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USISSN0013-872X
JANUARY & FEBRUARY, 1990 NO. 1

ENTOMOLOGICAL NEWS

Larval and pupal descriptions of Marilia fusca (Trichop-

tera: (Odontoceridae) H. Gene Drecktrah I

A new Xyela (Hymenoptera: Xyelidae) from western

United States David R. Smith 9

A new Rhadinoceraea (Hymenoptera: Tenthredinidae) feed-
ing on Zigadenus (Liliaceae) from southeastern
United States D.R. Smith, W. Dearman 13

A new species of Curtara (Homoptera: Cicadellidae)

from Bermuda Paul H. Freytag 20

Spider (Araneae) taxa associated with the immature
stages of Mantispa interrupta (Neuroptera: Mantis-
jidae) K.M. Hoffman, J.R. Brushwein 23

relation between hearing and flying in
crickets

Daniel Otte 29

The Leptoceridae (Trichoptera) of West

Virginia

J.B. Glover, D.C. Tarter 35

Some ectoparasites of bats from Halmahera Is.,
Indonesia

B.V. Peterson, L.A. Durden, J.E. Keirans, P.M. Taylor 39

Some ectoparasites of bats from Seram Island,
Indonesia

L.A. Durden, B.V. Peterson, N. Wilson, B. Christiansen 48

Annotated checklist of the Rhyacophiloidea and
Integripalpia (Trichoptera) of Alabama

S.C. Harris, P.K. Lago 57

Collections of Dufourea versatilis (Hymenoptera:

Halictidae) from Idaho

Sharon L. Walsh 67

SOCIETY MEETING OF NOVEMBER 29, 1989

THE AMERICAN ENTOMOLOGICAL SOCIETY

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Vol. 101, No. 1, January & February 1990

LARVAL AND PUPAL DESCRIPTIONS OF
MARILIA FUSCA (TRICHOPTERA: ODONTO-

CERIDAE)1

H. Gene Drecktrah2

ABSTRACT: The larva and pupa of Marilia fusca from Australia are described and
illustrated.

The genus Marilia is composed of two species in Australia: M. fusca
Kimmins and A/. 60/aMosely (Neboiss 1983, 1986). Additional species of
Marilia have been recorded from Oriental regions, China and South and
North America (Wiggins 1977). Both Australian species were reported to
have the same distribution: New South Wales and the southeast coastal
ranges of Australia (Neboiss 1983). M. fusca was described by Kimmins
(Mosely and Kimmins 1953) from adults taken at Epping, NSW, but no
descriptions of the immature stages were included. This is the first
recorded association of the immature stages of M. fusca with that of the
adult. Identification and association of the immature stages, based on
two pharate males, were made by Dr. A. Neboiss, Curator of the Entomol-
ogy Department, Museum of Victoria, Melbourne, Australia.

Ulmer (1955) described and illustrated the larvae and pupae of M
sumatrana Ulmer from Sumatra and Java (Sunda Islands). Wiggins
( 1977) provided generic characteristics and the description of M.flexuosa
from North America. The other North American Marilia, M. nobsca
Milne, has been associated with the adult (Wiggins 1977) but apparently
no descriptions nor illustrations were published. The pupae of M. major
Muller and M. minor Muller from South America were described by
Thienemann (1905). Unfortunately, the larva of the other Australian
species, M. bola, is not known. The larvae of M. fusca, M.flexuosa and M .
sumatrana share several characteristics: metanotal setal area 1 (=sa\)
sclerites large and rectangular; metanotal sal sclerites small and rec-
tangular; fore tibia approximately same length as tarsus; labrum with 6
long setae across central part; anal claw lacking dorsal accessory hook;
and lateral sclerite of anal proleg edged mesially with stout spines.

There are, however, several characteristics that can be used to dif-
ferentiate these three species. In M. fusca and M.flexuosa, the mesonotal
plate is subdivided into 3 sclerites, whereas in M. sumatrana the mesono-
tal plate is not subdivided. The ventral apotome of M.flexuosa and M.
sumatrana completely separates the genae, but in M. fusca, the ventral

Received April 21, 1989. Accepted June 21, 1989.

-Department of Biology and Microbiology, University of Wisconsin-Oshkosh, Oshkosh,
WI 54901

ENT. NEWS 101(1): 1-8, January & February, 1990

ENTOMOLOGICAL NEWS

apotome separates the genae to approximately the midpoint of the
ecdysial line. The anterolateral corner of the pronotum of M. flexousa
andM. sumatrana is rounded; in M.fusca it is slightly pointed but not pro-
duced into a sharp point as in two other genera (Parthina and Psilotreta)of
North American odontocerids (Wiggins 1977). In M.fusca and M.flex-
uosa, the lateral sclerite of the anal proleg possesses 3 very long setae on
the mesial margin, whereas M. sumatrana possesses a dense cluster of
setae at the apex of the hind tarsus; this cluster of setae is lacking in M.
flexuosa and M.fusca.

The pupae of M.fusca, M. sumatrana, M. major and M. minor can be
separated on the basis of the teeth (serrations) of the anal processes. In M.
sumatrana, several prominent teeth are located on the hooked apex of the
anal process. In M.fusca, a few low teeth are found just proximad of the
apical hook with one small tooth on the inner margin of the hook. The
anal process of M. major has several prominent teeth ending con-
siderably proximad of the apical hook. In M. minor prominent teeth
extend to the base of the apical hook. The pupal mandibles of these four
species are basically similar, i.e. thick basally with a flattened bladelike
distal portion which is strongly attenuated. In M.fusca the attenuated
portion is approximately equal in length to the distal bladelike portion;
in M. major and M. minor, the attenuated portion is less than half the
length of the bladelike portion. The tip of the attenuated portion of the
mandible of M. sumatrana is subdivided into several small points (or
teeth), whereas, in M.fusca, the tip of the mandible is simply pointed.

MATERIALS AND METHODS

The specimens of M.fusca used to describe the immature stages were
collected in the Yarra River near Reefton, Victoria, Australia. Illus-
trations were prepared from specimens preserved in 80% ethanol using a
squared ocular grid in a stereo binocular microscope. For greater detail,
larval and pupal structures were dissected, mounted in Hoyers mount-
ing medium on a microscope slide, and examined using a compound
microscope at 100X-400X magnifications. Measurements were made
with a calibrated ocular micrometer or the squared ocular grid.

LARVA

Overall length of final instar 10-12.5 mm.; body creamy-white (Figure 1).

Head (Figures 2-4): Head capsule (measured from anterior margin of frontoclypeus to
posterior margin of capsule) slightly longer (1.165 mm) than wide (1.0 mm), sides sub-
parallel. Color pattern distinct: very dark brown bands extending from anterior margin on
either side and mostly parallel to frontoclypeal and coronal sutures; mesal portions of
frontoclypeus lighter brown; lateral areas light yellow-brown or tan; ventral and lateroven-

Vol. 101, No. 1, January & February 1990

tral areas very dark brown with a few paler muscle scars on posteroventral areas. Eyes sur-
rounded by pale areas. Very weak lateral carina extending posteriad from near base of
mandibles just dorsad of eye and slightly beyond. Antennae small, just posteriad of dorsal
articulations of mandibles. Ventral apotome elongated, triangular, separating genae for
approximtely one-half their length. Mandibles (Figure 5) stout, pointed, with a few low
subapical teeth on dorsal cutting edge; setae and mesal brush lacking. Labrum (Figure 6)
slightly rounded anteriorly and narrowed posteriorly; 4 pairs of setae arising from dorsal
surface; 2 pairs of curved, pale setae arising from anterior margin.

Thorax (Figures 7-8): Pronotal plates heavily scerotized; dark brown band on either
side of mesal suture forming a distinct mid-dorsal stripe; each plate light brown with dark
brown margins; anterior margin of each plate with 2 long, dark setae and a few short, pale
setae; anterolateral margins slightly pointed, each with a long, dark seta; ventrolateral
margins each with 5-7 setae; a group of 7-9 setae just dorsad of mid-ventrolateral margin
arising from area slightly darker brown than ground-color; an oblique row of 4 long, dark
setae dorsad to this row; posterolateral margin very dark brown and somewhat sinuate.

Each mesonotal plate subdivided into 3 sclerites (anterior, posterior and lateral ) sepa-
rated by distinct sutures. Anterior plates subquadrate, light brown with dark brown mesal
bands; each plate with several short, pale setae along anterior margin and 4 long, dark setae
just posteriad. Posterior plates rectangular, light brown with dark brown mesal bands;
anterolateral corners dark brown and discontinuous with dark brown posterolateral cor-
ners of anterior plates; posterior margins very dark brown, somewhat serrated; each plate
with 5 long, dark setae just posteriad of anterior margin. Lateral plates elongated, dark
brown, each with a few long, dark setae and several shorter, pale setae on anterior half.

Metanotum with 2 pairs of heavily sclerotized plates. Larger, mesal plates subrec-
tangular. each with pale brown margin and lighter central area; each with 10-15 short, pale
setae along anterior margin. 4 long, dark setae just posteriad of anterior margin, and 6 long,
dark setae along posterior margin. Smaller, lateral plates elongate, narrow, each with
approximately 14 short setae.

Prosternite well-developed, brown and irregularly shaped: middle of anterior margin
broadly truncated and produced anteriorly; anterolateral margins pointed; lateral mar-
gins oblique; and posterior margin slightly concave. Mesosternum with 2 pairs of light-
brown, rectangular plates; anterior pair about 1 .5X wider than long; posterior pair narrow,
about 4X wider than long. Metasternum lacking sclerotized plates but with 2 pairs of lateral
and 1 mesal lightly pigmented areas.

Legs (Figures 9-11): Anterior legs short, slightly robust, protrochantin poorly developed,
bluntly rounded at apex from which arises a short seta. Middle legs slightly longer. Pos-
terior legs longer, more slender. All legs with numerous setae. Claws slender, curved, each
with a basal seta.

Abdomen (Figures 1,12): Creamy-white with distinct purple speckling on dorsum and
sometimes fainter purple speckling along midventral line. Dorsal spacing hump of seg-
ment I moderately developed with a pairof setae (1 long. 1 short) on each side. Lateral spac-
ing hump of I with well-developed, light-brown, irregular-shaped sclerite with about 10
vertical rows of filelike ridges; single setae dorsad and ventrad of sclerite. Segments II-VII
with several pairs of very short, pale setae visible only at high magnification. Segments II-
VII with mid-lateral, dense row of very fine, inconspicuous, pale hairs. Segment VIII with a
row of 23-30 bifid lateral tubercles (also termed bifid spiculesby Neboiss 1977) (Figure 12).
Segment IX with a pairof short, lateroventral setae; dorsal sclerite weakly-developed with 2
long, dark and 3 short, pale pairs of setae along posterior margin. Lateral sclerite of anal
proleg well-developed, slightly pigmented; 3 long, dark setae arising from posterior mar-
gin; a single shorter, dark seta dorsad of anal claw and a single, short pale seta dorsad of the
3 setae. Posterior margin of lateral sclerite distinctly serrated with approximately 10 very
dark teeth. Ventral sole plate with two setae. Anal claw lacking dorsal accessor) hook but

ENTOMOLOGICAL NEWS

Figure 1 . Mariliafusca larva, lateral. Figures 2-6: M.fusca larva. 2. head, dorsal; 3. head, ven-
tral; 4. head, lateral; 5. mandibles, ventral; 6. labrum, dorsal. Scale lines: 0.5 mm (Figs. 2-5),
0.1 mm (Fig. 6).

Vol. 101, No. 1, January & February 1990

Figures 7-12: M.fusca larva. 7. thorax, dorsal; 8. pronorum, lateral; 9-1 1. anterior, middle
and posterior legs; 12. terminal abdomen, lateral, A -enlarged lateral tubercles of segment
VIII; B - terminal segment enlarged. Scale lines: 0.5 mm (Figs. 7-12); 0.025 mm (Fig. 12A,
lateral tubercles).

ENTOMOLOGICAL NEWS

with 5-6 stout spines and 6 setae arising from mesal surface; basal two-thirds of claw with
sparse, very tiny spines.

Gills: Paired abdominal gills on segments II-VIII (Figure 1); single or double gills
(each gill arising independently) each with a few to several digitiform filaments, located at
the anterodorsal (AD), anterolateral (AL) or anteroventral (AV) positions depending on
the segment. Gill arrangement: segment II with double AD, AL and AV gills; segment III
with double AD and AV and single AL gills; segments IV and V with double AD and AV
gills; segments VI and VII with single AD and double AV gills (VII AD gill infrequently
absent); segment VIII with single AD (sometimes absent) and AV gills. The number of
digitiform filaments per gill is reduced on posterior segments.

Case (Figure 13): Maximum length 12 mm; anterior and posterior openings circular,
diameters approximately 2 and 1 mm, respectively. Constructed of small sand grains; con-
tour smooth, slightly curved and tapered posteriad. Posterior opening partially occluded
by vertical, silkened projection from ventral margin. Line of transverse discontinuity
(Figure 13, arrow) distinct in some cases, more frequently in cases of early instars.

PUPA

Head (Figure 14); Antennae very long, extending posteriad along body; distal portions
coiled around abdominal apex 5 times. Vertex with paired setae. Frontoclypeus with 2 pairs
of setae. Paired setae mesad of each eye. Labrum subquadrate, anterolateral angles some-
what rounded; 3 pairs of setae at each posterolateral angle, 1 setae along each lateral
margin, and 4 pairs of setae at each anterolateral angle. Mandibles (Figure 15) broad, thick
basally, each with a pair of setae; distally flattened and bladelike with strongly attenuated,
hooked apex; proximal portion of blade with numerous minute serrations on outer
margin.

Legs: Mesotibia and mesotarsi with fringe of long hairs. Fringe lacking on pro- and
metathoracic legs.

Abdomen (Figure 16): Creamy-white with faint purple speckling along mid-dorsum of
segments II-V( Figure 16, arrow, segment II). Dorsum of segment I with 4 pairs of setae and
a narrowband of short, stout denticles along posterior margin. Dorsum of segments II-VI
with 2 pairs of short setae; dorsum of segments VII-VIII each with a pair of short setae
anteriorly and 3 pairs of long, dark setae near posterior margin. Segments II-VII each with
a pair of dorsolateral, longitudinal, pigmented bars each with anteromesal projection;
those of segment VIII connected by anteromesal projections. Ventrolateral longitudinal,
pigmented bars similar but lacking anteromesal projections. Venter of segment VIII with 2
dense patches of hairs along posterolateral margins. Lateral abdominal fringe lacking.
Anterior hookplates (Figure 16.A) on segments III-VII small, oval, each with single,
caudally-directed hook; posterior hookplates (Figure 16.B) on segment V subrectangular.
each with 2 anteriorly-directed hooks. Anal processes (Figure 16.C) long, narrow, slightly
tapered and divergent distally; apices recurved and hooked; a few low serrations just prox-
imad of apex; conspicuous setae lacking but with sparse, short setae along length of
process.

