardeutica); Acta
Zool. Cracov. 27: 217. Holotype 4,
Cuba, Sierra del Cobra Oriente, Loma
del Gato, 2,600’, 24-30 September
1935, J. Acufia, S. Bruner & L. Scara-
muzza.

meligma Clarke, 1986 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 141. Holo-
type 6, Marquesas Islands, Fatu Hiva,
Tahuna, 2,000’, 22 March 1968, J. & T.
Clarke.

membrosa Heinrich, 1926 (Laspeyresia);
Bull. U.S. Natl. Mus. 132: 49. Holotype
2, USA, Texas, Bexar Co., San Anto-
Mower. w_rosopis.s 290 June, 1917, A-
Busck.

mendaciana Blanchard & Knudson, 1983
(Suleima); Proc. Entomol. Soc. Wash.
85: 848. Holotype 6, USA, Texas,
Brewster Co., Big Bend National Park,
Dugout Wells, 28 September 1981, E.
Knudson.

mendora Clarke, 1968 (Cochylis); Proc.
U.S. Natl. Mus. 125: 24. Holotype d,
Chile, Santiago Province, Cajon de
Maypo, Cordillera, El Canelo, 12-20
January 1948, T. Ramirez.

mengelana Fernald, 1894 (Sericoris); En-
tomol. News 5: 131: Lectotype ¢,
Greenland, McCormick Bay. Designat-
ed by Miller (1970).

meridionalis Yasuda & Kawabe, 1980 (Ar-
chips); Tinea 11: 9. Holotype 6, Japan,
Hatsuno, Is. Amami-oshima, 11—13 Au-
gust 1977, A. Seino.

meritana Heinrich, 1923 (Epinotia); Bull.
U.S. Natl. Mus 123: 226. Holotype ¢,
USA, Utah, Carbon Co., Hiawatha, rf.
Pinus em: July 1921, H.<Peck.

messalina Clarke, 1971 (Dichelopa),;
Smithson. Contrib. Zool. 56: 102. Ho-
lotype 2, Rapa Island, Maugaoa, 950°
@92sm)e7 Nov 1963.5. &. 1. Clarke:

metallica Busck, 1914 (Evetria); Proc. En-
tomol. Soc. Wash. 16: 146. Holotype 6,
USA, Montana, Missoula Co., Missou-
la, r.f. pitch of yellow pine, 2 May 1913,
J. Brunner.

metariana Heinrich, 1923 (Eucosma); Bull.

1047

U.S. Natl. Mus. 123: 133. Holotype 6,
USA, California, Siskiyou Co., Shasta
Retreat, “June 16—23.”

mexicana Busck, 1907 (Pharmacis); J. N.
Y. Entomol. Soc. 15: 30. Holotype ¢,
USA, New Mexico, Beulah, 8,000’, T.
Cockerell, USNM type 10238.

mezion Razowski 1984 (Ardeutica); Acta
Zool. Cracoye, 27; 4218, Holotype 6:
Cuba, Sierra del Cobre Oriente, Loma
del Gato, 2,600’, 24-30 September
1935, J. Acuna, S. Bruner & L. Scara-
muzza.

mesostigmatias Diakonoff, 1977 (Thauma-
tographa),; Zool. Verhandel. (Leiden)
158: 42. Holotype 6, Taiwan, Rantais-
an, 15 May 1933;,S. Issiki:

messalina Clarke, 1971 (Dichelopa);
Smithsonian Contrib. Zool. 56:102. Ho-
lotype 2, Rapa Island, Maugaoa, 950’
[292 m], 7 November 1963, J. & T.
Clarke.

microptera Clarke, 1953 (Endothenia),; J.
Wash. Acad. Sci. 43: 230. Holotype 6,
USA, Illinois, Putnam Co., 24 August
1949, M. Glenn.

mieae Kawabe, 1980 (Lobesia); Tinea 11:
20. Holotype 6, Japan, Narahara, Hach-
iooji-shi, Tokyo, 9 August 1978, A. Ka-
wabe.

migratana Heinrich, 1923 (Thiodia); Bull.
U.S. Natl. Mus. 123: 53. Holotype 4,
USA, California, Inyo Co., Olanche,
“Apr 24-30.”

mimica Clarke, 1976 (Herpystis); Insects of
Micronesia 9: 78. Holotype °, Micro-
nesia, Palau Islands, Koror Island, Ko-
ror, 29 May 1957, C. Sabrosky.

minor Brown 1990 (Macrochlidia); J. N. Y.
Entomol. Soc. 98: 373. Holotype ¢,
Venezuela, Aragua, Charoni Pass, 1,400
m, 20 January 1965, S. & W. Duck-
worth.

minorata Heinrich, 1924 (Eucosma gigan-
teana); J. Wash. Acad. Sci. 14: 388. Ho-
lotype ¢, USA, Texas, Liberty Co., Lib-
erty, em: 28 July 1922, FE Bottimer.

minutana Kearfott, 1905 (Eucosma); Proc.
U.S. Natl. Mus. 28: 356. Lectotype 6,

1048

USA, New Jersey, Essex Co., Montclair,
July 1908, W. Kearfott. Designated by
Blanchard (1979).

mira Heinrich, 1929 (Anchylopera); Proc.
U.S. Natl. Mus. 75: 17. Holotype 6,
USA, Colorado, Jefferson Co., Chim-
ney Gulch, Golden, April, Oslar.

mira Razowski, 1989 (Saphenista); SHI-
LAP Revista Lepidopterologia 17: 206.
Holotype 2, Guatemala, Volcan Santa
Maria, November, Schaus & Barnes.

miramundi Razowski, 1988 (Clarkenia);
Acta Zool. Cracov. 31: 407. Holotype
?, El Salvador, Cerro Miramundo, 233
January 1971, S. Steinhauser.

mirosignata Heinrich, 1929 (Eucosma);
Proc. U.S. Natl. Mus. 75: 11. Holotype
3, USA, Arizona, Pima Co., Baboqui-
vari Mountains, 15-30 October 1924,
O. Poling.

miscitata Heinrich, 1926 (Laspeyresia);
Bull. U.S. Natl. Mus. 132: 64. Holotype
3, USA, California, Shasta National
Forest, rf. Pinus jeffreyi, May 1912,
Hopk. 11414, J. Miller.

misturana Heinrich, 1923 (Thiodia); Bull.
U.S. Natl. Mus. 123: 54. Holotype 6,
Canada, Saskatchewan, Oxbow, 9 June
1907, E Knab.

mobilensis Heinrich, 1923 (Eucosma); Bull.
U.S. Natl. Mus. 123: 125. Holotype ¢,
USA, Alabama, Baldwin Co., near
Daphne, eastern shore of Mobile Bay,
‘from root cuttings of Chrysoma (Sol-
idago) pauciflosculosa ... during Sep-
tember (1920),”’ em: 3 October 1920, T.
van Aller.

mochana Busck, 1913 (Olethreutes); Insect.
Inscit. Menst. 1: 92. Holotype @, British
Guiana, Mocha, “‘bred from Vochysea
guianensis,’’ H. Moore.

modestana Busck, 1907 (Hysterosia); J. N.
Y. Entomol. Soc. 15: 32. Holotype <4,
USA, Pennsylvania, Allegheny Co.,
Pittsburgh, 2 July 1905, H. Engel.

moffatiana Fernald, 1905 (Proteoteras);
Canad. Entomol. 37: 16. Lectotype <6,
Canada, Ontario, London. Designated
by Miller (1970).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

molesta Busck, 1916 (Laspeyresia); J.
Agric. Res. U.S; Dept: Agric. 77°373:
Holotype 6, USA, Virginia, Arlington
Co., Arlington, rf. peach, em: 12 Oc-
tober 1916.

molybdaina Clarke, 1976 (Adoxophyes); In-
sects of Micronesia 9: 138. Holotype 6,
Micronesia, Ponape, Colonia, 7 January
19535 J:<Glarke:

molybdina Clarke, 1968 (Carolella); Proc.
U.S. Natl. Mus. 125: 53. Holotype 2,
Mexico, San Luis Potosi, Tamazun-
chale, 6 January 1947, Chapman.

monela Clarke, 1976 (Icelita); Insects of
Micronesia 9: 39. Holotype 2, Micro-
nesia, Kusaie, Mutunlik, 22 m, 29 Jan-
uary 1953, J. Clarke.

monetiferanum Riley, 1881 (Exartema);
Trans. St. Louis Acad. Sci. 4: 317. Ho-
lotype 6, USA, Alabama, Barbour Co.,
Eufala, G. Latimer. Riley described this
species from a single specimen.

monitorana Heinrich, 1920 (Eucosma);
Proc. U.S. Natl. Mus. 57: 58. Holotype
3, USA, Pennsylvania, Danville, rf.
cones of Pinus, em: 4 May 1916, A.
Champlain.

monstrata Razowski, 1984 (Aphalonia);
Ann. Zool. Warsz. 38: 276. Holotype 6,
Peru, Divisoria, 5,200’, 20-23 June
1982, C. Covell.

montana Busck, 1914 (Evetria); Proc. En-
tomol. Soc. Wash. 16: 147. Holotype 3,
USA. Montana, Elliston, rf. Pinus con-
torta, em: 10 December 1913, J. Brun-
ner.

montana Bartlett-Calvert, 1892 (Antithes-
ia); Univ. Santiago Chile, Univ. Ann.
84: 831. Neotype 6, Argentina, Neu-
quén, Chapelco, Techos, 1,400 m, 24
January 1984, M. & P. Gentili. Desig-
nated by Brown (1998).

monticolana Kawabe, 1964 (Clepsis); Tyo
to Ga 15: 1. Holotype ¢, Japan, Toyama
Pref., Honshu, Mt. Tateyama, 29 August
1962, A. Kawabe.

moriutit Kawabe, 1987 (Stenodes); Micro-
lepid. Thailand 1: 72. Holotype 6,
Thailand, Chiang Mai, Doi Pui, ca.

VOLUME 102, NUMBER 4

1,300 m, 26—27 October 1985, S. Mor-
iuti, T. Saito & Y. Arita.

mormonensis Heinrich, 1923 (Evetria);
Bull. U.S. Natl. Mus. 123: 44. Holotype
36, USA, Utah, Sal Lake Co., Salt Lake
City, C. Ainslie.

murtfeldtiana Riley, 1881 (Anchylopera);
Trans. St. Louis Acad. Sci. 4: 323.
“Type’’ ¢, USA, Missouri, rf. oak, em:
19 May 1875, M. Murtfeldt. Riley de-
scribed this species from three speci-
mens, only one of which could be lo-
cated by us.

musetta Blanchard & Knudson, 1983 (Pha-
neta); Proc. Entomol. Soc. Wash. 85:
845. Holotype 6, USA, New Mexico,
Socorro Co., Gran Quivara Natl. Mon-
ument, 6,600’, 1-3 July 1964, D. Davis.

mydros Obraztsov, 1966 (Idolatteria); Proc.
U.S. Natl. Mus. 119: 6. Holotype ¢, Ec-
uador, Environs de Loja, 1887.

myopori Clarke, 1971 (Dichelopa); Smith-
sonian Contrib. Zool. 56: 115. Holotype
6, Rapa Island, Pariati Bay, rf. Myopo-
rum rapensis, em: 30 Nov 1963, J. & T.
Clarke.

myoxa Razowski, 1984 (Polyortha); Acta
Zool. Cracov. 27: 220. Holotype ¢,
Brazil, Santa Catarina, Nova Teutonia,
November 1961, E Plaumann.

naevifera Razowski, 1984 (Polyortha);
Acta Zool. Cracov. 27: 223. Holotype
6, Venezuela, Aragua, Rancho Grande,
10-21 February 1969, Duckworth &
Dietz.

nakajimai Kawabe, 1992 (Acleris); Tinea
13: 172. Holotype 6, Taiwan, Hualien
Hsien, Houhuanshan, Luoying Lodge,
2,800 m, 31 December 1988, A. Ka-
wabe.

naoma Clarke, 1953 (Epiblema); J. Wash.
Acad. Sci. 43: 226. Holotype ¢, USA,
Illinois, Putnam Co., rf. Ratibida pin-
nata, 25 June 1950, M. Glenn.

nebulosana Packard, 1867 (Grapholitha);
Proc. Boston Soc. Nat. Hist. 11: 61.
Syntype 6, Canada, Labrador, Straw-
berry Harbor, 26—30 July. Although the

1049

original description does not indicate
how many specimens were examined
nor identify a type, it states that the spe-
cies “is not infrequent at Strawberry
Harbor ...”

negligens Kawabe, 1978 (Neoanathamna);
Tinea 10: 185. Holotype 6, Japan, Shi-
koku, Kagawa Pref., Mt. Zozu, 22 May
1968, H. Toshima.

negundana Dyar, 1902 (Cacoecia); in Cau-
dell, Proc. Entomol. Soc. Wash. 5: 78.
“Type” 6, USA, Colorado, El Paso
Co., Pike’s Peak, Dyar & Caudell. Dyar
briefly diagnosed this species in a report
by Caudell (1902) without mentioning
any specimens or designating a type.

neoclyta Razowski, 1988 (Clarkeulia); Acta
Zool. Cracov. 31: 405. Holotype 6, Co-
lombia, Narino, Volcan Galeras, 2,900
m, 13 January 1959, J. Clarke.

neomexicana Dyar, 1903 (Evetria); Proc.
Entomol. Soc. Wash. 5: 286. Holotype
3, USA, New Mexico, San Miguel Co.,
Las Vegas. This species was described
from the single male cited above.

nepotinana Heinrich, 1923 (Thiodia); Bull.
U.S. Natl. Mus. 123: 263. Holotype d,
USA, Utah, Juab Co., Eureka, 30 May
1911, T. Spalding.

nephelodes Clarke, 1968 (Amallectis); Proc.
U.S. Natl. Mus. 125: 27. Holotype ¢,
Bolivia, Cochabamba, Incachaca, tropi-
cal cloud area, 2,100 m, 27 August—5
September 1956, L. Pena.

nigra Miller, 1966 (Laspeyresia); J. Lepid.
Socs ;20:0251. Holotype sd.Mexico,
Tlaxcala, February 1964, “en semilla P.
ayacahuite.”’

nigralbana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 41. Syn-
type 6, USA, California, Mendocina
Co., 10 June 1871. This species was de-
scribed from two males and two fe-
males; the other syntypes are presumed
to be in the BMNH.

nigricolor Yasuda & Kawabe, 1980 (Ka-
wabeia); Tinea 11: 10. Holotype 6, Ja-
pan, Nagano Pref. Azumi V., Shimaji-

1050

ma-dani, 780 m, 23 October 1976, N.
Hirano.

nigrilineana Kawabe, 1963 (Acleris); Tyo
to Ga 14: 71. Holotype 6, Japan, Na-
gano Pref., Honshu, Asama-sanso, 4
May 1962, A. Kawabe.

nigriplagana Franclemont, 1986 (Archips);
Proc. Entomol. Soc. Wash. 88: 59. Ho-
lotype ¢, USA, New York, Tompkins
Co., McLean Bogs Reserve, 2 July
1953, J. Franclemont.

nigrivelata Walsingham, 1914 (Tortrix);
Biol. Centr.-Am., Lepid., Heter. 4: 283.
Holotype ¢, Panama, Canal Zone, Ta-
bernilla, A. Busck. Walsingham de-
scribed this species from a single male.

nigromaculata Issiki, 1930 (Simaethis);
Ann. Mag. Natl. Hist. 6: 423. Holotype
?, Japan, Honsyu, Iwawakisan, 9 Au-
gust 1918, S. Issiki.

nigropunctata Kawabe, 1985 (Geogepa);
Tinea 12: 7. Holotype d, Taiwan, Chiai
Hsien, Fenchihu, 1,400 m, 8 August
1971, Y. Shabata.

ninana Dyar, 1903 (Carpocapsa); List Lep-
id. North Am.: 471. Two syntypes (1 ¢,
1 2), USA, Arizona, Fort Grant, 18 July
1892. Authorship of this taxon (as Gra-
pholitha ninana) usually is attributed to
Riley; the name first appeared in Riley’s
contribution to Smith’s (1891: 93)
checklist of the Lepidoptera of North
America without a description or refer-
ence to specimens examined. Heinrich
(1926: 60) correctly recognized that
Dyar presented the first description,
based on Riley’s specimens, in a foot-
note in his checklist (Dyar 1903), and
hence should be considered the author.

nipponana Razowski, 1977 (Stenodes); Tyo
to Ga 28: 35. Holotype 6, Japan, Si-
koku, Asizuri-Saki, 10 May 1951, S. Is-
siki.

nipponica Kawabe, 1976 (Eucosma); Tinea
11: 40. Holotype 6, Japan, Kanagawa
Pref., Oyama, 13 May 1958, A. Kawa-
be.

nitida Clarke, 1955 (Orthocomotis); Trans.
Royal Entomol. Soc. London 107: 143.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Holotype ¢d, Guatemala, Cayuga, “4,”
Schaus & Barnes.

nitida Horak, 1984 (Heterochorista); Syst.
Entomol. 9: 429. Holotype 3, Papua
New Guinea, Morobe District, Wau,
1,200 m, mercury vapor light, 8-14 De-
cember 1976, G. Hevel & R. Dietz.

niveopunctata Dognin, 1905 (Polyortha);
Ann. Soc. Entomol. Belgium 49: 85.
Holotype 2, Ecuador, Environs de Loja,
1893.

niveosana Packard, 1867 (Sciaphila); Proc.
Boston Soc. Nat. Hist. 11: 55. Holotype
3d, Canada, Labrador, Hopedale, 3 Au-
gust. The original description appears to
have been based on a single male.

nota Kawabe, 1978 (Cryptophlebia); Tinea
10: 188. Holotype ¢, Japan, Honshu,
Ishikawa Pref., Anamizu, Bira, 9 Au-
gust 1961, A. Kawabe.

notialis Miller, 1985 (Eucosma); Ann. En-
tomol. Soc. Am. 78: 244. Holotype 6,
USA, Texas, Kerr Co., Kerrville, 6
June, Lacey.

novimundi Heinrich, 1920 (Laspeyresia);
Canad. Entomol. 52: 257. Holotype 3,
USA, Wisconsin, Door Co., Sturgeon
Bay, 14 July 1920, C. Fluke.

nymphana Blanchard & Knudson, 1983
(Gretchena); Proc. Entomol. Soc. Wash.
85: 851. Holotype 6, USA, Texas, An-
derson Co., Tennessee Colony, Gus En-
geling Wildlife Management Area, 15
April 1968, A. & M. E. Blanchard.

nuntia Heinrich, 1929 (Eucosma); Proc.
U.S. Natl. Mus. 75: 10. Holotype 3,
USA, Utah, Juab Co., Callao, 17 April
1922:

nythobia Clarke, 1971 (Cryptophlebia);
Smithsonian Contrib. Zool. 56: 133.
Holotype 2, Rapa Island, Haurei, rf.
Myoporum rapensis, 19 September
1963, J. & T. Clarke.

oachranta Diakonoff, 1974 (Polylopha);
Zool. Verhandel. (Leiden) 131: 72. Ho-
lotype ¢, Saipan Island, Marianas, 0.5
mi E Tanapag, 20 April 1945, H. Dybas.
obfuscana Riley, 1888 (Paedisca); Proc.

VOLUME 102, NUMBER 4

Entomol. Soc. Wash. 1: 33. ““Type” 3,
USA, rf. Solidago, 2 June 1884. Riley
did not indicate the number of speci-
mens examined.

obliqua Diakonoff, 1982 (Grapholita);
Zool. Verhandel. (Leiden) 193: 20. Ho-
lotype 6, Sri Lanka, Anuradhapura Dis-
trict, Wildlife Sanctuary Bungalow, Hu-
nuwilagama, Wilpattu, 200’, 10-19
March 1970, Davis & Rowe.

obnigrana Heinrich, 1923 (Proteoteras);
Bull. U.S. Natl. Mus. 123: 169. Holo-
type 6, USA, New Hampshire, Chesh-
ire Co., Dublin, A. Busck.

obnisa Heinrich, 1926 (Laspeyresia); Bull.
U.S. Natl. Mus. 132: 47. Holotype °&,
Canada, British Columbia, Fraser Mills,
16 June 1922, E. Blackmore.

occidentalis Heinrich, 1923 (Thiodia stria-
tana); Bull. U.S. Natl. Mus. 123: 58.
Holotype 6, USA, California, Siskiyou
Co., Shasta Retreat, “‘July 16—23.”

oceani Diakonoff, 1956 (Bactra); Zool.
Verhandel. (Leiden) 29: 59. Holotype
6, Fiji Islands, Nadi, at light, July, J.
Tllinoworth.

ochracea Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 144. Holotype d6, Costa Rica, Car-
tago Province, Juan Vinas, W. Schaus.

ochraceana Fernald, 1901 (Epiblema); J.N.
Y. Entomol. Soc. 9: 51. Holotype 4,
USA, Florida, Palm Beach Co., Palm
Beach. This species was described from
a single male.

ochroma Clarke, 1986 (Dichelopa); Smith-
sonian Contrib. Zool. 416: 138. Holo-
type ¢, Marquesas Islands, Nuku Hiva,
Tapuaooa, 2,500’, 30 January 1968, J.
& T. Clarke.

ochrosaphes Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 152. Holotype 6, Brazil, Santa Ca-
tarina, 27 December 1935, EK Hoffman.

ocliferia Heinrich, 1926 (Hemimene), Bull.
U.S. Natl. Mus. 132: 21. Holotype 6,
USA, Nevada, Washoe Co., Pyramid
Lake.

oenotherana 1881

Riley, (Conchylis);

1051

Trans. St. Louis Acad. Sci. 4: 316.
“Type”? 2, USA, Missouri, r.f. Oenoth-
era, M. Murtfeld. Riley refers to four
specimens in the original description;
we have been able to locate only the
specimen cited above.

ogasawaraensis Kawabe & Ku, 1978 (Me-
tendothenia); Tinea 10: 169. Holotype
36, Japan, Ogasawara Island, Omura,
Chichijima, 14 June 1972, Y. Watanabe.

okubiensis Kawabe, 1987 (Eucosma); Tinea
12: 141. Holotype ¢, Japan, Okinawa
Island, Okubi, Kin, r.f. Bruguiera gym-
norrhiza, 26 April 1983, S. Hori.

olivaceana Fernald, 1882 (Eccopsis); Trans.
Am. Entomol. Soc. 10: 71. Lectotype
3, USA, Pennsylvania. Designated by
Miller (1970).

olivaceana Riley, 1881 (Grapholitha);
Trans. St. Louis Acad. Sci. 4: 320. Ho-
lotype ¢, USA, Illinois, J. Muhleman.
Riley described this species from a sin-
gle specimen.

olivata Dognin, 1905 (Tortrix); Ann. Soc.
Entomol. Belgium 49: 85. Holotype <6,
Colombia, Popayan, 1895.

onychistica Diakonoff, 1982 (Age); Zool.
Verhandel. (Leiden) 193: 57. Holotype
6, Sri Lanka, Mannar District, Mannar
Island, 2 mi. NE Mannar, 15’, 22 March
1970, D. Davis & B. Rowe.

opistocapna Diakonoff, 1977 (Thaumato-
grapha); Zool. Verhandel. (Leiden) 158:
38. Holotype 6, New Guinea, Wareng,
12 July 1936, S. Issiki.

opposita Heinrich, 1931 (Epinotia); Proc.
U.S. Natl. Mus. 79: 7. Holotype 4,
Peru, Lima, D. Willie, ““No. 154.29.”

optimana Dyar, 1893 (Eucosma),; Proc. En-
tomol. Soc. Wash. 5: 180. Syntype ¢d,
USA, Colorado, Garfield Co., Glen-
wood Springs, ““Oct 1—7,’’ W. Barnes;
syntype 2, USA, Colorado, Eagle Co.,
T. Cockerell. The two specimens listed
comprise the type series.

orcula Razowski, 1988 (Oregocerata);
Acta Zool. Cracov. 31: 393. Holotype
3, Bolivia, Cochabamba, Incachaca,

1052

tropical cloud area, 2,100 m, 27 Au-
gust—5 September 1956, L. Pena.

oregonensis Heinrich, 1923 (Thiodia); Bull.
U.S. Natl. Mus. 123: 47. Holotype <4,
USA, Oregon, Klamath Co., Crater
Lake, “July 24-31.”

oribasus Razowski, 1988 (Oryguncus);
Acta Zool. Cracov. 31: 402. Holotype
3, Peru, Cusco, Machu Picchu, 2,700
m, 6 February 1959, J. Clarke.

ornatula Heinrich, 1924 (Thiodia); J. Wash.
Acad. Sci. 14: 385. Holotype 6, USA,
Pennsylvania, Allegheny Co., Oak Sta-
tion, 21 June 1909, FE Marloff.

orphnoxantha Obraztsov, 1966 (Pseudo-
meritastis); Proc. U.S. Natl. Mus. 118:
227. Holotype 6, Costa Rica, Cartago
Province, Tuis, 2,400’, W. Schaus.

osmundana Fernald, 1879 (Penthina); Can-
ad. Entomol. 11: 156. Lectotype °&,
USA, Maine, Penobscot Co., Orono,
July, r.f. O. regalis. Designated by Mill-
er (1970).

ostentatrix Heinrich, 1923 (Hystricophora);
Bull. U.S. Natl. Mus. 123: 270. Holo-
type 6, USA, California, Tulare Co.,
Mineral King, “‘Aug. 1-7.” Heinrich
(1923) consistently misspelled HAystri-
chophora in his revision.

oxybela Razowski, 1988 (Ptynognathosia);
Acta Zool. Cracov. 31: 394. Holotype
3, Colombia, Cauca, Paramo de Parace,
Lake San Rafael, 3,570 m, 29 January
1959, J. Clarke.

oxycoccana Packard, 1869 (Peronea);
Guide to the Study of Insects: 334.
“Type” 3d, USA, eastern states. This is
the only Packard specimen of this spe-
cies we could locate; it is clearly labeled
as the type. Packard did not indicate
how many specimens he examined.

oxygona Diakonoff, 1968 (Eudemis); Bull.
U.S. Natl. Mus. 257: 50. Holotype 2,
Philippine Islands, Los Banfios, 2 July
1932, A. Evangelista.

oxyropa Razowski, 1990 (Ptoseulia); SHI-
LAP Rev. Lepid. 18: 212. Holotype <4,
Costa Rica, Cartago Province, Turrialba,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

22-28 February 1965, S. & W. Duck-
worth.

ozonia Razowski, 1990 (Ptoseulia); SHI-
LAP Rev. Lepid. 18: 213. Holotype ?,
Bolivia, Cochabamba, Incachaca, tropi-
cal cloud area, 2,100 m, 27 August—5
September 1956, L. Pefia.

packardiana Fernald, 1886 (Tortrix); Bull.
U.S. Dept. Agric. Entomol. 12: 20. Two
syntypes (2 66), USA, Maine, Peaks
Island, Casco Bay; and USA, New
Hampshire; both specimens are labeled
““cotype.’’ Freeman (1958) correctly in-
dicated that the “‘type”’ is in the USNM,
listing Peaks Island as the type locality.
This could be interpreted as his desig-
nation of the Maine specimen as the lec-
toype.

pallens Kawabe, 1980 (Neoanathamna);
Tinea 11: 22. Holotype 6, Japan, Hi-
totsu-uchi, Shionoe, Kagawa, 2 June
1973, H. Tashima.

pallidibasalis Heinrich, 1920 (Laspeyre-
sia); Proc. U.S. Natl. Mus. 57: 60. Ho-
lotype 6, USA, Oregon, Kaolin Beds,
rf. cones of Abies concolor, 7 Septem-
ber 1915, Sergent & Patterson.

pallifasciata Powell, 1978 (Rhyacionia); in
Powell and Miller, U.S. Dept. Agric.,
Agric. Handb. 514: 28. Holotype 6,
USA, Arizona, Coconino Co., Fort Val-
ley, 2.5 mi NW Flagstaff, 10 July 1981,
R. Hodges.

palmetum Heinrich, 1928 (Laspeyresia);
Proc. Entomol. Soc. Wash. 30: 6. Ho-
lotype 36, USA, Florida, near Royal
Palms State Park, “‘II.4,’’ E Jones.

palpana Walsingham, 1879 (Paedisca); Il.
Lepid. Heter. Brit. Mus. 4: 54. Two syn-
types (2 66), USA, California, Siski-
you Co., Mount Shasta, August 1871.
This species was described from five
males and one female. All but the two
syntypes listed above are presumed to
be in the BMNH.

paracinderella Powell, 1964 (Acleris);
Univ. Calif. Publ. Entomol. 32: 96. Ho-

VOLUME 102, NUMBER 4

lotype 6, USA, California, Nevada Co.,
r.f., Prunus dumosa, C. Riley.

paracornuta Brown, 1991 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 55. Holotype 6, Vene-
zuela, Aragua, Rancho Grande, 1,100
m, 16—23 October 1966, S. & W. Duck-
worth.

paradisiae Heinrich, 1923 (Hystricophora);
Bull. U.S. Natl. Mus. 123: 255. Holo-
type 6, USA, Washington, Pierce Co.,
Mount Rainier, Paradise Valley, “‘July
24-30.” Heinrich (1923) consistently
misspelled Hystrichophora in his revi-
sion.

paragnoma Clarke, 1986 (Dichelopa);
Smithsonian Contrib. Zool. 416: 139.
Holotype °, Marquesas Islands, Hiva
Oa, Puamau, 8 February 1968, J. & T.
Clarke.

parallelana Walsingham, 1879 (Cochylis);
Ill. Lepid. Heter. Brit. Mus. 4: 28. Syn-
type 2, USA, California, Lake Co., 18
June 1871. This species was described
from two males and two females; the
other syntypes are presumably in the
BMNH.

parana Busck, 1911 (Tortrix); Proc. U.S.
Natl. Mus. 40: 228. Holotype 2, Brazil,
Parana, Castro, “6234 Wlsm. 1908.”

paraplesiana Blanchard, 1979 (Sonia); J.
Lepid. Soc. 33: 184. Holotype ¢, USA,
Texas, Harris Co., Houston, 5 June
1968, A. & M. Blanchard.

parmata Razowski, 1984 (Ardeutica); Acta
Zool. Cracov. 27: 216. Holotype ¢,
Costa Rica, Ojo de Agua, Rt. 2, km 75,
20 July 1967, O. Flint.

parva Brown, 1984 (Corticivora); Proc.
Entomol. Soc. Wash. 86: 279. Holotype
6, USA, Arkansas, Washington Co.,
Devil’s Den State Park, 24 June 1966,
R. Hodges.

parvana Kawabe, 1980 (Phalonidia); Tinea
11: 29. Holotype 6, Japan, Yamanashi
Pref., Kiyosato, 22 July 1972, A. Ka-
wabe.

passerana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 49. Syn-

1053

type 2, USA, California, Mendocino
Co., 9-10 June 1871. According to the
original description, this species was de-
scribed from four males. Because the
USNM specimen is almost certainly
part of the type series, Walsingham ap-
parently erred in his determination of
the sexes. There is a congeneric male
specimen in the type collection labeled
as ‘““Paedisca plumigerana Wlsm. Cal.
Type’’—apparently a Walsingham man-
uscript name.

pastigiata Heinrich, 1929 (Thiodia); Proc.
U:S® Natl) Mus#i75: . 32 Holotyperd:
USA, California, Tulare Co., Monachee
Meadows, 8,000’, “July 8-14.”

pediasios Miller, 1985 (Eucosma); Ann.
Entomol. Soc. Am. 78: 241. Holotype
3, USA, Texas, Terrel Co., Sanderson,
25 April 1981, E. Knudson.

penai Brown, 2000 (Lobogenesis); Proc.
Entomol. Soc. Wash. 102: 27. Holotype
36, Bolivia, Incachaca, Cochabamba,
2,100 m, tropical cloud forest area, 27
August—5 September 1956, L. Pefia.

penai Clarke, 1968 (Amallectis); Proc. U.S.
Natl. Mus. 125: 28. Holotype °, Boliv-
ia, Cochabamba, Incachaca, tropical
cloud area, 2,100 m, 27 August—5 Sep-
tember 1956, L. Pena.

penai Razowski, 1988 (nape); Acta Zool.
Cracov. 31: 395. Holotype 6, Bolivia,
Cochabamba, Incachaca, tropical cloud
area, 2,100 m, 27 August-5 September
1956, L. Pena.

pennsylvaniana Kearfott, 1907 (Proteopte-
ryx); Trans. Am. Entomol. Soc. 33: 48.
‘Type’? 6, USA, Pennsylvania, Beaver
Co., New Brighton, 22 March 1903, H.
Merrick [genitalia slide only; we have
been unable to find the pinned speci-
men]. The original description of penn-
sylvaniana consists of the final para-
graph of the description of Proteopertyx
albicapitana: “‘It is not impossible that
fuller knowledge of the life histories of
the western and eastern forms may
prove them to be separate and distinct,
in which case I would propose the name

1054

pennsylvaniana for the latter.’ Klots
(1942) makes no mention of this spe-
cies; i.e., presumably no Kearfott spec-
imens are in the AMNH.

pergandeana Fernald, 1905 (Eucosma);
Canad. Entomol. 37: 399. Lectotype 6,
USA, Virginia, 4 June 1882. Designated
by Miller (1970).

periculosana Heinrich, 1923 (Epiblema);
Bull. U.S. Natl. Mus. 123: 268. Holo-
type 6, Canada, British Columbia,
Mount McLean, 5,500’, August, A.
Hanham.

peroneana Barnes & Busck, 1920 (Tortrix);
Contrib. Nat. Hist. Lepid. North Am. 4:
214. Holotype 6, USA, Arizona, Co-
chise Co., Palmerlee [Huachuca Moun-
tians].

perplexana Fernald, 1901 (Epiblema); J. N.
Y. Entomol. Soc. 9: 51. Holotype ¢,
USA, Florida, Palm Beach Co., Palm
Beach, H. Dyar.

perpropinqua Heinrich, 1929 (Eucosma);
Proc. U.S. Natl. Mus. 75: 8. Holotype
3d, USA, Arizona, Pima Co., Indian Oa-
sis, Sells Post Office, 15-30 April 1923,
O. Poling.

persolita Heinrich, 1929 (Eucosma); Proc.
U. S. Natl. Mus. 75: 7. Holotype ¢,
USA, Texas, Cameron Co., San Benito,
*“March 16—23.”’

perspicua Diakonoff, 1968 (Sycacantha in-
odes); Bull. U.S. Natl. Mus. 257: 61.
Holotype 6, Philippine Islands, Luzon,
Mt. Makiling, Baker.

perspicuana Barnes & Busck, 1920 (Hys-
terosia); Contrib. Nat. Hist. Lepid.
North Am. 4: 218. Holotype 2, USA,
Arizona, Cochise Co., Paradise, August.

peruvianus Razowski, 1988 (Telurips);
Acta Zool. Cracov. 31: 391. Holotype
3, Peru, Cusco, Machu Picchu, 2,700
m, 6 February 1959, J. Clarke.

phaeostropha Clarke, 1976 (Duessa); In-
sects of Micronesia 9: 30. Holotype &,
Micronesia, Rota, Rota, 22 m, at light,
20 July 1946, Townes 805.

phalaenopa Diakonoff, 1968 (Argyropto-
cha); Bull. U.S. Natl. Mus. 257: 70. Ho-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

lotype ¢, Philippine Islands, Luzon, Mt.
Makiling, Baker.

pharetrata Razowski, 1987 (Terinebrica);
Tinea 12(suppl.): 136. Holotype 3, Ar-
gentina, Tucuman, Ciudad Universitar-
ia, 800 m, 17 February 1959, J. Clarke.

philerocherda Diakonoff, 1964 (Bactra);
Zool. Verhandel. (Leiden) 70: 31. Ho-
lotype ¢, British West Indies, Domini-
ca, Antrim, 1,000’, 13 March 1956, J.
Clarke.

physoptila Diakonoff, 1968 (Peridaedala);
Bull. U.S. Natl. Mus. 257: 75. Holotype
6, Philippine Islands, Davao Province,
Mindanao, Mt. Apo, 7,000’, at light,
November 1946, CNHM Philippine
Zool. Exped., Alcasid.

piceae Issiki, 1961 (Panoplia); in Issiki &
Mutuura, Shinyogu Kagai Shogarui: 36.
Holotype ¢, Japan, Honsyu, Sinano,
Omekura, 14 July 1957, rf. pupa on
German Picea, T. Yasuda.

piceaea Busck, 1916 (Olethreutes); Proc.
Entomol. Soc. Wash. 18: 151. Holotype
3d, USA, Colorado, El Paso Co., Colo-
rado Springs, “bred during May and
July in the Forest Insect Division from
Picea parryana and Picea engelman-
ni,’ J. Polloch & B. Harvey.

picicola Dyar, 1906 (Eucosma); J. N. Y. En-
tomol. Soc. 14: 108. Holotype 2, USA,
Washington, Pierce Co., Mt. Rainier,
Paradise Valley, rf. Abies lasiocarpa,
H. Burke. According to the original de-
scription, this species was described
from a single female.

pimana Busck, 1907 (Phalonia); J. N. Y.
Entomol. Soc. 15: 24. Holotype 9°,
USA, Arizona, Pima Co., Baboquivera
Mtns., August.

piperana Busck, 1906 (Hemimene); Proc.
Biol. Soc. Wash. 19: 177. Holotype 6,
USA, Washington, Whitman Co., Pull-
man, 19 July 1898.

pithecolobiae Busck, 1934 (Gymnandroso-
ma); Entomol. Am. 13: 156. Holotype
3d, Cuba, Havana Province, Santiago,
Vegas, 15 July 1931, A. Otero.

placidus Yasuda & Kawabe, 1980 (Acleris);

VOLUME 102, NUMBER 4

Tinea 11: 12. Holotype 6, Japan, Nip-
para, Tokyo, 11 April 1964, A. Kawabe.

platina Clarke, 1968 (Irazona); Proc. U.S.
Natl. Mus. 125: 45. Holotype 6, Costa
Rica, Alajuela Province, Mount Poas,
May.

platyxantha Clarke, 1986 (Dichelopa);
Smithsonian Contrib. Zool. 416: 131.
Holotype 2, Marquesas Islands, Fatu
Hiva, Mt. Teoiaua, 2,000’, 22 March
1968, J. & T. Clarke.

pleodontia Razowski, 1987 (Monortha),
Bull. Polish Acad. Sci. 35: 65. Holotype
36, Panama, Rio Trinidad, March 1912,
A. Busck.

pleurogramma Clarke, 1976 (Duessa); In-
sects of Micronesia 9: 28. Holotype 6,
Micronesia, Kusaie, Mutunlik, 22 m, 30
March 1953, J. Clarke.

plicata Brown, 1992 (Catastega); J. N. Y.
Entomol. Soc. 100: 224. Holotype 2°,
USA, Arizona, Santa Cruz Co., Santa
Rita Mountains, Madera Canyon, Bog
Spring Camp Ground, 5,100’, 10-26
July 1964, D. Davis.

plumbata Clarke, 1951 (Anchylopera); J.
Wash. Acad. Sci. 41: 296. Holotype ¢,
Argentina, Tigre, May 1944, E Bour-
quin.

plumbosana Kawabe, 1972 (Olethreutes);
Tinea 9: 244. Holotype ¢, Toyama
Pref., Ohmaki, 3 June 1951, M. Hoshi-
no.

plummeriana Busck, 1906 (Hemimene);
Proc. Biol. Soc. Wash. 19: 181. Holo-
type 6, USA, Maryland, Montgomery
Co., Plummers Island (in Potomac Riv-
er above Washington, D.C.), May 1906.

plummeriana Busck, 1907 (Phalonia); J. N.
Y. Entomol. Soc. 15: 24. Holotype 6,
USA, Maryland, Montgomery Co.,
Plummers Island, 1 May 1903, A.
Busck.

poecilogramma Clarke, 1976 (Adoxo-
phyes); Insects of Micronesia 9: 125.
Holotype 6, Micronesia, Kusaie, Hill
1010, 300 m, light trap, 13 April 1953,
J. Clarke.

pollexifera Razowski, 1989 (Tamboma-

1055

chaya); SHILAP Revta. Lepid. 17: 205.
Holotype 6, Peru, Cusco, Tamboma-
chay 3,690 m, 3 February 1959, J.
Clarke.

polysticta Clarke, 1976 (Cryptaspasma); In-
sects of Micronesia 9: 63. Holotype &,
Micronesia, Kusaie, Mt. Matante, 300
m, 1 February 1953, J. Clarke.

ponderosa Powell, 1968 (Eucosma); Hil-
gardia,, 39:13.) Holotype, (GasUSA.
Oregon, Lake Co., Embody’s Mill, nr.
Silver Lake, r.f. Pinus ponderosa, 5 Au-
gust 1915,,P Sergent.

ponderosana Obraztsov, 1962 (Choristo-
neura lambertiana); Am. Mus. Novit.
2101: 14. Holotype ¢, USA, Colorado,
Sugarloaf, rf. Pinus ponderosa, 7 Au-
gust 1937, J. Beal:

ponera Walsingham, 1914 (Tortrix); Biol.
Centr.-Am., Lepid., Heter. 4: 279. Ho-
lotype 3d, Mexico, Puebla, Popocateptl
Park, 13,000’, W. Schaus. Walsingham
clearly identifies the USNM male as
‘““Type”’ in the original description.

poolei Brown, 1991 (Bidorpitia); in Brown
and Powell, Univ. Calif. Publ. Entomol.
111: 73. Holotype 6, Venezuela, Ara-
gua, Rancho Grande, 1,100 m, 8—14
August 1967, R. Poole.

populana Busck, 1916 (Laspeyresia); Proc.
Entomol. Soc. Wash. 18: 151. Holotype
2, USA, Montana, Missoula Co., Mis-
soula, 15 June 1914, J. Brunner.

postica Bae, 1993 (Lobesia); Japan. J. En-
tomol. 61: 522. Holotype 2, Taiwan,
Pintung Hsien, Kenting Park, 100 m,
26-27 March 1984, A. Kawabe.

pottsi Clarke, 1976 (Dudua); Insects of Mi-
cronesia 9: 94. Holotype 6, Micronesia,
Truk, Wena (Moen), Civil Administra-
tion Area, 6 March 1949, R. Potts.

praesumptiosa Heinrich, 1923 (Epiblema);
Bull. U.S. Natl. Mus. 123: 143. Holo-
type 6, USA, Texas, Cameron Co.,
Brownsville, 27 May 1917, A. Busck.

prochaldera Clarke, 1955 (Orthocomotis);
Trans. Royal Entomol. Soc. London
107: 145. Holotype ¢, Equador, Envi-
rons de Loja, 1890.

1056

proxima Clarke, 1976 (Dudua); Insects of
Micronesia 9: 91. Holotype ¢, Micro-
nesia, Ponape, Colonia, 9 January 1953,
J. Clarke.

pruni Heinrich, 1923 (Anchylopera burges-
siana); Bull. U.S. Natl. Mus. 123: 239.
Holotype 6, USA, New Jersey, Cald-
well, 17 May 1903, W. Kearfott.

psathyra Diakonoff, 1989 (Lopharcha);
Tinea 12: 202. Holotype 6, Japan, Kyu-
syu, Hikosan, 15 July 1937, S. Issiki.

pseudogamma Brown, 1990 (Paraptila); J.
Lepid. Soc. 44: 265. Holotype 6, El
Salvador, Santa Tecla, 28-29 October
1967, E. Todd.

pseudolivata Clarke, 1955 (Orthocomotis);
Trans. Entomol. Soc. London 107: 147.
Holotype 6, Brazil, New Bremen, 7
October 1936, E Hoffman.

pseudomalesana Clarke, 1986 (Cydia);
Smithsonian Contrib. Zool. 416: 164.
Holotype ¢, Marquesas Islands, Fatu
Hiva, Omoa, 16 March 1968, J. & T.
Clarke.

pseudoplesia Brown, 1991 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 60. Holotype 3, Vene-
zuela, Aragua, Rancho Grande, 1,100
m, 22-31 July 1967, R. Poole.

pseudotsugana Kearfott, 1904 (Cydia);
Canad. Entomol. 36: 110. Holotype 6,
Canada, British Columbia, Kaslo, rf.
Pseudotsuga, larva collected 24 June.
Although the specimen bearing the
Kearfott type label (Type No. 7788) has
no locality data, the pupal shell and as-
sociated plant material indicate that it is
the specimen referred to as ‘“‘also bred
by him [Dyar] from larvae taken June
24 on Pseudotsuga sp.”

ptycta Razowski, 1987 (Popayanita); Tinea
12(suppl.): 126. Holotype 6, Colombia,
Cauca, 17 km SE Popayan, 2,000 m, 10
January 1959, J. Clarke.

ptygma Razowski, 1993 (Apotoforma);
Acta Zool. Cracov. 36: 186. Holotype
2, Mexico, San Luis Potosi, 8 August
1966, O. Flint. Although the specimen
is not labeled as the holotype, Razows-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ki’s description was clearly based on
this single female specimen.

pulchella Kawabe, 1963 (Acleris); Tyo to
Ga 14: 71. Holotype 6, Japan, Honshu,
Tokyo, Tama Hills, 27 June 1958, M.
Suzuki.

pulcheria Clarke, 1971 (Dichelopa); Smith-
sonian Contrib. Zool. 56: 120. Holotype
6, Rapa Island, Perau, 1,900’, 15 Oc-
tober 1963, J. & T. Clarke.

pulcherrima Razowski, 1971 (Acleris);
Acta Zool. Cracov. 16: 548. Holotype
?, Taiwan, Hassenzan, 5 June 1942, S.
Issiki.

pulsatillana Dyar, 1903 (Eucosma); Proc.
Entomol. Soc. Wash. 5: 297. Nineteen
syntypes, USA, Colorado, Jefferson
Co., “‘foothills at Boulder and Golden,”’
rf. Pulsatilla hirsutissima. According to
the original description, this species was
described from 20 males and eight fe-
males.

pulveratana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 45. Syn-
type 6, USA, California, San Francisco,
16 May 1871. This species was de-
scribed from four males, three of which
are presumed to be in the BMNH.

punctidiscanum Dyar, 1904 (Gymnandro-
soma); Proc. Entomol. Soc. Wash. 6:
60. Two syntypes (2 ¢¢), USA, Wash-
ington, D.C., May, L. Howard; Penn-
sylvania, New Brighton. The two spec-
imens comprise the original type series.

purpuratus Kawabe, 1965 (Archips); Tyo to
Ga 16: 16. Holotype 3, Japan, Gunma
Pref., Mt. Akagi, 19-21 July 1964, A.
Kawabe.

purpurea Dognin, 1904 (Atteria); Ann. Soc.
Entomol. Belgium 48: 132. Holotype @,
Ecuador, Environs de Loja, 1893.

quadrifasciana Fernald, 1882 (Tortrix);
Trans. Am. Entomol. Soc. 10: 67. Two
syntypes (2 66), USA, Maine. Accord-
ing to the original description, Fernald
had specimens from Maine, New
Hampshire, Massachusetts, New York,
and Illinois. Although the number of

VOLUME 102, NUMBER 4

specimens examined was not given in
the description, Fernald’s unpublished
list of types donated to the USNM in-
dicates three “‘cotypes.”’

quercana Fernald, 1882 (Cenopis); Trans.
Am. Entomol. Soc. 10: 69. Lectotype ?
(designated here), USA, Texas, Dallas
Co., Dallas, 29 May. Selected by Powell
in 1965. Fernald described this species
from four males and four females from
New York, Missouri, and Texas.

quercifoliana Fitch, 1858 (Tortrix); New
Work Agric: (Rept;,.826 “Type” «<,
USA, New York [no locality data on
specimen label]. Fitch did not indicate
the number of specimens he examined;
this is the only Fitch specimen of this
species we could find.

radicana Heinrich, 1923 (Griselda); Bull.
U.S. Natl. Mus. 123: 186. Lectotype ¢,
Canada, British Columbia, on spruce,
G. Hewitt. Designated by Obraztsov
(1965).

rafaeliana Razowski, 1989 (Saphenista);
SHILAP Revta. Lepid. 17: 206. Holo-
type 6, Colombia, Cauca, Paramo de
Parace, Lake San Rafael, 3,570 m, 27
January 1959, J. Clarke.

rana Busck, 1907 (Phalonia); J. N. Y. En-
tomol. Soc. 15: 22. Holotype 6, USA,
Pennsylvania, Allegheny Co., Pitts-
burgh, 30 July 1905, H. Engel.

razowskii Kawabe, 1963 (Tortricodes) Tin-
ea 6: 7. Holotype 6, Japan, Chiba Pref.,
Narashino, 4 March 1958, A. Kawabe.

razowskit Kawabe, 1992 (Clepsis); Tinea
13: 178. Holotype 3, Taiwan, Hualien
Hsien, Kuanyuan, 2,400 m, 7-8 August
1987, A. Kawabe.

reprobata Clarke, 1976 (Lobesia); Insects
of Micronesia 9: 104. Holotype °, Mi-
cronesia, Guam, Pt. Oca, 4 June 1945,
Bohart and Gressitt.

rescissoriana Heinrich, 1920 (Eucosma);
Proc. U.S. Natl. Mus. 57: 58. Holotype
3, USA, Oregon, Sprague River, rf.
cones of Pinus murrayana, 30 May
1914, P. Sergent.

1057

retrusa Razowski, 1993 (Acleris); Acta
Zool. Cracov. 36: 191. Holotype °&,
Mexico, Jalapa, November 1963, rf.
Rubus, N. Krauss.

rhodographa Clarke, 1971 (Dichelopa);
Smithsonian Contrib. Zool. 56: 103.
Holotype ¢, Rapa Island, Perau, 1000’
(307 m), rf. Veronica rapensis, em: 18
November 1963, J. Clarke.

rhoifructana Kearfott, 1904 (Polychrosis);
Trans. Am. Entomol. Soc. 30: 296. Ho-
lotype ¢, USA, Ohio, Wayne Co., Woo-
ster, Insectory, 29 January 1884, Murt-
feldt.

rigidana Fernald, 1880 (Retinia); in Com-
stock, Annu. Rep. Dept. Agric. 1880:
237. Lectotype 2, USA, New York,
Tompkins Co., Ithaca. Designated by
Miller (1970).

rosaocellana Knudson, 1986 (Eucosma); J.
Lepid. Soc. 40: 322. Holotype 36, USA,
Texas, Hemphill Co., Gene Howe
WMA, 18 May 1985, E. Knudson.

rubra Kawabe, 1992 (Neopotamia); Japan.
Hetero. J. 170: 358. Holotype 3, Tai-
wan, Huslien Hsien, Hohuanshan, 3,100
m, 30 July—1 August 1983, A. Kawabe.

ruidosana Heinrich, 1923 (Epinotia); Bull.
U.S. Natl. Mus. 123: 216. Holotype 6,
USA, New Mexico, Lincoln National
Forest, Ruidosa Canyon, larvae mining
leaves of “‘Alum root”? [Heuchera woo-
toni], 8 May 1917, C. Heinrich.

ruptimacula Dognin, 1904 (Tortrix); Ann.
Soc. Entomol. Belgium 48: 132. Holo-
type °, Ecuador, Cayanuma, Loja.

russata Heinrich, 1924 (Epinotia cruciana);
J. Wash. Acad. Sci. 14:391. Holotype
3, Canada, British Columbia, Victoria,
24 June 1923.

russeola Heinrich, 1929 (Eucosma); Proc.
U.S. Natl. Mus. 75: 6. Holotype 6,
USA, California, Los Angeles Co., 11
June 1921, K. Coolidge.

sabiniana Kearfott, 1907 (Evetria); Trans.
Am “Entomols'Soech 33:2. siypey ci,
USA, California, Sacramento Co., rf.
Pinus sabiniana, 9 August 1885. Kear-

1058

fott described this species from two
specimens, one in the USNM collection
and one in his personal collection. Al-
though the Kearfott collection is depos-
ited at the AMNH, Klots (1942) makes
no mention of a specimen of this spe-
cies there.

saetigera Razowski, 1987 (Terinebrica);
Tinea 12(suppl.): 137. Holotype 6, Bo-
livia, Cochabamba, Incachaca, tropical
cloud area, 2,100 m, 27 August—5 Sep-
tember 1956, L. Pena.

sagax Razowski, 1984 (Polyortha); Acta
Zool. Cracov. 27: 221. Holotype ¢, Bo-
livia, Cochabamba, Incachaca, J. Stein-
bach.

saileri Clarke, 1987 (Cryptophlebia); Acta
Entomol. Chilean 14: 7. Holotype 6,
Chile, Iquique Province, Region Tara-
paca, Pampa Tamarugal, Fundo Refres-
co, rf. Prosopis tamarugo, 29 June
1985, D. Bobadilla.

salaciana Blanchard & Knudson, 1981 (Eu-
cosma); J. Lepid. Soc. 35: 176. Holo-
type 6, USA, Texas, Nueces Co., North
Padre Island, 13 October 1979, A. & M.
Blanchard.

saliciana Clemens, 1864 (Hedya); Proc.
Entomol. Soc. Philadelphia 3: 515. Syn-
type 36, USA, [probably eastern U.S.;
no data on specimen label or in original
description], rf. Salix longifolia, Walsh.
Although Clemens indicated the num-
ber of specimens examined and the de-
pository (e.g., “Coll. Ent. Soc. Phila-
delphia’’) for most species described in
this work, he did not do so for this or
the following species, H. salicicolana.

salicicolana Clemens, 1864 (Hedya); Proc.
Entomol. Soc. Philadelphia 3: 515. Two
syntypes (1 6, 1 2), USA, [probably
eastern U.S.; no data on specimen labels
or in original description], r.f. Salix hu-
milis, Walsh.

salmicolorana Heinrich, 1923 (Thiodia);
Bull. U.S. Natl. Mus. 123: 62. Holotype
3, USA, Utah, Tooele Co., Stockton, 30
July 1913, T. Spalding.

salmonicolor Powell, 1978 (Rhyacionia); in

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Powell and Miller, U.S. Dept. Agric.,
Agric. Handb. 514: 15. Holotype 6,
USA, Texas, Jeff Davis Co., Davis
Mtns., Mt. Locke, 19 May 1971, A. &
M. Blanchard.

salvadorana Brown & Powell, 2000 (Ano-
pina); Univ. Calif. Publ. Entomol. 120:
57. Holotype 6, El Salvador, Cerro Mir-
amundo, 2,300 m, 11 January 1969, S.
Steinhauser.

salweenensis Miller, 1977 (Petrova); J.
Lepid. Soc. 31: 136. Holotype 6, Thai-
land, Baw Luang, 20 March 1972, D.
Chaiglom.

sambuci Clarke, 1952 (Polychrosis); J.
Wash. Acad. Sci. 43: 228. Holotype 36,
USA, Illinois, Putnam Co., rf. Sambu-
cus canadensis, 5 July, M. Glenn.

santacrucis Obraztsov, 1963 (Acleris);
Proc. U.S. Natl. Mus. 114: 225. Holo-
type 6, USA, California, Santa Cruz
Co., Santa Cruz, 29 October 1932, W.
Tilden.

santamariana Razowski, 1988 (Ortogna-
thosia); Acta Zool. Cracov. 31: 392.
Holotype ¢, Guatemala, Volcan Santa
Maria, Barnes & Schaus.

sapodilla Heppner, 1981 (Dichrorampha);
Florida Entomol. 64: 271. Holotype 3d,
USA, Florida, Dade Co., Homestead,
rf. Achras zapota, 14 April 1974, R.
Baronowski.

saracana Kearfott, 1907 (Cenopis); Trans.
Am. Entomol. Soc. 33: 68. “Type” 6,
USA, New Jersey, Essex Co., Essex
County Park, hills above Montclair, July
1907, W. Kearfott. Kearfott described
this species from one male and two fe-
males, two field-collected and one
reared from sassafras. Klots (1942) does
not mention this species in his treatment
of Kearfott types in the AMNH.

satoi Kawabe, 1978 (Pseudohedya); Tinea
10: 174. Holotype 6, Japan, Hokkaido,
Mt. Tentosan, Abashiri, 4—5 August
1975, R. Sato.

sayonae Kawabe, 1985 (Acleris); Tinea 12:
5. Holotype 6, Taiwan, Nantou Hsien,

VOLUME 102, NUMBER 4

Nanshanchi, 25—26 July 1983, A. Ka-
wabe.