Gills: Abdominal gills on segments II-VII similar to larval gills. Gill arrangements:
segment II with double AD, AL, and AV gills: segment III with double AD and AV and
single AL gills; segment IV with double AD and AV gills; segment V with double (some-
times single) AD and AV gills; segment VI with double AV gills; segment VII with double
(sometimes single) AV gills. The number of digitiform filaments per gill is highest on
anterior segments (e.g. 30 filaments on AD gill of segment II) and decreases on posterior
segments (e.g. 10 on AV gill of segment VII).

Case: Similar to larval case but slightly less tapered and curved. Anterior margin

Vol. 101, No. 1. January & February 1990

Figure 13: Larval case, lateral, with posterior opening; Figures 14-17: M.fusca pupa. 14
head, anterior; 15. left mandible; 16. abdomen, dorsal, A - anterior bookplate, B - posterior
bookplate. C- anal process; 1 7. case, A -anterior opening, B- posterior opening. Scale lines:
1.4 mm (Fig. 13); 0.5 mm (Figs. 14. 16); 0.1 mm (Figs 14 A,B - bookplates): 1.0 nun
(Figs. 17 A. B).

ENTOMOLOGICAL NEWS

(Figure 1 7,A) slightly flared, with opening partially closed by slightly convex, silkened par-
tition attached just inside opening; sand grains embedded in partition; partition not
attached to case ventrally leaving a narrow, crescent-shaped opening; anteroventral mar-
gin of case with numerous teethlike projections. Posterior opening (Figure 17, B) partially
closed by a silkened partition, the ventral margin of which is deflected inward and not
attached to case leaving a narrow, crescent-shaped opening; posteroventral margin of case
also with numerous, teethlike projections. Attachment of case by anterior and posterior
silkened filaments. Pupae frequently encountered in cracks and crevices of submerged
limbs or other objects.

SPECIMENS EXAMINED

Victoria: 8 larvae and 8 pupae, 8 Nov. 1980; 28 larvae, 1 3 Nov. 1980; 2 1 larvae and 29 pupae,
27 Nov. 1980. All specimens collected in Yarra River near Reefton, Victoria, Australia by G.
Drecktrah. Voucher specimens deposited in Department of Entomology, Museum of Vic-
toria, Melbourne, Australia and Department of Biology/Microbiology, University of Wis-
consin, Oshkosh, WI.

ACKNOWLEDGMENTS

I wish to thank Arturs Neboiss, Curator, Department of Entomology. Museum of Vic-
toria, Melbourne, for his assistance and guidance in this study and for identifying the
specimens. Appreciation is extended to the Board of the Museum of Victoria for providing
me the opportunity to work in the Department of Entomology. I also wish to thank John
Morse, Clemsen University, and Glenn Wiggins. Royal Ontario Museum, for providing
suggestions in preparing this manuscript.

LITERATURE CITED

Mosely, M. E. and D. E. Kimmins. 1953. The Trichoptera (Caddis-Hies) of Australia and

New Zealannd. British Museum of Natural History. London. 550 pp.
Neboiss, A. 1977. Atriplectididae, a new caddisfly family (Trichoptera: Atriplectididae).

Pp. 67-73 //; M. 1. Crichton. editor, Proc. of the 2nd Int. Symp. on Trichoptera. Dr. W..

Junk Publishers. The Hague.
Neboiss, A. 1983. Checklist and Bibliography of the Australian Caddis-flies (Trichoptera).

Australian Society for Limnology. Special Publication 5. 132 pp.
Neboiss, A. 1986. Atlas of Trichoptera of the SW Pacific-Australian Region. Dr. W. Junk

Publishers. Dordrecht. 286 pp.

Thienemann, A. 1905. Biologie der Trichopteren-Puppe. Zool. Jb. Syst. 22:523-534.
Ulmer, G. 1955. Kocherfiiegen (Trichoptera) von den Sunda-Inseln. Arch. Hydrobiol.

suppl. 21(3/4):408-608.
Wiggins, G. B. 1977. Larvae of the North American Caddisfiy Genera (Trichoptera).

University of Toronto Press. Toronto. 401 pp.

Vol. 101, No. 1, January & February 1990

A NEW XYELA (HYMENOPTERA: XYELIDAE)
FROM WESTERN UNITED STATES1

David R. Smith2

ABSTRACT: Xyela lata, n. sp., is described from Colorado, Nevada, and Oregon. Adults
were collected from Pinus flexilis and Pinus monophylla.

Larvae of Xyela feed on the developing pollen of the male strobili of
pines. Species of the genus probably occur wherever pines are found.
Adults are small and short-lived and may be found on the larval host
plant or on nearby pollen sources such as the catkins of willow, alder, or
birch. Because of their small size and habits, there are undoubtedly a
number of species yet to be discovered. One unusual new species has
recently come to my attention and is described here.

Burdick's (1961) revision of Xyela included 15 species in the North
American fauna and notes on the biology of some. Since then, two
species have been added (Smith, 1979). Smith (1978) listed 32 world
species.

Xyela lata Smith, new species

Figs. 1-4

Female. — Body length, 2.6-3.3 mm; sheath length, 2. 1-2.3 mm; forewing length 4.0-4.3
mm. Antenna yellowish with 1st segment, inner surface of 3rd segment, and apical filament
usually more brownish. Background color of head yellow with interocellar area, pos-
tocellar area, spot between postocellar area and eye, lines from anterior ocellus to each
antennal socket, line extending from anterior ocellus to interantennal area, clypeal suture,
and apical margin of clypeus black to dark brownish. Background color of thorax yellow
with blackish to dark brown on mesosternum. cervical sclerities. and most of dorsum;
usually yellowish spots on mesonotal lateral lobes and anterior portion of mesoscutellum.
Abdomen blackish with narrow apical margins of segments yellow; ventrally and laterally
dark brownish to partly or mostly yellowish. Legs with coxae, trochanters and most of
femora dark brownish; extreme apices of femora, tibiae and tarsi yellowish. Wings hyaline;
veins and stigma amber. Third segment of maxillary palpus longer than 1st antennal seg-
ment (as 1.3:0.7) and as broad or slightly broader than 3rd antennal segment. Sheath broad,
laterally flattened, with lateral, longitudinal ridge; in lateral view narrow at extreme base
but immediately broadening, ventral margin straight, apex with dorsal margin curving
downward to rounded apex near ventral margin ( Figs. 1,2). Lance and lancet without teeth:
apex of lancet with apico-ventral notch, extreme apex narrowed and pointed at apex, with
about 7 distinct annuli (Figs. 3, 4).

Male. — Unknown.

Holotype. — Female. Nederland. Boulder Col. .Colorado, labeled "Nederland. Colo..
Science Lodge. 9500', 27-VI-61, W.R.M. Mason, Pinus flexilis." In the Canadian National

^Received May 20, 1989. Accepted June 13, 1989.

"Systematic Entomology Laboratory, PS1, Agricultural Research Service. U.S. Depart-
ment of Agriculture, c/o National Museum of Natural History. Washington, D.C. 20560.

ENT. NEWS 101(1): 9-12. January & February. 1990

ENTOMOLOGICAL NEWS

Figs. 1, 2. Xyela lata. 1, Lateral view. 2, Apex of abdomen and sheath.

Vol. 101. No. 1. January & February 1990

Figs. 3, 4. Xyela lata. 3, Lance and lancet. 4, Apex of lancet.

Collection, Ottawa.

Paratypes. — COLORADO: Same data as holotype ( 1 9); same data as holotype except
date. 29-VT-61 (one without host data) (2 9); Nederland, 8500', 18-V1-61, W.R.M. Mason (1
9); Estes Park, 7500'. 19-VI-61. B.H. Poole ( 1 9). NEVADA: Mt. Springs summit, Clark Co.,
V-26-1961, elev. 5400', Pinus monophylla, R.C. Bechtel (1 9). OREGON: Mt. Washington
[Linn Co.), July 13. 1963 ( 1 9). In the Canadian National Collection and National Museum
of Natural History, Washington, D.C.

Hosts. — Adults were collected from Pinus flexilis James and P. monophvlla Torr. &
Frem. Hosts may include several species of pine, as is the case for some other Xyela
species.

DISCUSSION

In Burdick's (1961) key, this species runs to bakeri Konow, buibakeri
is commonly mostly black, the lancet has no apical notch or distinct
annuli, and the sawsheath is less than 2 mm long. The extremely broad,
straight, laterally flattened sheath and the constricted apex of the lancet
are unique loX. lata and are not known in any other Xyela. These unusual
characteristics will immediately distinguish this new species.

Some variation occurs in the amount of yellow, especially on the dor-
sum of the thorax and venter of the abdomen. The mesonotum may be
almost all black to dark brown, or the yellow spots on the lateral lobes
and scutellum may be extensive; the abdomen may be mostly black ven-
trally and laterally, or mostly yellowish.

Collections are from high elevations, as indicated by label data about
5400' or above.

The specific name is from the Latin "latus", referring to the unusually
broad sawsheath.

ACKNOWLEDGMENTS

My appreciation is extended to Henri Goulet, Biosystematics Research Centre, Agri-
culture Canada. Ottawa, for allowing study of the specimens and for reviewing the manu-

12 ENTOMOLOGICAL NEWS

script, and to the following for reviewing the manuscript: H. R. Wong, Canadian Forestry
Service, Edmonton, Alberta; R. E. White and E. E. Grissell, Systematic Entomology
Laboratory, U.S. Department of Agriculture, Washington, D.C.

LITERATURE CITED

Burdick, D J. 1961. A taxonomic and biological study of the genus Xyela Dalman in North

America. Univ. Calif. Publ. Ent. 17: 285-356.
Smith, D.R. 1978. Family Xyelidae, pp. 1-27. In van der Vecht, J. and R.D. Shenefelt, eds.,

Hymenopterorum Catalogus, pars 14. Dr. W. Junk B.V., The Hague.
. 1979. Symphyta, pp. 3-137. In Krombein, K. V. et a/., eds. Catalog of

Hymenoptera in America North of Mexico, Vol. 1. Smithsonian Institution Press,

Washington, D.C.

SOCIETY MEETING OF NOVEMBER 29, 1989
NORTH AMERICAN BIRD BLOW FLIES AND MAGGOTS

(PROTOCALLIPHORA)

Dr. Curtis W. Sabrosky, Speaker

About a quarter of all species of North American perching birds have been found infested
with maggots of blow flies. These blood sucking parasites feed on nestling birds, engorging
themselves two or three times before pupating in the bottom of the nest or falling to the
ground to pupate. Although the iridescent blue-black or coppery-colored adults are rarely
seen except when reared, they have a very long life span and probably overwinter in this
stage even in the north of Canada and Greenland. Dr. Curtis W. Sabrosky, retired from the
USDA Systematic Entomology Laboratory and former president of the Entomological
Society of America, described these little known flies at the society meeting at the Academy
of Natural Sciences of Philadelphia. He has recently coauthored a book on the taxonomy,
biology and ecology of the Protocalliphora.

Twenty-six species of bird blow flies are recognized in North America; however, subtile
key characters make their identification difficult. While the different species are somewhat
stratified vertically in a woodland habitat and have distinctive geographical distributions,
there is little host specialization. For example, one species, P. braueri (Hendel), has been
found in the nests of 42 bird species and nine Protocalliphora have been found in robin
nests. Once 1200 maggots were found in a single hawk nest. Only ground nesting, shore
birds and birds of the lower Mississippi Valley and Gulf Coast seem to be spared from these
parasites.

Dr. Sabrosky 's lecture was attended by 19 members and 6 guests who came early and
stayed late to talk about insects. Perhaps it was the mulled cider or refreshments that put
everyone in a good mood. Among a number of topics discussed in the open forum before
the featured talk was the preservation of habitats by Jane Ruffin. Cape May. New Jersey has
long been recognized as a place where migrating birds and insects pause to feed before con-
tinuing their annual southward migration. The rapid development in Cape May County is
reducing the cover and resources normally used by migrants such as the monarch butter-
fly. Mrs. Ruffin urged members to write letters supporting efforts to preserve existing old
growth and to create new backyard habitats that will sustain migrant birds and insects in a
residential setting.

(Continued on page 19)

Vol. 101, No. 1, January & February 1990 13

A NEW RHADINOCERAEA (HYMENOPTERA:

TENTHREDINIDAE) FEEDING ON ZIGADENUS

(LILIACEAE) FROM SOUTHEASTERN

UNITED STATES1

David R. Smith2, Will McDearman

ABSTRACT: Rhadinoceraea (Veratra)zigadenusae, n. sp., is described from the southeas-
tern United States. Young larvae feed on the developing stamens and pistils and older lar-
vae consume open flowers of Zigadenus densus and Z. leimanthoides. The species is
described and illustrated and notes on its life history are given. Plants of the tribe Veratrae
(Liliaceae) possess alkaloids toxic to many animals, and species of Rhadinoceraea (Veratra)
are among the few herbivores of this tribe.

Sawfly larvae feeding on flowers of Zigadenus densus(Descr.)Fema\d
and Z. leimanthoides A. Gray were discovered by the junior author dur-
ing studies on the relationship between plant fecundity and herbivores.
Examination of the adults associated with these larvae revealed a new
species of sawfly of the genus Rhadinoceraea. This new species resembles
the only other eastern North American species of the genus, Rhadi-
noceraea nubilipennis (Norton).

Eight North American species of Rhadinoceraea were treated by
Smith (1969), four in the typical subgenus and four in the subgenus
Veratra. All occur west of the Rockies except for R. nubilipennis. Known
hosts are Calochortus sp. for R. (R.) nigra (Rohwer) and Veratrum spp. for
R. (V.) aldrichi (MacGillivray), R. (V.) insularis (Kincaid), and R. (V.)
nubilipennis. Zigadenus represents a new host plant for sawflies. All hosts
are in the family Liliaceae.

Rhadinoceraea (Veratra) zigadenusae Smith, new species

Figs. 2, 4, 6, 8, 10

Female. — Length, 7.0-8.0 mm. Black, only outer surfaces of forefemur and foretibia
brownish. Wings uniformly, darkly, black infuscated; veins and stigma black. Third anten-
nal segment slightly longer than 4th. Tarsal claw simple. Sheath as in Figs. 2, 4; 3rd valvula
(sheath) only slightly longer than 2nd valvifer (basal plate). Lancet as in Figs. 2, 6. with
about 10 serrulae, each serrula with 2 or 3 anterior and 5 or 6 posterior subbasal teeth.

^Received April 6, 1989. Accepted April 24. 1989.

"Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart-
ment of Agriculture, c/o National Museum of Natural History, Washington. D.C. 20560.
-Mississippi Museum of Natural Science, 1 1 1 N. Jefferson Street. Jackson, Mississippi
39202.

ENT. NEWS 101(1): 13-19. January & February, 1990

14 ENTOMOLOGICAL NEWS

Male. — Length, 5.5-6.5 mm. Coloration as for female. Third antennal segment subequal
in length to 4th. Genitalia as in Fig. 8.