saxicolana Walsingham 1879 (Cochlyis);
Ill. Lepid. Heter. Brit. Mus. 4: 29. Syn-
type 3d, USA, southern Oregon, end of
May, 1872. Walsingham described this
species from three males; the other two
are presumed to be in the BMNH.

scalaris Diakonoff, 1968 (Petrova); U.S.
Natl. Mus. Bull 257: 86. Holotype &,
Philippine Islands, Mountain Province,
Luzon, base of Mt. Data, 5,500’, 22
April 1946, CNHM Philippine Zool.
Exped. 1946—47, H. Hoogstraal.

scardiana Dognin, 1905 (Cnephasia); Ann.
Soc. Entomol. Belgium 49: 86. Holo-
type 2, Ecuador, El Monje de Loja,
1894.

schizodelta Diakonoff, 1982 (Grapholita);
Zool. Verhandel. (Leiden) 193: 18. Ho-
lotype ¢, Sri Lanka, Ratnapura District,
Uggalkaltota, 350’, Irrigation Bunga-
low, 31 January—8 February 1970, D.
Davis & B. Rowe.

schwarziana Busck, 1907 (Phalonia); J. N.
Y. Entomol. Soc. 15: 25. Holotype °&,
USA, Maryland, Montgomery Co.,
Plummers Island (in Potomac River
above Washington, D.C.), July 1903, A.
Busck.

scleroductus Brown, 1991 (Punctapinella);
Los Angeles Co. Mus. Contrib. Zool.
423: 4. Holotype °, Brazil, Rio de Jan-
iero. 3 July 1929, J. Zikan-

sedatana Busck, 1906 (Hemimene); Proc.
Biol. Soc. Wash. 19: 177. Holotype <6,
USA, Colorado, South Park, 27 August
1905.

segregata Heinrich, 1924 (Thiodia); J.
Wash. Acad. Sci. 14: 388. Holotype 6,
USA, California, Tulare Co., Monachee
Meadows, “July 16—23.”

seiugata Razowski, 1987 (Terinebrica),
Tinea 12(suppl.): 138. Holotype °,
Peru, Cusco, Machu Picchu, 2,385 m, 4
February 1959, J. Clarke.

semicirculana Fernald, 1882 (Tortrix);
Trans. Am. Entomol. Soc. 10: 68. Lec-
totype 6, USA, Colorado, Morrison,

1059

1880. Designated by Brown and Powell
@g91).

senatrix Heinrich, 1924 (Exentera); J.
Wash. Acad. Sci. 14: 390. Holotype ¢,
USA, Arizona, Cochise Co., Paradise,
*“March 8-15.”

senilis Razowski, 1987 (Silenis); Tinea
12(suppl.): 128. Holotype 6, Bolivia,
Cochabamba, Incachaca, tropical cloud
area, 17 August-5 September 1956,
2,100 m, L. Pena.

seorsa Heinrich, 1924 (Epinotia); J. Wash.
Acad. Sci. 14: 392. Holotype 6, Cana-
da, British Columbia, Vavenby, 16 Sep-
tember 1922, T. Moilliet.

separationis Heinrich, 1923 (Epiblema
praesumptiosa); Bull. U.S. Natl. Mus.
123: 143. Holotype ¢, USA, Texas,
Cameron Co., Brownsville, 27 May
1917, A. Busck.

seraphicana Heinrich, 1923 (Hystricophora
asphodelana); Bull. U.S. Natl. Mus.
123: 258. Holotype 6, USA, Washing-
ton, Whitman Co., Pullman, 13 May
1898, C. Piper. Heinrich (1923) consis-
tently misspelled Hystrichophora in his
revision.

serapicana Heinrich, 1923 (Eucosma);
Bull. U.S. Natl. Mus. 123: 266. Holo-
type 6, USA, Montana, Cascade Co.,
Great Falls, 8 July 1921, H. Dyar.

serena Clarke, 1968 (Cochylis); Proc. U.S.
Natl. Mus. 125: 20. Holotype 6, Brazil,
Santa Catarina, Nova Teutonia, July
1963, E Plaumann.

sertula Diakonoff, 1982 (Licigena); Zool.
Verhandel. (Leiden) 193: 16. Holotype
6, Sri Lanka, Kandy, Udawattekelle,
15-24 January 1970, D. Davis & B.
Rowe.

shastana Walsingham, 1879 (Paedisca); Il.
Lepid. Heter. Brit. Mus. 4: 46. Syntype
6, USA, California, Siskiyou Co.,
Mount Shasta, August 1871. This spe-
cies was described from two males: one
syntype in the USNM and the other ap-
parently in the BMNH.

shastensis McDunnough, 1955 (Anchylo-
pera columbiana); Am. Mus. Novit.

1060

1725: 12. Holotype 6, USA, California,
Siskiyou Co., Shasta Retreat, “‘June 8—
Le

shibatai Kawabe, 1985 (Archips); Tinea 12:
1. Holotype 3, Taiwan, Chiai Hsien,
Fenchihu, 1,400 m, 3 August 1973, Y.
Shibata.

shikokuensis Kawabe, 1984 (Epinotia); Tin-
ea 11: 187. Holotype 6, Japan, Shiko-
ku, Kagawa Pref., Oku-shio-iri, 5 Au-
gust 1972, H. Toshima.

shimekii Kawabe, 1974 (Zeiraphera); Tyo
to Ga 25: 97. Holotype 6, Japan, Gun-
ma Pref., Yunotaira, 26 July 1968, A.
Kawabe.

sicaria Diakonoff, 1982 (Acroclita); Zool.
Verhandel. (Leiden) 193: 55. Holotype
36, Sri Lanka, Galle District, Kanneliya,
200’, 15—17 October 1976, G. Hevel, R.
Dietz, R. Karunaratne & D. Balasoori-
ya.

sierrae Blanchard & Knudson, 1983 (Eu-
cosma); Proc. Entomol. Soc. Wash. 85:
850. Holotype ¢, USA, Texas, Culber-
son Co., Sierra Diablo Wildlife Man-
agement Area, 6,400’, 30 August 1970,
J. Franclemont.

sierrae McDunnough, 1955 (Anchylopera
simuloides); Am. Mus. Novit. 1725: 14.
Holotype 6, USA, California, Tulare
Co., Mineral King, “‘June 24-30.”

signiferana Heinrich, 1923 (Epinotia); Bull.
U.S. Natl. Mus. 123: 232. Holotype 6,
USA, California, San Diego Co., San
Diego, 14 November 1911, W. Wright.

silvertonana Obraztsov, 1962 (Anopina);
Am. Mus. Novit. 2082: 19. Holotype ¢d,
USA, Colorado, San Juan Co., Silver-
ton, “July 24-31.”

silvertoniensis Heinrich, 1923 (Epinotia);
Bull. U.S. Natl. Mus. 123: 214. Holo-
type 6, USA, Colorado, San Juan Co.,
Silverton, “July 16—23.”

simplex Diakonoff, 1968 (Ebodina); Bull.
U.S. Natl. Mus. 257: 35. Holotype <4,
Philippine Islands, Luzon, Mt. Makil-
ing, Baker.

simplex McDunnough, 1925 (Eucosma);
Canad. Entomol. 57:21. Holotype °&,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

USA, Iowa, Woodbury Co., Sioux City,
7 May 1916.

simpsoni Busck, 1903 (Carpocapsa pomo-
nella); Proc. Entomol. Soc. Wash. 5:
236. Holotype 6, USA, Idaho, Ada Co.,
Boise, 8 August 1902, C. Simpson.

simulata Heinrich, 1928 (Rhyacionia),;
Proc. Entomol. Soc. Wash. 30: 62. Ho-
lotype °, Japan, Yokahoma, r.f. Pinus
thunbergii, em: 5 July 1924, A. Kariya.

sinistra Heinrich, 1926 (Bactra); Bull. U.S.
Natl. Mus. 132: 87. Holotype 2, USA,
Louisiana, Sabine River Ferry, opposite
Orange, Cornell University Lot 542,
Sub 20, 20 June 1917.

siskiyouensis Heinrich, 1923 (Epinotia pul-
satillana); Bull. U.S. Natl. Mus. 123:
202. Holotype 6, USA, California, Sis-
kiyou Co., Shasta Retreat, “Aug 16—

skinnerana Heinrich, 1923 (Suleima); Bull.
U.S. Natl. Mus. 123: 157. Holotype 6,
USA, Arizona, Cochise Co., Huachuca
Mountain, Carr Canyon, August 1905,
H. Skinner.

sonae Clarke, 1949 (Eulia); Acta Zool. Lil-
loana (Tucuman) 7: 587. Holotype ¢,
Brazil, Santa Catarina, 12 July 1935, E
Hoffmann.

sotipena Brown, 1986 (Epinotia); J. Lepid.
Soc. 40: 341. Holotype 2, USA, Mary-
land, Montgomery Co., Plummers Is-
land, Potomac River, 7 April 1962, R.
W. Hodges.

spaldingiana Obraztsov, 1961 (Argyrotaen-
ia); Am. Mus. Novit. 2048: 20. Holo-
type 6, USA, Utah, Utah Co., Provo, 4
August 1908, T. Spalding.

spartinana Barnes & McDunnough, 1916
(Phalonia); Canad. Entomol. 48: 144.
Holotype 36, USA, South Dakota,
Union Co., Elk Point, rf. Spartina, 30
July 1915, C. Ainslie.

spectra Brown, 1992 (Catastega); J. N. Y.
Entomol Soc. 100: 211. Holotype ¢,
USA, Texas, Jeff Davis Co., Mt. Locke,
McDonald Observatory, 19 July 1971,
A. Blanchard.

spinacea Razowski, 1994 (Psiathovalva);

VOLUME 102, NUMBER 4

SHILAP Revta. Lepid. 22: 69. Holotype
3d, Venezuela, Aragua, Rancho Grande,
1,100 m, 11—15 January 1966, S. & W.
Duckworth.

spinea Razowski, 1999 (Netechma); Polski
Pismo Entomol. 68: 93. Holotype 6,
Bolivia, Cochabamba, Incachaca, 2,100
m, tropical cloud area, 27 August—5
September 1956, L. Pena. Razowski
(1999) incorrect identifies the disposi-
tion of the type as NHMW.

spiraeifoliana Heinrich, 1923 (Polychros-
is); Proc. Entomol. Soc. Wash. 25: 106.
Holotype 6, USA, Pennsylvania, Luz-
erne Co., Hazelton, rf. Spiraea salici-
yolia, -5/29-19e%,

spirographa Diakonoff, 1968 (Bubonox-
ena); Bull. U.S. Natl. Mus. 257: 66. Ho-
lotype ¢, Philippine Islands, Luzon, Mt.
Makiling.

stadiana Barnes & Busck, 1920 (Peronea);
Contrib. Nat. Hist. Lepid. North Am. 4:
217. Holotype 6, Canada, Ontario, Ot-
tawa, 18 September 1905, C. Young.

stauroma Diakonoff, 1968 (Polemograp-
tis); Bull. U.S. Natl. Mus. 257: 33. Ho-
lotype 6, Philippine Islands, Negros
Oriental, Canlaon, 915 m, light trap, 20
December 1959, L. Quate.

stellifera Kawabe, 1978 (Hiroshiinoueana);
Tinea 10: 181. Holotype ¢, Japan, Yak-
ushima Island, Kurio, 7 August 1972, T.
Watanabe.

stenampyx Diakonoff, 1982 (Ancylis); Zool.
Verhandel. (Leiden) 193: 64. Holotype
36, Sri Lanka, Kandy District, Udawat-
tekelle, 3 November 1966.

stenotes Clarke, 1976 (Alcina); Insects of
Micronesia 9: 21. Holotype 3, Micro-
nesia, Guam, Tenjo, rf. Styphelia, 16
May 1936, R. Usinger.

stevensi Powell, 1980 (Decodes); Pacific
Insects 22: 90. Holotype ¢, USA, Col-
orado, Larimer Co., Owl Canyon, 12
km NW Fort Collins, June 1977, R. Ste-
vens.

straminoides Grote, 1873 (Conchylis); Bull.
Buttalo Soc. Natl Sek (ll66 Type od;
USA, New York, Buffalo. According to

1061

Fernald’s unpublished list, two Grote
specimens of this species were received
by the USNM; we found only one. The
single Grote specimen cited above is
clearly labeled as the type.

strianus Fernald, 1905 (Archips); Canad.
Entomol. 37: 399. ““Type’’ 6, Canada,
Quebec, A. Hanham. Fernald mentions
3 specimens in his description: a male
from London, Ontario; a female from
New Hampshire; and a male from Que-
bec. He indicated that the former two
were in his personal collection and the
last was in the USNM.

striatana Brown, 1999 (Dimorphopalpa);
Pan-Pac. Entomol. 75: 88. Holotype ¢,
Venezuela, Aragua, Rancho Grande,
1,100 m, 21—25 January 1966, S. & W.
Duckworth.

striatanoides Brown, 1999 (Dimorphopal-
pa); Pan-Pac. Entomol. 75: 88. Holo-
type 6, Ecuador, Carci, Maldonado,
2,200 m, 6—11 January 1963, V. Becker.

strigatella Brown, 1992 (Catastega); J. N.
Y. Entomol. Soc. 100: 216. Holotype d,
USA, Texas, Brewster Co., Big Bend
National Park, Chisos Basin, 29 July
1982, E. Knudson.

strigosa Heinrich, 1926 (Aphania); Bull.
U.S. Natl. Mus. 132: 118. Holotype <4,
USA, Alaska, Dawson, 16 June 1916,
B. Clark.

strigulana Brown & Obrazstov, 1991 (Dor-
ithia); in Brown and Powell, Univ. Ca-
lif. Publ. Entomol. 111: 46. Holotype &,
Mexico, Veracruz, Las Vegas [ = Las
Vigas], 8,000’, 1887, W. Schaus.

stygiana Dyar, 1903 (Thiodia); Proc. Ento-
mol. Soc. Wash. 5: 230. Five syntypes
(3 66,2 22), USA, Colorado, Jeffer-
son Co., Golden, May—June, Dyar &
Caudell; syntype 6, USA, Arizona,
Williams. This species was described
from the six specimens cited above.
One male from Colorado was placed in
the type collection, presumably by J.
Clarke; the other specimens are in the
main collection.

suadana Heinrich, 1923 (Eucosma); Bull.

1062

U.S. Natl. Mus. 123: 130. Holotype 6,
USA, Utah, Vineyard, 6 August 1912,
T. Spalding.

subcandida Heinrich, 1929 (Thiodia for-
mosana); Proco\U:S: Nath Mus...75;: 1.
Holotype ¢, Canada, Alberta, Bilby, 12
June 1924, O. Bryant.

subditiva Heinrich, 1929 (Eucosma tahoen-
sis): sProc: US: Nath Muss 75219» Ho-
lotype d6, USA, California, Lake Tahoe,
Deer Park Springs, “‘June 24-30.”

subelectana Kawabe, 1976 (Olethreutes);
Tinea 10: 46. Holotype ¢, Japan,
Ohmaki, Toyama Pref., 3 June 1951, M.
Hoshino. Although the specimen is la-
beled “Olethreutes pseudoereciana,”’ it
is clearly the specimen referred to in the
original description.

subflavana Walsingham, 1879 (Paedisca);
Ill. Lepid. Heter. Brit. Mus. 4: 48. Syn-
type d, USA, California. According to
the original description, this species was
described from two males and one fe-
male from Rouge River, Oregon, end of
May 1871. The USNM specimen bears
a typical Walsingham type label and is
presumed to be part of the original se-
ries; we cannot explain the discrepancy
in the locality data.

subminimana Heinrich, 1923 (Thiodia);
Bull. U.S. Natl. Mus. 123: 61. Holotype
3, USA, California, San Diego Co., San
Diego, ““Aug 1-7.”

subolivata Clarke, 1955 (Orthocomtis);
Trans. Royal Entomol. Soc. London
107: 148. Holotype 6, Costa Rica, Car-
tago Province, Tuis, 5,800’, 28 August
1908, W. Schaus.

subretracta Kawabe, 1976 (Olethreutes);
Tinea 10: 45. Holotype 6, Japan, To-
kyo, Mt. Takao, 11 June 1952, M.
Hoshino & T. Haga.

subsolana Miller, 1978 (Larisa); J. Lepid.
Soc. 32: 258. Holotype 6, USA, Arkan-
sas, Washington Co., Devil’s Den State
Park, 26 June 1966, R. Hodges.

subretiniana Obraztsov, 1962 (Choristo-
neura); Am. Mus. Novit. 2101: 9. Ho-
lotype 6, USA, California, Tulare Co.,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Monachee Meadows, 8,000’, “July 16—
23.7

substitutionis Heinrich, 1923 (Gypsonoma);
Bull. U.S. Natl. Mus. 123: 163. Holo-
type d, Canada, Manitoba, Aweme, 27
July 1905, Criddle.

subtropica Miller, 1960 (Rhyacionia); J.
Lepid. Soc. 14: 231. Holotype 2, USA,
Florida, Okaloosa Co., Valparaiso, rf.
Pinus, em: 26 May 1927, E. Gemner.

subviridis Heinrich, 1929 (Epinotia); Proc.
U.S. Natl. Mus. 75: 15. Holotype <d,
USA, California, San Diego Co., San
Diego, 5 July 1921 [genitalia slide only;
pinned specimen has not been found].
A paratype specimen is also in the type
collection.

sugii Kawabe, 1976 (Epiblema); Tinea 10:
42. Holotype ¢6, Japan, Tokyo, Seta-
gaya, 16 June 1963, A. Kawabe.

sugii Kawabe, 1989 (Dichrorampha); Tinea
12: 196. Holotype d, Taiwan, Hualien
Hsien, Kuanyuan, 2,400 m, 7-8 August
1987, A. Kawabe.

sulawesiensis Kawabe, 1993 (Bathypluta);
Tyo to Ga 43: 237. Holotype 2, N Su-
lawesi, Tandano, December 1988.

sulphurica Razowski, 1999 (Netechma);
Polski Posmo Entomol. 68: 95. Holo-
type 2, Brazil, Santa Catarina, 21 Jan-
uary 1936, EK Hoffman. Razowski
(1999) incorrect identifies the disposi-
tion of the type as NHMW.

superba Razowski, 1988 (Clarkenia); Acta
Zool. Cracov. 31: 406. Holotype 3, Co-
lombia, Cauca, Paramo de Parace, Lake
San Rafael, 3,570 m, 27 January 1959,
J. Clarke.

symphyla Razowski, 1984 (Polyortha);
Acta Zool. Cracov. 27: 222. Holotype
3, Bolivia, Cochabamba, Incachaca, J.
Steinbach.

synchysis Pogue, 1986 (Apolychrosis); in
Cibrian-Tovar et al., Cone and Seed In-
sects of the Mexican Conifers: 27. Ho-
lotype 6, Mexico, Tlaxcala, Municipio
de Terrenate, Villareal, rf., Abies reli-
giosa, em: 10 January 1985.

synneurana Barnes & Busck, 1920 (Amor-

VOLUME 102, NUMBER 4

bia); Contrib. Nat. Hist. Lepid. North
Am. 4: 211. Holotype 2, USA, Arizo-
na, Redington.

taedana Miller, 1978 (Petrova); Ann. En-
tomol. Soc. Am. 71: 337. Holotype °,
USA, Florida, Seminole Co., Geneva,
21 May 1967, R. Leuschner.

taetera Razowski, 1984 (Histurodes); Acta
Zool. Cracov. 27: 214. Holotype 6,
Guatemala, Cerro Zunil, 4,000—5,000’,
Champion.

tahoensis Heinrich, 1923 (Eucosma); Bull.
U.S. Natl. Mus. 123: 112. Holotype <6,
USA, California, Lake Tahoe, Deer
Park Springs, “July 8—15.”

taiwana Kawabe, 1986 (Zeiraphera); En-
tomol. Pap. Pres. Kurosawa, Tokyo: 79.
Holotype ¢3, Taiwan, Chiai Hsien, Al-
ishan, 2,200 m, 12 August 1974, Y.
Kishida.

taiwana Kawabe, 1992 (Acleris); Tinea 13:
174. Holotype 6, Taiwan, Nantou
Hsien, Tayuling, forest, 2,500 m, 9-18
June 1980, D. Davis.

taiwana Kawabe, 1992 (Neopotamia cryp-
tocosma); Japan. Hetero. J. 170: 359.
Holotype 6, Taiwan, Hualien Hsien,
Kuanyuan, 2,400 m, 7-8 August 1987,
A. Kawabe.

taiwanensis Kawabe, 1985 (Archips); Tinea
12: 3. Holotype ¢, Taiwan, Nantou
Hsien, Lushan Spa, 1,200 m, 27-29
July 1983, A. Kawabe.

taiwanica Razowski, 1977 (Aethes cni-
cana); Tyo to Ga 28: 37. Holotype 6,
Taiwan, Tattaka, 6 June 1943, S. Issiki.

taleana Grote, 1878 (Grapholitha); Canad.
Entomol. 10: 54. Holotype d (only a
forewing), USA, Illinois, T. Bean. The
original description appears to have
been based on a single male specimen.

tamaensis Kawabe, 1974 (Epinotia); Tyo to
Ga 25: 96. Holotype 6, Japan, Tokyo,
Tama Hill, 22 April 1958, A. Kawabe.

tambomachaya Razowski, 1988 (Seticosta);
Acta Zool. Cracov. 31: 401. Holotype
2, Peru, Cusco, Tambomachya, 3,690
m, 3 February 1959, J. Clarke.

1063

tantilla Heinrich, 1926 (Satronia); Bull.
U.S. Natl. Mus. 132: 17. Holotype 6,
USA, Florida, Alachua Co., Archer, 4
May 1882.

tayulingensis Kawabe, 1986 (Dichroram-
pha); Entomol. Pap. Pres. Kurosawa,
Tokyo: 83. Holotype ¢, Taiwan, Hu-
alien Hsien, Tayuling, 2,560 m, 2—3 Au-
gust 1983, A. Kawabe.

taxifoliella Busck, 1914 (Evetria); Proc.
Entomol. Soc. Wash. 16: 146. Holotype
2, USA, Montana, Missoula Co., hill at
canyon east of Missoula, rf. P. taxifolia,
24 April 1913, J. Brunner.

telesocia Razowski, 1994 (Cylichneulia);
SHILAP Revta. Lepid. 22: 68. Holotype
3, Venezuela, Aragua, Rancho Grande,
1,100 m, 16—19 January 1966, S. & W.
Duckworth.

temerana Busck, 1907 (Phalonia); J. N. Y.
Entomol. Soc. 15: 28. Holotype ¢d,
USA, Pennsylvania, Allegheny Co.,
Oak Station, 21 May 1906, E Marloff.

tenebrica Heinrich, 1929 (Anchylopera);
Proc. U.S. Natl. Mus. 75: 17. Holotype
6, Canada, Alberta, Bilby, 12 June
1924, O. Bryant.

tenebrica Razowski, 1987 (Terinebrica);
Tinea 12(suppl:): 135. Holotype 6,
Peru, Cusco, Machu Picchu, 2,385 m, 6
February 1959, J. Clarke.

terminana Busck, 1907 (Hysterosia); J. N.
Y. Entomol. Soc. 15: 33. Holotype 6,
USA, Pennsylvania, Allegheny Co.,
Pittsburgh, H. Engel.

tetropsis Busck, 1913 (Eucosma); Insect.
Inscit. Menst. 1: 91. Holotype 6, British
Guiana, Kirby, rf. guava, H. Moore.

teutoniana Brown, 1999 (Dimorphopalpa);
Pan-Pac. Entomol. 75: 89. Holotype 6,
Brazil, Santa Catarina, Nova Teutonia,
27°11'S, 52°23'W, 300-500 m, Septem-
ber 1963, EK Plaumann.

texasana Blanchard & Knudson, 1984 (An-
opina); Proc. Entomol. Soc. Wash. 86:
449. Holotype 6, USA, Texas, Jeff Da-
vis Co., Mt. Locke, 6,700’, 26 April
1981, E. Knudson.

thelmae Diakonoff, 1968 (Capua); Bull.

1064

U.S. Natl. Mus. 257: 23. Holotype ¢,
Philippine Islands, Mindanao, Bukid-
non, Mt. Katanglad, 1,480 m, 27-31
October 1959, L. Quate & C. Yashi-
moto. Although the holotype is labeled
“Capua thalmae,”’ the original descrip-
tion uses the correct spelling of “‘thel-
mae.”

theodora Clarke, 1976 (Herpystis); Insects
of Micronesia 9: 76. Holotype 36, Mi-
cronesia, Kusaie, Mutunlik, 22 m, 24
January—30 April 1953, J. Clarke.

theta Brown, 1991 (Punctapinella); Los
Angeles Co. Mus. Contrib. Zool. 423:
4. Holotype @, Venezuela, Aragua,
Rancho Grande, 1,100 m, 15—21 August
1967, R. Poole.

thylacophora Diakonoff, 1968 (Peridaeda-
la); Bull. U.S. Natl. Mus. 257: 76. Ho-
lotype <6, Philippine Islands, Davao
Province, Mindanao, E. slope Mt. Mc-
Kinley, 17 September 1946, CNHM
Philippine Zool. Exped., H. Hoogstraal.

tocullionana Heinrich, 1920 (Eucosma);
Proc. U.S. Natl. Mus. 57: 59. Holotype
3, USA, Connecticut, Lyme, rf. cones
of Picea, em: 16 May 1916, A. Cham-
plain.

tokui Kawabe, 1974 (Didrimys); Tyo to Ga
25: 101. Holotype ¢, Japan, Yakushima
Island, Aiko-dake, 30 March 1972, T.
Watanabe.

tokui Kawabe, 1974 (Eudemopsis); Kontyu
42: 389. Holotype 6, Japan, Yakushima
Island, Mt. Aiko-dake, 8 June 1972, T.
Watanabe.

tonsilis Razowski, 1984 (Ardeutica); Acta
Zool. Cracov. 27: 217. Holotype 4,
Costa Rica, Cartago Province, Juan Vi-
nas.

tonto Powell, 1980 (Decodes); Pacific In-
sects 22: 93. Holotype 2, USA, Arizo-
na, Pinal-Gila Co. line, Miami Superior
Highway, 4,790’, 1-15 April 1925, O.
Poling.

toreuta Grote, 1873 (Penthina); Bull. Buf-
falo Soc. Nat. Sci. 1: 92. Holotype ¢,
USA, Pennsylvania.

tornosema Clarke, 1968 (Phalonia); Proc.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

U.S. Natl. Mus. 125: 12. Holotype 9,
Guatemala, Volcan Santa Maria, June,
Schaus and Barnes.

torostoma Clarke, 1972 (Laspeyresia);
Proc. Entomol. Soc. Wash. 74: 467. Ho-
lotype 2, Costa Rica, Cartago Province,
Turrialba, r.f. string bean stems, 8
March 1965, L. Bonnefil.

toshimai Kawabe, 1974 (Eudemopsis);
Kontya 42: 389. Holotype ¢, Japan,
Kochi Pref., Mt. Ashizuriyama, Cape
Ashizuri, 27 May 1973, H. Toshima.

toshimai Kawabe, 1978 (Griselda); Tinea
10: 186. Holotype 3, Japan, Shikoku,
Kagawa Pref., 15 July 1972, T. Toshi-
ma.

towadaensis Kawabe, 1978 (Statherotis);
Tinea 10: 176. Holotype 3, Japan, Hon-
shu, Aomori Pref., Towada, 3 August
1969, Sato & Oshima.

trema Brown & Obraztsov, 1991 (Cuprox-
ena); in Brown and Powell, Univ. Calif.
Publ. Entomol. 111: 56. Holotype 6,
Mexico, Guerrero, December 1920, R.
Muller.

triangulana Brown, 1992 (Catastega); J. N.
Y. Entomol. Soc. 100: 220. Holotype 6,
USA, Arizona, Gila Co., 4 mi ESE Pine,
4,500 ft, 1 September 1961, R. W.
Hodges.

triangulana Kearfott, 1908 (Tortrix); J. N.
Y. Entomol. Soc. 16: 179. Lectotype <6,
USA, California, San Diego Co., San
Diego, 31 July 1907. Designated by Ob-
raztsov (1962).

tricia Clarke, 1976 (Heleanna physalodes);
Insects of Micronesia 9: 17. Holotype
6, Micronesia, Kapingamarani Atoll,
Hare Island, rf. Callophyllum inophyl-
lum, 3 August 1946, Townes.

tridens Razowski, 1988 (Seticosta); Acta
Zool. Cracov. 31: 400. Holotype 3, Co-
lombia, Cauca, Paramo de Parace, Lake
San Rafael, 3,570 m, 27 January 1959,
J. Clarke.

trifida Adamski, 1986 (Apotomis); Canad.
Entomol. 118: 666. Holotype ¢, USA,
Washington, Ferry Co., Sherman Pass,
23 July 1962, J. Clarke.

VOLUME 102, NUMBER 4

triopis Diakonoff, 1959 (Cryptaspasma);
Zool. Verhandel. (Leiden) 43: 32. Ho-
lotype ¢, Guam Island, Commander
Mariana’s Hill, March 1949, K. Maeh-
ler.

triphera Brown & Obraztsov, 1991 (Cu-
proxena); in Brown and Powell, Univ.
Calif. Publ. Entomol. 111: 56. Holotype
6, Panama, Cabima, May 1911, A.
Busck.

triquetra Obraztsov, 1964 (Proeulia); Proc.
U.S. Natl. Mus. 116: 189. Holotype <d,
Chile, Nuble, Chillan, 10 November
1961, H. Lister.

tristriata Kearfott, 1907 (Sparganothis);
Trans. Am. Entomol. Soc. 33: 67. Syn-
type ¢, USA, Minnesota, Duluth. Un-
published lectotype selected by R. Lam-
bert, labeled by J. Powell. Klots (1942)
indicated that there was a female “‘co-
type’’ in the AMNH, but did not des-
ignate it as the lectotype.

tsugae Issiki, 1961 (Epagoge); in Issiki and
Mutuura, Microlepidoptera injurious to
coniferous plants in Japan: 36. Holotype
6, Japan, Osaka, Ikeda Pref., 23 May
1958, T. Kodama. Komai (1999) unnec-
essarily designated a lectotype for this
species; apparently he was unaware that
the holotype was deposited in the
USNM.

tsuifengana Kawabe, 1992 (Acleris); Tinea
13: 175. Holotype 3, Taiwan, Nantou
Hsien, Tsuifeng, 2,400 m, 29 December
1989, A. Kawabe.

tsushimaensis Kawabe, 1978 (Hedya); Tin-
ea 10: 173. Holotype ¢, Japan, Tsush-
ima Island, Nenbutsuzaka, 27 June
1973, T. Watanabe.

turbula Clarke, 1968 ([razona); Proc. U.S.
Natl. Mus. 125: 40. Holotype °, Gua-
temala, Volcdn Santa Maria, July,
Schaus and Barnes.

turialba Busck, 1920 (Hysterosia); Insect.
Inscit. Menst. 8: 86. Holotype ¢, Costa
Rica, Cartago Province, Juan Vinas,
February, W. Schaus.

twila Clarke, 1955 (Orthocomotis); Trans.
Royal Entomol. Soc. London 107: 152.

1065

Holotype 6, Brazil, Santa Catarina,
Nova Teutonia, February 1948, F Plau-
mann.

ulteriorana Heinrich, 1920 (Evetria); Proc.
U.S. Natl. Mus. 57: 55. Holotype <4,
USA, Oregon, Waldo, r.f. Psuedotsuga
taxifolia, 16 August 1914, Keen & Ser-
gent.

umbrana Barnes & Busck, 1920 (Spargan-
othis); Contrib. Nat. Hist. Lepid. North
Am. 4: 212. Holotype 6, USA, Colo-
rado, Adams Co., Denver, Oslar.

umbraticana Heinrich, 1923 (Thiodia);
Bull. U. S. Natl. Mus. 123: 70. Holo-
type 6, USA, Colorado, Jefferson Co.,
foothills above Golden, 13 March 1901,
Dyar & Caudell.

unciana Dognin, 1904 (Atteria); Ann. Soc.
Entomol. Belgium 48: 133. Holotype °,
Ecuador, Zamora, September 1886.

undata Walsingham, 1914 (Tortrix); Biol.
Centr.-Am., Lepid., Heter. 4: 286. Ho-
lotype 6, Mexico, Puebla, Popocatepetl
Park, 8,000’, June 1906, W. Schaus.

unguiculus Clarke, 1951 (Episimus); J.
Wash. Acad. Sci. 41: 297. Holotype °,
Argentina, Tigre, March 1946, E Bour-
quin.

unica Heinrich, 1923 (Epinotia); Bull. U.S.
Natl. Mus. 123: 221. Holotype ¢, USA,
Tennessee, Knox Co., Knoxville, rf.
Bradburya virginiana, G. Ainslie.

urbana Busck, 1912 (Tortrix); Proc. Ento-
mol. Soc. Wash. 14: 86. Holotype <6,
Mexico, Mexico City, November 1911,
R. Muller.

ustulatana Blanchard & Knudson, 1983
(Eucosma); Proc. Entomol. Soc. Wash.
85: 849. Holotype 6, USA, Texas,
Washington Co., Brenham, 4 June 1979,
E. Knudson.

usuiana Kawabe, 1976 (Epibactra); Tinea
10: 39. Holotype 6, Japan, Gunma
Pref., Usui Pass, 5 June 1951, M. Hosh-
ino.

usurpata Razowski, 1987 (Utrivalva); Bull.
Polish Acad. Sci. 35: 67. Holotype 6,

1066
Guatemala, Volcan Santa Maria, W.
Schaus.

uta Clarke, 1953 (Eucosma); J. Wash.
Acad. Sci. 43: 226. Holotype 3, USA,
Illinois, Putnam Co., 10 July 1938, M.
Glenn.

vaccinii Clarke, 1971 (Dichelopa); Smith-
sonian Contrib. Zool. 56: 107. Holotype
3, Rapa Island, Maugaoa, 950’ (292
m), rf. Veronica rapensis, 5 November
1963, J. & T. Clarke.

vagana Heinrich, 1923 (Epinotia); Bull.
U.S. Natl. Mus. 123: 230. Holotype ¢,
USA, Washington, Liaga, rf. Pyrus ri-
vularis [wild crab apple], 20—24 July
1918, em: 20 July 1918, E. Heckard.

variana Fernald, 1886 (Teras); Bull. U.S.
Dept. Agric. Entomol. 12: 17. ““Type”’
3, USA, Maine. This is the only ex-
ample of this species we could locate
from the Fernald Collection. Fernald
did not indicate how many specimens
he examined.

varnicosa Brown, 2000 (Lobogenesis);
Proc. Entomol. Soc. Wash. 102: 36. Ho-
lotype 2, Argentina, Tucuman, Ciudad
Universitaria, 17 February 1959, J.
Clarke.

venatana Kawabe, 1992 (Acleris); Tinea
13: 178. Holotype 6, Taiwan, Nantou
Hsien, Tsuifeng, 2,400 m, 29 December
1989, A. Kawabe.

verecundana Blanchard, 1979 (Phaneta); J.
Lepid. Soc. 33: 210. Holotype ¢, USA,
Texas, Hemphill Co., Canadian, 15 Au-
gust 1971, A. & M. Blanchard.

versicolor Powell, 1978 (Rhyacionia); in
Powell and Miller, U.S. Dept. Agric.,
Agric. Handb. 514: 30. Holotype dé,
USA, Arizona, Coconino Co., Fort Val-
ley, 7.5 mi NW Flagstaff, 29 June 1961,
R. Hodges.

vesta Clarke, 1968 (Phalonia); Proc. U.S.
Natl. Mus. 125: 14. Holotype ¢, Ven-
ezuela, Aragua, Rancho Grande, 1,100
m, 16—23 October 1966, S. & W. Duck-
worth.

victoriana Busck, 1921 (Cacoecia); Canad.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Entomol. 53: 278. Holotype ¢, Canada,
British Columbia, Victoria, 22 July
1920, E. Blackmore.

vietnamna Kawabe, 1993 (Cerace); Tyo to
Ga 44: 99. Holotype 2, North Vietnam,
Mt. Tam Dao, 20-31 July 1990, Mizun-
uma.

villana Busck, 1907 (Hysterosia); J. N. Y.
Entomol. Soc. 15: 34. Holotype 6,
USA, Colorado, Adams Co., Denver, 21
June 1905, Oslar.

violetana Kawabe, 1964 (Philedone); Tyo
to Ga 15: 3. Holotype ¢, Japan, Hon-
shu, Tokyo, Nippara, 16 September
1963, A. Kawabe.

virga Clarke, 1949 (Eulia); Acta Zool. Lil-
loana (Tucuman) 7: 585. Holotype <6,
Brazil, Santa Catarina, Nova Teutonia,
2711'S, 52°23’ W, February 194835E
Plaumann.

virginiana Busck, 1914 (Evetria); Proc. En-
tomol. Soc. Wash. 16: 145. Holotype @,
USA, Virginia, Fairfax Co., Falls
Church, rf. Pinus virginiana, em: 4
May 1914, C. Heinrich.

viridana Barnes & Busck, 1920 (Platyno-
ta); Contrib. Nat. Hist. Lepid. North
Am. 4: 213. Holotype 6, USA, Arizo-
na, Yavapai Co., Dewey, ““May 1-7.”

virifloscula Brown, 1991 (Cuproxena); in
Brown and Powell, Univ. Calif. Publ.
Entomol. 111: 67. Holotype 3, Vene-
zuela, Aragua, Rancho Grande, 1,100
m, 8-14 August 1967, R. Poole.

vitrana Walsingham, 1879 (Grapholitha)
Ill. Lepid. Heter. Brit. Mus. 4: 65. Syn-
type 6, USA, Oregon (northern), April
1872. According to the original descrip-
tion, this species was described from
four males; the other syntypes are pre-
sumed to be in the BMNH.

wellingtoniana Kearfott, 1907 (Olethreu-
tes); Trans. Am. Entomol. Soc. 33:13.
Lectotype 6, Canada, British Columbia,
Wellington, 19 May, G. W. Taylor. Des-
ignated by Blanchard & Knudson
(1984).

wielgusi Clarke, 1990 (Argyrotaenia); J.

VOLUME 102, NUMBER 4

Res. Lepid. 28: 97. Holotype 2, USA,
Arizona, Cochise Co., Huachuca Mts.,
Pueblo del Sol, 6 November 1986, R.
Wielgus.

willettana J. Comstock, 1939 (Carolella
busckana); Bull. South. Calif. Acad.
Sein 353 115. Holotype *6 “USA, (Cali-
fornia, Riverside Co., Chuckwalla
Springs, 25 December 1937, G. Willett.

worthingtoniana Fernald, 1878 (Paedisca);
Canad. Entomol. 10: 83. Lectotype <4,
USA, ‘North Illinois.”’ Designated by
Miller (1970).

wrighti Brown & Powell, 2000 (Anopina),
Univ. Calif. Publ. Entomol. 120: 98.
Holotype 6, USA, Colorado, Chaffee
Co., 2.5 mi ESE Buena Vista, 23 Au-
gust 1997, D. Wright.

yama Kawabe, 1974 (Olethreutes bipunc-
tana); Tyo to Ga 25: 102. Holotype 4,
Japan, Hokkaido, Mt. Daisetsu, 23 July
1959, M. Suzuki.

yasudai Kawabe, 1972 (Cryptophlebia);
Tinea 9: 242. Holotype 6, Japan, Sai-
tama Pref., Owa, 7 August 1952, T. Ha-
ruta.

yasutoshii Kawabe, 1985 (Acleris); Tinea
12: 9. Holotype 6, Taiwan, Taichung
Hsien, Lishan, 24 August 1970, Y. Shi-
bata.

yuccatana Busck, 1907 (Phalonia); J. N. Y.
Entomol. Soc. 15: 23. Holotype 2°,
USA, Texas, Nueces Co., Nuecestown,
Ei “Yucca baccata; W2»April 1897, C.
Marlatt.

yakushimensis Kawabe, 1974 (Notocelia),
Tyo to Ga 25: 96. Holotype ¢, Japan,
Yakushima Island, Onoaida, 7 October
1972, T. Watanabe.

zacualpana Busck, 1913 (Phtheochroa), In-
sect. Incit. Menst. Wash. 1: 141. Holo-
type 2, Mexico, Zacualpan, July 1913,
R. Muller.

zeteles Razowski, 1987 (Chicotortrix); Tin-
ea 12(suppl.): 124. Holotype 6, Colom-
bia, Bogota, Chico, 25 January 1959, J.
Clarke.

zimapanus Powell, 1980 (Decodes); Pacific

1067

Insects 22: 107. Holotype 2, Mexico,
Hidalgo, 3 mi E Zimapan, 6,400’, 31
July—1 August 1963, W. Duckworth &
D. Davis.

zona Clarke, 1986 (Dichelopa); Smithson-
ian Contrib. Zool. 416: 144. Holotype
36, Marquesas Islands, Nuku Hiva, Tap-
uaooa, 2,500’, 30 January 1968, J. & T.
Clarke.

ACKNOWLEDGMENTS

This list was begun by Brown in 1988
during a one-year Smithsonian Post-doctor-
al Fellowship, with the encouragement and
assistance of the late J. KE Gates Clarke.
From 1989 to about 1993 Clarke continued
to supply the names of type specimens as
they slowly surfaced from the USNM col-
lection or were returned by colleagues. Jack
had a strong ‘“‘collection-bias,’’ and we be-
lieve he would have been pleased to see the
USNM type collection thoroughly orga-
nized and catalogued.

We thank the following for helpful com-
ments on the list: William Miller, Univer-
sity of Minnesota, St. Paul; Eric Grissell
and David Smith, Systematic Entomology
Laboratory, USDA, National Museum of
Natural History, Washington, DC; Richard
Brown, Mississippi Entomological Muse-
um, Mississippi State University, Mississip-
pi; Kevin Tuck, The Natural History Mu-
seum, London, England.

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Brown, J. W. 1998. A new genus of tortricid moths
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. 1962. Anopina, a new genus of the Cnephas-

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na. Insects. British Museum, London. 482 pages.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1070-1071

NOTE

Fitchia aptera Stal (Hemiptera: Reduviidae):
Seasonal History and Habits in Mid-Appalachian Shale Barrens

Fitchia aptera Stal is an infrequently col-
lected though widely distributed harpactorine
reduviid. Adults are 9-14 mm long, elongate,
and dull yellow, with a dark longitudinal
stripe on either side of the abdominal mid-
line; this dorsal stripe is conspicuous in the
brachypterous morph. Slater and Baranowski
(1978. How to Know the True Bugs, W. C.
Brown, Dubuque, Iowa. 256 pp.) illustrated
the brachypterous and macropterous forms.
Fitchia aptera can be confused with the only
other member of this Nearctic genus, F. spi-
nosula Stal; characters traditionally used to
distinguish the two sympatric species—in-
cluding presence or absence of prothoracic
spines—are unreliable (McPherson et al.
1992. Florida Entomologist 75: 222-230).
McPherson et al. (1992) redescribed the
adults and provided additional characters to
separate these species.

DeCoursey (1963. Bulletin of the Brook-
lyn Entomological Society 58: 151—156)
described and illustrated the immature stag-
es and determined their duration by rearing
F. aptera in the laboratory (“‘room temper-
ature,’’ RH not stated); nymphs were fed
small flies and homopterans. DeCoursey
(1963) reported mean development (egg to
adult) of 60 days (range 51—67; n not stat-
ed) and found that F. aptera has only four
nymphal instars. Swadener and Yonke
(1973. Journal of the Kansas Entomological
Society 46: 123-136) reared this reduviid
in the laboratory (21°C night, 29°C day;
40—60% RH), reporting that total develop-
ment averaged 70 days (range 60.5—74.5; n
= 134 eggs, n = 3 instar IV). They also
confirmed the presence of only four instars.

In contrast to the availability of biological
data from laboratory studies, little is known
about the habits of F. aptera in nature. Rea-
dio (1927. University of Kansas Science

Bulletin 17: 5—291) did not treat the bionom-
ics of this species in his studies on reduviid
biology. This mainly ground-dwelling assas-
sin bug occurs by day at the bases of grass
clumps in old fields, the adults overwinter-
ing under boards, stones, or at roots of grass-
es. Adults occasionally can be swept from
tall grasses in early evening. Nymphs are
found on the ground, mainly in grassy areas
(DeCoursey 1963). Hagerty and McPherson
(2000. Great Lakes Entomologist 32: 133-—
160) reviewed additional collection records
and in Illinois reported an adult at the roots
of a grass clump, three adults from sweeping
herbaceous vegetation, and another adult
from sweeping a grassy field.

Here I give notes on the seasonality of
F. aptera in shale barrens and outcrops in
the mid-Appalachians. Collections and ob-
servations were made while inventorying
the plant bug (Hemiptera: Miridae) fauna of
moss phlox (Phlox subulata L.) during
1989-1995. Mats of this prostrate perennial
(family Polemoniaceae) were shaken over a
white tray (Wheeler. 1995a. Proceedings of
the Entomological Society of Washington
97: 435—451), and adults and all instars of
F. aptera were collected; nymphs were de-
termined to instar in the laboratory, using
DeCoursey’s (1963) descriptions. Voucher
specimens, including nymphs, are deposited
in the National Museum of Natural History,
Smithsonian Institution, Washington, D.C.

Unless otherwise stated, F. aptera was
collected from mats of moss phlox at the
following localities. The numbers of
nymphs (Roman numerals = instars) and
adults (A) are given parenthetically after
other collection data.

Maryland: Allegany Co., Country Club
shale barren, 2.5 km NE of Cumberland, 2
July 1994 (1 IID). Pennsylvania: Bedford

VOLUME 102, NUMBER 4

Co., shale slope, Rt 56 E of Fishertown, 24
June 1990 (1 I); shale slope, Rt. 96 NE of
Ryot, 24 June 1990 (2 I) & 10 May 1992
(1 A). Virginia: Bath Co., Fort Lewis shale
barren, Rt. 678 at Cowpasture River, 26 June
1994 (1 I); Highland Co., Head Waters shale
barren nr. jct. Rts. 250 & 616, 23 June 1990
(7 I-III), 25 Aug. 1990 (1 IV; 2 A ex undet.
sprawling plant), 10 July 1993 (3 II; 2 II ex
base of Eriogonum allenii S. Wats), 1 Aug.
1935021 tt), 26 June 1994 1.1.1,
6 Aug. 1994 (1 I, 1 ID, 26 Aug. 1994 (1
IV); Rockbridge Co., Furnace Mountain
shale barren, ca. 3 km S of Natural Bridge
Station, 24 May 1995 (1 I, 1A ex Antennaria
plantaginifolia (L.) Richardson); Rocking-
ham Co., shale slope, For. Rd. 87, W of
Fulks Run, 19 May 1994 (1 A; 1 A at base
of Tephrosia virginiana (L.) Pers.), 25 June
1994 (1 I); Shenandoah Co., Short Mountain
shale barren, ca. 4.6 km SE of Mount Jack-
son, 10 July 1993 (1 III). West Virginia:
Greenbrier Co., shale slope, Whites Draft
Rd., Alvon, 4 July 1994 (1 II); Hampshire
Co., shale bank, Rt. 29, 5.6 km N of Sla-
nesville, 30 June 1990 (1 II); Hardy Co.,
shale bank, Mathias, 30 June 1990 (1 ID).

Insect-plant associations in shale barrens
have been little studied (e.g., Wheeler
1995a; Braunschweig et al. 1999. pp. 83-
98. In Anderson, R. C. et al., eds., Savan-
nas, Barrens, and Rock Outcrop Commu-
nities of North America. Cambridge Uni-
versity Press, UK), but these xeric com-
munities of high soil-surface temperatures
(sometimes 50—60°C; Braunschweig et al.
1999) harbor great insect richness (Wheeler
1995a, b. Virginia Journal of Science 46:
148). The predacious F. aptera can be add-
ed to the diverse insect fauna of moss phlox
in mid-Appalachian shale barrens and out-
crops (Wheeler 1995a, b). All collections
were from the Valley and Ridge physio-
graphic province except the one in Rock-
bridge Co., Va., which was from the Blue
Ridge province. According to McPherson et
al. (1992), Maryland, Virginia, and West
Virginia are new state records.

Only small numbers of F. aptera were

1071

found during any 45—60 minutes’ sampling
of moss phlox, the seven early instars on 23
June 1990 representing the most individuals
in any sample. Adults, all brachypters (n =
6), were found at four sites where nymphs
were present; nymphs at other sites proba-
bly also were F. aptera rather than F. spi-
nosula.

Overwintered adults were found under
mats of moss phlox, at the base of Tephro-
sia virginiana, or among stolons of Anten-
naria plantaginifolia during May in shale
barrens and outcrops. Except for the late-
May collection in Rockbridge Co., Va., first
instars were not observed before late June
despite intensive sampling of moss phlox
during April, May, and early June (Wheeler
1995a). An early instar was observed feed-
ing on an immature isopod at the Head Wa-
ters shale barren in Virginia in late June.
Third and fourth instars were present from
early July to late August, suggesting a rath-
er late-season, univoltine life cycle. Two
adults found in late August likely were
those of a new, rather than overwintered,
generation. No nymphs or adults were ob-
served during sampling of moss phlox in
September and early October.

In Connecticut, DeCoursey (1963) re-
ported that overwintered adults of F. aptera
become active in April. An adult collected
in early April in Missouri did not oviposit
in the laboratory until mid-May and contin-
ued to lay eggs until mid-July (Swadener
and Yonke 1973). Such observations, cou-
pled with mean development of 60 (De-
Coursey 1963) or 70 days (Swadener and
Yonke 1973), also suggest that F. aptera is
a relatively late-season reduviid.

I thank Peter Adler (Clemson University)
for comments that improved the manu-
script, and Aaron Hagerty (Clemson Uni-
versity) and Jay McPherson (Southern Illi-
nois University) for allowing me to refer to
information in a paper in press.

A. G. Wheeler, Jr., Department of Ento-
mology, Clemson University, Clemson, SC
29634, U.S.A. (e-mail: awhlr@clemson.
edu)

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1072-1073

NOTE

First Host Plant and Parasitoid Record for Anastrepha spatulata Stone
(Diptera: Tephritidae)

Anastrepha spatulata Stone has been re-
ported in the U.S. (S. Texas), Mexico, Cen-
tral America (Guatemala S. to Panama),
Venezuela and Tobago (Norrbom et al.
1999. Myia 9: 65-251 [p. 82 for A. spatu-
lata]). In Mexico, it is commonly captured
in McPhail traps placed in commercial fruit
orchards (Aluja et al. 2000. Proceedings of
the Entomological Society of Washington
102: 802-815). In the past 60 years, there
have been considerable, but until now un-
successful, efforts to discover its host plant.
This species, together with A. alveata
Stone, A. alveatoides Blanchard, A. distans
Hendel, A. interrupta Stone, A. manihoti
Lima, A. montei Lima, A. pickeli Lima, and
A. umbrosa Blanchard, form the spatulata
species group (Norrbom et al. 2000. pp.
343-362. In Aluja and Norrbom, eds., Fruit
Flies (Diptera: Tephritidae): Phylogeny and
Evolution of Behavior, CRC Press).

We report here on A. spatulata infesta-
tions in Schoepfia schreberi Gmelin fruit
(Olacaceae). This bushy plant is locally
known in Veracruz as “guayabillo” or
‘“‘palo meco.”’ Fully ripe fruits measure, on
the average, 1.92 + 0.06 cm (diameter, n =
25) and weigh 185 + 0.061 mg (n = 25).
A mature plant measures ca. 2.10 m in
height. Floral buttons of S. schreberi were
observed from January 20th to February
Sth. The flowering period was recorded
from the Ist to the 12th of February and
the fruiting season occurred between the
10th and the 20th of February (all during
1998). Fly larvae were always found feed-
ing in the seed of the fruit (there is one seed
per fruit). There was always only one larva
per fruit. Given the small size of the host,
fully mature larvae, 8.6 + 0.12 mm (n =
8) in length, looked very compressed inside
the host. We collected 3 batches (different

collection dates) of ripe fruit from the
ground adjacent to four S. schreberi bushes.
A total of 583 g of fruit were collected on
February 2, 9, and 18, 1998. Fruit was han-
dled as described by Aluja et al. (2000).
Pupae were harvested between 1 and 6 days
after the fruit collecting date. The 583 g of
fruit yielded 579 pupae (0.993 pupae/g of
fruit). Mean pupal weight was 12 + 0.001
mg (n = 13). We divided pupae into two
batches. One was kept under field condi-
tions and another was transported to the
laboratory. The collection site was Llano
Grande, Municipio de Teocelo, Veracruz,
Mexico, at 720 m altitude (19°22’N and
26°53'W). Native vegetation around this
site is a tropical deciduous and subdeci-
duous forest (Castillo-Campos. 1995. Ecol-
ogia del Paisaje del Municipio de Jalco-
mulco, Veracruz, MS Thesis, UNAM, Mex-
ico City).

Between February 29 and March 5, 454
A. spatulata adults (223 females, 231
males), 17 Doryctobracon areolatus (Sze-
pligeti) (10 females, 7 males) and 2 Doryc-
tobracon toxotrypanae (Muesebeck) (2 fe-
males) emerged. We note that the D. areo-
latus individuals emerging from A. spatu-
lata were uniformly more darkly colored
than those individuals of the same species,
parasitizing, in the same region, A. alveata,
A. obliqua (Macquart), A. striata Schiner
and A. fraterculus (Wiedemann) (Lopez et
al. 1999. Biological Control 15: 119-129).
Length of the pupal period for flies was 22
+ 3 and 27 + 9 days under laboratory and
field conditions, respectively (mean tem-
perature of 26 + 1°C and 23 + 3°C under
laboratory and field conditions, respective-
ly). In the case of the parasitoids, the length
of the pupal period for D. areolatus was 25
+ 9 and 26 + 5 days under laboratory and

VOLUME 102, NUMBER 4

field conditions, respectively (same temper-
ature conditions as flies). For D. toxotry-
panae it was 21 days (laboratory conditions
only). Another 9 D. areolatus (8 females, 1
male) entered diapause and emerged after
ca. 11 months (range of 308-341 days).
Diapause by D. areolatus in the same re-
gion has been recently reported by Aluja et
al. (1998. Annals of the Entomological So-
ciety of America 91: 821-833).

Schoepfia schreberi had been previously
reported as a host plant of A. interrupta
(McClanahan. 1951. State Plant Board of
Florida Biennial Report No. 18, p. 44; as S.
chrysophylloides). Interestingly, the host
plant of another fly belonging to the same
species group (A. alveata), also belongs to
the family Olacaceae (Piedra et al. 1993.
Proceedings of the Entomological Society
of Washington 95: 127). Based on the few
host records for the spatulata group (Norr-
bom. 2000. Diptera Data Dissemination
Disk 2, in press), it appears that flies in this
assemblage of species are specialized on
only two plant families, Olacaceae and Eu-
phorbiaceae.

Our discovery of D. toxotrypanae para-
sitizing a species of Anastrepha is note-
worthy since this parasitoid had been pre-
viously reported parasitizing only T. curvi-
cauda Gerstaecker (Wharton and Marsh.
1978. Journal of the Washington Academy
of Sciences 68: 147-167). Toxotrypana
curvicauda is common in the study region
where it has been recently reported infest-
ing Gonolobus niger (Cav.) R. Br. (Ascle-
piadaceae) fruit (Aluja et al. 2000). It is
thus likely that the D. toxotrypanae individ-
uals parasitizing A. spatulata stemmed from
parasitized T. curvicauda. We note, how-
ever, that D. toxotrypanae is only distin-
guished from D. crawfordi on the basis of
color differences (D. toxotrypanae is dark-
er) (R. A. Wharton, pers. comm.). Based on

1073

the latter, the question remains open as to
which of the two above mentioned Doryc-
tobracon species was actually parasitizing
A. spatulata in our study region.

Voucher specimens of S. schreberi and A.
spatulata are placed in the herbarium and
insect collection of the Instituto de Ecolo-
gia, A.C. in Xalapa, Veracruz, Mexico. An-
astrephe spatulata individuals were also
sent to the National Museum of Natural
History, Washington, DC (A. L. Norrbom)
and D. areolatus and D. toxotrypanae in-
dividuals were sent to the parasitoid collec-
tion of Texas A&M University, College
Station, TX (R. A. Wharton).