Larva. — Very similar to that described for R. nubilipennis by Smith (1969), differing only
by the shape of the body tubercles. The body tubercles are lower and more rounded ( Fig. 10)
than the higher and more conical tubercles of R. nubilipennis (Fig. 9).
Holotype. — 9, "Mississippi, Jackson Co., ca. 1 1 mi. N. Pascagoula, on Big Point Road, 2
mi. E Hwy 63, 21 April 1987, Will McDearman"; mesic pine savanna, adult collected on
Zigadenus densus. Deposited in the National Museum of Natural History, Washington,
D.C. by permission of W. McDearman.

Paratypes. — MISSISSIPPI: Same data as for holotype (6 9, 6 cf ); Pearl River Co., Crosby
Arboretum, Dead Tiger Creek Savanna, ca. 2.5 mi. E Picayune, 2 mi. S Hwy 43, 28 April
1987, Will McDearman, mesic pine savanna and pitcher plant bog, adults collected on
Zigadenus densus ( 1 9, 5 cf ); Hancock Co., Crosby Arboretum, Hillside Bog, ca. 3.5 mi. E
Picayune, 0.5 mi S Hwy 43, 28 April 1987, Will McDearman, hillside pitcher plant bog,
adults collected on Zigadenus densus ( 1 9, 1 cf ). Deposited in the following collections; Mis-
sissippi Museum of Natural Science, Jackson; Mississippi Entomological Museum, Mis-
sissippi State University, Mississippi State; National Museum of Natural History, Wash-
ington, D.C.; and Canadian National Collection, Ottawa.

Distribution. — In addition to the above Mississippi localities based on adults, larvae
have been collected from the following localities by WMcD: ALABAMA: Mobile Co., ca. 1
mi. E Hwy 193 and 1 mi N Hwy 193-163 junction on Old Rangeline Rd., 14 May 1987, on Z.
densus in recently burned mesic longleaf pine savanna. NORTH CAROLINA: Caldwell
Co., Grandfather Mtn., ca. 1/2 mi. N Grandfather Mtn. Visitor Center, on trail between
McRae Peak and Attic Window Peak, 5,800 ft. elev., 7 August 1986, on Z. leimanthoides in
heath barren, host plant populations small and widely scattered, ca. 300 plants. SOUTH
CAROLINA: Jasper Co., ca. 1/2 mi. W of Hwy 17 at Switzerland, 17 May \9S1, on Z. densus
in recently burned longleaf pine savanna. WEST VIRGINIA: Tucker Co., Monongahela
National Forest, Dolly Sods, ca. 1 mi. W Red Creek Campground, 3500 Ft. elev. 9 August
1986, on Zigadenus leimanthoides in heath barren and bog, extensive host plant population
of over 2,000 plants.

Remarks. — This species runs to R. nubilipennis in Smith's ( 1969) key to
North American Rhadinoceraea. In general habitus, color, and size it is
similar to that species, but close examination reveals differences in the
female sheath and ovipositor and the male genitalia. The female sheath
is much smaller than that of R. nubilipennis (Figs. 1-4, photographs and
drawings to same scale from similar sized individuals). This is difficult
to compare without having representatives of both species, but valvula 3
(sheath) of R. nubilipennis is about 1 .4 times the length of valvi fer 2 (basal
plate). Valvula 3 of R. zigadenusae is only slightly longer than the length of
valvifer 2. The ovipositor of R. zigadenusae is much shorter than that of/?.
nubilipennis, the former having about 10 serrulae, the latter about 14
(Figs. 1, 2, 5, 6, photographs and drawings to same scale from similar
sized individuals). The serrulae of each species are very similar. The
male is somewhat more difficult to differentiate, and the only differences
noted are in the genitalia. The harpe of/?, zigadenusae has a more distinct
indentation on its inner margin, the apex of the parapenis is somewhat
flatter, and the penis valve is somewhat broader apically than in R.

Vol. 101, No. 1, January & February 1990

nubilipennis (compare Figs. 7, 8).

The specific name is based on the host plant genus.

Hosts and biological notes

Rhadinoceraea nubilipennis occurs only in the northeastern U.S. and
southeastern Canada, south in the Appalachians to North Carolina. Its
host, Veratrum virideAit. is a northern plant and is found mostly at higher
elevations further south. Zigadenus densus and Z. leimanthoides occur in
bogs, flatwoods and mesic savanna primarily on the lower coastal plain
from Louisiana east through Florida, and from southern peninsular
Florida north through New Jersey. Disjunct populations are isolated in
central Tennessee and the mountains of North Carolina and West Vir-
ginia. The junior author has collected larvae of R. zigadenusae on the
coastal plain from Mississippi to South Carolina and from the disjunct
host populations in North Carolina and West Virginia. Host pop-
ulations in the Applachicola basin of Florida and surrounding vicinity

Figs. 1.2. Apex of abdomen, sheath, and exerted saw of female. \, Rhadinoceraea nubilipen-
nis. 2, R. zigadenusae. Photographs of same magnification.

ENTOMOLOGICAL NEWS

^**J'"!-»«-^^?WJ-RWKK^jS)S:b^fjr^;^--rff^

Figs. 3,4, Female sheaths. 5,6, Female ovipositors. 7,8, Malegenitalia.9, 10, Body tubercles
of larvae., 3, 5, 7, 9. Rhadinoceraea nubilipennis. 4. 6, 8, 10, R. zigadenusae.

Vol. 101, No. 1, January & February 1990 17

were surveyed for two years without producing any evidence of sawflies.
Peninsular Florida populations and those from North Carolina north to
Delaware were not surveyed. Adult sawflies were collected only in
Mississippi.

Nothing is known about the life history of/?, nubilipennis except for
its host. However, a related species, /?.fl/Jnc/7/(MacGillivray) in western
North America, oviposits in and feeds on the foliage of Veratrum spp.
This is very different from the habits of/?, zigadenusae which oviposits in
the inflorescence stalk and feeds on the developing stamens and pistils
or open flowers of Zigdenus.

In southern Mississippi, R. zigadenusae emerged and flew in the sp-
ring, usually from late April to early May, which coincided with the
period when the host plant was just beginning to flower. Females ovi-
posited a single egg by inserting the ovipositor into subdermal and corti-
cal plant tissues on the primary inflorescence stalk. Of 20 ovipositing
females observed, all deposited one egg per plant visit and then flew to
another plant. The position and number of eggs was identifiable by a
nodule of swollen plant tissue. In Mississippi, 14 eggs was the maximum
number observed on a single plant. Larvae hatched within 5 to 10 days
and emerged by boring a small exit hole in the stalk. Young larvae
typically moved up the inflorescence and entered a flower bud where
they fed on the developing stamens and pistils. Older larve consumed
open flowers, and secondarily fed on fruits. When all flowers were con-
sumed, larvae ate the dermal tissue of the inflorescence stalk before dis-
persing. In plant populations where the period of ovipositing and hatching
occurred before the inflorescence was fully developed, larvae consumed
unopened flowers as well as most of the inflorescence stem. As a last
resort, larvae would attempt to feed on basal leaves, but less than 1% of all
observed infested plants displayed leaf damage and less than 5% of the
leaf area was damaged. When mature, in mid to late May on the coastal
plain, larvae burrowed shallowly into soil to pupate. Only one genera-
tion was produced each year.

Although larvae did not defoliate host plants or cause plant mor-
tality, floral herbivory caused a direct reduction in plant fecundity. Saw-
fly larvae were observed in 12 of 17 host plant populations in southern
Mississippi and literally all flowers were consumed in the majority of
these host populations. Zigadenus densus and Z. leimanthoides often
occur in the same habitat on the coastal plain with the sympatric Z.
glaberrimus Michx. However, sawflies did not utilize Z. glaberrimus as a
host. The flowering interval between the host species and Z. glaberrimus
is about three months.

Rhadinoceraea zigadenusae cannot be described as a monophagous
tloral herbivore since two host species are known, but the taxonomic dis-

18 ENTOMOLOGICAL NEWS

tinction between Zigadenus densus and Z. leimanthoides has recently
been questioned (McDearman, 1984). The nature of character differen-
tiation among these two species as well as whether Z. leimanthoides
should be reduced to an infraspecific taxon or treated as a synonym is the
subject of ongoing research. Nevertheless, the ecological specialization
of R. zigadenusae is much more restricted than the current host plant
nomenclature may suggest.

The alkaloids of the plant tribe Veratrae, which includes Zigadenus,
Veratrum, and four other genera, are the subject of an extensive literature
(e.g., Kupchan eta!., 1961; Tomko and Voticky, 1973). These compounds,
commonly known as the "veratrum" akaloids, can physiologically act as
neurotoxins (Narahashi, 1975; Jones et al., 1970), as teratogens (Van
Kampen era/., 1969; Keeler, 1971), and as vaso-dilators (Anon, 1975; Page
and Sidd, 1973). Crosby (1971) has reviewed the extremely poisonous
group of ester-alkaloids, their toxicity to a wide variety of insects, and the
history of their agricultural applications.

Considering the ecological implications of veratrum alkaloid research
as well as the role of defensive alkaloids in other plants (Levin, 1976;
Miller and Feeny, 1983; Zuniga et al., 1985) it is remarkable that only a
few of the 64 species of the Veratrae have been reported as hosts to her-
bivorous insects. Moreover, the known herbivores in North America
consist of four species of Rhadinoceraea. The Veratrae-Rhadinoceraea
( Veratra) association suggests that the distinctive alkaloid phytochemis-
try may be involved with sawfly speciation in Rhadinoceraea. Future
studies are planned by the junior author to address fundamental ques-
tions regarding host plant phytochemistry.

ACKNOWLEDGMENTS

We extend our thanks to the following who reviewed the manuscript: H. Goulet, Biosys-
tematics Research Centre, Agriculture Canada, Ottawa; H. R. Wong, Canadian Forestry
Service, Edmonton, Alberta; and R. D. Gordon and E. E. Grissell, Systematic Entomology
Laboratory, USDA, Washington. D.C.

LITERATURE CITED

Anonymous. 1975. The clinical trial of anti-hypertensive drugs. Therapie (Paris): 30(5):

631-652.
Crosby, D.G. 1971. Minor insecticides of plant origin, pp. 177-231. In Jacobsen, M. and

D.G. Crosby, eds.. Naturally Occurring Insecticides. 585 pp., Marcel Dekker, Inc.,

New York.
Jones, S.F., J. Brennan, and J.G. McLeod. 1970. The effect of germine diacetate on

neuromuscular transmission. Proc. Aust. Assoc. Neurol. 7: 61-65.
Keeler, R.F. 1971. Teratogenic compounds of Veratrum califomicum Part 13: Structure of

muldamine. Steroids 18(6): 741-752.
Kupchan, S.M., J.H. Zimmerman, and A. Afonso. 1961 . The alkaloids and taxonomy of

Veratrum and related genera. Lloydia 24: 1-27.

Vol. 101, No. 1, January & February 1990 19

Levin, D.A. 1976. Alkaloid-bearing plants: An ecogeographic perspective. The American

Naturalist 110:261-264.
McDearman, W. 1984. Systematics ofZigadenus densus and Z. leimanthoides and phylo-

genetic implications of breeding systems in the Veratrae. Assoc. Southeast. Biol. Bull.

31(2): 71.
Miller, J.S. and P. Feeny. 1983. Effects of benzylisoquinoline alkaloids on the larvae of

polyphagous Lepidoptera. Oecologia 58: 332-339.
Narashashi, T. 1975. Toxins and drugs that depolarize nerve membranes, pp. 214-261. In

Narashashi, T., ed.. Cellular pharmacology of excitable tissues. 535 pp. Charles C.

Thomas, Springfield, Illinois.
Page, L.B. and JJ. Sidd. 1973. Medical management of primary hypertension. New

England Journal of Medicine Medical Progress Series, New Eng. J. of Medicine.
Smith, D.R. 1969. Nearctic sawflies I. Blennocampinae: Adults and larvae (Hymenoptera:

Tenthredinidae). U. S. Dept. Agric. Tech. Bull. 1397, 179 pp., 19 pis.
Tomko, J.andZ. Voticky., 1973. Steroid alkaloids: The Veratrum and Buxus Groups, pp. 1-

82. In Manske, R.H.F., ed.. The Alkaloids: Chemistry and Physiology. Vol. 14. Academic

Press, New York.
Van Kampen, K..R. etal. 1969. Early embryonic death in ewes given Veratrum californicum.

Amer. J. Vet. Res. 30(4): 517-519.
Zuniga, G.E., M.S. Salgado, and LJ. Corcuera. 1985. Role of an indole alkaloid in the

resistance of barley seedlings to aphids. Phytochemistry 24(5): 945-947.

SOCIETY MEETING OF NOVEMBER 29, 1989

(Continued from page 12)

Kenneth Frank. M.D. commented on the dramatic increase in Ixodes dammini. the tick
vector of Lyme disease. Nymphs of this species were uninvited guests at the AES Insect
Field Day in the New Jersey Pine Barrens last June. A number of participants were
unknowing hosts, although none developed the annular rash associated with Lyme di-
sease. Dr. Frank cautioned that entomologists, who are frequently in the field, are more
likely to contract the disease than the general public and should be aware of the prevention,
recognition and treatment of the disease. He distributed copies of a recent review on Lyme
disease that appeared in the August 31 issue of the New England Journal of Medicine.

Paula Haines described her observations on strepsipterans she had found parasitizing
Polistes wasps. Her request for more information was answered by Dr. Paul Schaeffer who
had worked with Xenos peckii in the past.

Mr. and Mrs. Chaiken, operators of the gift shop at the Academy of Natural Sciences,
displayed a number of insect-related books and gifts with insect motifs. Also available for
examination and purchase were beautifully mounted specimens of the large and showy
insects of the world.

Harold B. White.
Corresponding Secretary

20 ENTOMOLOGICAL NEWS

A NEW SPECIES OF CURTARA (HOMOPTERA:
CICADELLIDAE) FROM BERMUDA 1.2

Paul H. Freytag^

ABSTRACT: A new species of gyponine leafhopper, Curtara bermudensis, is described
from Bermuda and compared with closely related species. This is the first gyponine known
from these islands.

A new species ofCurtara from Bermuda was sent to me for identifica-
tion, and is described at this time and compared with the closely related
species. I wish to thank M. R. Wilson, CAB International Institute of
Entomology, London, for making this series available. This species
belongs in the subgenus Curtara.

The Genus Curtara was described by DeLong and Freytag (1972),
with 76 species included (1976). DeLong and others (DeLong 1977, 1979,
1980, 1983; DeLong and Foster 1982; Delong and Triplehorn 1978, 1979;
and DeLong and Wolda 1978, 1982, 1984) have added an additional 67
species. This new species makes a total of 144 species known for this
genus.

Curtara bermudensis n. sp.

(Figures 1-5)

Length of males 6.4-6.8 mm., females 7.8-8.6 mm. Crown broadly rounded, twice as wide
between eyes at base as median length, margin foliaceous.

Color: Males brown with many dark brown spots on head and pronotum. Wings dark
brown spotted with lighter brown. Females yellow brown with few spots of darker brown on
head, pronotum and forewings.