Specimen identification was by the fol-
lowing: A. spatulata (A. L. Norrbom, Sys-
tematic Entomology Laboratory, ARS,
USDA, % National Museum of Natural
History, Washington, DC); D. areolatus
and D. toxotrypanae (Robert Wharton, Tex-
as A & M University, College Station), and
Schoepfia schreberi (Carlos M. Duran-Es-
pinoza, Instituto de Ecologia, A.C., Xalapa,
Veracruz, Mexico). Financial support was
provided by the Campania Nacional contra
las Moscas de la Fruta, Comisién Nacional
para el Conocimiento y Uso de la Biodiv-
ersidad (CONABIO) (Grant No. H296),
Sistema Regional de Ciencia del Golfo
(CONACyT-SIGOLFO) (Grant No. 96-01-
003-V) and the U.S. Department of Agri-
culture (ARS, USDA) (Grant Numbers
198-23 and 58-6615-3-025).

Martin Aluja, Emanuel Herrera, Maurilio
Lépez, and John Sivinski (MA, EH, ML)
Instituto de Ecologia, A.C., Apartado Post-
al 63, 91000 Xalapa, Veracruz, Mexico (e-
mail: alujam @ ecologia.edu.mx); (JS)
Center for Medical, Agricultural, and Vet-
erinary Entomology, Agricultural Research
Service, U.S. Department of Agriculture,
P.O. Box 14565, Gainesville, FL 32604,
U.S.A.

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, p. 1074

BoOoK REVIEW

Aphids on the World’s Crops, An Iden-
tification and Information Guide, Second
Edition, by R. L. Blackman and V. EF Ea-
stop. John Wiley & Sons, Baffins Lane,
Chichester, Sussex PO19 1UD, England.
466 pp. 2000. $145.00.

The first edition of Aphids on the
World’s Crops was published in 1984 and
brought together information available pre-
viously only in widely-distributed papers
and publications containing species’ de-
scriptions and keys. The information in the
keys, in the systematic reviews, and in the
photographs, will allow a non-specialist to
identify, with a good degree of accuracy,
aphids colonizing (feeding and producing
young) the crops treated.

The second edition has seven sections.
Section A (pages 1—24) contains general in-
formation on systematics, life cycles, host-
plant relationships, geographical distribu-
tion, and morphology and key characters.
Section B (pages 25-214) begins with a
statement on the limitations and use of the
keys and is devoted to crops and their
aphids. This section includes an alphabeti-
cal listing of the Latin names of crops with
their common names followed by an alpha-
betical listing of the crops by common
names, a list of the aphids reported to col-
onize the crop, and keys to those aphids.
Section C (pages 215-362) covers a dis-
cussion of the aphids and includes a brief
introduction followed by an alphabetical
listing by genus and species of the aphids
treated. This section is a wealth of infor-
mation on aphids and includes field char-
acters, host plants, virus transmission, dis-
tribution, and biology. Section D (pages
363-365) covers techniques on collecting,
preserving and mounting, and labeling and
storage. Section E (pages 367-373) con-
tains sources of information on regionally
classified faunal publications; general biol-

ogy; morphology, anatomy, and physiolo-
gy; genetics and development; migration
and dispersal; relationships with other in-
sects; host-plant relationships; control; and
bibliographies. Section F (pages 375—414)
is a list of references. Section G (pages
415—466) is a photographic guide to slide-
mounted aphids.

While both editions are 466 pages in
length, the second edition is about % larger
in size, covers 38 more species, and con-
tains more information in the keys and the
systematic section. While the basic format
is unchanged, the arrangement of the text
and the collection of the illustrations in
plates rather than their being shown with
each key couplet is a more efficient use of
space, is more pleasing to the eye, and does
not detract from the value of or ease in fol-
lowing the information. The biggest im-
provement, from a user’s standpoint, is that
the type in the section with the keys is larg-
er and more legible. The photomicrographs
and illustrations are much improved with
the target characters easier to see. Having
the names of the species with each photo-
graphic plate is useful, but I missed having
the listing of all species at the beginning of
the photographic guide.

Aphids on the World’s Crops remains the
only current, world-wide reference with
general and specific information on aphids
colonizing crops and is a must have for
anyone working in this field. Everyone who
has a copy of the first edition will want to
keep it as a reference (mine is well worn)
but will want to purchase a copy of the sec-
ond edition as a valuable working tool.

Manya B. Stoetzel, Systematic Entomol-
ogy Laboratory, PSI, Agricultural Research
Service, U.S. Department of Agriculture,
Bldg. 005, BARC-West, 10300 Baltimore
Avenue, Beltsville, MD 20705, U.S.A. (e-
mail: mstoetze @ sel.barc.usda.gov)

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1075-1076

Book REVIEW

Classification of Lepidoptera. Part 1. In-
troduction, by John B. Heppner. Holarc-
tic Lepidoptera, Volume 5, Supplement 1.
Published by the Association for Tropical
Lepidoptera, Gainesville, FL. 1998. 148
pp. + 6 pp. index. Soft bound, 8.5 X 11
in. ISBN 1070-4140. $25.00

This volume, the first of a proposed
three-part ‘“‘Manual of Lepidoptera,” is
chuck full of interesting factoids, keys, il-
lustrations, and references. It presents a
“new” higher classification of the order
with novel (and sometimes unusual) group-
ings of families based on the author’s ex-
pertise and biases. Included are a tabular
summary of Heppner’s (1991) previous
work on species richness and biogeographic
distribution of each of the superfamilies;
‘schematic phylogenies” of most of the
major groups (apparently based on intuition
rather than analysis); an impressive data
matrix (Table 2) of 24 morphological char-
acters of adults, larvae, and pupae across all
the superfamilies; an extensive key to
adults of the major families, previously
published by Heppner (1996); 33 pages of
illustrations of wing venation of all major
groups (never before compiled and pre-
sented in such an easily browsed manner);
and references, lots of them, subdivided by
category (e.g., taxonomy, morphology, bi-
ology, biogeography). The volume brings
together widely scattered concepts and lit-
erature, synthesized in Heppner’s unmistak-
able style.

If this book had been published 10 years
ago, it might have stood for a decade as the
standard reference on the order. Unfortu-
nately, any impact that it potentially may
have had likely will be overshadowed by
the recently published Handbook of Zool-
ogy (Kristensen 1999). Heppner’s timing
could not have been worse. In contrast to
Heppner’s single-authored manual, the edi-
tor of the Handbook enlisted the assistance

of 29 leading authorities in particular Lep-
idoptera families, creating a work of grand
proportions.

Aside from being scooped by the Hand-
book, Heppner’s work is plagued by a few
quirky shortcomings and/or inconsistencies.
Based on my knowledge of Tortricoidea (I
have limited knowledge of the higher clas-
sification of the entire order), I perused the
volume for details of this superfamily. Ta-
ble 1 indicates that it includes 6,683 spe-
cies, with 250 species occurring in the Ethi-
opian (Afrotropical) region. Actually, there
are over 8,000 described species of Tortri-
cidae (based on a checklist I have com-
piled), and Diakonoff, in various publica-
tions (e.g., Diakonoff 1960, 1970, 1973,
1981, 1983), treated over 250 species of
tortricids from Madagascar alone. Also, of
ten genera of Tortricidae for which wing
venation is illustrated (page 78), Heppner
uses older, antiquated names for two: *“‘Tor-
trix’ for Xenotemna (X. pallorana) and
‘““Clysiana” for Eupoecilia (E. sp.). Given
these minor errors in the group with which
I am most familiar, I am skeptical about the
quality of information presented for other
superfamilies.

In some cases, Heppner appears to have
disregarded the published findings of other
specialists. For example, Epstein’s (1996)
analysis of the limacodid-group of families
provides compelling evidence that Megal-
opygidae, Aididae, Dalceridae, and Lima-
codidae form a monophyletic group within
Zygaenoidea. Yet in Heppner’s classifica-
tion, Megalopygidae (including Aidinae) is
placed in Zygaenoidea, while Dalceridae
and Limacodidae are included in Cosso-
idea.

Despite its incongruities and shortcom-
ings, Heppner’s compendium is a valuable
source of information on the classification
of Lepidoptera, and comparisons with the
Handbook of Zoology are premature since

1076

parts 2 and 3 of Heppner’s work presum-
ably will present morphological and biolog-
ical details on each of the superfamilies,
one of the strengths of the Handbook. Com-
parisons with the Handbook also are unfair
given the considerable difference in price—
Heppner’s part 1 sells for about $25.00; the
Handbook goes for a whopping $250! Be-
cause currently proposed classifications of
the Lepidoptera exhibit little consensus, all
must be viewed as viable and reasonable
alternative interim solutions. Heppner’s
contribution will not go unnoticed.

LITERATURE CITED

Diakonoff, A. 1960. Tortricidae from Madagascar. Part
I. Tortricinae and Chlidanotinae. Verhandelingen
der Koninkluke Nederlandse, Akademie van We-
temschappen, Afd. Natuurkunde 53(1): 1—209.

. 1970. Lepidoptera Tortricoidea from Tsaratan-

ana Range [North Madagascar]. Mémoires

O.R.S.T.O.M. 37: 103-150.

. 1973. Tortricidae of the Andringitra Range,

Central Madagascar (Lepidoptera). Part 1. Tortri-

cinae. Bulletin de Muséum National d’ Historie

Naturelle, no. 108, Zoologie 82: 105-143.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

. 1981. Tortricidae from Madagascar, Part 2.
Olethreutinae, 1. Annales de la Société Entomo-
logique de France (new series) 17: 7—32.

. 1983. Tortricidae from Madagascar. Part 2.
Olethreutinae, 2. Annales de la Société Entomo-
logique de France (new series) 19: 291—310.

Heppner, J. B. 1991. Faunal regions and the diversity
of Lepidoptera. Tropical Lepidoptera (Gaines-
ville), Vol. 2, supplement 1. 85 pp.

. 1996. Keys to Families of Lepidoptera. Trop-
ical Lepidoptera (Gainesville), Vol. 4, supplement
3. 28 pp.

Epstein, M. E. 1996. Revision and Phylogeny of the
Limacodid-group Families, with Evolutionary
Studies on Slug Caterpillars (Lepidoptera: Zyg-
aenoidea). Smithsonian Contributions to Zoology
582. 102 pp.

Kristensen, N. P., ed. 1999. Handbook of Zoology, Vol.
IV Arthropoda: Insects, Part 35 Lepidoptera,
Moths and Butterflies. Walter de Gruyter, Berlin.

John W. Brown, Systematic Entomology
Laboratory, PSI, Agricultural Research
Service, U.S. Department of Agriculture,
c/o National Museum of Natural History,
Washington, DC 20560-0168, U.S.A. (e-
mail: jbrown@ sel.barc.usda.gov)

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1077-1087

OBITUARY

Clyde Fuhriman Smith
(1913-2000)

Clyde Fuhriman Smith enjoyed a distin-
guished, multi-faceted career in entomolo-
gy, with an enduring interest in the taxon-
omy of aphids and their host-plant associ-
ations. He was born 10 August 1913, in
Franklin County, Idaho, where his early ed-
ucation began. Smith went on to study en-
tomology, earning degrees at Utah State
Agricultural College (now Utah State Uni-
versity: B.S. 1935, M.S. 1938: George FE
Knowlton, major advisor) and the Ohio
State University (Ph.D. 1939: Alvah Peter-
son, major advisor). His entomological ed-

ucation extended far beyond course work
with part-time or short, full-time periods as
research assistant, teaching assistant, exten-
sion assistant, and research fellow, expos-
ing him to a wide range of insects and
crops.

In September 1939, Smith accepted a po-
sition in Raleigh as an Assistant Entomol-
ogist at the North Carolina Agricultural Ex-
periment Station and N.C. State College
(now University) and subsequently pro-
gressed to Associate Professor (1943-
1950), Professor and Department Head

1078

(1951-1963), Professor (1964-1979), and
Professor Emeritus (1980—2000). Smith
worked on insects and mites associated
with nearly every major crop plant in North
Carolina as documented by his publications
(listed below). Under his leadership, the
Department of Entomology (initially a sec-
tion of the Division of Biological Sciences)
more than doubled in number of faculty.
After his tenure as head, he returned to re-
search on cucumber and peach insects, no-
tably the management of the pickleworm, a
complex problem requiring intensive and
complete control. As a professor emeritus,
he continued publishing on aphid system-
atics and served on the NCSU Insect Col-
lection’s Museum Council.

Stretching far beyond his microscope and
boxes of prepared specimens, Smith’s lab-
oratory embraced North Carolina’s or-
chards, forests, farms, gardens, pest-infest-
ed buildings, and roadside vegetation. He
excelled at working with and gaining the
trust of farmers and orchardists, talents vital
to conducting meaningful scientific re-
search in the days before university-sup-
ported experimental research farms.

Smith had an extraordinary ability to
bring together scientific colleagues and oth-
ers to exchange information. The present
names of organizations he founded or
helped to found are the Southeastern Peach
Workers Conference (initiated 1948), N.C.
Agricultural Chemicals School (1949), N.C.
Pest Control Technicians School (1950),
N.C. Pest Control Association (1950), N.C.
Entomological Society (1956, first Presi-
dent), and Heliothis Conference (late 1950s,
now meeting with the Entomological So-
ciety of America). In 1972, Smith received
the N.C. Entomological Society’s Entomol-
ogist of the Year award. Furthermore, he
held membership in several other scholarly
societies: Entomological Society of Amer-
ica (President of Southeastern Branch in
1972), Entomological Society of Washing-
ton, Society of Systematic Zoology, Geor-
gia Entomological Society, Gamma Sigma
Delta, Society of the Sigma Xi, and Phi

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Kappa Phi. Smith was also instrumental in
the passage of the N.C. Structural Pest Con-
trol Act of 1955 and served as chair of the
N.C. Structural Pest Control Commission
from 1955 to 1967.

Internationally known as a leading re-
searcher on the systematics of aphids and
their parasites, Smith deposited his large re-
search collection of aphids at the NCSU In-
sect Collection. It is rich in material from
North Carolina and Utah, but includes spec-
imens from around the world as well as
Smith’s collection notebooks (including
field notes). His collection of small parasitic
wasps (Hymenoptera: Braconidae: Aphidi-
inae) is also held at NC State. Additionally,
the W. FE Barr Entomological Collection,
University of Idaho, Moscow, and the En-
tomological Museum of Utah State Univer-
sity, Logan, both hold significant numbers
of aphid slides donated by Smith. The pri-
mary types for most (75) of the aphid spe-
cies that Smith authored (or coauthored) are
deposited at the U.S. National Collection of
Insects and Mites housed at Beltsville,
Maryland (www.sel.barc.usda.gov/aphid/
aphframe.htm).

Among Smith’s numerous publications
on aphids are: Aphididae of Puerto Rico
(1963, with L. F Martorell and M. E. Pérez-
Escolar), Bibliography of the Aphididae of
the World (1972b), Keys to and Descrip-
tions of the Genera of Pemphigini in North
America (1974), An Annotated List of Aphi-
didae of North America (1978, with C. S.
Parron), An Annotated List of Aphididae of
the Caribbean Islands and South America
(1979, with M. M. Cermeli [Cermeli-Lol-
lini]), and A Key to Many of the Common
Alate Aphids of North Carolina (1992, with
R. W. Eckel and E. Lampert). Smith’s doc-
toral dissertation, which treated a group of
small wasps parasitic on aphids, was the ba-
sis for his Aphidiinae of North America
(1944c).

The extensive collection of papers Smith
used to compile his bibliographies, check-
lists and taxonomic works is deposited with
the NCSU Libraries’s Special Collections

VOLUME 102, NUMBER 4

Department as the Clyde FE Smith Papers
(Collection Number MC 221). These pa-
pers, some of which are rare, encompass
literature, reprints, pamphlets, and books,
primarily pertaining to the insect family
Aphididae. Associated with the papers are
approximately 288,000 index cards on
Aphididae with indexes that include: an an-
notated generic and specific index to all lit-
erature cited in Smith’s 1972 Bibliography
of Aphididae, a host-plant index indicating
the aphids associated with each host, the
source citing this association, and an index
of aphid-parasitoid associations. Together,
the 1972 Bibliography and the 1978 An-
notated List of Aphididae of North America
provide indexes to the Smith Papers.

His hobbies included gardening, hunting,
fishing, lapidary, and genealogical research.
Another more unusual pastime was watch-
ing Hopkins’ Bioclimatic Law play itself
out among the plants and associated arthro-
pods of North Carolina, a state with ex-
tremes in topography. This empirical law,
which Smith taught in his fruit insect
course, states that:

“Other conditions being equal, the
variation in the time of occurrence of
a given periodical event in life activity
in temperate North America is at the
general average rate of 4 days to each
I degree of latitude, 5 degrees of lon-
gitude and 400 feet of altitude, later
northward, eastward and upward in
the spring and early summer, and the
reverse in late summer and autumn.”
(Hopkins, A. D. 1918. United States
Department of Agriculture, Monthly
Weather Review Supplement 9: 7).

Over a 10-year period, Smith observed the
seasonal development of a scrawny flow-
ering peach tree near his office. He used it
as an index to the first appearance in the
spring of the adults of the plum curculio,
whose grubs would later infest the peaches
of the North Carolina Sandhills. Smith had,
of course, observed and predicted other
phenological events across the state corre-

1079

lated with the vernal development of that
peach tree. He then would cajole his grad-
uate students into bets of soft drinks con-
cerning what they would find at the re-
search site when they arrived. Smith lost a
few sodas, but not many.

Frequently, he suggested graduate stu-
dent research projects that ranged far from
his own expertise, expecting and inspiring
students to learn on their own, as he had
throughout his life. His graduate students
included: R. Bastida, C. S. Black, Jr, C. S.
Parron, and L. Van Balen (all M.S.); M. M.
Cermeli-Lollini, H. L. Comroy, M. H. Far-
rier, J. Graham, H. B. Moore, Jr., A. T. Ol-
ive, M. E. Pérez-Escolar, and C. G. Wright
(all Ph.D.). Smith genuinely enjoyed ob-
serving and puzzling out nature, including
the complex life histories of his beloved
aphids.

On 13 February 2000, his long, produc-
tive life drew to a close, following a period
of ill-health initiated by a stroke in January
1999.

Clyde FE Smith will be long remembered
for his devotion to family, community, and
church (Church of Jesus Christ of Latter
Day Saints) as well as his professional
achievements. He will be missed by his
many friends, co-workers, and family mem-
bers, particularly his ever supportive wife,
Crystle; three children, Clara Beth, Caro-
lynn, and Clyde Leslie, and their spouses;
15 grandchildren; and 14 great-grandchil-
dren.

PUBLICATIONS BY CLYDE F. SMITH!

Knowlton, G. F, J. S. Stanford, and C. F
Smith. 1934. Birds as predators of the
beet leafhopper. Journal of Economic
Entomology 27: 1196—1197.

Knowlton, G. FE and C. E Smith. 1935a. To-

! Dr. Smith maintained a card index of his publica-
tions with each item numbered. The 156 items in that
index, plus nine others, are listed below, as far as pos-
sible following Smith’s numerical ordering. Our efforts
to verify the citations of certain older Extension pub-
lications and a movie have so far proven unsuccessful.
Two further papers are in review.

1080

mato fruitworms. Utah State Agricultural
Experiment Station Leaflet 61: 1—4.

Knowlton, G. FEF and C. E Smith. 1935b.
The desert gridiron-tailed lizard. Copeia
2: 102-103.

Knowlton, G. E and C. E Smith. 1935c.
Toads in the control of auto-camp in-
sects. Journal of Economic Entomology
28: 496.

Knowlton, G. EF and C. FE Smith. 1935d.
Currant and gooseberry aphids. Utah
State Agricultural Experiment Station
Leaflet 63: [1—4].

Knowlton, G. FE and C. E Smith. 1935e.
Beet leafhopper predators—Birds. Pro-
ceedings of the Utah Academy of Sci-
ences, Arts, and Letters 12: 249-253.

Knowlton, G. FE and C. E Smith. 1935f.
Notes on Utah Scarabaeidae and Chry-
somelidae (Coleoptera). Entomological
News 46: 241-244.

Knowlton, G. FE and C. E Smith. 1936a.
Lizard aids in range insect control. Jour-
nal of Economic Entomology 29: 461.

Knowlton, G. EF and C. E Smith. 1936b.
The aphid genus Epameibaphis in Utah.
Proceedings of the Entomological So-
ciety of Washington 38: 89-92.

Knowlton, G. EF and C. E Smith. 1936c.
Capitophorus aphids infesting Chryso-
thamnus. Canadian Entomologist 68:
107-113.

Knowlton, G. E and C. E Smith. 1936d.
Notes on intermountain aphids. Ento-
mological News 47: 210-213.

Knowlton, G. E and C. EF Smith. 1936e.
Rose insects. Proceedings of the Utah
Academy of Sciences, Arts, and Letters
13: 263-267.

Knowlton, G. E and C. E Smith. 1936f.
Strawberry insects. Proceedings of the
Utah Academy of Sciences, Arts, and
Letters 13: 289-292.

Knowlton, G. EK and C. E Smith. 1936g.
Capitophorus aphids infesting Artemi-
sia. Canadian Entomologist 68: 229—
234.

Knowlton, G. EK and C. E Smith. 1936h.
Descriptions of some rabbit brush and

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

willow aphids. Annals of the Entomo-
logical Society of America 29: 776—
778.

Knowlton, G. E and C. FE Smith. 19361.
Aphid studies. Festschrift zum 60. Ge-
burtstage von Professor Dr. Embrik
Strand (Riga, Latvija) 1: 487—492, plate
VIII.

Smith, C. EF 1937. The aphid genus Fla-
bellomicrosiphum in Utah. Pan-Pacific
Entomologist 13: 127-129.

Smith, C. E and G. E Knowlton. 1937. Ma-
crosiphum aphids infesting Chryso-
thamnus and Gutierrezia. Canadian En-
tomologist 69: 269-272.

Knowlton, G. EF and C. E Smith. 1937.
Some aphids of the genus Capitopho-
rus. Canadian Entomologist 69: 150—
152:

Smith, C. FE and G. EF Knowlton. 1938. The
aphid genus Microsiphum in Utah and
Idaho. Annals of the Entomological So-
ciety of America 31: 162-166.

Knowlton, G. FE and C. EF Smith. 1938a.
Notes on western conifer aphids (Ho-
moptera: Aphididae). Entomological
News 49: 65-69.

Knowlton, G. E and C. E Smith. 1938b.
The aphid genus Pseudoepameibaphis.
Journal of the New York Entomological
Society 46: 217-222.

Knowlton, G. F, C. E Smith, and FE C.
Harmston. 1938. Pea aphid investiga-
tions. Proceedings of the Utah Academy
of Sciences, Arts, and Letters 15: 71—
80.

Smith, C. EF 1939a. The digestive system of
Macrosiphum solanifolii (Ash.) (Aphi-
dae: Homoptera). Ohio Journal of Sci-
ence 39: 57-59.

Smith, C. E 1939b. Hymenopterous para-
sites of aphids with special reference to
the Aphidiinae. Abstracts of Doctoral
Dissertations (Ohio State University,
Columbus) 30: 169-174.

Smith, C. FEF and G. FE Knowlton. 1939.
Three intermountain aphids. Canadian
Entomologist 71: 241-243.

Smith, C. E 1940. Notes on some Ohio

VOLUME 102, NUMBER 4

aphids. Ohio Journal of Science 40:
139-142, plate I.

Smith, C. E and G. E Knowlton. 1940.
Three aphids of the genus Brevicoryne
van der Goot. Annals of the Entomo-
logical Society of America 33: 404—
405.

Smith, C. EF 194la. Control studies of the
woolly apple aphid. Journal of Econom-
ic Entomology 34: 590.

Smith, C. E 1941b. A new species of hy-
menopterous parasite of the pea aphid
(Macrosiphum pisi Kaltentbach). An-
nals of the Entomological Society of
America 34: 537-538.

Smith, C. FE 1941c. The genus Drepanaphis
Del Guercio east of the Rocky Moun-
tains. Journal of the Elisha Mitchell So-
ciety 57: 226—242.

Smith, C. E 1941d. Trichogramma and the
Oriental fruit moth. Journal of Econom-
ic Entomology 34: 590.

Smith, C. E 1942. The use of cyanide in
controlling the root form of the woolly
apple aphid. Journal of Economic En-
tomology 35: 908-910.

Smith, C. E 1943a. North Carolina, p. 6. In
State by State Summary of 1943 Or-
chard Pest Problems. American Fruit
Grower 63(2): 6.

Smith, C. E 1943b. Scale insects harm
peach trees in North Carolina. Research
and Farming (North Carolina Agricul-
tural Experiment Station, Raleigh) 1(2):
12:

Smith, C. EF 1943c. Experiments with the
peach tree borer in North Carolina.
Journal of Economic Entomology 36:
215-218.

Smith, C. E 1943d. Peach tree borer control
recommendations, 1943. North Carolina
Agricultural Experiment Station, 1943.
[mimeograph printing]

Smith, C. E and G. FE Knowlton. 1943. The
aphid genus Drepanaphis Del Guercio.
Journal of the Elisha Mitchell Society
59: 171-176, plate 22.

Smith, C. EF 1944a. Drepanaphis tissoti, a

1081

new species of aphid from Florida. Flor-
ida Entomologist 27: [55]-57.

Smith, C. E 1944b. Peach insect control.
Proceedings of the Southern Workers’
Conference on Entomology (18th An-
nual Meeting, February 1—3, 1944) 18:
63-67.

Smith, C. E 1944c. The Aphidiinae of
North America (Braconidae: Hymenop-
tera). Ohio State University Press, Co-
lumbus, xii + 154 pp.

Smith, C. E 1944d. Peach tree borer, control
recommendations for North Carolina.
[alternate title: The peach tree borer in
North Carolina]. North Carolina Agri-
cultural Extension Circular 277: [1]-8.

Smith, C. E 1944e. Controlling the plum
curculio on peaches. Research and
Farming (North Carolina Agricultural
Experiment Station, Raleigh) 2(2): 2-3.

Smith, C. E 1945. Reduced concentrations
of ethylene dichloride for peachtree bor-
er control. Journal of Economic Ento-
mology 38: 500-501.

Smith, C. E 1946a. Curculio control rec-
ommendations for North Carolina.
North Carolina State College, January
14, 1946. [lithograph printing]

Smith, C. FE 1946b. Controlling soybean in-
sects. Research and Farming (North
Carolina Agricultural Experiment Sta-
tion, Raleigh) 4(3): 7, 9.

Smith, C. E 1946c. Plum curculio, post har-
vest spraying. North Carolina Agricul-
tural Experiment Station, August 1,
1946. [mimeograph printing]

Smith, C. E 1946d. Peach insects, North
Carolina—1946. Proceedings of the
Cumberland-Shenandoah Fruit Work-
ers’ Conference (23rd Annual Meeting)
239) 20" 0r22 1:

Smith, C. E 1947a. Controlling pea aphids.
North Carolina Agricultural Extension
Service, March 1947. [mimeograph
printing]

Smith, C. E 1947b. Burning to control the
curculio. North Carolina Agricultural
Experiment Station, March 12, 1947.
[mimeograph printing]

1082

Smith, C. EF 1947c. Emergence of the plum
curculio. North Carolina Agricultural
Experiment Station, March 20, 1947.
[mimeograph printing]

Smith, C. E 1947d. Curculio emergence
and petal fall spray. North Carolina Ag-
ricultural Experiment Station, April 7,
1947. [mimeograph printing]

Smith, C. EF 1947e. Emergence of the plum
curculio. North Carolina Agricultural
Experiment Station, April 16, 1947.
[mimeograph printing]

Smith, C. E 1947f. Wormy drops. North
Carolina Agricultural Experiment Sta-
tion, April 28, 1947. [mimeograph
printing]

Smith, C. EF 1947g. Notes on hexaethyl te-
trophosphate. North Carolina Agricul-
tural Experiment Station, June 3, 1947.
[mimeograph printing]

Smith, C. FE and C. N. Clayton. 1947. First
generation curculio and brown rot con-
trol. North Carolina Agricultural Exper-
imental Station, June 9, 1947. [mimeo-
graph printing]

Smith, C. F 1947h. Plum curculio; post har-
vest spraying. North Carolina Agricul-
tural Experiment Station, August 19,
1947. [mimeograph printing]

Smith, C. FE 19471. Things to do and not to
do; peach borer control; scale insects
and burning. North Carolina Agricultur-
al Experiment Station, September 1947.
[mimeograph printing]

Clayton, C. N. and C. FE Smith. 1947. Apple
spray suggestions for 1947. North Car-
olina Agricultural Experiment Station,
1947. [mimeograph printing]

Smith, C. FE and O. Veerhoff. 1947. Ethyl-
ene dichloride injury to peach. Journal
of Economic Entomology 40: 588.