Male genitalia: Pygofer bluntly pointed at apex. Plate four times as long as wide, trun-
cate at apex. Style broad at base, narrowed to hatchet-shaped apex, ventral margin finely
serrate. Aedeagus with shaft long, tubular, with pair of processes extending along shaft
about half length of shaft, then curving dorsad; paraphysis broadened near middle, with
dorsally hooked apex.

Female genitalia: Seventh sternum produced medially with very small median
emargination.

Holotype male: Paget Parish, Paget Marsh, 14-22 VII 1988. M. R. Wilson & D. J.
Hilburn, in the British Museum. Paratypes: one female, same data as holotype, in the
British Museum; one male; same data as holotype, in the University of Kentucky Collec-
tion; one male, Devonshire Parish, Devonshire Marsh, 14-22 VII 1988, M. R. Wilson & D. J.
Hilburn. in the Bermuda Collection; one male St. George's Parish, 4 VI 1987, D. J. Hilburn,
Ferry Reach on G. Beating; one female, St. George's Parish, 16 II 1967, F. M. Collector, Non

Received May 15, 1989, Accepted June 10, 1989.

-This paper is published with the approval of the Director of the Kentucky Agricultural
Experiment Station as journal article No. 89-7-94.
^Department of Entomology, University of Kentucky, Lexington, KY 40546-0091.

ENT. NEWS 101(1): 20-22, January & February, 1990

Vol. 101, No. 1, January & February 1990

Figures 1-5, Male genital structures of Curtarabermudensisn.sp. 1. ventral viewofaedeagus.
2, lateral view of aedeagus. 3. lateroventral view of style. 4. ventral view of plate. 5. lateral
view of pygofer, plate and valve. All drawn to the same scale, which equals 0.5 mm.

22 ENTOMOLOGICAL NEWS

Such Island, in grass, Bda. Dept. Agr. Coll. No. 323, both in the U.S. National Museum; one
female, Non Such Island, 26 VIII 1966, F. M. Collector, collected on Conocarpus erecta. No.
323.; one female. Pembroke Parish, 22 IX 1966, F. M. Collector, Mount Hill, at light. No.
323; one male, Sandy's Parish, 19 IX 1987, D. Hilburn, General Beating; one male. Smith's
Parish. 6 IX 1987. D. Hilburn. Spittal Pond, Sweeping, all in the Bermuda Collection.

Notes: This species is closely related tosamera DeLong and Freytag
but differs from it by having the males smaller, much darker in color and
the aedeagus with longer processes which curve dorsally. Also, it resem-
bles compacta DeLong but differs from this species by being larger and
the male style being more robust near the middle.

LITERATURE CITED

DeLong, D. M. 1977. Four new species of British Guiana and Brazil Curtara (Homoptera:

Cicadellidae). J. Kansas Entomol. Soc. 50(1): 23-26.
DeLong, D. M. 1979. Species of Gyponinae (Homoptera: Cicadellidae) described by

Spangberg. Brenesia 16: 159-168.
DeLong, D. M. 1980. New species of Curtara (Homoptera: Cicadellidae) from Central and

South America. Brenesia 17:179-214.
Delong, D. M. 1983. New species Curtara (Homoptera: Cicadellidae) from Central and

South America. Proc. Entomol. Soc. Wash. 85(3 ):60 1-606.
DeLong, D. M. & Foster, D. R. 1982. New Species of Bolivian Gyponinae (Homoptera:

Cicadellidae). Entomol. News 93(4): 114-1 18.
DeLong, D. M. & Freytag, P. H. 1972. Studies of the Gyponinae: A key to the known

genera and descriptions of five new genera. J. Kansas Entomol. Soc. 45(2): 218-235.
DeLong, D. M . & Freytag, P. H. 1976. Studies of the world Gyponinae (Homoptera:

Cicadellidae): A synopsis of the Genus Curtara. Brenesia 7: 1-97.
DeLong, D. M. & Triplehorn, B. W. 1978. Four new species of Gyponinae (Homoptera:

Cicadellidae) from Paraguay. Entomol. News 89(7 & 8): 179-182.
DeLong, D. M. & Triplehorn, B. W. 1979. New species of Gyponinae (Homoptera:

Cicadellidae) from Peru. Brenesia 16: 175-188.
DeLong, D. M. & Wolda, H. 1978. New species of Polana and Curtara (Gyponinae)

(Homoptera: Cicadellidae) from Panama. Entomol. News 89 (9 & 10): 227-230.
DeLong, D. M. & Wolda, H. 1982. New species of Curtara. Polana and Acuera (Homop-
tera: Cicadellidae: Gyponinae) from Panama, Peru, Bolivia and Venezuela. Entomol-

ogia Scandinavica 13(3): 301-311.
DeLong, D. M. & Wolda, H. 1984. New Panamanian gyponine leafhoppers (Homoptera:

Cicadellidae) belonging to the Genera Polana and Curtara. Uttar Pradesh J. Zool. 4(1)

22-30.

Vol. 101, No. 1. January & February 1990 23

SPIDER (ARANEAE) TAXA ASSOCIATED WITH

THE IMMATURE STAGES OF MANTISPA
INTERRUPTA (NEUROPTERA: MANTIS PI DAE)1'2

Kevin M. Hoffman , Jeffrey R. Brushwein

ABSTRACT: The immature stages ofMantispa interrupta are associated with 10 species of
spiders, not previously recorded as hosts, from the families Gnaphosidae, Lycosidae,
Ctenidae, and Pisauridae. First-instar mantispids were found on adult and juvenile spiders of
both genders, while later-instar mantispids were located inside spider egg sacs. Larvae on
spiders were located on the edge of the carapace, on the dorsal, ventral, and lateral surfaces
of the pedicel, and inside book lungs. The locations on spiders occupied by M. interrupta
first instars are compared with those occupied by first instars of other mantispine species.

Adult Mantispinae have been reared exclusively from spider egg sacs
(Redborg and MacLeod 1985, Brushwein 1986, Hoffman and Brushwein
1989). Mantispine larvae develop through three instars by feeding on
spider eggs. Pupation occurs within the egg sac and pharate adults exit
both their own cocoons and the spider egg sacs before adult eclosion
occurs. First instars procure spider eggs either by locating and entering
preexisting egg sacs or by locating and boarding female spiders and sub-
sequently entering egg sacs as they are deposited by the spiders. Recent
studies on the spiders associated with particular mantispine species sug-
gest that these associations may be indicative of the egg procurement
methods used by first instars of those mantispine species (Redborg and
MacLeod 1985, Hoffman and Brushwein 1989).

Mantispa interrupta Say has been recorded throughout the eastern
United States and westward into Texas and Arizona (Throne 1972), and
adults have been reared from the egg sacs of three spider species. Smith
( 1 934) reported the emergence of two pupae from two spider egg sacs, one
of which was associated with a female Eris militaris (Hentz) [as Philaeus
militaris (Hentz)]. Subsequently, adult emergence has been reported
from egg sacs of Gnaphosa muscorum (L. Koch) and Lycosa rabida
Walckenaer (Kaston 1940, Rice 1985). In addition to the above records,
M. interrupta has been incorrectly reported as being associated with two
other spider species, but both associations were actually based on species
other than M. interrupta. Eason et al. (1967) associated M. intermpta with
Philodromus aureolus (Clerck) and attributed the association to Auten

^Received February 6, 1989. Accepted May 2, 1989.

-This is Technical Contributon No. 2936 of the South Carolina Agricultural Experiment
Station, Clemson University.

• Department of Entomology, Clemson University, Clemson, SC 29634-0365.
University of Florida, IFAS, Citrus Research and Education Center. 700 Experiment Sta-
tion Road, Lake Alfred. FL 33850.

ENT. NEWS 101(1): 23-28. January & February. 1990

24 ENTOMOLOGICAL NEWS

(1925). However, Auten (1925) reported on the chrysopid Chrysopa inter-
rupta Schneider, notM. interrupta. Killebrew(1982) reported that a man-
tispid which emerged from an egg sac of Peucetia viridam (Hentz) was
possibly M. interrupta. However, this specimen was recently examined by
one of us (KMH) and proved to be a pharate adult of Mantispa sayi
Banks.

Viets ( 194 1 ) supplied the only indication that M. interrupta larvae can
board spiders in order to procure eggs. The laboratory boarding of an
unidentified female lycosid by several first instars was followed by
deposition of an egg sac by the spider and subsequent emergence of an
adult mantispid. Both Redborg and MacLeod (1985) and Brushwein
(1986) reported that M. interrupta has been reared using techniques
developed for two other mantispine species, but no information on
spider associations or boarding was supplied. The present paper reports
further spider associations for M. interrupta, confirms larval boarding
behavior in the field by documenting the presence of larvae on collected
spiders, and compares the locations on spiders of M. interrupta larvae
with those reported for other mantispine species.

METHODS

Eight of the 13 spiders associated with M. interrupta were collected in
pitfall traps from 12 June to 1 August 1988, four were collected by visual
searching, and the remaining association was based on museum speci-
mens. The pitfall traps were located in the Clemson University Experi-
mental Forest surrounding Lake Issaqueena, Pickens County, South
Carolina. The traps were constructed of 2 or 3 liter soft drink bottles
whose tops were cut off and inverted to form funnels. Holes were
punched in the bottoms to aid in water drainage and a few curled leaves
were supplied in each trap to provide harborage sites for trapped spiders.
Forty-four traps were buried to ground level at six locations within a
mixed hardwood-pine forest and checked one to three times each week.
The female Schizocosa saltatrix (Hentz) and egg sac were collected in the
vicinity of Lake Issaqueena on 14 May 1988, while the specimens of
Gladicosa gulosa (Walckenaer) and Varacosa avara (Keyserling) were
collected on 24 July 1988 along a tributary of Toxaway Creek at County
Road 88, Oconee County, South Carolina. The female Pisaurina brevipes
(Emerton) and egg sac were collected on 15 November 1988 near Cork-
screw, Lee County, Florida. The association of M interrupta with Sosip-
pusfloridanus Simon was based on specimens located in the entomo-
logical collections of the Museum of Comparative Zoology (MCZ), Har-
vard University. The spider and its egg sac had been collected by P. J.

Vol. 101, No. 1, January & February 1990 25

Cone in Collier County, Florida, at the junction of 840A and 846, approx-
imately 5 miles east of Immokalee. The pharate adult emerged on 16
April 1968, but collection date of the egg sac was unrecorded.

Identities of the immature stages of M. interrupta were confirmed by
comparisons of first instars with those previously obtained from an adult
female and by allowing later instars to complete development within the
egg sacs in which they were found. Spider egg sacs were identified by
determination of the particular female spider accompanying each sac.
The specimen of V. avara was reared to maturity in order to facilitate a
species-level identification. Spiders were identified both by the use of
selected taxonomic references (Brady 1962, 1979, Carico 1972, Platnick
and Shadab 1975, Dondale and Redner 1978, Peck 1981, Roth 1985) and
with the assistance of A. R. Brady and C. D. Dondale. Voucher speci-
mens of M. interrupta first instars, adults, and associated spiders are
deposited in the Clemson University Arthropod Collection (CUAC),
Department of Entomology.

RESULTS AND DISCUSSION

Eighteen M. interrupta immatures were associated with 13 spiders
representing ten spider species from the Gnaphosidae, Lycosidae,
Ctenidae, and Pisauridae, thereby bringing the total spider taxa associ-
ated with M. interrupta to 13 species in nine genera from five families
(Table 1). These species are predominantly ground-wandering hunters,
although both P. brevipes and E. tnilitaris are somewhat more arboreal
wanderers than the others and S. floridanus builds funnel webs near
ground level.

First instars of M interrupta were associated with adult and juvenile
spiders of both genders, whereas later instars were found inside spider
egg sacs (Table 1). The locations of first instars aboard spiders included
the edge of the carapace, the dorsal, ventral, and lateral surfaces of the
pedicel, and the book lungs. Two of the nine spiders boarded by first
instars had more than one larva. Larvae on these spiders occupied either
different regions of the same structure, e.g., opposite sides of a pedicel, or
similar regions of separate structures, e.g., the left and right book lungs.
Two first instars were dead when collected; one was found in a book lung
along with a live larva and the other was alone in the book lung of a dif-
ferent spider. In the only previous report of M. interrupta first instars
aboard a spider, Viets ( 1941 ) noted that larvae crawled over the body of
the spider upon boarding and that the area between and around the
spinnerets seemed to be preferred. However, the positions of the several
larvae which remained on the spider after 15 days was not mentioned,
and it is possible that some larvae resided in locations similar to those

ENTOMOLOGICAL NEWS

reported above. The presence of first-instar M. interrnpta on spiders,
coupled with the predominantly wandering-mode behavior of the as-
sociated spiders, supports the hypothesis of Hoffman and Brushwein
(1989) that mantispine species which use spider boarding to gain access
to spider eggs will be associated generally with wandering spiders.

Table 1. Spider taxa associated with the immature stages ofMantispa interrupts Super-
families and families are arranged according to the taxonomic list presented by Shear
(1986). (juv. = juvenile).

SUPERFAMILY

Family
Species

CLUBIONOIDEA

Gnaphosidae

Gnaphosa fontinalis

Keyserling
Gnaphosa muscorum

(C.L. Koch)
LYCOSOIDEA
Lycosidae

Gladicosa gulosa

(Walckenaer)
Lycosa acompa

Chamberlin
Lycosa georgicola Wal-
ckenaer

Lycosa rabida Wal-
ckenaer
Lycosa sp., helluo

group
Schizocosa ocreata

(Hentz)
Schizocosa saltatrix

(Hentz)
Sosippus floridan us

Simon
Varacosa avara

(Keyserling)
undetermined
undetermined
Ctenidae

Anahita punctulata

(Hentz)
Pisauridae

Pisaurina brevipes

(Emerton)
SALT1COIDEA
Salticidae

Eris militaris

(Hentz)

Developmental

stage & gender M. interrupta immatures
of spider Instar Number Location

female
egg sac

pupa?

pedicel
egg sac

Reference

this report
Kaston 1940

juv. male

book lung

this report

male

book lung

this report

juv. male
egg sac

1
pupa

5
1

pedicel &
book lungs
egg sac

this report
Rice 1985

juvenile

pedicel

this report

egg sac

this report

egg sac

this report

egg sac

this report

juv. female

carapace

this report

juvenile
juvenile

1
1

pedicel
pedicel

this report
this report

male

pedicel

this report

egg sac

pupa

egg sac

this report

egg sac

pupa

egg sac

Smith 1934

Vol. 101, No. 1, January & February 1990

Of the four other mantispine species known to board spiders, only
first instars of M. sayi [as Mantispa uhleri Banks and Mantispa fuscicornis
Banks, both recently synonymized with M. sayi by Hoffman (1989)]
occupy a range of resting sites on spiders similar to that of M interrupta
first instars (Redborg and MacLeod 1985, Rice 1986). In contrast, first
instars of the other three species occupy a more restricted range of sites.
First instars of Climaciella brunnea (Say) have been found only on the
sternum and the edge of the carapace (Redborg and MacLeod 1983,
LaSalle 1986), while those of both Perlamantispa perla (Pallas) and Man-
tispa pulchella (Banks) occupy only the dorsal surface of the pedicel
(Lucchese 1955, 1956, Hoffman and Brushwein 1989).