Smith, C. FE and W. H. Rankin. 1947. Does
fertilization increase grain aphids? Re-
search and Farming (North Carolina
Agricultural Experiment Station, Ra-
leigh) 6(1): 6.

Smith, C. FE 1947}. Plum curculio control in
North Carolina—1947. Proceedings of
the | Cumberland-Shenandoah _ Fruit

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Workers’ Conference (24th Annual
Meeting) 24: 26-29.

Smith, C. EF and C. N. Clayton. 1948. Peach
spray information for 1948. North Car-
olina Agricultural Experiment Station
Special Circular 5. [pagination un-
known]

Clayton, C. N. and C. F Smith. 1948. Apple
spray information for 1948. North Car-
olina Agricultural Experiment Station
Special Circular 6. [pagination un-
known]

Smith, C. E 1948a. Pea aphid control in
North Carolina. North Carolina Agri-
cultural Experiment Station Special Cir-
cular 7: [1-7].

Smith, C. EF 1948b. A new aphid on sweet
potato. Florida Entomologist 31: 24-26.

Smith, C. E 1948c. Plum curculio control
in North Carolina. Journal of Economic
Entomology 41: 220-227.

Smith, C. EF 1948d. A new aphid on devil
shoe string (Tephrosia virginiana, L.
Pers.). Annals of the Entomological So-
ciety of America 41: 384-386.

Smith, C. F and C. N. Clayton. 1949. Peach
spray information for 1949. North Car-
olina Agricultural Experiment Station
Special Circular 5(revised): 1-8.

Clayton, C. N. and C. EF Smith. 1949. Apple
spray information for 1949. North Car-
olina Agricultural Experiment Station
Special Circular 6(revised): 1-8.

Smith, C. F, I. D. Jones, and J. A. Rigney.
1949. Effect of insecticides on the fla-
vor of peaches—1948. Journal of Eco-
nomic Entomology 42: 618-623.

Smith, C. E 1949. DDT-parathion: new
weapon for peach growers. Research and
Farming (North Carolina Agricultural
Experiment Station, Raleigh) 7(3): 3.

Smith, C. FE and C. Black. 1950. Peach bor-
er paradox. Research and Farming
(North Carolina Agricultural Experi-
ment Station, Raleigh) 8(3): 5, 7.

Smith, C. FE, I. D. Jones, and L. D. Calvin.
1950. Effect of insecticides on the fla-
vor of peaches—1949. Journal of Eco-
nomic Entomology 43: 179-181.

VOLUME 102, NUMBER 4

Smith, C. FE and C. N. Clayton. 1950. Peach
spray information. North Carolina Ag-
ricultural Experiment Station Special
Circular 5(revised): 1-[11].

Clayton, C. N. and C. EK Smith. 1950. Apple
spray information. North Carolina Ag-
ricultural Experiment Station Circular
6(revised): 1—12.

Randall, G. O., H. R. Garriss, and C. E
Smith. 1951. Successful rose culture.
North Carolina Agricultural Extension
Service, Extension Circular 200(re-
vised): [1—26].

Smith, C. E and C. N. Clayton. 1951. Peach
spray information. North Carolina Ag-
ricultural Experiment Station Special
Circular 12: 1-11.

Smith, C. E and L. Bennett. 1951. Some
tobacco insects and their control. Mov-
ie, 16 mm., color, sound. North Carolina
State College, Raleigh, December 1951.

Smith, C. E and C. N. Clayton. 1953a.
Peach spray information, 1953. North
Carolina Agricultural Experiment Sta-
tion Special Circular 17: 1-11.

smith, (E> By and'C, N. Clayton: 1953b.
Peach spray information, 1953. North
Carolina Agricultural College Plant Pa-
thology Information Note 31: 10—15.

Turnipseed,.G., FE. and ‘C.F Smith. 1953.
Life history and control of scales on ap-
ples in North Carolina. Journal of Eco-
nomic Entomology 46: 969-972.

Smith, C. EF, G. D. Jones, and R. L. Rabb.
1953. Tobacco insect control. North
Carolina Agricultural Extension Ser-
vice, Raleigh, 4 pp.

Smith, C. FE and C. N. Clayton. 1954. Peach
spray information. North Carolina Ag-
ricultural Experiment Station Special
Circular 20: 1—12.

Smith, C. EF and R. L. Rabb. 1954. The ef-
fects of insecticides on the flavor of to-
bacco. Proceedings, Association of
Southern Agricultural Workers (51st
Annual Convention, Dallas, Texas, Feb-
ruary 1—3, 1954) 51: 107.

Randall, G. O., H. R. Garriss, and C. FE
Smith. 1954. Successful rose culture.

1083

North Carolina Agricultural Extension
Circular 200(revised): [1—24].

Clayton, C. N., H. C. Fink, C. EK Smith, and
G. E Turnipseed. 1954. Apple spray in-
formation. North Carolina Agricultural
Experiment Station Special Circular 19:
1-16.

Rabb, R. L., E E. Guthrie, H. E. Scott, and
C. E Smith. 1955. Tobacco insects of
North Carolina and their natural ene-
mies. North Carolina Agricultural Ex-
periment Station Bulletin 394: [1]-35.

Harris, J. H., E A. Hassis [Haasis], and C.
FE Smith. 1955. Azaleas and camellias.
North Carolina Agricultural Extension
Circular 246: [1-32].

Clayton, C. N. and C. FE Smith. 1956. Apple
disease and insect control recommen-
dations—1956. North Carolina Pesti-
cide Manual 1956: 27-28.

Smith, C. FE and C. N. Clayton. 1956. Peach
insect and disease control recommen-
dations—1956. North Carolina Pesti-
cide Manual 1956: 29-31.

Smith, C. EF 1956. Zeno Payne Metcalf, dis-
tinguished entomologist. Science 123:
1022.

Metcalf, Z. P. and C. EK Smith. 1956. Intro-
duction, pp. ii—vil. Jn Metcalf, Z. P,
ed., General Catalogue of the Homop-
tera. Fascicle IV. Fulgoroidea. Part 18.
Eurybrachidae and Gengidae. North
Carolina State College, Raleigh.

Metcalf, Z. P and C. E Smith. 1957. Intro-
duction, pp. ui—vill. In Metcalf, Z. P,,
ed., General Catalogue of the Homop-
tera. Fascicle IV. Fulgoroidea. Part 13.
Flatidae and Hypochthonellidae. North
Carolina State College, Raleigh.

Clayton, C. N., G. E Turnipseed, and C. E
Smith. 1957. Apple spray information.
North Carolina Extension Circular 406:
1-20.

Smith, C. EK and C. N. Clayton. 1957. Peach
spray information. North Carolina Ex-
tension Circular 407: 1—16.

Harris, J. H., E A. Hassis [Haasis], and C.
E Smith. 1957. Azaleas and camellias.

1084 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

North Carolina Agricultural Extension
Circular 246 (revised): [1—32].

Metcalf, Z. P. and C. EF Smith. 1958. Intro-
duction, pp. ili-vu. In Metcalf, Z. P,
ed., General Catalogue of the Homop-
tera. Fascicle IV. Fulgoroidea. Part 15.
Issidae. North Carolina State College,
Raleigh.

Smith; <G) B.-L: EF Martorell;and*M~. E:
Pérez-Escolar. 1958. Myzus persicae
(Sulzer) in Puerto Rico. Journal of Ag-
riculture of the University of Puerto
Rico 42: 263-266.

Smith, C. EF 1958. What’s new in insecti-
cides? Agricultural Chemicals 13(3):
43, 45.

Smith, C. EF 1959. A new species of aphid
on sugar maple (Aphidae: Homoptera).
Annals of the Entomological Society of
America 52: 647-649.

Smith, C. FE and C. N. Clayton. 1959. Peach
spray information. North Carolina Ag-
ricultural Extension Circular 407(tre-
vised): 1-15.

Smith, C. EF 1960a. Aphids (Aphidae: Ho-
moptera) on “‘cacao”’ in the Dominican
Republic. Journal of Agriculture of the
University of Puerto Rico 44: 154-156.

Smith, C. E 1960b. New species of Aphi-
dae: Homoptera from Puerto Rico. Jour-
nal of Agriculture of the University of
Puerto Rico 44: 157-162.

Smith, C. FE and N. Tuatay. 1960. The genus
Microparsus Patch (Aphidae: Homop-
tera). Annals of the Entomological So-
ciety of America 53: 735-742.

Smith, C. EF 1960c. North Carolina finds 3-
year old inspection form still accept-
able. Pest Control 28(9): 40—41.

Harris, J. H., EF A. Haasis, and C. E Smith.
1960a. Azaleas and camellias. North
Carolina Agricultural Extension Circu-
lar 246(revised): [1—32].

Harris, J. H., E A. Haasis, and C. E Smith.
1960b. How to grow azaleas and ca-
mellias. North Carolina Agricultural
Extension Service, Extension folder.

Smith, C. E 1961. p.194. In Comments of
the proposed use of the plenary powers

to designate a type-species for Eucera-
phis Walker, 1870. Z.N.(S.) 1363. Bul-
letin of Zoological Nomenclature 18:
194.

Smith, C. FE 1962. p. 196. In Discussion of
the case “Aphis Linnaeus, 1758, its
type-species, and the family-group
name derived from it (Insecta: Hemip-
tera). Z.N.(S) 881”. Bulletin of Zoolog-
ical Nomenclature 19: 195-198.

Smith, C. E and O. E. Heie. 1963. Me-
gouroparsus, new genus, related to Mi-
croparsus Patch and Megoura Buckton
(Homoptera: Aphidae). Annals of the
Entomological Society of America 56:
401—406.

Fields, H. M., G. C. Klingman, C. FE Smith,
and C. N. Clayton. 1963. Pesticides,
facts on their use in North Carolina. Na-
tional Agricultural Chemicals Associa-
tion News and Pesticide Review 21(4):
8—10.

Smith, C. E, L. E Martorell, and M. E.
Pérez-Escolar. 1963. Aphididae of
Puerto Rico. University of Puerto Rico
Agricultural Experiment Station Tech-
nical Paper 37: [1]-121.

Smith, C. F 1965a. Tiliphagus lycoposugus,
new genus, new species (Aphididae:
Homoptera) from Tilia americana and
Lycopus virginicus. Annals of the En-
tomological Society of America 58:
781-786.

Smith, C. F 1965b. Book review: Ento-
mological techniques. How to work
with insects, by Alvah Peterson. Bulle-
tin of the Entomological Society of
America 11: 96.

Smith, C. E and G. EK Knowlton. 1966. The
genus Aspidaphium Borner (Homop-
tera: Aphididae). Pan-Pacific Entomol-
ogist 42: 20-24.

Smith, C. FE and J. Graham. 1967. Life his-
tory, synonymy, and description of Neo-
prociphilus aceris (Homoptera: Aphi-
didae). Annals of the Entomological So-
ciety of America 60: 67-72.

Smith, C. F and D. G. Dillery. 1968. The
genus Drepanaphis Del Guercio (Ho-

VOLUME 102, NUMBER 4

moptera: Aphididae). Annals of the En-
tomological Society of America 61:
185-204.

Smith, C. E and J. O. Pepper. 1968. Gryl-
loprociphilus frosti, new genus, new
species, from the eastern United States
(Homoptera: Aphididae). Proceedings
of the Entomological Society of Wash-
ington 70: 57—60.

Clayton, C. N., G. T. Weekman, and C. E
Smith. 1968. Peach spray program.
North Carolina Pesticide Manual 1968:
a6:

Smith, C. EK 1969a. Pemphiginae associated
with the roots of conifers in North
America (Homoptera: Aphididae). An-
nals of the Entomological Society of
America 62: 1128-1152.

Smith, C. E 1969b. Controlling peach scale.
Research and Farming (North Carolina
Agricultural Experiment Station, Ra-
leigh) 28(1—2): 12.

Smith, C. E and C. N. Clayton. 1970. Peach
disease and insect control in North Car-
olina. North Carolina Agricultural Ex-
tension Circular 407: 1-17.

Smith, C. FE 1970a. Emergence of the peach
tree borer in North Carolina. Journal of
Economic Entomology 63: 1700-1701.

Smith, C. EF 1970b. Notes on the genus Pic-
turaphis and related genera with a new
species of Picturaphis from Puerto Rico
(Aphididae: Homoptera). Journal of Ag-
riculture of the University of Puerto
Rico 54: 683-688.

Clayton, C. N., C. E Smith, and G. T.
Weekman. 1970. Peach spray program.
North Carolina Agricultural Chemicals
Manual 1970: 112-113.

Smith, C. F 1970c. Sexual activity and egg
fertility of the peach tree borer. Journal
of Economic Entomology 63: 1909—
1910.

Clayton, C. N., C. E Smith, and K. A. Sor-
ensen. 1971. Peach spray program.
North Carolina Agricultural Chemicals
Manual 1971: 124-125.

Smith, C. F, S. M. Gaud, L. E Martorell,
and M. E. Pérez-Escolar. 1971. Addi-

1085

tions and corrections to the Aphididae
of Puerto Rico. Journal of Agriculture
of the University of Puerto Rico 55:
192-258.

Smith, C. E and G. EF Knowlton. 1971.
Pleotrichophorus tetradymiae, a new
species of aphid from Utah (Homoptera:
Aphididae). Proceedings of the Ento-
mological Society of Washington 73:
320-323.

Smith, C. E 1971. The life cycle and re-
description of Mordvilkoja vagabunda
(Homoptera: Aphididae). Proceedings
of the Entomological Society of Wash-
ington 73: 359-367.

Smith, C. FE and H. L. G. Stroyan. 1972.
The probable identity of Prociphilus
(Pulvius) probosceus (Homoptera:
Aphididae). Annals of the Entomologi-
cal Society of America 65: 804-807.

Smith, C. E and C. N. Clayton. 1972. Peach
disease and insect control in North Car-
olina. North Carolina Agricultural Ex-
tension Circular 407(revised): [1]-17.

Clayton, C. N., C. EF Smith, and K. A. Sor-
ensen. 1972. Peach spray program.
North Carolina Agricultural Chemicals
Manual 1972: 124-125.

Smith, C. EF 1972a. Synonymy, redescrip-
tion, and biology of Neoparacletus cor-
rugatans (Homoptera: Aphididae). An-
nals of the Entomological Society of
America 65: 1328-1331.

Smith, C. E 1972b. Bibliography of the
Aphididae of the world. North Carolina
Agricultural Experiment Station Tech-
nical Bulletin 216: [1]-717.

Clayton, C. N., C. E Smith, and K. A. Sor-
ensen. 1973. Peach spray program.
North Carolina Agricultural Chemicals
Manual 1973: 221-222.

Clayton, C. N., C. E Smith, and K. A. Sor-
ensen. 1974. Peach spray program.
North Carolina Agricultural Chemicals
Manual 1974: 251-252.

Smith, C. EF 1974a. The genus Phloeomyzus
with the description of P. dearborni n.
sp. from Populus tremuloides Michx.
(Homoptera: Aphididae). Proceedings

1086

of the Entomological Society of Wash-
ington 76: 66—72.

Smith, C. E and S. M. Gaud. 1974. The
alate viviparae of Picturaphis Blanchard
(Homoptera: Aphididae). Journal of Ag-
riculture of the University of Puerto
Rico 58: 381-383.

Smith, C. E 1974b. Keys to and descrip-
tions of the genera of Pemphigini in
North America (Homoptera: Aphididae:
Pemphiginae). North Carolina Agricul-
tural Experiment Station Technical Bul-
letin 266: [i]—vi, [1]-61.

Smith, C. EF and G. E Knowlton. 1975.
Moss aphids in the United States (Ho-
moptera: Aphididae). United States De-
partment of Agriculture Cooperative
Economic Insect Report 25: 423—431.

Smith, C. FE and A. G. Robinson. 1975. The
genus Myzodium with the description of
M. knowltoni, new species (Homoptera:
Aphididae). Proceedings of the Ento-
mological Society of Washington 77:
48 1—486.

Clayton, C. N., C. E Smith, and K. A. Sor-
ensen. 1975. Peach spray program.
North Carolina Agricultural Chemicals
Manual 1975: 251-252.

Smith, C. E and C. N. Clayton. 1976a.
Peach disease and insect control in
North Carolina. North Carolina Agri-
cultural Extension Circular 407(re-
vised): 1-19.

Stary, P. and C. E Smith. 1976. The Nearc-
tic distribution, host plants and nomen-
clature of Monoctonus (Homoptera:
Aphididae). Proceedings of the Ento-
mological Society of Washington 78:
171-175.

Smith, C. EF and C. N. Clayton. 1976b.
Peach spray program. North Carolina
Agricultural Extension Circular 407(sup-
plement): 1—4.

Smith, C. E and G. FE Knowlton. 1977. The
genus Rhopalosiphoninus Baker (Ho-
moptera: Aphididae) in North America.
United States Department of Agricul-
ture Cooperative Plant Pest Report 2:
75-80.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Smith, C. E and C. N. Clayton. 1978a.
Peach spray information. North Caroli-
na Agricultural Extension Service Cir-
cular 407(supplement, revised): [1—4
(folded leaflet)].

Smith, C. E and C. N. Clayton. 1978b.
Peach disease and insect control in
North Carolina. North Carolina Agri-
cultural Extension Service AG-146 [re-
vision of former Circular 407]: [i—1i, 1]-
YS)

Smith, C. E and C. S. Parron. 1978. An
annotated list of Aphididae (Homop-
tera) of North America. North Carolina
Agricultural Experiment Station Tech-
nical Bulletin 255: [iJ—viii, 1—428.

Smith, C. EF and W. V. Campbell. 1979.
Charles Henry Brett 1909-1978. Jour-
nal of Economic Entomology 72: 157.

Smith, C. E and M. M. Cermeli [Cermeli-
Lollini]. 1979. An annotated list of
Aphididae (Homoptera) of the Carib-
bean Islands and South and Central
America. North Carolina Agricultural
Research Service Technical Bulletin
259: [iii], 1-131.

Cermeli L[ollini], M. [M.], and C. E Smith.
1979. Keys to species of the genus Pic-
turaphis (Homoptera: Aphididae) with
descriptions of new species. Proceed-
ings of the Entomological Society of
Washington 81: 611—620.

Smith, C. E 1980a. A fourth species of Tox-
opterella Hille Ris Lambers (Homop-
tera: Aphididae) from North America
with a key to species. Proceedings of
the Entomological Society of Washing-
ton 82: 276—283.

Smith, C. E 1980b. Notes and keys to the
species of Carolinaia (Homoptera:
Aphididae). Proceedings of the Ento-
mological Society of Washington 82:
312-318.

Smith, C. E 1982. A key to the species of
Hyalomyzus (Homoptera: Aphididae) in
North America, with description of a
new species. Proceedings of the Ento-
mological Society of Washington 84:
325-331.

VOLUME 102, NUMBER 4

Smith, C. EK and H. A. Denmark. 1982. Tri-
chosiphonaphis polygoni (van der Goot)
(Homoptera: Aphididae), a genus and
species new to the United States. Flor-
ida Entomologist 65: 381-382.

Smith, C. E 1983. A new species of aphid
(Homoptera: Aphididae) from Potentil-
la canadensis. Proceedings of the En-
tomological Society of Washington 85:
64-68.

Smith, C. E and G. FE Knowlton. 1983. A
key to the species of aphids (Homop-
tera: Aphididae) on wild Geranium spp.
in the United States, with description of
a new species. Proceedings of the En-
tomological Society of Washington 85:
686-690.

Smith, C. F and H. A. Denmark. 1984. Life
history and synonymy of Grylloproci-
philus imbricator (Fitch) (Homoptera:
Aphididae). Florida Entomologist 67:
430-434.

Smith, C. E 1985. Pemphiginae in North
America, pp. 277—302. In Szelegiewicz,
H., ed., Evolution and Biosystematics of
Aphids. Proceedings of the International
Aphidological Symposium at Jablonna,
5-11 April 1981. Zaklad Narodowy im
Ossolinskich Wyawnictwo, Warsaw.

Smith, C. FE, R. W. Eckel, and E. Lampert.
1992. A key to many of the common
alate aphids of North Carolina (Aphi-
didae: Homoptera). North Carolina Ag-
ricultural Research Service Technical
Bulletin 299: [i]—vi, 1—92.

Smith, C. E and R. W. Eckel. 1996. Aphis
crassicauda n. sp. (Homoptera: Aphi-

1087

didae), with a key to the alate species
of Aphis on Viburnum spp. in the Unit-
ed States. Proceedings of the Entomo-
logical Society of Washington 98: 44—
49.

Clements, K. M., B. M. Wiegmann, C. E.
Sorenson, C. FE Smith, P. A. Neese, and
R. M. Roe. 2000. Genetic variation in
the Myzus persicae complex (Homop-
tera: Aphididae): evidence for a single
species. Annals of the Entomological
Society of America 93: 31—46.

ACKNOWLEDGMENTS

For useful suggestions and information,
we thank: Herbert H. Neunzig, Charles G.
Wright (both Professors Emeriti), Carol S.
Parron (former Curator of the NCSU Insect
Collection), James D. Harper, John R. Mey-
er, Michael R. Roe, and Clyde E. Sorenson
(Department of Entomology, North Caroli-
na State University, Raleigh); Caroline
Weaver and Linda A. McCormick (Special
Collections Department, North Carolina
State University Libraries); Pamela E. Pur-
year (Agricultural Communications—Tobac-
co Literature Service, North Carolina State
University); Randi V. W. Eckel (RVWE
Consulting, Frenchtown, New Jersey); and
Manya B. Stoetzel, (Systematic Entomolo-
gy Laboratory, Agricultural Research Ser-
vice, U.S. Department of Agriculture,
Beltsville, Maryland).

Maurice H. Farrier and Lewis L. Deitz,
Department of Entomology, North Carolina
State University, Raleigh, NC 27695-7613,
U.S.A. (email: lewis.deitz@ ncsu.edu)

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1088-1092

SOCIETY MEETINGS

1044th Regular Meeting—January 6, 2000

The 1044th regular meeting of the En-
tomological Society of Washington (ESW)
was called to order in the Cathy Kerby con-
ference room (CE-340) of the National Mu-
seum of Natural History, Washington, D.C.,
by incoming President Dave Furth at 7:35
pm. The meeting was attended by 20 mem-
bers and 10 guests. Stu McKamey read the
minutes of the 1043rd meeting, which were
approved without modification.

Chris Thompson called a point of order,
that the ESW audit was not presented. Don
Anderson reported that the audit was signed
by committee members and provided a
summary of the status: ESW is doing fine.
Thompson moved to accept the report and
the motion was quickly seconded and the
audit approved unanimously. There were no
new applicants for membership and no new
members were present. One visitor was in-
troduced. In miscellaneous business, there
was short discussion about the ESW web-
site, which has not yet materialized.

For exhibits, Dave Furth announced the
Washington Academy of Science website
(http://www.washacadsci.org), displayed
the calendars of the Entomological Society
of America and the Organization for Trop-
ical Studies, displayed insect toys recently
collected without permits in Canada and
Mexico, insect tacks (is nothing safe from
the IRS?) and even a bee-motif shower cur-
tain. Warren Steiner added to his ‘global
warming’ insect sighting list: buckeye and
sulpher butterflies spotted on December 12,
to which a January 2nd monarch eclosion
was added from the audience. Warren noted
the insects included in the revamped
NMNH elephant exhibit and lastly an-
nounced a new publication: the Proceedings
of the Appalachian Biogeography Sympo-
sium, edited by ESW member Ralph Eck-
erlin. Edd Barrows completed the exhibits
with a butterfly mobile calendar.

John Brown introduced the evening’s
speaker, past ESW President Mike Schauff,
who took us on a trip “Out and About
Down Under: Collecting Tiny Wasps in
Australia.’ In truth, three of his trips were
cleverly merged into one tour, all beauti-
fully documented with slides of cities, wa-
terfalls, panoramic vistas, beaches, charis-
matic megafauna, and not-so-charismatic
megafauna: crocodiles, snakes, and huge
leeches. We started our trip in subtropical
SE Australia, which proved the best for his
parasitic chalcidoid wasps. We traveled
northward into the understory of the
Queensland tropical rainforest, which was,
as expected, poor collecting. From there we
moved inland to the edge of the more arid
central plateau, heavily agricultural but
with open forests and drainage areas that
yielded many chalcidoids. Then, drawn like
a moth to flame, we headed into the ‘red
center’ desert of the continent. Collecting
was excellent in the shaded areas around
Ayer’s Rock, Uluru National Park, where
water persists year-round. We continued
south to Adelaide, Bookmark Biosphere
Reserve in the Murray River watershed,
and Kangaroo Island, collecting wasps and
enjoying the abundant marsupials, some-
times scaring them away from the best
shade spots for a break from the sun, other
times sharing a research station with them.
Eventually we had to return to Canberra,
but couldn’t leave without collecting on
Black Mountain, located just behind CSI-
RO headquarters. Collecting permits for the
trip were arranged by state and by National
Park and required a couple months’ ad-
vance planning.

The meeting was adjourned at 8:50 pm.
Refreshments were provided by John
Brown.

Respectfully submitted,
Stuart H. McKamey
Recording Secretary

VOLUME 102, NUMBER 4

1045th Regular Meeting—February 3,
2000

The 1045th regular meeting of the En-
tomological Society of Washington (ESW)
was called to order in the Cathy Kerby con-
ference room (CE-340) of the National Mu-
seum of Natural History, Washington, D.C.,
by President Dave Furth at 7:35 pm. The
meeting was attended by 22 members and
4 guests. Stu McKamey read the minutes of
the 1044th meeting, which were approved
with minor modification.

Steve Lingafelter read the names of four
new applicants for membership: Kevin
O’Neill, Wayne Wehling, Qiao Gexia, and
Daniel E. Perez-Gelabert. Two new mem-
bers and two visitors were introduced.

For exhibits, Chris Thompson called at-
tention to a recent Nature article on the pu-
tative forces driving the evolutionary origin
of butterflies, an excellent new handbook
for taxonomy (Describing Species: Practi-
cal Taxonomic Procedure for Biologists),
and a new, world catalog of beeflies. Edd
Barrows noted a special issue of the Amer-
ican Entomologist on biodiversity and had
one on display. Ralph Eckerlin displayed a
new Spanish language entomology text: Jn-
sectos y Otros Artrépodos de Importancia
Médica y Veterinaria, by E. Méndez. Dan-
iel Perez had on hand two new publications
from the Dominican Republic: a catalog of
Dominican amber fossils and an Occasional
Publication on taxonomy from the national
museum. Dave displayed a recent Science
article on ‘hire classification’ (i.e., charging
for patronyms), a new curation book (Care
and Conservation of Natural History Col-
lections), a new book on the systematics of
Western North American butterflies, and
“Bug Bites,”’ chocolate candies with insect
trading cards.

Stu McKamey introduced the meeting’s
speaker, Dr. Steven Lingafelter, ESW mem-
ber and Research Scientist with the USDA
Systematic Entomology Laboratory. He
spoke to us about “‘Asian Longhorned Bee-
tles of the Genus Anoplophora: a New

1089

Threat to the United States.”’ First detected
in New York in 1996 but probably present
for 10 years, these large, showy beetles
have received wide media coverage owing
to their destruction of living ornamental
trees. The larvae are the real culprits, bor-
ing in the live wood for up to a year. Con-
trol here and even in Asia often consists of
eradication: destroying any infested trees.
Lingafelter’s connection to Anoplophora is
his taxonomic expertise on longhorns (Cer-
ambycidae) and the need to resolve species
delimitations in the genus, with about 50
nominal species, to aid control and inter-
ception efforts. Additionally, all distribution
records will be databased and an identifi-
cation guide will be produced. Most records
of the genus are from Japan, Korea, and
China, where Lingafelter visited recently to
examine institutional and private collec-
tions, and to do more collecting. The Chi-
nese portion was coordinated through the
joint U.S.A.-China Institute of Biological
Control and the Chinese Forestry Institute.
The showy patches of color on the beetles’
elytra, traditionally used to delimit species,
are actually quite variable among and even
within populations. The genitalia are, sur-
prisingly, not very useful either. The species
appear to fall into five morphological
groups defined by minor structural, external
features. Quarantine controls have been
strengthened and continued vigilance is re-
quired to avoid additional introductions of
Anoplophora.

The meeting was adjourned at 8:53 pm.
Refreshments were provided by John
Brown.

Respectfully submitted,
Stuart H. McKamey
Recording Secretary

1046th Regular Meeting—March 2, 2000

The 1046th regular meeting of the En-
tomological Society of Washington (ESW)
was called to order in the Cathy Kerby con-
ference room (CE-340) of the National Mu-
seum of Natural History, Washington, D.C.,

1090

by President Dave Furth at 7:37 pm. The
meeting was attended by 23 members and
10 visitors. M. Alma Solis read the minutes
of the 1045th meeting, which were ap-
proved without modification.

Steve Lingafelter read the names of three
new applicants for membership: Ken Karns,
Tam Nguyen, and Art Evans. One new
member was introduced.

For exhibits, Ralph Eckerlin brought the
book entitled Kinabalu, edited by K. M.
Wong & A. Phillipps and Chris Thompson
brought Biology Systematics: Principles
and Applications by Randall T. Schuh.
Dave Furth brought the following books
and articles: Ants at Work by Deborah Gor-
don; Ecology of Insects: Concepts and Ap-
plications, edited by M. R. Speight, M. D.
Hunter & A. D. Watt; Forester Moths by
K. A. Efetov & G. M. Tarmann; Insect
Plant Relationships edited by S. J. Simp-
son, A. J. Mordue & J. Hardie; The Johns
Hopkins University of School of Hygiene
and Public Health, Lloyd E. Rozeboom
Mosquito Collection (1999. Amer. Mosqui-
to Control Assoc. 15: 526-551), and two
articles by Ashley Gurney entitled A Short
History of the Entomological Society of
Washington (1976. Proc. Entomol Soc.
Wash. 78: 225-239), and Who Designed the
Schmitt Box? (1975. Bull. Entomol. Soc.
Amer. 21: 225-228). Edd Barrows brought
some sawflies from Dyke Marsh. Gaby
Chavarria advertised the film, Pollinators in
Peril, to air on TBS Superstation, Monday,
March 27, 11:00 pm/et., 8:00 pm/pt., and
modeled a “‘pollinator’’ blouse worn by her.

John Brown introduced the speaker, Dr.
Gabriella Chavarria, National Fish and
Wildlife Foundation (NWF) and Research
Associate at the National Museum of Nat-
ural History. Her talk was entitled ‘‘Polli-
nation Conservation: the Truth About the
Birds and the Bees.’ She spoke generally
on pollination biology, flower morphology,
kinds of bees, and co-evolution between
flowers and pollinating organisms. At the
NWFE she has been instrumental in a “Save
the Pollinators”’ program. Every 3rd bite of

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

food is brought to humans by a pollinator.
The best pollinators are bees (72.3%),
among many others, including vertebrates
such as birds and bats. But these pollinators
are threatened by habitat destruction and by
habitat and chemical habitat fragmentation.
The major threats to Apis mellifera have
been parasitic mites, bee diseases and pests,
competition with africanized bees, and pes-
ticide use. Fortunately, there are native bees
to serve as alternative pollinators such as
mason bees and bumblebees. Some solu-
tions to the threats are: pollination research
and monitoring, international agreements to
protect migrating species, incorporation of
wildlife corridors between agricultural are-
as, environmentally safe golf courses, plant-
ing native plants in backyards, and placing
‘bee condominiums.”’ She suggested read-
ing The Forgotten Pollinators by Steve
Buckman & Gary Nabhan.

The meeting was adjourned at 8:35 pm.
Refreshments were provided by John
Brown.

Respectfully submitted,
M. Alma Solis

1047th Regular Meeting—April 6, 2000

The 1047th regular meeting of the En-
tomological Society of Washington (ESW)
was called to order in the Cathy Kerby con-
ference room (CE-340) of the National Mu-
seum of Natural History, Washington, D.C.,
by President Dave Furth at 7:32 pm. The
meeting was attended by 20 members and
1 guest. Stu McKamey read the minutes of
the 1046th meeting, which were approved
with minor modification.

Steve Lingafelter read the names of five
new applicants for membership: Netta Dor-
chin, James Kruse, Gordon E Pratt, Gale E.
Ridge-O’Connor, and David Campbell. No
new members were present.

For exhibits, Ed Cohen displayed the No-
vember 1999 issue of The American Mu-
seum of Natural History containing an ar-
ticle on the flight mechanics of insects.
Dave Furth announced BugFest on the Mall

VOLUME 102, NUMBER 4

May 20th, encouraging ESW support; dis-
played a new CD on the NHB East Court
and the entomology compactor construc-
tion; had the three latest issues of Food In-
sects Newsletter from Montana State Uni-
versity; announced CDs on beetles by John
Lawrence, CSIRO; a world catalog of Cas-
sidinae; The Cicadidae of China, in Chinese
but full of photos [Ilustrataj Insect-faunoj:
2, Chou & Lei (eds.)]; and a new Peterson
field guide to western butterflies by P. Opler
& A. Bartlett Wright. He also solicited
opinions for an ESW website, should the
project get started. Lastly, Furth announced
the annual ESW banquet, tentatively sched-
uled for June 8 with guest speaker Dave
Grimaldi.

John Brown introduced the night’s speak-
er, Jim Whitfield (University of Arkansas/
University of Illinois) who presented his re-
search on the ““Phylogeny of Microgastroid
Braconid Wasps, and What it Tells Us
About Polydnavirus Evolution.”’ Like other
insects with endoparasitoid larvae, these
wasp larvae battle the hosts’ immune sys-
tems. Their special tools are polydnavirus-
es: viral genes incorporated into their own
genome and therefore inherited between
generations. The expression of these genes
suppresses the host’s immune response and
can also alter its nutrition and developmen-
tal rate. Dr. Whitfield investigated the origin
of this virus-wasp relationship by compar-
ing their phylogenies at three taxonomic
levels: subfamilies, genera, and species.
Looking at wasp subfamilies, evidence
from morphology or molecules or both, and
analyzed by parsimony or maximum like-
lihood all confirmed a single origin of the
association, with no subsequent loss. At the
generic level, all data sets were congruent
concerning which branches were short vs.
long, but despite all attempts to wring out
well supported hypotheses of relationship,
all they could show was a rapid radiation
of genera. That is, the problem seemed to
be the evolutionary pace of the microgas-
troid wasp genera, which is confirmed in-
dependently by fossils, rather than flaws in

1091

the methods or data. At the species level,
analysis is difficult because the viral genes
are meshed into the wasp genome, yet sep-
arated from each other. Whitfield and col-
leagues so far have found two viral genes
and their phylogenies match closely that of
their wasp hosts. Dr. Whitfield concluded
that the viral genes were probably incor-
porated 60 to 80 million years ago and were
followed by rapid diversification at the ge-
neric level.

The meeting was adjourned at 8:55 pm.
Refreshments were provided by John
Brown.

Respectfully submitted,
Stuart H. McKamey
Recording Secretary

1048th Regular Meeting—May 4, 2000

The 1048th regular meeting of the En-
tomological Society of Washington (ESW)
was called to order in the Cathy Kerby con-
ference room (CE-340) of the National Mu-
seum of Natural History, Washington, D.C.,
by President Dave Furth at 7:36 pm. The
meeting was attended by 18 members and
4 guests. Stu McKamey read the minutes of
the 1047th meeting, which were approved
without modification.

Steve Lingafelter, the membership chair,
was in Nepal so no new applicants for
membership were read. No new members
were present.

For exhibits, Warren Steiner passed
around a live caterpillar hunter (Callasoma,
Carabidae), John Brown announced the
ESW Annual Banquet, June 8 at the Uni-
formed Services University of Health Sci-
ences in Bethesda, Maryland. The feature
will be a talk about the ““New Jersey Mud
Pits and the Glories of Fossil Insect Work,”’
presented by Dave Grimaldi of the Ameri-
can Museum of Natural History, New York.
Tickets are $24 per person, $12 for students
and half-people. Dave Furth dominated the
rest of the exhibits: notes from the ESW
executive e-meeting, at which it was decid-
ed that starting in October the regular meet-

1092

ings would be held at 7:00 pm; announced
the BugFest on the Mall and again invited
members to participate; announced the
opening of a one-year exhibit on ‘‘Vanish-
ing Pollinators” at the National Zoo in the
Amazonia Science Gallery; displayed some
new books: /nsects and Their Spermatozoa
and Their Phylogeny; Catalog of the Het-
eroptera of the Palearctic—Miridae;, The
Preying Mantids; and The Butterflies of
Hong Kong, which in addition to the usual
photographs gives details on their mor-
phology, larvae, and parasites.

John Brown announced the evening’s
speaker, Michael G. Pogue of the System-
atic Entomology Laboratory, based in the
National Museum of Natural History. He
spoke to us about “The Armyworm Genus
Spodoptera Guenée (Lepidoptera: Noctui-
dae): a Multimedia Review.’ Spodoptera
has 30 species and includes the single most
serious pest of Lepidoptera; other species of
the genus are also serious pests. The genus
is mainly subtropical and tropical but some
migrate into temperate regions. The genus
is particularly abundant in Africa, Southeast
Asia, Borneo, and South America. Dr. Po-
gue’s monographic goals are to treat every
species comprehensively, estimate their
phylogeny, build a specimen database, and
provide an interactive key. Features he uses
to delimit species are taken from wing col-
or, genitalia of both sexes, and larvae. The
database actually consists of six linked da-
tabases: specimens (more than 10,000 re-

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

cords), a gazetteer (more than 4,000 Spo-
doptera localities with latitude and longi-
tude), larval hosts, parasitoids, predators,
and a bibliography. The latter four are now
available on the WWW (http://160.111.87.
10:591/entomology/spodoptera/spodintro.
html). The Expert System is in progress.
Although primarily a key, the system is de-
signed to give experts additional informa-
tion once an identification is attained. The
system uses the LucID windows program
(University of Queensland and CSIRO),
which is very flexible. The builder can code
character states as polymorphic or “‘likely
misinterpreted,’ options difficult to incor-
porate into traditional, couplet-based keys.
The user can choose characters in any order,
display characters ranked by difficulty, and
see existing images of any character states.
Each choice produces a shorter list of pos-
sible species. A traditional key for 30 spe-
cies will have 29 couplets, so to identify the
majority of the species a user passes
through 15 couplets. In comparison, iden-
tification paths using LucID are much
shorter. For example, any species of Spo-
doptera can usually be identified within five
character state choices. Dr. Pogue plans to
continue adding to the database as time per-
mits.

The meeting was adjourned at 8:55 pm.
Refreshments were provided by John
Brown.

Respectfully submitted,
Stuart H. McKamey
Recording Secretary

PROC. ENTOMOL. SOC. WASH.
102(4), 2000, pp. 1093-1099

PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
of
WASHINGTON

Volume 102

OFFICERS FOR THE YEAR 2000

President David G. Furth
President-Elect John W. Brown
Recording Secretary Stuart H. McKamey
Corresponding Secretary Hollis B. Williams
Treasurer Michael G. Pogue
Program Chair Theodore R. Schultz
Membership Chair Steven W. Lingafelter
Custodian Jon A. Lewis
Editor David R. Smith
Past President Michael E. Schauff

Published by The Society
WASHINGTON, D.C.
2000

TABLE OF CONTENTS, VOLUME 102

ARTICLES
ABRAHAMSON, WARREN G.—See MELIKA, GEORGE ..........---...s ss eee cece sees cece tenes 198
ADAMSKI, DAVID and ERIC H. METZLER—A new species of Glyphidocera Walsingham
from southwestern Ohio (Lepidoptera: Gelechioidea: Glyphidoceridae) ..........-....-++..+++5 301
ADLER. PETER H:i=—See PAYSEN, BRIG SF oiii es. s cece eee n cc cee es soe ce cm seecimneecimenenmerccs sc 843
/NUICIQOIN|, ID) ——$"See TIE, VS Soke ssoncesosneoobosbenconasapeposaeccodanecogesossce0s0sec dosccoboc 688
ALONSO-ZARAZAGA, MIGUEL A.—See HOEBEKE, E. RICHARD .............--...---++555 151

ALUJA, MARTIN, JAIME PINERO, MAURILIO LOPEZ, CESAR RUIZ, ALBERTO ZUNIGA,
ENRIQUE PIEDRA, FRANCISCO DIAZ-FLEISCHER, and JOHN SIVINSKI—New host plant
and distribution records in Mexico for Anastrepha spp., Toxotrypana curvicauda Gerstacker, Rha-
goletis zoqui Bush, Rhagoletis sp., and Hexacheta sp. (Diptera: Tephritidae) .........-.----++++++5 802
ANTOLIK, CHRISTOPHER—See ROONEY, THOMAS P. .........--. 22-222 cece eee ee eee eees 308
ARCHANGELSKY, MIGUEL—Immature stages of Neotropical Hydrophilidae (Coleoptera):
Hydramara argentina (Knisch, 1925) and Hemiosus bruchi Knisch, 1924 ..................... 280

1094 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ARCHANGELSKY, MIGUEL—See BRANHAM, MARC@ Av core ec ceene cece ccc saecueanes cece 869
ASSIS-PUJOL, CRISTIANE VIEIRA DE and MICHEL LECOQ—Comparative study of sper-
mathecae in eleven Rhammatocerus Saussure 1861 grasshopper species (Orthoptera: Acri-

didae: Gomphocernae:S cyMlinini) hts 2 5cee en. dose ete ee eee An eee EEE en ae 120
BAPTISTA, ALESSANDRA R. P. and WAYNE N. MATHIS—Notes on the genus Cyamops

Melander (Diptera: Periscelididae), including description of ten new species .................. 481
BARROWS: EDWARD INVE——Sceei CIC BRO MAR Ye Re meeea ee Cee er EEe etree eee te ere eEreaaee 50
BLANCO:METZLER, HEEGA— See GAGNES RAYMOND S. spe.o502: sue se ee eee 831
BORKENT, ART—The larva and pupa of Schizonyxhelea forattinii Wirth and Grogan (Diptera:

Ceratopogonidae) with a discussion of the phylogenetic relationships of the genus ........... 862
BOYDSTONWE==Sée PIRI, 6S (SP sccariccch tac ccn ademas Bese ee eee Taare A GER EEE eee ee 688

BRANHAM, MARC A. and MIGUEL ARCHANGELSK Y—Description of the last larval in-
star and pupa of Lucidota atra (G. A. Olivier 1790) (Coleoptera: Lampyridae), with a dis-

cussion of abdominal segment homology across life stages. ...............0..cceeeeeceee cece sees 869
BRASWELL, W. EVAN and JAMES R. OTT—The biology of Doa ampla (Grote) (Lepidop-

tera: Doidae) on its host plant Stillingia texana (Euphorbiaceae) ..................... eee e eee e eee 507
BRIGHT, DONALD E.—See VANDENBERG, NAWTATIA Jr o.oo oc ecenenesecccscseennene ceases seen 62
BROWN, JOHN W.—Revision of Lobogenesis Razowski and Odonthalitus Razowski (Lepi-

doptera: Tortricidae: Tortricinae), with comments on their monophyly .......................5. 72)

BROWN, JOHN W. and JON LEWIS—Catalogue of the type specimens of Tortricidae (Lep-
idoptera) in the collection of the National Museum of Natural History, Smithsonian Institu-
trons WashinstomiyDNC), Rew rccieaececncomnh atane eaaika canis ensuu soa sass ac neeenias ance mertre meeen 1014
BROWER, ANDREW V. Z.—On the validity of Heliconius tristero Brower Heliconius mel-
pomene mocoa Brower, with notes on species concepts in Heliconius Kluk (Lepidoptera:

IN Aion (0) 02 WG 21) oe ote are or ere tor en Seren Re Aer Senora Sac Penene ic nadonauondocaimaposcoe oC 678
BUTLER, LINDA and JOHN STRAZANAC—Macrolepidopteran larvae sampled by tree bands

in temperate mesic and xeric forests in eastern United States ................. 0.0 cece eee eee e ee eee 188
BYERS ROBERT At Sees HORBERE, ES RICHARD a9. 2a. eee eee eee eee eee 151
CICERO, MARY R. and EDWARD M. BARROWS—Milesine flower flies (Diptera: Syrphidae)

in a central Appalachian broadleaf forest: Abundances, flight periods, and diflubenzuron. ........ 50
CONTRERAS-RAMOS, ATILANO—A new species of Chloronia Banks (Megaloptera: Cor-

ydalidae) from southeastern Brazil, with a key to the species of Brazil ........................ 919

CRYAN, JASON R. and LEWIS L. DEITZ—Review of the New World treehopper tribe Ste-
gaspidini (Hemiptera: Membracidae: Stegaspidinae): HI: Flexocentrus Goding, Stylocentrus
Staleand"Wimibelligertus MD GitZs conc cece cass Se eee ae ee EERE ee 82

DARSIE, RICHARD E, JR.—Description of the pupae of five species in the subgenus Armi-
geres, genus Armigeres Theobald, with a key to species of the known pupae of the subgenus
(Diptera GultcrGae) MRS x kee terre hse ese cid este ROS AT RSet ERE 108

DARSIE, RICHARD EF, JR.—The pupae of three species in Minomyia Theobald from Nepal
and a key to the known pupae of the genus in the Oriental and Australasian regions (Diptera:

(GU LKec EVO) Seta meecmrcen ane aoc Gana accd mien ecu en EO mnan cr aae nom ane ma coaeommnnnboeranmacce noc 625

DARSIE, RICHARD FEF, JR.—Description of the pupa of Armigeres (Leicesteeria) omissus
(Edwards) and a key to the larvae and pupae of the Armigeres occurring in Nepal (Diptera:

(SULT ARTO) eee ae ee aeRO et Berens at Seen EO Me See Shen | EPP ERE Aas taaaace cnosGoacdcoous 964
DAVIDSON; JOHN ‘A.——See) POLAVAIR ABU Ss 7 ieet oecct ast cence otc car er senieee eee cereeeee 549
DEIVZ, LEWISjL.—See'CRYAN, JASON Reiiks a.p.cbh asa aeaeton- usleceaaeae teahi ae eee eee 82
DIAZ-FLEIS CHER. See ALUIA,, MARTIN. 4. cere oe eae cae x eee ee 802
DIETRICH, C. H.—See NOVIKOV,.D! Vis .ns2Sohaaserens os teangctnteedan 6 15 setie deemee eee eee 170
ECKERLIN, RALPH P. and HARRY E PAINTER—New records of fleas (Siphonaptera) from

eastern West Virginia’ 3...) 00.60 cama nae sas gohan olaumiene rence maemisece eee eee eee ae eee eee 969
EDMISTON, JAMES F——SeesMiATHITS SWAYINIESINets-se-e- oe ea deaeieece de ene he aea-eeeee eee 652
EDMONSONER> E-=Scé) WALES ON AR ID Seer ea ee neo ee enolase och nee Lee eee ee 360
ETIENNE SBAN —=SeeiGAGNEVRAY MOND: 205.254 s.o. cee ee oe ee 831
EVANS, HOWARD E.—Three new species of Dipogon Fox (subgenus Dipogon) (Hymenop-

tera: Pompilidae) from central and western North America ..................0.seeeee cece eres es 1010
FOOLE, BENJAMIN A:——See NORRBOMEPAIBIER Noe aepe-peeee eee aae eee eee eee 142

FOSTER, GEORGE A. and WAYNE N. MATHIS—Notes on Neotropical species of Tethina
Haliday (Diptera: Tethimidae)) eee hn ee ae ae ees ees see aE 542

VOLUME 102, NUMBER 4

FROESCHNER, RICHARD C.—See HENRY, THOMAS J. ...... 2.02... c cece eect meee eset e eee
GAGNE, RAYMOND J., HELGA BLANCO-METZLER, and JEAN ETIENNE—A new Neo-
tropical species of Clinodiplosis (Diptera: Cecidomyiidae), an important new pest of culti-
vated peppers (Capsicum spp.: Solanaceae) ............ 6. cece eee cece eee e teen eee eees
GAIMARI, STEPHEN D. and MICHAEL E. IRWIN—Revision of the mexicana-group of the
cycloteline genus Ozodiceromyia Bigot (Diptera: Therevidae) .................-s2. ee eeeee seen ees
GATES, MICHAEL W.—A new species of Cirrospilus Westwood (Hymenoptera: Eulophidae)
fromthe southwesterm United (States! and! MEXICO) foc ee cre crclete ore -iociatnie s)eisteialnla = ole eteteter= lores ete elelerotaa =
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota sig-
nifera (Coquillett) (Diptera: Tephritidae) on Hemizonia pungens (Hooker and Arnott) Torrey
and A. Gray (Asteraceae) in southern Califormia .................2.:.eseseeseeeeeecerccssce estes
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota aenig-
ma Friedberg and Mathis (Diptera: Tephritidae) on Erigeron divergens Torrey and Gray
(Asteraceae) mms outherns@alifomialpeeeeerreee eee eres ee tere eee neat eee pee eer rye er reer
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota ap-
pendiculata Freidberg and Mathis (Diptera: Tephritidae) on Machaeranthera canescens
(Pursh) A. Gray (Asteraceae) in southern California ...................+.ssecceeneneecc seme ecerees
GOEDEN, RICHARD D.—Life history and description of immature stages of Neaspilota pu-
bescens Freidberg and Mathis (Diptera: Tephritidae) on Lessingia filaginifolia (Hooker and
Arnott) M.A Wane (Asteraceae) ini southern Califormia cose ote ate ornins oa e)oele eialeieteletele oieiel= ==
(GIRN EMG = Ses THUMB KS Se) ohpnooranecasboasasnsaceA00 10060 9600000 cons conn apadaabauposspeeapodooonoce
GUILBERT, ERIC—Revision of the genus Parada Horvath (Hemiptera: Tingidae) with cla-
GUSTIN. AUTEN. 5 Ata aa Me cteiooic Secon OA BAAS aR oee tas SRS en ano esac So aCe oro cee Sessa ocr Sanaa: Oras
EU ANIKS SIEAWIREINGE IME Sees MOORES ROBERT Greece cree eee eee reer ase eer:
BUN RIBVACISE RUNILIPIEL 1, See IMO SITES. IDIOISIING ZN, 30254590000000 o0nosoerenodus sass ganoeaccasHonGs
HASTRITER, MICHAEL W.—Jordanopsylla becki (Siphonaptera: Ctenophthalmidae), a new
species of flea from the Nevada Test Site ............... 1... e ese e cece cece eect e eee ee tenes
HASTRITER, MICHAEL W. and EUSTORGIO MENDEZ—A review of the flea genera Hec-
topsylla Frauenfeld and Rhynchopsyllus Haller (Siphonaptera: Pulicidae) .............-....-.--
HENRY, THOMAS J. and RICHARD C. FROESCHNER—Corrections and additions to the “Cat-
alog of the Stilt Bugs, or Berytidae, of the World (Insecta: Hemiptera: Heteroptera)” ............
HOEBEKE, E. RICHARD and A. G. WHEELER, JR.—Telmatophilus typhae (Fallén) (Cole-
optera: Cryptophagidae), a Palearctic cattail specialist established in the Canadian maritime
[DRO LE (CE SinGtilaeiee wigs cde OG REE DUS onc Hone En a Tne Peaa Sa RaA See TOS OC USSR OSS Ba REAR EenmEr ooo Praag P9aF
HOEBEKE, E. RICHARD, ROBERT A. BYERS, MIGUEL A. ALONSO-ZARAZAGA, and
JAMES E STIMMEL—/schnopterapion (Chlorapion) virens (Herbst) (Coleoptera: Curcu-
lionoidea: Brentidae: Apioninae), a Palearctic clover pest new to North America: Recognition
featuness GIStmbUtOMe and) DIOMOIMMIGS cece nee cesses eee elelelciclalars ciate le etet eye ole ter=l= ele) =Jelelalal-le(oleieroinietciei=
HOWARD, THERESA M.—See MOSES, DUSTY A. ...............0s sees eect eee eee e cece cece
HUNG, AKEY C. E and HACHIRO SHIMANUKI—Nucleotide sequence and restriction site anal-
yses in three isolates of Kashmir bee virus from Apis mellifera L. (Hymenoptera: JN GES) sesooc
HUSBAND, ROBERT W.—Redescription of Eutarsopolipus desani Cooreman and description
of E. mirifica, n. sp. (Acari: Podapolipidae) from Chlaenius spp. (Coleoptera: Carabidae)
MRO Sieur /NGOISEL Bon bb dduaoosoewosns boodacudeoon codeennanudoaunopodeonoTusno0es sesoHeD aude DpeeCdoUscoOs
IRWIN, MICHAEL E.—See GAIMARI, STEPHEN D. ...........-- +--+ +--+ 22s eeeee teen tenet
JAMESON, MARY LIZ—Synopsis of the Mexican and Guatemalan genera Rutelisca Bates
and Metapachylus Bates (Coleoptera: Scarabaeidae: Rutelinae) with comments on classifi-
cation of the subtribe Rutelina ................2..... cece eect eect ee etter cece tenet ees cc eee eeceecs
JENSEN, ANDREW S.—Eight new species of Macrosiphum Passerini (Hemiptera: Aphididae)
from western North America, with notes on four other poorly known species ..........-..--.
JOHNSON, NORMAN F—See MUSETTI, LUCIANA ............... 2022222 s eee teen eee n eee
KEIPER, J. B. and W. E. WALTON—Biology and immature stages of Ochrotrichia quadrispina
Denning and Blickle (Trichoptera: Hydroptilidae), a spring-scraper .........-.++-+++eeeseeeeeee
KIMSEY, LYNN S.—The western Australian genus Oncorhinothynnus Salter: New species and
relationships (Hymenoptera: Tiphiidae: Thynninae) ........... 6... .60s sees eee e eee eee eet
KOCH, FRANK and DAVID R. SMITH—Nematus oligospilus Forster (Hymenoptera: Ten-
thredinidae), an introduced willow sawfly in the southern hemisphere ..............--...+++.++-
KONSTANTINOV, ALEXANDER S.—See KOROTYAEV, BORIS A. ............------2222055:

561

69

384

519

1003

398

178

1096 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

KOROTYAEV, BORIS A., ALEXANDER S. KONSTANTINOV, and CHARLES W.
O’ BRIEN—A new genus of the Orobitidinae and discussion of its relationships (Coleoptera:

Curculionidae) siti 2 sasinbass date Ricken Stale = oie saeale Reet ete ASE to a ein eats eet eee O29
IKROMBEIN, KARE RY.—_SceawAIs, RAYMOND ite eaeeeereceee ree atte cee Een re enaeee 271
KRUSE, JAMES J.—Archips goyeriana, n. sp. (Lepidoptera: Tortricidae) an important pest of

baldcypress (Taxodiaceae) in Louisiana and Mississippi ...............-.....ceeeeeeccceeeseeceaes 741
LAPIERRE, LOUIS M.—Prey selection and diurnal activity of Holcocephala oculata (F.) (Dip-

tera: “Asilidae) in | COStamRiGa rab osticb Soo ae eA eae Ee ne Ae Pe RE 643
LECOOs MICHELE See ASSIS-PUIOENERISTIANESVIER AIDE a eee ne ane een eee re 120
LEMMON CAROTR==sees MATER. GCHRISME Wieen eee eeeeee ee eee eee aeaee ae aes 747
EESTER;, DiGi =—SeesWwileSON: Az Di iin d5ccdten cc Ee, SE A nee 360
BEWIS; VON ==SceeiBiRO WNI ORIN OWS eeiecnccsncescocne sae rises ceereniosaceionn i biarirosactee ie aan 1014
LOPEZ MAURIIO™=s ce FATWA SIAR TING: Oe eC ee ee ee 802
LOPEZ-MARTINEZ, VICTOR—See WHARTON, ROBERT A. ............0:cc0secceeeeeeeereees 794

MAIER, CHRIS T. and CAROL R. LEMMON—Discovery of the small Japanese cedar lon-
ghorned beetle, Callidellum rufipenne (Motschulsky) (Coleoptera: Cerambycidae), in live

arborvitaemn"Gonnecticut™ 2: A dtace lane ones nen one fone beac ewer sete mgt banades ae eenres son aeenre anaes 747
MARSHALL, S. A.—Chespiritos, a new genus of Limosininae (Diptera: Sphaeroceridae) from