In previous reports of M interrupta spider associations, the develop-
mental stages of the mantispids at the time of egg sac collection were not
known. However, some estimates can now be made by comparing the
intervals between egg sac collections and emergences of adult mantis-
pids with the durations of appropriate developmental stages of M inter-
rupta and other North American mantispines. During the present study,
the two M. interrupta which were reared under ambient conditions from
third instars to adult required 17 to 22 days from the onset of cocoon
spinning to adult eclosion. Under standard laboratory conditions. C.
brunnea, M. sayi, and Mantispa viridis Walker spend an average of 14 to 20
days as prepupae and pupae (Redborg and MacLeod 1983, 1985, Brush-
wein 1986). Both Smith (1934) and Rice (1985) reported the emergences
of adult mantispids within four days of egg sac collections, and therefore
these records surely represent field-collected pupae and are listed as
such in Table 1. Because of the 17 day lag between egg sac collection and
adult emergence, the mantispid collected by Kaston (1940) could have
been either a late-instar larva or pupa. Unfortunately, the collection date
of the S.floridanus egg sac was unrecorded, and therefore the develop-
mental stage of the mantispid when initially collected can not be deter-
mined.

ACKNOWLEDGMENTS

Gratitude is extended to Allen R. Brady. Hope College, Michigan, and Charles D. Don-
dale. Biosystematics Research Centre, Ottawa, Ontario, for their assistance in identifica-
tion of the lycosids. We are also grateful to Don W. Killehrew, The University of Texas at
Tyler, and Scott R. Shaw. MCZ, Harvard University, for the opportunity to examine
specimens under their care. We thank Joseph D. Culin and John C. Morse. Clemson
University, Marlin E. Rice, Iowa State University, and one anonymous reviewer for their
helpful comments on this manuscript. Portions of this study were conducted while on
grants from the Ernst Mayr Fund of the MCZ and from the South Carolina Heritage Trust
Program, and this support is gratefully acknowledged.

28 ENTOMOLOGICAL NEWS

LITERATURE CITED

Auten, M. 1925. Insects associated with spider nests. Ann. Entomol. Soc. Am. 18: 240-250.

Brady, A. R. 1962. The spider genus Sosippus in North America, Mexico and Central
America (Araneae: Lycosidae). Psyche 69: 129-164.

1979. Nearctic species of the wolf spider genus Trochosa (Araneae: Lycosidae).

Psyche 86: 167-212.

Brushwein, J. R. 1986. Bionomics of Mantispa viridis Walker (Neuroptera: Mantispidae).
Ph.D. dissertation, Clemson University, Clemson, South Carolina.

Carico, J. E. 1972. The Nearctic spider genus Pisaurina (Pisauridae). Psyche 79: 295-310.

Dondale, C. D. and J. H. Redner. 1978. Revision of the Nearctic wolf spider genus
Schizocosa (Araneida: Lycosidae). Can. Entomol. 110: 143-181.

Eason, R. R., W. B. Peck, and W. H. Whitcomb. 1967. Notes on spider parasites,
including a reference list. J. Kans. Entomol. Soc. 40: 422-434.

Hoffman, K. M. 1989. Taxonomic status ofMantispa sayi, Mantispa fuscicomis. and Man-
tispa uhleri (Neuroptera: Mantispidae). Proc. Entomol. Soc. Wash. 91: 637-639.

Hoffman, K. M. and J. R. Brushwein. 1989. Species of spiders (Araneae) associated with
the immature stages of Mantispa pulchella (Neuroptera: Mantispidae). J. Arachnol. 17:
7-14.

Kaston, B. J. 1940. Another Mantispa reared. Bull. Brooklyn Entomol. Soc. 35: 21.

Killebrew, D. W. 1982. Mantispa in a Peucetia egg case. J. Arachnol. 10: 281-282.

LaSalle, M. W. 1986. Note on the mantispid Climaciella brunnea (Neuroptera: Mantis-
pidae) in a coastal marsh habitat. Entomol. News 97: 7-10.

Lucchese, E. 1955. Ricerche sulla Mantispa perla Pallas (Neuroptera Planipennia - Fam.
Mantispidae). Nota preventiva su nuovi reperti concernenti 1'etologia della larva della
la eta. Ann. Fac. Agrar. Univ. Stud. Perugia 11: 242-262.

1956. Ricerche sulla Mantispa perla Pallas (Neuroptera Planipennia - Fam.

Mantispidae). 11° Contribute su nuovi reperti biologici e morfologici concernenti
1'uovo, la larva della la eta e la completa evoluzione di questa nella sua sede defmitiva.
Ann. Fac. Agrar. Univ. Stud. Perugia 12: 83-213.

Peck, W. B. 1981. The Ctenidae of temperate zone North America. Bull. Am. Mus. Nat.
Hist. 170: 157-169.

Platnick, N. I. and M. U. Shadab. 1975. A revision of the spider genus Gnaphosa
(Araneae: Gnaphosidae) in America. Bull. Am. Mus. Nat. Hist. 155: 1-166.

Redborg, K. E. and E. G. MacLeod. 1983. Climaciella brunnea (Neuroptera: Mantis-
pidae): a mantispid that obligately boards spiders. J. Nat Hist. 17: 63-73.

1985. The developmental ecology of Mantispa uhleri Banks (Neuroptera: Man-
tispidae). 111. Biol. Monogr. 53, 130 pp.

Rice, M. E. 1985. Spiderling survival in a Mantispa (Neuroptera, Mantispidae) infested egg
sac. J. Arachnol. 13: 139-140.

1986. Communal oviposition by Mantispa fuscicomis (Say) (Neuroptera: Man-
tispidae) and subsequent larval parasitism on spiders (Arachnida: Araneida) in south
Texas. J. Kansas. Entomol. Soc. 59: 121-126.

Roth, V. D. 1985. Spider Genera of North America. Am. Arachnol. Soc. 176 pp.

Shear, W. A. 1986. Taxonomic Glossary, pp. 403-432. In Spiders - Webs, Behavior, and
Evolution. W. A. Shear (ed.). Stanford University Press, Stanford.

Smith, R. C. 1934. Notes on the Neuroptera and Mecoptera of Kansas, with keys for the
identification of species. J. Kans. Entomol. Soc. 7: 120-145.

Throne, A. L. 1972. The Neuroptera - suborder Planipennia of Wisconsin. Part III - Man-
tispidae, Ascalaphidae, Myrmeleontidae and Coniopterygidae. Great Lakes Entomol.
5: 119-128.

Viets, D. 1941. A biological note on the Mantispidae (Neuroptera). J. Kans. Entomol. Soc.
14: 70-71.

Vol. 101, No. 1. January & February 1990 29

THE RELATION BETWEEN HEARING
AND FLYING IN CRICKETS1

Daniel Otte2

ABSTRACT: Hearing and flying are closely coupled functions in true crickets (Gryl-
loidea). Although the auditory tympana have been lost many times independently in cric-
kets, they are virtually never lost in species that can fly. Since crickets migrate at night it
seems likely that tympana are retained to avoid bat predation.

The ancestor to modern crickets (Grylloidea) probably possessed
tibial auditory tympana and tegminal stridulatory devices similar to
those of modern species. Subsequently the stridulum (and therefore
acoustical communication) has been lost many times. In Australia, for
example, where 103 species (or 18.9% of the known fauna) lack a stri-
dulum, the stridulum was lost at least 27 times. In Africa it was lost at
least 17 times. It was lost a number of times on Pacific islands as well (i.e.
New Caledonia, Hawaii, Fiji, and Lord Howe — Otte, Alexander and
Cade 1988, Otte and Rentz 1985, Otte and Rice in prep.)

Although the stridulum has not been lost in any United States species,
calling behavior has been lost in at least seven species (Gryllus ovisopis
[Gryllinae], Oligocanthropus prograptus [Mogoplistinae], Tafalisca lurida
and Hapithus brevepennis [Eneopterinae], Falcicula hebardi [Trigoni-
diinae], Scapteriscus abbreviates [Gryllotalpinae] and northern pop-
ulations of Hapith us agitator [Eneopterniae]) (Walker 1974). Absence of a
calling song must be a precurser to the loss of the stridulum; therefore
examination of these species should give one clues as to the selective forces
causing muteness and deafness.

Walker (1974) notes also that Gryllus fultoni on Key Largo lacks a
functional calling song. And in the sibling pair Hapith usmelodius and//.
brevipennis, the former species retains both calling and courtship songs,
while the latter has never been heard to produce either song, even though
it appears to have a functional stridulum.

The circumstances which cause non-acoustical methods of com-
munication to entirely replace acoustical modes is open to speculation.
Do acoustical signals lose their directionality, and therefore effective-
ness, in certain situations (caves, burrows), or become ineffective in the
presence of noise (sea shores), or less effective than other modes of signal
transmission (pheromones, substrate vibration, visual signals) under
certain circumstances (burrows, caves, or on grasses)?

Received June 6, 1989. Accepted July 12, 1989.

~The Academy of Natural Sciences. 19th and the Parkway. Philadelphia. Pennsylvania
19103

ENT. NEWS 101(1): 29-34, January & February. 1990

30 ENTOMOLOGICAL NEWS

Walker (1974), noting that all U.S. mute species are flightless, specu-
lates that muteness is associated with their higher degree of sedentari-
ness: "Sedentary populations characteristically occupy relatively per-
manent habitats and are not subject to the extreme fluctuations in den-
sity of breeding adults characteristic of temporary habitats. Consequen-
tly, chance encounters or short-range signals become more dependable
pair-forming techniques." He suggests two other possible causes: acous-
tically orienting predators could select against singing males; or the loss
of song could evolve if it caused reduction in interbreeding with a closely
related species possessing a nearly identical song (as may have hap-
pened to Gryllus ovisopis when it speciated from G.fultoni).

In Australia mute species are best represented among the following
groups: a) burrowing crickets from rain-forests and open woodlands
(most belong to the genus Apterogryllus [Brachytrupinae] and all pro-
bably derive from a single mute and wingless ancestor); b) crickets
inhabiting lush grasses along water courses (all are small Trigonidiinae,
but the stridulum may have been lost 3 or 4 times in this group); c) cric-
kets inhabiting canopy foliage in rain forest or seasonally wet woodland
(these belong to three groups: Trigonidiinae (Amusurgus, Metiochodes,
Cyrtoxiphoides); Podoscirtinae (Mundeicus, Aphonoides); and Pentacen-
trinae; d) crickets inhabiting seasonally moist grasslands (Euscyrtinae
— probably lost at least twice); e) crickets inhabiting ant nests (Myr-
mecophilinae — muteness probably evolved but once in this group,
perhaps outside Australia); 0 forest floor (leaf litter) species (Nemo-
biinae and Phalangopsinae — stridulum perhaps lost three times in the
Nemobiinae and two or three times in the Phalangopsinae); g) shore-
inhabiting species (Nemobiinae, Apternonemobius — since this genus is
widespread over the Pacific ocean, the stridulum may have been lost out-
side Australia; (h) cave-inhabiting species (Nemobiinae, Nambungia}.

A survey of the Australian crickets shows that auditory tympana are
often retained after the tegminal stridulatory mechanism is lost, that is,
they continue to hear after becoming mute (Amusurgus, Metiochodes
[Trigonidiinae] Pentacentrus [Pentacentrinae] Mundeicus, Umbulgaria,
Aphonoides, [Podoscirtinae], Euscyrtus, and Patiscus [Euscyrtinae]. Par-
tial loss of the stridulum is seen in Hemiphonus, Unka (Podoscirtinae),
Trigonidomorpha (Trigonidiinae) and Merrinella (Euscyrtinae). Since a
loss of the sound producing mechanism is probably usually accom-
panied (or followed) by a loss in the listening mechanism, one must pre-
sume that some kind of selection pressure opposes the loss of a tympanum
in these species.

In my survey of the Australian fauna (based on Otte and Alexander
1983) I noticed that virtually all flying species retain a tympanum, even
those species that have lost the stridulum. The only non-hearing crickets

Vol. 101, No. 1, January & February 1990 31

are ones that cannot fly. We can look at the relation between singing:',
hearing4, and flying5 in the Australian fauna more closely. All the possi-
ble combinations of these three characters are shown in Figure 1 and
next to them the number and the percentage of species possessing the
condition. Two very common conditions occur in this fauna: A large
number of species can sing, hear, and fly. But a much larger number of
species can sing and hear, and are flightless (B). Of the remainder, 57
species (or 20% of the total) cannot fly, cannot hear and cannot sing (H);
46 species (10%) cannot sing but can both fly and hear (D). Notice that
two of the possible combinations have no representative species: There
are no species which can sing and fly but cannot hear (C) and there are
no mute and deaf species which can fly (G). Condition C may be absent
for two reasons: a) Perhaps a species which cannot hear will not retain its
song mechanism. This is probably true in most cases, but rare instances
do occur in which males continue to sing even though a stridulum is lost
(as in condition F — Evans 1988). b) The condition is rare because flying
species are selected against if they are deaf. The latter reason is probably
true, given that there are no species which can fly and which cannot also
hear (see also condition G).

Few species have conditions E and F; both conditions are probably
transitional between B and H. Outside Australia I know of no species
with either of these conditions. Of the four Australian species which
have lost the stridulum but retain the tympanum, one species has a tiny,
apparently rudimentary, organ; another species is geographically vari-
able with western Australian specimens retaining a tympanum, and eas-
tern Australian ones having lost it (perhaps in this species occasional
individuals are macropterous).

Condition F, in which the stridulum is retained in a species that has
lost its tympana, is also exceptional. Fortunately the acoustical behavior
of one of these species is known. Males ofBalamarogidya have a peculiar
mode of signalling (Evans 1988). In the presence of females they tap the
grass on which they rest with the abdomen, presumably transmitting
information to the females through vibration of the subtrate. Males tap
in pairs, and during the first tap of each pair they stridulate. We do not
know what B. gidyas nearest relatives do. B. marroo possesses both a
stridulum and well-developed tympana. B. albovittata from eastern Aus-
tralia has no stridulum and no tympana; in western Australia this species
has no stridulum but retains a small tympanum. We speculate that in the
lineage leading to B. gidya males called females by stridulating; later,
males began to vibrate the substratum (grass blades on which both were

have a tegminal stridulatory mechanism
^have a tihial auditory tympanum
-possess long hind wings

ENTOMOLOGICAL NEWS

perched) during stridulation; gradually tapping began to predominate
as the information carrier, and tympana were lost when the acoustical
component of the signal disappeared; the stridulation now heard faintly
during the first tap is perhaps a vestige of the original call. Once it disap-
pears, as it may already have done in B. albovittata, the stridulum may
also be lost.

One is led to conclude that hearing is important to flying crickets.

SINGING

HEARING

FLYING

loss of

flying

wings

SINGING

HEARING

no flying

SINGING

no hearing

FLYING

46 (9.3)

no singing

HEARING

FLYING

no singing

no hearing

FLYING

no singing

HEARING

no flying

loss of
tympanum

SINGING

no hearing

no flying

57 (11.5)

no singing

no hearing

no flying

loss of
stridulum

Figure 1 . All possible combinations of singing, hearing, and flying in the known Australian
cricket fauna. Singing means possession of a stridulatory mechanism; hearing means
possession of a tibial auditory tympanum', flying means possession of hind wings long
enough to make flight possible. Numbers in the upper right indicate the number (and per-
centage) of species possessing the condition. Arrows indicate the presumed direction of
evolutionary change. Parallel lines indicate evolutionary transformations which have pro-
bably not occurred — a and b because absence of hearing in flying species is perhaps
strongly selected against by bats or other predators; c, d, e, and f because once a particular
complex mechanism is lost it is highly unlikely to evolve again. Conditions E and F are
absent in African and other faunas, suggesting that the transition from B or D to H is
rapid.

Vol. 101, No. 1, January & February 1990 33

Africa and Pacific island crickets hold to this pattern precisely, though
we have discovered a single species of Adenopterus (Loyalty Islands)
which possesses long hind wings and lacks tympana (Otte, Alexander
and Cade 1987).

The association between hearing and flying is also evident within
some Australian species which show developmental flexibility in both
characteristics — further confirming a functional coupling between
them. Euscyrtus hemelytrus (Euscyrtinae) always possesses tympana, but
the outer tympanum is sometimes obsolete in micropterous specimens.
In general, macropterous individuals have larger tympana. We noted the
following variation in this species (Otte and Alexander 1983): Microp-
terous individuals are quite variable, with the outer tympanum some-
times obsolete and represented only by a dimple. Macropterous males
had very large and conspicous inner and outer tympana. A macrop-
terous male and female from Upper Burma have very large inner and
outer tympana, but a micropterous female has inner and outer tympana
barely visible. A micropterous female from Manila, Philippines has an
outer depression and a conspicuous inner tympanum. Two microp-
terous males and a female from Assam (Bangladesh) have an inner tym-
panum and the other tympanum represented by a dimple. Four other
females from the same place are macropterous and have large inner and
outer tympana.

Metioche vittaticollis (Trigonidiinae) are similar to Euscyrtus. Macrop-
terous individuals posses large tympana, while micropterous individuals
have either small inner and no outer tympana, or small outer and larger
inner tympana, or no tympana at all. Of 61 individuals examined only
the 45 macropterous individuals had prominent inner and outer tym-
pana.

We also noted dimorphism in wings and tympana in Trigonido-
morpha sjostedti (Otte and Alexander 1983). In this species wingless
males and females usually have small dimples instead of tympana:
occasionally a small inner tympanum is visibile. Evans (1988) has fur-
ther studied wing length and tympana in this species. She crossed the two
phenotypes (winged/ +tympana and wingless/ -tympana) and found
that the dimorphism does not result from a simple Mendelian 1-locus. 2-
allele mechanism, since all crosses produced both phenotypes in both
sexes. Field collected wingless/-tympana adults produced winged/
+ tympana offspring. She also found that both morphs possessed tym-
panal organs with well developed scolopidia, attachment cells, and
accessory cells, in close proximity to the anterior tympana trachea; but
the tympana of the wingless morph were hidden beneath a layer of
cuticle.

34 ENTOMOLOGICAL NEWS

Several experimental studies have shown that crickets can hear bat
sounds and take evasive action in the presence of bat sounds (Griffin
1958, Popov and Mrkovich 1982, Moiseff and Hoy 1983, Nolen and Hoy
1984, Doherry and Hoy 1985).

It has also been suggested that the development of receptors sensitive
to aerial sounds may have been favored by selection if they enabled
individuals to stay within a dispersing group by responding to the flight
sounds of conspecifics (Evans 1988, and references therein).

LITERATURE CITED

Alexander, R.D. 1968. Life cycle origins, speciation, and related phenomena in crickets.
Quarterly Review of Biology. 43:1-41.

Alexander, R.D. and G.H. Meral. 1967. Seasonal and daily chirping cycles in the
northern spring and fall field crickets, Gryllus veletis and G. pennsylvanicus. Ohio Jour-
nal of Science. 67:200-209.

Doherty, J. A. and R. R. Hoy. 1985. Communication in insects. III. The auditory behavior
of crickets: some views of genetic coupling, song recognition, and predator detection.
Quarterly Review of Biology. 60:457-472.

Evans, A. 1988. Mating systems and reproductive strategies of some Australian crickets
(Orthoptera: Gryllidae). Ph.D. Thesis, Univ. of Melbourne.

Griffin, D. R. 1958. Listening in the Dark. New Haven, Yale University Press.

Moiseff, A. and R. R. Hoy. 1983. Sensitivity to ultrasound in an identified auditory inter-
neuron in the cricket: a possible neural link to phonotactic behaviour. Journal of Com-
parative Physiology. 152: 155-167.

Nolan, T. C. and R. R. Hoy. 1984. Initiation of behavior by single neurons; the role of
behavioral context. Science. 226:992-994.

Otte, D. 1977. Communication in Orthoptera. How Animals Communicate. Blooming-
ton, Indiana University Press.

Otte, D. 1989. Speciation in Hawaiian crickets. Speciation and its Consequences. Sun-
derland, Mass., Sinauer Associates.

Otte, D. and R. D. Alexander. 1983. The Australian Crickets. Academy of Natural
Sciences of Philadelphia Monograph 22:1-477.

Otte, D. and R. D. Alexander and W. Cade. 1987. The crickets of New Caledonia
(Gryllidae). 139:375-475.

Otte, D. and D. C. Rentz. 1985. The crickets of Lord Howe and Norfolk Islands (Orthop-
tera: Gryllidae). Proceedings of the Academy of Natural Sciences of Philadelphia.
137:79-101.

Otte, D. and R. Rice, in prep. The Hawaiian Crickets: Systematics, Biogeography and
Speciation. Special Publications, The Academy of Natural Sciences, Philadelphia.

Popov, A. V. and A. M. Markovich. 1982. Auditory interneurons in the prothoracic
gangion of the cricket, Gryllus bimaculatis. II. A high frequency ascending neuron
(HF IAN). Journal of Comparative Physiology. 146:351-359.

Walker, T. J. 1974. Gryllus ovisopis, n.sp.: a taciturn cricket with a life cycle suggesting
allochronic speciation. Journal of Comparative Physiology. 57:13-22.

Vol. 101, No. 1, January & February 1990 35

THE LEPTOCERIDAE (TRICHOPTERA) OF
WEST VIRGINIA1

James B. Glover, Donald C. Tarter2

ABSTRACT: Over 9,000 adult leptocerids were identified from light trap samples through-
out West Virginia. A total of 27 species, including 25 new state records, are reported for West
Virginia. The most common species in the state included Cerac/ea cancellata, C. maculata.
Oecetis avara, and O. inconspicua. Species that were abundant only in the larger rivers of
the western portion of the state included C. /7ava, C. macu/afa, Nectopsyche pavida, and
Triaenodes ignitus. Cerac/ea ne/77 was most abundant in the smaller, cooler, fast-flowing
streams at the higher elevations in the eastern part of the state. Cerac/ea ophioderus (IL.
NC, SC), C. slossonae (FL, GA. PA, VA), and C. wetzeli (MI, PA) were important range
extensions into West Virginia. The collection period ranged from 30 April (O. inconspicua
and T. ignita) to 17 October (O. c/nerascens).

Prior to this study, only two species ofleptocerids had been reported
from West Virginia: Oecetis avara (Banks) (Ross, 1944) and Mystacides
sepulchralis (Walker) (Yamamoto and Wiggins, 1964). Based on the
examination of 9,136 adult leptocerids from black light and Malaise
traps, 27 species, including 25 new state records, are recorded from
West Virginia.

Three important range extensions are noted: Cerac/ea ophioderus
(Ross) (IL, NC, SC); C. slossonae (Banks) (FL, GA, PA, VA); and C. wet-
zeli (Ross) (MI, PA).

The collection period ranged from 30 April, Oecetis inconspicua
(Walker) and Triaenodes ignitus (Walker), to 17 October, O. cinerascens
(Hagen). The only species that was collected exclusively in the early part
of the year (30 April- 6 June) was T. ignitus. One male of C. wetzeli was
collected on 14 May. Some species had extended flight periods begin-
ning in May and extending into September.

Detailed collecting data are found in Glover (1988). All adults are
stored in the West Virginia Benthological Survey at Marshall Univers-
ity.

Collecting Stations

Collecting stations are listed alphabetically by county (in caps) and
each station is assigned a number. The station numbers given below are
shown on Figure 1. Stations number 34 and 47 are listed only as county
because of incomplete information and are not shown on map. An

^Received December 12, 1988. Accepted May 20, 1989.
^Department of Biological Sciences, Marshall University. Huntington,
West Virginia 25701

ENT. NEWS 101(1): 35-38, January & February, 1990

36 ENTOMOLOGICAL NEWS

annotation of the months of collection for each species is given in the
species list.

BOONE: 1. Coal River; 2. Madison.

BRAXTON: 3. Burnsville Lake; 4. Falls Mill; 5. Little Kanawha River; 6. Sutton Lake.

CABELL: 7. Greenbottom Swamp; 8. Guyandotte River; 9. Ona; 10. Salt Rock.

GRANT: 1 1. North Fork of the South Branch of Potomac River; 12. South Branch of the

Potomac River.

HAMPSHIRE: 13. Romney.

HANCOCK: 14. Newel.

HARDY: 15. Howards Lick Run; 16. Lost River; 17. Wardensville.

HARRISON: 18. Bridge Port.; 19. East View.

JACKSON: 20. Ravenswood; 21. Ripley.

JEFFERSON: 22. Harpers Ferry; 23. Kearneysville; 24. Shanandoah Junction.

KANAWHA: 25. Charleston; 26. Coal River; 27. Guthrie; 28. London Locks; 29.

Marmet Locks.

MASON: 30. Flatfoot Creek; 31. Gallipolis; 32. Lakin; 33. McClintic Pond #16; 34. Ohio

River; 35. Point Pleasant.

McDOWELL: 36. lager.

MONONGALIA: 37. Triune.

MONROE: 38. Hollywood; 39. Union.

MORGAN: 40. Berkely Springs (Yamamoto and Wiggins, 1964).

NICHOLAS: 41. Gauley River; 42. Summersville Lake.

PENDLETON: 43. Seneca Rock; 44. Smoke Hole Camp.

POCAHONTAS: 45. Cranberry Glades; 46. Durbin; 47. Greenbrier River; 48. Marling-

ton

PRESTON: 49. Cranesville.

PUTNAM: 50. Hurricane; 51. Winfield Locks.

RALEIGH: 52. Beckley: 53. Daniels.

RANDOLPH: 54. Valley Bend.

RITCHIE: 55. Hughes River.

SUMMERS: 56. Bluestone River; 57. Bluestone Lake; 58. Hinton; 59. Pipestem.

TAYLOR: 60. Tygart River.

TYLER: 61. Middlebourne.

WAYNE: 62. Dickson dam; 63. Fort Gay; 64. Prichard; 65. Twelvepole Creek.

List of Species

Ceraclea cancellata (Betten). Stations 1, 3, 6, 1 1, 14, 21, 22, 23, 24, 25, 26, 28,

29, 30, 31, 34, 38, 43, 46, 47, 48, 50, 52, 58, 62. 4 June - 14 August.
Ceraclea diluta (Hagen). Station 46. 5 July; one male.

Ceraclea flava (Banks). Stations 25, 27, 28, 32, 42. 6 June - 23 July.
Ceraclea maculata (Banks). Stations 3, 6, 14, 20, 21, 22, 23, 25, 26, 27, 28, 29,

30, 32, 34, 37, 38, 39, 50, 51, 58, 62, 63, 64. 8 June - 15 September.
Ceraclea neffi (Resh). Stations 5, 6, 43, 46, 57, 58, 59. 6 June - 26 August.
Ceraclea ophioderus (Ross). Station 58. 6 June - 22 August.

Ceraclea slossonae (Banks). Station 13. Collection date unknown.
Ceraclea tarsipunctata (Vorhies). Stations 4, 24, 25, 26, 27, 28, 58, 62, 63, 64. 7
June - 22 July.

Vol. 101, No. 1, January & February 1990

•39 •—

Fig. 1. Collection sites for adult Leptoceridae in West Virginia.

38 ENTOMOLOGICAL NEWS

Ceraclea transversa (Hagen). Stations 4, 26, 28, 31, 38, 42, 46, 58. 29

June - 1 September.

Ceraclea wetzeli (Ross). Station 44. 14 May; one male.
Leptocerus americanus (Banks). Station 55. 14 July; one male.
Mystacides sepulchralis (Walker). Stations 6, 13, 40. 5 July - 4 August.
Nectopsyche Candida (Hagen). Station 31.2 June - 20 August.
Nectopsycheexquisita (Walker). Stations 6, 8, 23, 27, 34, 38, 43, 58, 59, 62, 63.

1 June - 10 August.
Nectopsyche pavida (Hagen). Stations 5, 22, 34, 50, 63. 3 July - 22

August.
Oecef/jovara (Banks). Stations 4, 6, 12, 17, 22, 25, 27, 28, 38, 41,42, 43, 44, 56,

58, 59. 5 June - 20 September.

Oecetis cinerascens (Hagen). Stations 22, 33, 34, 54. 7 July - 17 October.
Oecetis inconspicua (Walker). Stations 1, 2, 6, 7, 9, 10, 12, 13, 15, 16, 18, 19,

22, 23, 24, 26, 27, 30, 33, 34, 35, 36, 37, 38, 39, 42, 43, 44, 45, 49, 50, 52, 53,

54, 58, 59, 60, 61, 62, 63, 65. 30 April - 23 September.
Oecetis noctuma Ross. Stations 11, 26, 28, 30, 43, 46, 64. 26 June - 10

September.
Oecetis persimilis (Banks). Stations 6, 9, 20, 46, 58, 59, 62. 1 June - 17

August.

Setodes incerta (Walker). Stations 58, 59. 25 June - 26 August.
Triaenodes flavescens Banks. Stations 38, 59. 30 July - 31 August.
Thaenodes ignitus (Walker). Stations 7, 62. 30 April - 6 June.
Triaenodes injustus (Hagen). Stations 27, 50, 54, 58. 20 June - 2 August.
Triaenodes marginatus Sibley. Stations 58, 59. 1 July - 19 August.
Triaenodes pernus Ross. Station 58. 18 June; one male.
Triaenodes tardus Milne. Station 50. 5 August; two males.