GOStAMRIC AM en. sass eso: ote as Stee Seele eee eae EEE ce coher E ae Sein eetes Hon ae SoM an emer me ea 609
MATHIS, WAYNE N.—See BAPTISTA, ALESSANDRA R.P. 122.2202... 22sec. ccccceesecceee esse 481
MIAUENIS SWAIN INS ees OSMERS GEORGE TA ec saeco eee eae ener een ae ee eeeee cn eeee 542
MATHIS, WAYNE N. and JAMES EF EDMISTON—A revision of the shore-fly genus Lem-

naphilat@ressony(Dipteray Ee phy dndac)t-.-eacce ace see sec eecis see eee e ee eee eee eer 652

McKAMEY, STUART H.—A review of the New World leafhopper subgenus Texananus (lo-
wanus Ball) (Hemiptera: Cicadellidae: Deltocephalinae) with a checklist and distribution sum-

IMAaby 1Oh SPEClESMM te GENUS acne eneren cen on tren omar connie etnies ame tm seria eee aeienen eer DN
MELIKA, GEORGE and WARREN G. ABRAHAMSON—Review of the cynipid gall wasps

of the genus Loxaulus Mayr (Hymenoptera: Cynipidae) with descriptions of new species... 198
MENDEZ, EUSTORGIO—See HASTRITER, MICHAEL W. .-...0..-.5-c0..)21s cess eee 613
METZEBR. ERIC Hi See ADAMSRIn DA VID en nc se seecesnprecls see Sec ae Aee Oe ace eee Seen 301
MICE RS DOUGEASS R——see ROLAVARARUS Saeccaeise ee rec ee eee ener eee eee eeeree 549
IVIIGTE EAR RE aS CO PUIG RS ca crane ee rsiacerctea Menara eactaltevarcias ie se svn seg hue ePeveas acefol ae orate eI ee ae sea eee ORE 688
TAA OU Ce) EH Saeed eS ereiel 2 UI Sa ahs aes Noein a nearer arene seacmangaedaacdcade suodanoseroeanacoos 688
MOORE, ROBERT G. and LAWRENCE M. HANKS—Avian predation of the evergreen bag-

worm (kepidopteta: ESVChidae) Gana waeracerreeeeteee ace n ore cere eeeCe eer ee Peete eens 350
IMMORVAING, IMUANIMIU SUEY IDK =e IROLOINDE NG INSKOIMUNS) 12. Gonasccoocadooenscnosononovcsdaadedcanceecs 308
MORON, MIGUEL ANGEL and ANGEL SOLIS—New Costa Rican species of Phyllophaga

Flarnish (Coleoptera Melolonthidac-)Melolonthinae) saa-eere eee eeeee eee ere een eee cree eee 901
MOSES, DUSTY A., THERESA M. HOWARD, and RALPH E. HARBACH—A new species

of the subgenus Sabethoides of Sabethes (Diptera: Culicidae) from Venezuela and Brazil .. 991
MUSETTI, LUCIANA and NORMAN FE JOHNSON—First documented record of Monomachidae

(Hymenoptera: Proctotrupoidea) in New Guinea, and description of two new species ............ QSi7/
NEUNZIG, H. H.—Uncitruncata leuschneri, a new genus and species of Phycitinae (Lepidop-

teraweyralidac) proms Calitoniarandi@Ores oniee-eeeeer eee eee eee eerGeereeeet eee Eee eee ens 408
NEUNZIG, H. H.—New species of Puerto Rican Phycitinae (Lepidoptera: Pyralidae) ......... 838
NORRBOM, ALLEN L. and BENJAMIN A. FOOTE—A new subgenus, Footerellia, and new

distribution records of Neaspilota Osten Sacken (Diptera: Tephritidae: Terelliini) ............ 142
NOVELO-GUTIERREZ, RODOLFO—Description of the larva of Hetaerina infecta Calvert

(Odonata: Calopteryeidae)) (o.xtee de saswjac enue. qeae-erre ne discs <ckn-oa scene seem tere eer ee 99
NOVIKOV, D. V. and C. H. DIETRICH—New megophthalmine leafhoppers (Homoptera: Ci-

cadellidae) strom) Mexico; withiaskeystosNews Worldsspeciesiys..--ems-teeteaseeeee eee eee eee 170
© BRIENT CHARLES W:—seesKORODMYAEV, BORIS “Ag fe) sepa eae ee eee eee ee ere eee 929
OTT, JAMES R:— Sees BRASWELL AW HENAN) \.c:restecerocen tease ae ere eacee eeecie: eoeeecteene 507
PAINTER, HARRY F——See BEMERIEINGRVAISP He Psa eee eee acne eee a eee eee 969

PAPP, JENO and SCOTT R. SHAW—A study of the genus Falcosyntretus Tobias from the
New World with five new species and a key to known species (Hymenoptera: Braconidae:
Euphorinae)) s,s.scsea stipe cbse ee Se ee ae EE Sa ee Eee 634

VOLUME 102, NUMBER 4 1097

PAYSEN, ERIC S. and PETER H. ADLER—Taxonomy and polytene chromosomes of Simu-

lium parnassum Malloch (Diptera: Simuliidae) .................6.20 ss eee cee cece ee cee eee eee e cee ees 843
PEDROSA-MACEDO, J. H.—Biology and behavior of the strawberry guava sawfly, Haplos-

tegus epimelas Konow 1901 (Hymenoptera: Pergidae), in southern Brazil ...............-....- 129
PIEDRASENRIOUE—See.A EUSANIARIBING)..2:: <aib eae deniers neat eee ee 802
PIKE, K. S., P STARY, T. MILLER, G. GRAF D. ALLISON, L. BOYDSTON, and R. MILL-

ER—Aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) of northwest USA .......... 688
EINERO.VAIME See AWUTA: MARTIN: o6.0:.c8h stasis tie sel Ghee TE vrs an de ee 802
POGUESMICHAE LI G:——Sceer SALUKES SANDRAOWV. co ie cen crecena-eeeteceoce caste ernneer aa ee 233

POLAVARAPU, S., JOHN A. DAVIDSON, and DOUGLASS R. MILLER—Life history of
the Putnam scale, Diaspidiotus ancylus (Putnam) (Hemiptera: Coccoidea: Diaspididae) on
blueberries (Vaccinium corymbosum, Ericaceae) in New Jersey, with a world list of scale

INSECtSKONMDIMEDE RGLES reenact oer ers Oras 2 nue mee ealoiQak ale aiere lays clenals Measeielbvare stato Ne erstelattstee 549
POLHEMUS, DAN A.—A revision of the endemic Hawaiian reduviid genus Saicella Usinger,

with descriptions of four new species (Heteroptera: Reduviidae: Emesinae) ................--. 1
QIAO, GE XIA and GUANG XUE ZHANG—A taxonomic review of the genus Delphiniobium

Mordvilke,(@iemeptera: Aphididae)an (Chimay S722 censen eee 222 eee eierolenemeleeinccine reser 892
RABAGLIA, ROBERT J.—See VANDENBERG, NATALIA J. 22.2.0... 5.000222 ce ccc cece esse cece 62
ROBBINS, ROBERT K.—The New World hairstreak genus Arawacus Kaye (Lepidoptera: Ly-

CASMCESS INocelteies IK SN) pone sogd>quacodsenodeseacone s4d0e ce cetacodenouaeabsenopsopscopadoasun 162
ROONEY, THOMAS P., CHRISTOPHER ANTOLIK, and MATTHEW D. MORAN—The im-

pact of salamander predation on Collembola abundance ................... 222s. see eeeeee eee ee ees 308
RIZE SAR See VAT UNA INMANROTAING 0 bat tee ans encr eres eetean eed atcae 3 are ae ieee one 802

RUSSELL, LOUISE M.—Notes on the family Aleyrodidae and it subfamilies: Redescription
of the genus Aleurocybotus Quaintance and Baker and description of Vasdavidius, a new

genus (Homoptera: Allyerodidae) 2.2. vce recimersiecins seem cles ore femelle ole elses sieve vis tine nice 374
S/ATINIL IMUAILIRIUNIT S—=Gyae, WANSIUR YG cacacoucaocsndosoncogaqoodasdndssepene coon oosnRooNcoD doo BasdcoORC 601
SALUKE, SANDRA V. and MICHAEL G. POGUE—Resolution of the Elaphria festivoides

(Guenée)) species complex (Lepidoptera: Noctuidae) ........... 02. iiii eee eee ee eee 233

SCARBROUGH, A. G.—Two additional species of robber flies of the genus Ommatius Wie-
demann (Diptera: Asilidae) from the Bahamas and with replacement names for two other

5) DSLETER) 5 ce done OS RIBIe ae 9 SORTA Cre SDD A FB RES AR TRCADO dre Gh Ec Loc ge dap. 7 cGE RO OnOCanA gp aaa opedoee oF 912
SCHAUFE MICHAEL E.—A new genus and species of Eulophidae (Hymenoptera) from Costa

RUA Symlilal TMOWES Cin HS SSNS, 0450000 cosdocssacnedeasesseboeunD on CSusscauuODSUdODeodoHseacAboAneauacac 403
SHAWASCO IE IR: —See- PARP? JENO 6 .x).5 ccc cea ren scl ents te oo nosed dation athe Aevaetes HeseaO ae 634
SHIMIANUIKG. HA CHIR @— See EIUINGay AIK (© isteresereteererreretete toler s)o ro = eroded =letetel- fatale tate tte) 178
SITES, ROBERT W. and MICHAEL R. WILLIG—Morphometric variation among populations

of Ambrysus mormon Montandon (Heteroptera: Naucoridae) .............-.-+++see esses sees eee 533
SIVINSKLVIOHN= See AlUNAs MARTIN, 35.case¥ eer nets eS ele SR 802
SIMIMOSE IDVAUD) IRi——Syee IM(OLEI8 F JERVAINILG Go cooduasessncocenencnesanees coodbbenHSbaccseodnsacace ono 292
SIMICMaL IDZAWALD) IR Seo WANSIOR WG ccvosccaconscsb0osonccns caus eon bocGndnocunoposeccocdonooToTecRosont 601
SIMIGTEEs AWAD RES ee) ZIIN @) VIE VSPA EGG Gos Freee eters erste evoke ele rele oleate let dedeleke fale let elated =r stalet=tele 852
SIMMS E IDYANA0D) Re Srees ZAIN(O UIE, ZENE EN'C (Gi oa6 GondacaasenaccadoopuaoddeSdencspsecescnuancc 974
SOLIS S ANGEL — See: MORON) MIGUEL 2ANGED iwc. .cacyah sen sens she ete thence ones aeincm tener 901
SPIRO GP" See DIK Ke. Sn 8 A or eerye rete isi: eee. (Sauda Sacto coe eee 688
STARVAZZ/NINUNGS JOVSIN Ses IBIUMNLIENRS ILINIDYA 5 ocooncse0ccne0sconbeoongrogscodogpeuadnavacnssesabnan 188
SEIMMETDT, JAMES! E—See HOEBEKE, Es RICHARD) Jo 5225. cae -- acters lee =e ese ame 151
THOMPSON, FE CHRISTIAN and MANUEL A. ZUMBADO—Flower flies of the subgenus

Octyptamus (Mimocalla Hull) (Diptera: Syrphidae) ............. 22... eee eeee eee eee ee eee eee e eens V3
TOGASHI, ICHIJI—Description of a new species of the genus Stiricorsia Konow (Hymenop-

ieee OSS Ch) ison EVEN Soocscuoscnnnaenseadobo0bpoo Sb occduonEscenasuddaSsseoaesAboconAen ooasqeocc 105
TOGASHI, ICHIHI—Japanese sawflies of the japonicus group of the genus Taxonus Hartig

(Hymenoptera: Tenthredinidae) .................. ee eeeeeeee eee eee eee e eee eee een e eee eeenencees 33
VANDENBERG, NATALIA J., ROBERT J. RABAGLIA, and DONALD E. BRIGHT—New

records of two Xyleborus (Coleoptera: Scolytidae) in North America ..........--..+-.+00e005- 62
VASU, V., DAVID R. SMITH, and MALKIAT S. SAINI—Review of the Asian sawfly genus

Anisoarthra Cameron (Hymenoptera: Tenthredinidae) .................+. 22s eee eee e reese eee eee 601

WAHIS, RAYMOND and KARL V. KROMBEIN—A new Machaerothrix Haupt from Sri
Lanka with notes on the genus (Hymenoptera: Pompilidae: Pepsinae: Ageniellini) ........... NWA

1098 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

WALTON WB: Seer RBIPBRe 7 Bit Sion. gaiacas tn dene sete som neers oooh TL an ener 183
WELLS, SAMUEL A.—Two new species of Horistonotus Candeze (Coleoptera: Elateridae),
new synonymies, and a key to the species of the United States and Canada .................. 412

WHARTON, ROBERT A. and VICTOR LOPEZ-MARTINEZ—A new species of Triaspis
Haliday (Hymenoptera: Braconidae) parasitic on the pepper weevil, Anthonomus eugenii

Canox(ColeopteraxGurculionidae)) Seco. secon eect lee ee Sar ren RES eRe SEERA eee eee 794
WHEBRER (ASG; JRe— Sees HOEBE KEE MRICHARD Ieeenaseen eee nae ee eee eee eee 398
WIETETG VMIGHABIOR Sees iES. ROBE RU Wiareesseaeeaseeeeeee een ehhe eee eee eeeeee 5535
WILSON, A. D., D. G. LESTER, and R. E. EDMONSON—Live oaks, new hosts for Odon-

tocynips nebulosa Kieffer (Hymenoptera: Cynipidae) in North America ....................... 360
WOODLEY, NORMAN E.—A new Afrotropical species of Allognosta Osten Sacken (Diptera:

S rat Onnyii Gd aAS) Per ata cer es ee a Se ne eee 924
YASUNAGA, TOMOHIDE—Anthocorid bugs of the tribe Oriini (Heteroptera: Anthocoridae)

of the /@Ocasiwarai(Bonin) islands Japaniytes.t a2 Sige a Bo, ae Paes 353
ZEHANGIGUANGEXUE—=Scel@ VA ON GEUXTAG Wyse eee ae ee one en eae 892
ZINOVJEV, ALEXEY G. and DAVID R. SMITH—Types of sawflies described in the genus

Pontania A. Costa (Hymenoptera: Tenthredinidae) in the Illinois Natural History Survey ... 852
ZINOVJEV, ALEXEY G. and DAVID R. SMITH—Sawflies (Hymenoptera: Tenthredinidae)

described by Benjamin D. Walsh, with notes on their hosts and biology .....................+5 974
ZUMBADO; MANUEBIE A See DHOMPSON, EF GHRISTIAN 252220) cee ee eee eee a3
ZUINIGA:-Al-BERTO=See AWUIA. MARTIN 2:4,20 125.0 ERO ott ea eee 802

NOTES
ALUJA, MARTIN, EMANUEL HERRERA, MAURILIO LOPEZ, and JOHN SIVINSKI—First

host plant and parasitoid record for Anastrepha spatulata Stone (Diptera: Tephritidae) ...... 1072
COCA-ABIA,, MILAGROS-—“See GARCIA=PARIS, MARION... 42 Len alse oe 473
GARCIA-PARIS, MARIO, GABRIELA PARRA-OLEA, and MILAGROS COCA-ABIA—

First records of the genus Noserus LeConte (Coleoptera: Zopheridae) in Mexico ............. 473
IRI BININISRVAS JaIMUANNIOI BU Syeret JNILIUIUAS, IMIR, soca nagocs 900 asoovanopbvconcancooonauocenncscocass 1072
JACOBS, EWGENE Ay——See FARTS TEPHENL IS csctvantatass seheconsaioaehiereee neon eee eae 75
KERSTNER ;CORENNAD==Seev PAPEL) STEPHEN: laus.cciee: Maeda aecee eet ee eee eee 2D
LAZZARI, SONIA MARIA NOEMBERG—See ZONATA DE CARVALHO, REGINA ...... 757
LOPEZ, MARUILIO “See /AbUIAM MAR TING 2 hace 25.5528 s eee en ee 1072
PARRA-OLEA, GABRIELA—See GARCIA-PARIS, MARIO) .....5.0..¢0.5.0005b2eccsc teehee estes 473
PEASE VENGG—SceROBBINS RICHARDG A fest asta e ee eer cee renee 225

ROBBINS, RICHARD G. and STEVEN G. PLATT—First report of Amblyomma clypeolatum
Neumann (Acari: [xodida: Ixodidae) from the Union of Myanmar, with two new records from

LOMEOISES 5258 stains eitee tha erie Seis ok by a arene enter ele eS a So Ee ee RR oes on St A eae 225
SIVINSKE: JOHINSee- AL UIA. MARTIN 555m tise ovo nation lise PASO Ee AOR ee rae 1072
TAFT, STEPHEN J., CORENNA D. KERSTNER, and EUGENE A. JACOBS—Ectoparasitic

insects from migrating saw-whet owls (Aegolius acadicus) in central Wisconsin ............. IS
WHEELER, A. G., JR.—New distributional and first specific host-plant records for Thionia

acuta Doering and T. producta Van Duzee (Auchenorrhyncha: Fulgoroidea: Issidae) ........ 759
WHEELER, A.G., JR.—Fitchia aptera Stal (Hemiptera: Reduviidae): Seasonal history and

habitsan: mid-Appalachian:shale: barrens)... kasssanene eh ose eee oe ee eee eee 1070

ZONTA DE CARVALHO, REGINA and SONIA MARIA NOEMBERG LASSARI—First re-
cord of the California pine needle aphid, Essigella (Essigella) californica (Essig) (Homoptera:
Aphicidacwleachninae) sine SOUtherny nazar e eee eres ae nee eee US

BOOK REVIEWS

BROWN, JOHN W.—Classification of Lepidoptera. Part 1. Introduction, by John B. Heppner ... 1075
GAGNE, RAYMOND, J.—ldentification of North American Porricondyline Larvae (Diptera:
Cecidomytidaamiby: Jolin’ D;, Plakidas: 17. cee ee asa ce Leon Ae eee eee 227
ROBBINS, RICHARD G.—The Genus Rhipicephalus (Acari, Ixodidae): A Guide to the Brown
Ticks of the World, by Jane B. Walker, James E. Keirans, and Ivan G. Horak ................ 768

VOLUME 102, NUMBER 4 1099

SMITH, DAVID R.—Pflanzenwespen Deutschlands (Hymenoptera, Symphyta) Kommentierte

Bestandsaufnahme, edited by Andreas Taeger and Stephan M. Blank .......................... 229
STOETZEL, MANYA B.—Aphids on the World’s Crops, An Identification and Information
Guide, Second Edition, by R. L. Blackman and V. FE Eastop ..................0cce cece cece eeees 1074
OBITUARIES
ARNE TM ROSS) He JR —SeerS REINER] WARRENGES DIRe sacceesceereceee seen cere esas 761
DETEZS BE WIS le —See*PARRIERS MAURICE, He 325 coccncekan satecen oo os tiies tate sis meiseesaulsieer ec 1077
FARRIER, MAURICE H. and LEWIS L. DEITZ—Clyde Fuhriman Smith (1913-2000) ...... 1077
IKINGSOLVIERSJOHNeME——See Ss MEINERS WARREN) Ex IRQ aaaecseetecee cea ote sei see errr 761

STEINER, WARREN E., JR., JOHN M. KINGSOLVER, and ROSS H. ARNETT, JR.—A

HRLOLTNES (1) eral SyovllonBn SooseeeaconeccasncoosososocsoubogsgnGpnaadobo0ccpene spobogococcosDEsoDDLSNEDdOC 761
MISCELLANEOUS

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i eeO RG Cit ONES RS Geeennanepecns csanedo sue cooc ecb sorcontncr CoEBe Hest tae or enn aan scab 1c Oro dcEr ocr 475

SOS IRIN SSIES sacra aosipon7 ase iecee cbce nooo eegoncan Stes sce uo aa oU sEac acerca spncedcnn asa 477, 1088

EAblero tl GOntents sn, O LUTE wil De praceee ariel. esa etaeva rate rats erate No cde arralcvo id ioe ce aIsre eee eter Sevare eeieerareiets 1093

1100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Entomological Society of Washington

Publications.—Available Miscellaneous Publications and Memoirs of the Society are
listed on the inside back cover of each issue.

Instructions for Authors.—Instructions were last published in 2000, 102(3): 771-772.

Bylaws.—The bylaws were last published in 1966, 98(3): 610-614. Several amend-
ments were proposed at the December 3, 1998 regular meeting and passed at the January
7, 1999 regular meeting. See minutes of the Society meetings, 1999, 101(2): 467 (1034th
regular meeting), and 1999, 101(4): 920 (1035th regular meeting).

Membership.—A membership list is published every three years and was last published
in 1999, 101(4): 914-919.

Seal.—The history of the official seal of the Society was published by Jon L. Herring,
1964, 66(1): 1, and was reprinted in 1997, 99(1): 208.

History.—The following articles relate to the history of the Society:

Wade, J. S. 1936. The officers of our Society for fifty years (1884—1934). Proceedings
of the Entomological Society of Washington 39: 121-132.

Gurney, A. B. 1976. A short history of the Entomological Society of Washington.
Proceedings of the Entomological Society of Washington 78(3): 225—239.

Spilman, T. J. 1984. Vignettes of 100 years of the Entomological Society of Washington.
Proceedings of the Entomological Society of Washington 86(1): 1—10.

Stoetzel, M. B. 1984. ESW past-presidents for the years 1884 through 1983, photo-
graphs and support officers. Proceedings of the Entomological Society of Wash-
ington 86(1): 11-35.

The papers by Spilman and Stoetzel were part of the centennial issue of the Society,
1984, 86(1). The centennial banquet was held at the University of Maryland, College
Park, March 12, 1984; see Society Meetings, 1984, 86(4): 975-976.

The 100th regular meeting of the Society was celebrated February 2, 1995 and was
held at the Beltsville Agricultural Research Center, Beltsville, MD. It was attended by
115 members and guests and written up in the minutes of Society meetings 1995, 97(4):
897-898.

PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON

MISCELLANEOUS PUBLICATIONS

A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera), by E. Eric Grissell and Michael E.

Schauff. 85 pp. 1990

A Handbook of the Families of Nearctic Chalcidoidea (Hymenoptera): Second Edition, Revised, by E. Eric

Grissell and Michael E. Schauff. 87 pp. 1997

MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Memoirs 2, 3, 7, 9, 10, 11, and 13 are no longer available.

No. 1.

No. 21.

No. 22.

The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 ___.........-
A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952
A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 1957 __..

The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
EEE TE TSP Es) UEP CT oR 120) I Maa ACE eh aN ale oe eM aA Shes MGM A eek STONE Os SING BL

The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cerato-
pogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 _

. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.

OOS GMS NES elk AST Se Sg alae, 22 AMS BCU A aM AMO CA Dea Ry. PN «COR Co BUPA Eira as

. Biology and Phylogeny of Curculionoidea, edited by R. S. Anderson and C. H. C. Lyal. 174

TTS te Rk ERS ag ES ENGR cele) a A Se a LON | AOL aI eal es eT, SE SSAA SATE

. A Revision of the Genus Ceratopogon Meigen (Diptera: Ceratopogonidae), by A. Borkent

CUR Ags Wig Soy ee Pa Pa So CS fo ca Re RP RRS, PST CARN EH iE See Ne Zane

. The Genera of Beridinae (Diptera: Stratiomyidae), by Norman E. Woodley. 231 pp. 1995 ___.
. Contributions on Hymenoptera and Associated Insects, Dedicated to Karl V. Krombein, edited

ys. 2_ Norden. and) ALS Menke (21 G'ps: 1096 eal Be ae A he ra

. Contributions on Diptera, Dedicated to Willis W. Wirth, edited by Wayne N. Mathis and

Pyatiaganle: (srogan, Jn 2Onppr hoo hi ets ek ee kU aE Trek okt) ae nah

. Monograph of the Stilt Bugs, or Berytidae (Heteroptera), of the Western Hemisphere, by

Noistnas:) benty C40 psi QO | eat ake BRO Ia pn de re ce! Ree

. The Genera of Elaphidiini Thomson 1864 (Coleoptera: Cerambycidae), by Steven W. Lin-

“SLT S Egy RYT ea IED? RR ee eee oO COOMERA ar a ALPEN a AIDS a Stemi MAR RM as ban ir

New World Blepharida Chevrolat 1836 (Coleoptera: Chrysomelidae: Alticinae), by David G.
Jay Tae UN Mao Bid LS 2: Sian A RRL Seep cee ae a ane) ae al We AER. FORGO, SQ eS UEC SAR teal EE aN som, Sdn Le

Systematics of the North American Species of Trichogramma Westwood (Hymenoptera:
iiachosrammatidae)sby John) Pinto 287. pp. L999 sane ae ee eee

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Back issues of the Proceedings of the Entomological Society of Washington are available at $60.00 per volume
to non-members and $25.00 per volume to members of the Society.

Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10
percent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with the
Custodian, Entomological Society of Washington, % Department of Entomology, Smithsonian Institution,
Washington, D.C. 20560-0168.

CONTENTS

(Continued from front cover)
+
HENRY, THOMAS J. and RICHARD C. FROESCHNER—Corrections and additions to the
“Catalog of the Stilt Bugs, or Berytidae, of the World (Insecta: Hemiptera: Heteroptera)” ..

KOROTYAEV, BORIS A., ALEXANDER S. KONSTANTINOV, and CHARLES W. O’BRIEN—
A new genus of the Orobitidinae and discussion of its relationships (Coleoptera: Curculi-
onidae)

MORON, MIGUEL ANGEL and ANGEL SOL{S—New Costa Rican species of Phyllophaga
Harris (Coleoptera: Melolonthidae: Melolonthinae)

MOSES, DUSTY A., THERESA M. HOWARD, and RALPH E. HARBACH—A new species of
the subgenus Sabethoides of Sabethes (Diptera: Culicidae) from Venezuela and Brazil

MUSETTI, LUCIANA and NORMAN F. JOHNSON—First documented record of Monomachidae
(Hymenoptera: Proctotrupoidea) in New Guinea, and description of two new species
NEUNZIG, H. H.—New species of Puerto Rican Phycitinae (Lepidoptera: Pyralidae)

PAYSEN, ERIC S. and PETER H. ADLER—Taxonomy and polytene chromosomes of Simulium
parnassum Malloch (Diptera: Simuliidae)

QIAO, GE XIA and GUANG XUE ZHANG—A taxonomic review of the genus Delphiniobium
Mordvilko (Homoptera: Aphididae) in China

SCARBROUGH, A. G.—Two additional species of robber flies of the genus Ommatius Wiedemann
(Diptera: Asilidae) from the Bahamas and with replacement names for two other species ....

THOMPSON, F. CHRISTIAN and MANUEL A. ZUMBADO—Flower flies of the subgenus
Ocyptamus (Mimocalla Hull) (Diptera: Syrphidae)

WHARTON, ROBERT A. and VICTOR LOPEZ-MARTINEZ—A new species of Triaspis Haliday
(Hymenoptera: Braconidae) parasitic on the pepper weevil, Anthonomus eugenii Cano (Cole-
optera: Curculionidae)

WOODLEY, NORMAN E.—A new Afrotropical species of Allognosta Osten Sacken (Diptera:
Stratiomyidae)

ZINOVJEV, ALEXEY G. and DAVID R. SMITH—Types of sawflies described in the genus
Pontania A. Costa (Hymenoptera: Tenthredinidae) in the Illinois Natural History Survey ....

ZINOVJEV, ALEXEY G. and DAVID R. SMITH—Sawflies (Hymenoptera: Tenthredinidae) de-
scribed by Benjamin D. Walsh, with notes on their hosts and biology

NOTES

ALUJA, MARTIN, EMANUEL HERRERA, MAURILIO LOPEZ, and JOHN SIVINSKI—
First host plant and parasitoid record for Anastrepha spatulata Stone (Diptera: Tephritidae) ....

WHEELER, A. G., JR.—Fitchia aptera Stal (Hemiptera: Reduviidae): Seasonal history and habits
in mid-Appalachian shale barrens

BOOK REVIEWS
BROWN, JOHN W.—-Classification of Lepidoptera. Part 1. Introduction, by John B. Heppner ....

STOETZEL, MANYA B.—4phids on the World's Crops, An Identification and Information Guide,
Second Edition, by R. L. Blackman and V. F. Eastop

OBITUARY

FARRIER, MAURICE H. and LEWIS L. DEITZ—Clyde Fuhriman Smith (1913-2000)
MISCELLANEOUS

Society Meetings

Table of Contents, Volume 102

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