ACKNOWLEDGMENTS

The authors are grateful to the following persons and institutions for the loan of
specimens: Linda Butler, West Virginia University; Fred Kirchner. U.S. Army Corps of
Engineers; Jan Hacker and Brian Hagenhuch, West Virginia Department of Agriculture;
and John Morse, Clemson University. A special note of thanks to Oliver S. Flint. Jr..
Curator of Neuropteroids, U.S. National Museum, for help in the identification ofthecad-
disflies. We thank Dean Adkins and Weldon Burrows for critically reviewing the manu-
script. The authors are thankful to Lu Ann South for typing the manuscript.

LITERATURE CITED

Glover, J.B. 1988. A taxonomic and distributional study of the adult caddisflies of the

family Leptoccridae (Insecta: Trichoptera) of West Virginia. Unpuh. M.S. thesis,

Marshall Univ.. Huntington, WV.
Ross, H.H. 1944. The caddis flies, or Trichoptera. oflllinois. Bull. Illinois Nat. Hist. Surv.

23:1-326.
Yamamoto, T., and G.B. Wiggins. 1964. A comparative study of the North American

species in the caddisfly genus Mystacides (Trichoptera: Leptoceridae). Canad. J. Zool.

42:1105-1126.

Vol. 101, No. 1, January & February 1990 39

SOME ECTOPARASITES OF BATS FROM
HALMAHERA ISLAND, INDONESIA1

B.V. Peterson2, L.A. Durden3, J.E. Keirans4, P.M. Taylor5

ABSTRACT: New host and distribution records are given for some ectoparasites of the
families Nycteribiidae. Streblidae, Ischnopsyllidae(Insecta).Argasidae, Laelapidae. Spin-
turnicidae and Trombiculidae (Acari) removed from eight species of bats collected on the
Indonesian island of Halmahera.

The ectoparasites upon which this report is based were collected as
part of an ethnographic field research study conducted by P.M. Taylor
among Tobelo-speaking peoples of Halmahera Island, Indonesia, from
December 1980 to November 1981, and was sponsored locally by
Khairun University, Ternate, Indonesia. The study was carried out with
the cooperation of the Indonesian Institute of Sciences, and the Indone-
sian Nature Conservancy. Biological specimens were collected through-
out the area inhabited by the Tobelo people in order to record local
information on the native classification and uses of animals and plants
and to provide material for zoological and botanical investigations
(Taylor, 1985, in press). Halmahera (sometimes referred to as "Jilolo" or
"Gilolo") is the largest island of the Moluccas. It lies on the Equator and
is situated southeast of the Philippines, west of the western tip of Irian
Jaya, and North of Seram. Knowledge of the ectoparasitic fauna of bats
from the Moluccas and surrounding territories, and especially Halma-
hera, is meager, therefore the records from this survey are valuable for
inventory purposes and also for elucidating our knowledge of host-
parasite associations in this part of the world.

MATERIALS AND METHODS

Ectoparasites taken during this study were removed from 1 55 bats of
8 species, plus 16 unidentified bats, mist-netted on the Indonesian island
of Halmahera (KampungPasirPutihJailolo District, aKraK 127°41T£)
by P.M. Taylor and A.C. Messer in 1981. An additional specimen of

'Received April 3, 1989. Accepted May 23. 19X9.

-17229 Founders Mill Drive. Rockville. MD 20855.

-Department of Entomology, Museum Support Center. Smithsonian Institution.
Washington. D.C. 20560

4National Institute of Allergy and Infectious Diseases, National Institutes of Health, c/o
Department of Entomology. Museum Support Center, Smithsonian Institution, Wash-
ington, D.C. 20560

-Department of Anthropology. National Museum of Natural History. NHB-112 Wash-
ington. D.C. 20560

ENT. NEWS 101(1): 39-47. January & February. 1990

40 ENTOMOLOGICAL NEWS

Pteropus personatus Temminck was collected by P.M. Taylor in 1984. As
field conditions allowed, care was taken to prevent contamination of
ectoparasite collections from different host individuals. All ectopara-
sites collected were placed in 70% ethanol for later processing and iden-
tification. The bat flies and fleas are deposited in the National Museum
of Natural History, and the ticks are deposited in the U.S. National Tick
Collection, Museum Support Center, Smithsonian Institution, Wash-
ington, D.C. Bat nomenclature follows Honacki et al. (1982).

RESULTS

The bats examined for ectoparasites were as follows: Dobsonia cren-
ulata K. Anderson, Eonycteris spelaea (Dobson), Macroglossus minimus
(E. Geoffroy), Nyctimene albiventer (Gray), Pteropus conspicillatus Gould,
Pteropus personatus, Rousettus amplexicaudatus (E. Geoffroy) (family
Pteropodidae), and Rhinolophus euryotis Temminck (family Rhinolo-
phidae). Table 1 depicts the host associations and the numbers of ecto-
parasites collected in this survey. In addition, 2 1 nycteribiids, 2 streblids,
208 larval ticks, and a mixture of 13 nymphal and adult spinturnicid
mites were removed from 16 bat specimens from which the identification
labels became detached. These ectoparasites could not be matched with
individual bat species (see Table 1).

DISCUSSION

Of the 1 1 species of bat flies collected during this survey (7 nyc-
teribiids^ streblids) the following represent new records for Halmahera:
Archinycteribia actena Speiser, Eucampsipoda inermis Theodor, Cyclo-
podia albertisii Rondani, Leptocyclopodia (Oncoposthia) macrura (Speiser),
Phthiridium phthisicum (Speiser) (Nycteribiidae), and Megastrebla
(Megastrebla} gigantea (Speiser), M. parvior Maa, and Raymondia pseu-
dopagodarum Jobling (Streblidae). Two species, Cyclopodia species B
(Nycteribiidae) and Brachytarsina species A (Streblidae), apparently are
undescribed but cannot be treated further until specimens of previously
described, related species are obtained for comparative purposes. The
ischnopsyllid flea, Thaumapsylla longiforceps Traub, and the laelapid
mite, Neolaelaps spinosa (Berlese), are also new records for Halmahera.

Nycteribiidae

Archinycteribia actena has been recorded from Sulawesi (Celebes)
and Seram east to the Solomon Islands and Australia (Maa, 1962, 1971;
Theodor, 1967, 1968; Durden et al., 1990). In the present survey it was

Vol. 101, No. 1, January & February 1990 41

collected from 3 hosts (Table 1 ), always in association with Cyclopedia
tennis and either Eucampsipoda inermis or Megastrebla (M.) gigantea.
Species of Dobsonia Palmer serve as primary hosts of Archinycteribia
actena throughout the range of this fly with D. crenulata the primary host
on Halmahera.

Our specimens of Eucampsipoda inermis from Halmahera apparently
constitute the first record of the genus from the Molucca Islands. This
species has been reported from Burma, Thailand, Malaya, Java, Sumba,
the Philippines, and Papua New Guinea (Theodor, 1955, 1963, 1967,
1968; Maa, 1962, 1977). Its primary hosts are Rousettus amplexicaudatus
and Eonycterisspelaea. On Halmahera the species also occurred primarily
on these 2 hosts; however, a few specimens were collected from Dobsonia
and Macroglossus. The records, other than those from the 2 primary host
species, probably represent accidental occurrences.

Cyclopodia tennis is known from Malaya, Java, Borneo and the
Moluccas (Theodor, 1959, 1967; Maa, 1966, 1977). It was first reported
from Halmahera by Theodor (1959) from Macroglossus minimus. Maa
(1966) stated that Theodor's record of a single female from Halmahera
needed confirmation. We are now able to verify this distribution record.
This was the second most common species in our collection. On Halma-
hera, Cyclopodia tennis was found almost entirely on Macroglossus min-
imus (originally identified as M. lagochilus but Honacki et al. (1982) list
lagochilus as a synonym of minimus). This parasitic fly was taken once
from Rousettus amplexicaudatus, once from Eonycteris spelaea, and once
from Nyctimene albiventer. The latter 3 host records probably represent
accidental occurrences.

Cyclopodia albertisii is known from the Moluccas (Goram), Belau
(Palau) Islands. Papua NewGuinea, and Australia (Theodor. 1959, 1967;
Maa. 1962, 1971). Our collection from Halmahera contains 9 specimens
all from a single host specimen of Pteropus conspicillatus. There were no
other associated ectoparasites on this host specimen.

Twenty-two specimens of Cyclopodia species B were taken from 4
specimens of Pteropus personatus. This fly probably is undescribed even
though it runs to Cyclopodia bougainvillensis Theodor in Theodor's 1967
keys. There are some differences between features in Theodor's descrip-
tion and those of our specimens but. until comparative material ot
bougainvillensis becomes available, it is impossible to be sure of the cor-
rect identity of our specimens.

Thirteen specimens of Leptocyclopodia (O.) macrura were removed
from 10 specimens of Dobsonia crenulata, and another male was found
on 1 unidentified bat. In addition to the range previously recorded for
this distinctive species (Durden, et al.. 1990), we now add Halmahera.
Associated species are listed in Table 1.

42 ENTOMOLOGICAL NEWS

Three individual specimens of Phthiridium phthisicum were collected
from 3 separate individuals of Rhinolophus euryotis, and always in as-
sociation with at least 1 other species, Raymondia pseudopagodarum or
Brachytarsina species A. The female of Phthiridium phthisicum was ori-
ginally described from Rhinolophus euryotis, from Amboina (Speiser,
1907). The male was later described from a specimen from Seram taken
from an unidentified species of Rhinolophus Lacepede, ". . . which ac-
cording to the form of the thorax and other characters resembles the
female closely [and] is considered to belong to this species." (Theodor,
1 968). The identity of our female specimen is reasonably certain, but that
of our 2 males is less certain. Our males do not completely coincide with
Theodor's description or match his figures. It is possible that our, or
Theodor' s association of the sexes is wrong. Only when both sexes from a
single host individual are available can a decision be made. Our speci-
mens might be one of several subspecies mentioned by Maa ( 1986) but,
again, comparative material is needed to be sure.

Streblidae.

Megastrebla (M.)gigantea is a widespread bat fly (Durden et al., 1990).
Seven specimens were collected from 3 species of bats and always in
association with 1 to 3 species of Nycteribiidae (Table 1).

A single female of Megastrebla (M.)parvior was taken from an Eonyc-
teris spelaea in association with Eucampsipoda inermis and Cyclopedia
tenuis.

Five specimens of RaBrachytarsina (here referred to as species A)
were collected from Rhinolophus euryotis. This fly closely resembles
Brachytarsina modesta Jobling and B. trinotata Maa, but comparative
material is not available to help confirm its identity.

Two males and 2 females of an apparently undescribed species of
Brachytarsina (here referred to as species A) were collected from Rhino-
lophus euryotis. This fly closely resembles Brachytarsina modesta Jobling
and B. trinotata Maa, but comparative material is not available to help
confirm its identity.

Ischnopsyllidae

Thaurnapsylla longiforceps was the only species of flea retrieved dur-
ing this survey. Although it has been collected from several unrelated bat
hosts (Hopkins and Rothschild, 1956), it appears to show a preference
for various species of the genus Rousettus( Holland, 1969). In Java, Hadi
et al. (1983) recorded 'Thaurnapsylla sp/ from Rousettus leschenaulti (Des-
marest) and Eonycteris spelaea', the latter was the only host species from
which T. longiforceps was recovered in this survey. Thaumapsylla longifor-

Vol. 101, No. 1, January & February 1990 43

ceps has been reported from Java, Borneo and the Philippines southeast
to New Guinea (Hopkins and Rothschild, 1956). In addition, we have
seen 1 19 specimens from 21 previously unreported collections of T. lon-
giforceps, all identified by R. Traub (pers. comm.): 73 specimens from
Rousettus amplexicaudatus (Sumatra (8), the Philippines (41), Sulawesi
(2),Flores (7) and Timor ( 1 5) ); 2 specimens from Rousettus sp. (the Philip-
pines); 3 specimens from Cynoptems brachyotis (Muller) (Sumatra (2) and
Java (1); 1 specimen from Eonycteris major (K. Anderson) (the Philip-
pines); and 40 specimens from unidentified bats (the Philippines (39)
and New Guinea (1)).

Argasidae

All ticks collected were larvae belonging to the genus Ornithodoros
Koch, subgenus Reticulinasus Schulze. All known species of this sub-
genus parasitize cave-dwelling megachiropteran bats (Dumbleton,
1958; Hoogstraal, 1962; Hoogstraal and Aeschlimann, 1982). Species of
Ornithodoros (Reticulinasus} are known from the Near East, Africa and
Madagascar eastward to India, Indo-Australia and the Solomon islands
(Dumbleton, 1958; Wirorenoe/ar/., 1979; Hoogstraal and Aeschlimann.
1982). The ticks collected from Halmahera represent 3 undescribed
species: most specimens belong to the species here designated as species
# 1 , but 1 specimen removed from Dobsonia crenulata belongs to a second
species (species #2), and 10 taken from an individual Rousettus amplex-
icaudatus include specimens of species # 1 and examples of a third taxon,
species #3. It is hoped that future collecting on Halmahera will produce
postlarval specimens of these 3 undescribed ticks.

Laelapidae

Neolaelaps spinosa was the only laelapid mite collected in this survey.
It occurred in low numbers on both Dobsonia crenulata and Pteropusper-
sonatus. This mite typically parasitizes several species ofPteropus and is
known from Sri Lanka eastward through Indo-Australia to New Ireland
(Strandtmann and Garrett, 1967). Our record from Dobsonia crenulata is
considered to represent an accidental infestation.

Spinturnicidae and Trombiculidae

The remainder of the ectoparasites collected in this survey consisted
of an estimated 6 to 9 species of spinturnicid mites, and 2 species of larval
chigger mites (Trombiculidae). The host associations of these two families
are shown in Table 1. Both mite groups are frequent ectoparasites of
pteropodid bats, particularly in Indo-Australia (Beck, 1971; Hadi et al..

44 ENTOMOLOGICAL NEWS

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Laelapidae:

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Spinturnicidae

Tromhiculidae

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03
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46 ENTOMOLOGICAL NEWS

1983; Durden et al., 1990). The chiggers showed a strong host preference
forDobsonia crenulata with all but 2 of the 76 specimens collected being
taken from this host. Spinturnicid mites were common on D. crenulata,
Eonycteris spelaea and Rousettus amplexicaudatus, and scarce on Mac-
roglossus minimus; none were taken from the other 4 bat species exam-
ined in this survey.

ACKNOWLEDGMENTS

Gratitude is extended to A.C. Messer, a former research assistant. Department of
Anthropology, National Museum of Natural History, Washington, D.C. (now at Cornell
University, Ithaca, New York), who assisted P.M. Taylor in making the bat and ectoparasite
collections; to L.K Gordon and D.F. Schmidt, Department of Mammalogy, National
Museum of Natural History, Washington, D.C., who identified the bats; and R. Traub, Flea
Unit, Department of Entomology, Museum Support Center, Smithsonian Institution,
Washington, D.C., who confirmed the flea identifications and supplied additional dis-
tribution records. We also are grateful to R.G. Robbins, National Institue of Allergy and
Infectious Diseases, National Institutes of Health, c/o Department of Entomology,
Museum Support Center, Smithsonian Institution, Washington, D.C., and J.O. Whitaker,
Jr., Department of Life Sciences, Indiana State University, Terre Haute, IN, for reviewing
the manuscript.

LITERATURE CITED

Beck, AJ. 1971. A survey of bat ectoparasites in West Malaysia. Jour. Med. Entomol. 8:
147-152.

Dumbleton, L.J. 1958. Bat infesting Ornithodoros (Ixodoidea - Argasidae) of the Oriental-
Australian region. Proc. Linn. Soc. N.S.W. 83: 303-308.

Durden, L.A., B.V. Peterson, N. Wilson and B. Christiansen. 1990. Notes on some
ectoparasites of bats from Seram Island, Indonesia. Entomol. News 101(l):48-56.

Hadi, T.R., S. Sarbini and RJ. Brown. 1983. Small mammalian ectoparasites from Mt.
Bromo area, east Java, Indonesia. S.E. Asian Jour. Trop. Med. Publ. Hlth. 14: 422-
425.

Holland, G.P. 1969. Contribution towards a monograph of the fleas of NewGuinea. Mem.
Entomol. Soc. Can. 61: 1-77.

Honacki, J.H., K.E. Kinman, and J.W. Koeppl (eds.). 1982. Mammal species of the
world: a taxonomic and geographic reference. Allen Press, Inc. & Association of Sys-
tematic Collections, Lawrence, Kansas, x + 694 Pages.

Hoogstraal, H. 1962. Description of Ornithodoros (Reticulinasus) madagascariensis n. sp.
(Ixodoidea, Argasidae). Acarologia 4: 185-189.

Hoogstraal, H. and A. Aeschlimann. 1982. Tick-host specificity. Bull. Soc. Entomol.
Suisse 55: 5-32.

Hopkins, G.H.E. and M. Rothschild. 1956. An illustrated catalogue of the Rothschild
collection of fleas (Siphonaptera) in the British Museum (Natural History). With keys
and short descriptions for the identification of families, genera, species and subspecies
of the order. Vol. II. Coptopsyllidae, Vermipsyllidae, Stephanocircidae, Macrop-
syllidae, Ischnopsyllidae, Chimaeropsyllidae, Xiphiopsyllidae. Brit. Mus. (Nat. Hist.),
London, xi + 445 Pages, 32 plates.

Maa, T.C. 1962. Records and descriptions of Nycteribiidae and Streblidae (Diptera).
Pacif. Insects 4: 4 17-436.

Vol. 101, No. 1. January & February 1990 47

Maa, T.C. 1966. Partial revision of the Cyclopodiinae (Diptera: Nycteribiidae). Pacif.

Insects 8: 648-685.
Maa, T.C. 1971. Revision of the Australian hatflies (Diptera: Strehlidae and Nyc-

teribiidae). Pacif. Insects Monogr. 28: 1-118.
Maa, T.C. 1977. Family Streblidae, Pages 432-435. In M.D. Delfinado and D.E. Hardy

(eds.). A catalog of the Diptera of the Oriental Region. Volume III. Suborder Cyclor-

rhapha (excluding Division Aschiza). University Press of Hawaii. Honolulu.
Maa, T.C. 1986. Three new Diptera Pupipara from Sulawesi, Indonesia. Jour. Taiwan

Mus. 39: 93-98.
Speiser, P. 1907. XXIV.-Preliminary descriptions of three new Nycteribiidae from India.

Rec. Indian Mus. 1: 295-296.
Strandtmann, R.W. and L.E. Garrett. 1967. Neolaelaps palpispinosus, a new species of

laelapid mite from fruit bats in New Guinea (Acarina: Laelapidae). Jour. Med.

Entomol. 4: 237-239.
Taylor, P.M. 1985. Ethnobiology and the ethnography of Tobelo-speaking peoples of

Halmahera Island, Indonesia. Nat. Geogr. Soc. Res. Repts. 21: 475-480.
Theodore, O. 1955. On the genus Eucampsipoda Kol. and Dipseliopoda n.g. (Nycteribiidae.

Diptera). Parasitology 45: 195-229.

Theodore, O. 1959. A revision of the genus Cvclopodia (Nycteribiidae, Diptera). Parasitol-
ogy 49: 242-308.
Theodor, O. 1963. Philippine batflies of the family Nycteribiidae (Diptera, Pupipara).

Fieldiana Zool. 42: 151-192.
Theodor, O. 1967. An illustrated catalogue of the Rothschild Collection of Nycteribiidae

(Diptera) in the British Museum (Natural History), with keys and short descriptions for

the identification of subfamilies, genera, species and subspecies. Brit. Mus. (Nat. Hist.)

Publ. 655: viii + 506 Pages, 5 plates.
Theodor, O. 1968. New species and new records of Nycteribiidae from the Ethiopian,

Oriental and Pacific Regions. Parasitology 58: 247-276.
Wiroreno, W., S. Kadarsan and H.B. Munaf. 1979. Larval characters of some argasid

ticks (Acarina: Argasidae) of Indonesia. Biotrop. Spec. Publ. 6: 47-54.

48 ENTOMOLOGICAL NEWS

SOME ECTOPARASITES OF BATS FROM
SERAM ISLAND, INDONESIA1

L.A. Durden2, B.V. Peterson3, N. Wilson4, B. Christiansen5

ABSTRACT: New host and distribution records are provided for some ectoparasites of the
families Streblidae, Nycteribiidae, Ischnopsyllidae (Insecta), Ixodidae, Argasidae, Lae-
lapidae, Spinturnicidae and Macronyssidae (Acari) removed from 9 species of bats collect-
ed in Manusela National Park, on the island of Seram, Indonesia.

The Indonesian island of Seram (Ceram) is situated in the Moluccan
Archipelago between Sulawesi and Irian Jaya. Zoogeographically, it is
part of the Wallacean subregion (the faunal transition zone between
Asia and Australasia) and while part of the terrestrial fauna is endemic,
much of it extends to the east and west in varying degrees. No endemic
species of bats are known from Seram and this fauna shows Asian, Aus-
tralasian, Malesian (Melesia: the biogeographical region stretching
from Sumatra and the Malay Peninsula east to the Bismarck Archi-
pelago (Whitmore, 1981, 1987)) or Moluccan distributions. Bat ecto-
parasite records from Seram are few and therefore valuable not only for
inventory purposes but also for elucidation of host-parasite associations.

MATERIALS AND METHODS

Ectoparasites were collected by visual searches from 36 bats of 9
species mist-netted in Manusela National Park, Seram (3° 15'S, 129°38'E),
during July and August, 1987. Ectoparasite material was stored in 70%
ethanol until it could be processed for identification.

RESULTS AND DISCUSSION

The 9 bat species collected and the numbers of each examined for
ectoparasites were as follows (bat nomenclature follows Honacki et al.,
1982): Dobsonia viridis (Huede) (Icf, 1699), D. moluccensis (Quoy and
Gaimard) (Icf, 19), Rousettus amplexicaudatus ( E. Geoffrey) (2c?cf, 599),

Deceived March 16, 1989. Accepted May 13, 1989.

"Department of Entomology, Smithsonian Institution, Museum Support Center. Wash-
ington, D.C. 20560, U.S.A.

Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, c/o
National Museum of Natural History, NHB-168, Washington, D.C. 20560, U.S.A
Department of Biology, University of Northern Iowa, Cedar Falls, IA, 50614, U.S.A.
56 Clifton Road. Kingston-upon-Thames, Surrey KT2 6JP, United Kingdom.

ENT. NEWS 101(1): 48-56, January & February. 1990

Vol. 101. No. 1, January & February 1990 49

Syconycteris australis (Peters) ( 1 cf , 19), Pteropus temmincki Peters (1 cf , 1 Q )
(family Pteropodidae), Miniopterus australis Tomes (2cfcf) (family Ves-
pertilionidae), Rhinolophus euryotis Temminck (Icf) (family Rhinolo-
phidae), Chaerephon jobensis (Miller) (19), and Mormopterus beccarii
Peters (299) (family Molossidae).

Table I lists the ectoparasites collected during this survey. The bat
flies and the single flea are deposited in the U.S. National Museum of
Natural History, Washington, D.C., and the ticks are in the U.S.
National Tick Collection, Smithsonian Institution, Washington, D.C.
(accessioned under RML 1 19,061-1 19,075).

The ectoparasites recorded here from Seram bats are fairly typical for
this group of hosts. The following species represent new records for
Seram: Brachytarsina (Brachytarsina} amboinensis Rondani, Megastrebla
(Magastrebla) gigantea (Speiser), M. (M.)pan'ior (Maa), Raymondia pseu-
dopagodarum Jobling (Streblidae); Archinycteribia actena Speiser, Nyc-
teribia parilis Walker (Nycteribiidae); Thaumapsylla breviceps Rothschild
(Ischnopsyllidae), and all of the ticks and mites.

Most of the bats and ectoparasites retrieved during this survey are
known to have wide but contrasting geographical distributions. With
respect to the streblid bat flies, Brachytarsina (B.) amboinensis was only
taken from Miniopterus australis which ranges from India to China
(Hainan) and northeastern Australia. The known range of Brachytarsina
(B.) amboinensis is similar extending from India and Sri Lanka east to
Australia, Vanuatu and New Caledonia (Jobling, 1951; Hiregaudar and
Bal, 1956; Maa, 1962, 197 la, 1977; Maa and Marshall, 1981). This bat fly
is represented by several subspecies in various parts of its range, and
specimens from Seram appear to belong to the nominate subspecies.
Brachytarsina (B.) amboinensis is principally parasitic on various species
of Miniopterus Bonaparte.

The streblid, Megastrebla (M.) gigantea was collected from both
species of Dobsonia Palmer. Dobsonia viridis is distributed throughout the
Moluccan and Banda Islands and the Philippine island of Negros while
D. moluccensis extends from the Moluccas eastward to Aru, New Guinea,
and northern Queensland. Megastrebla (M.) gigantea has been recorded
from India east to New Britain (Jobling, 1951; Hiregaudar and Bal, 1956;
Maa, 1962). However, Maa (1971 b) later stated that this species with its 3
subspecies (gigantea gigantea, g. kalawawae Maa, g. solomonis Maa)
ranges only from Sumba east to the Solomon Islands, and that all other
records must be held suspect. This is the largest known species in the
genus and is probably restricted to bats of the genus Dobsonia. The
specimens from Seram are of the nominate subspecies.

Megastrebla (M.) parvior was collected from Rousettus amplex-

50 ENTOMOLOGICAL NEWS

icaudatus. This bat ranges throughout Malesia whereas Megastrebla par-
vior has a known distribution extending from India and Burma east to
the Philippines, south to Sumatra, Sumba and New Guinea (Maa, 1962,
197 Ib, 1977). The specimens from Seram are of the nominate subspecies.
Maa (1962) recorded the Sumba series of specimens from Rousettus
amplexicaudatus, and the remaining type series from Eonycteris spelaea
(Dobson). Rousettus Gray is the primary host of the nominate subspecies
(Maa, 1971b),but it has also been recorded from a number of other hosts.
A second subspecies, Megastrebla parvior papuae Maa, was described
from New Guinea from Rousettus amplexicaudatus and Dobsonia moluc-

censis.

Raymon dia pseu dopago daru m was removed from a male Rhinolophus

euryotis. This bat ranges from Sulawesi east to the Moluccas, Timor, New
Guinea, the Bismarck Archipelago and adjacent small islands whereas
its tiny parasite ranges from Burma, Thailand, China, east to the Philip-
pines, and south to Timor and Seram (Jobling, 1951; Maa, 1962, 1977).
Primary hosts for Raymondia pseudopagodarum seem to be species of
Rhinolophus Lacepede and Hipposideros Gray, although a few other sus-
pect hosts have been listed for the species.

With respect to the nycteribiid bat flies, Archinycteribia actena was
collected only from Dobsonia viridis. This fly has previously been record-
ed from Ambon, New Guinea, the Solomon Islands and Australia (Maa,
1962, 1971a;Theodor, 1967). In4ofthecollections,/l.tf<:'tert<7 was taken in
association with Leptocyclopodia (Oncoposthia) sp. A, once with Megas-
trebla (M.)gigantea, and twice by itself.

The single specimen of Cyclopedia sp. collected from Syconycterisaus-
tralis is in poor condition, but from the characters that can be seen it
could be Cyclopodia sycophanta Maa that parasitizes the same host, or it
might represent a new species in the C. sycophanta group. Additional
specimens will be needed before the identity of this species can be deter-
mined with certainty.

Leptocyclopodia (Oncoposthia) macrura Speiser was collected only
from Dobsonia moluccensis; this bat fly was previously reported from D.
moluccensis on the islands of Biak and Owi off the northern coast of New
Guinea (Hadi et al., 1980). Leptocyclopodia (O.) macrura is a distinctive
species and has also been recorded from Sumbawa, Ambon, Seram, New
Guinea, and New Britain (Maa, 1962, 1966, 1968, 1975; Treodor, 1959,
1967) which approximates the distribution of Dobsonia moluccensis.

Leptocyclopodia (Oncoposthia) sp. A was collected mainly from Dob-
sonia viridis although 2 specimens were taken from Syconycteris australis.
This latter bat species ranges from the Moluccas eastward to New
Guinea, the Bismarck Archipelago and adjacent small islands and
south to Australia. These bat fly specimens probably represent a new

Vol. 101, No. 1, January & February 1990 51

species since neither sex runs successfully in Maa's key (1975) nor do
they fully match the descriptions or illustrations of the other species of
this subgenus. The true status of this fly can be determined only when
comparative material of other closely related species is available.

Nycteribia parilis was found only on a male Miniopterns australis. The
same host individual yielded 8 specimens of the streblid Brachytarsina
(B.) amboinensis. Nycteribia parilis is known from Ambon, Batchian,
Timor, and other Moluccan islands. New Guinea and Australia prin-
cipally from various species of Miniopterus (Maa. 1962, 197 la; Theodor.
1967). There are 2 subspecies of this bat fly: the nonimate subspecies over
most of its range, and p. vicaria Maa from Australia.

Thaumapsylla breviceps, the only flea collected, has a very broad
geographical range that includes much of the Ethiopian and Oriental
Regions. However, 2 subspecies are known: the nominate subspecies in
Africa, and b. orientalis in Southeast Asia (Hopkins and Rothschild.
1956). Although it is most likely that the latter taxon was collected in this
survey, the 2 subspecies can only be separated by male characters and
the Seram specimen is a female. This flea typically parasitizes Rousettus
bats, which agrees with the R. amplexicaudatus record documented here
for Seram. Thaumapsylla breviceps has hyper-developed pronotal combs
that may facilitate attachment to its volant hosts and partly explain why
this species has such a large geographical distribution (Traub. pers.
comm.).

The tick, Ixodes (Eschatocephalus) simplex Neumann, occupies a vast
geographical area within the Palaearctic, Ethopian, Oriental and Aus-
tralasian Regions (Arthur, 1 956; Wilson, 1 970; Clifford et al., 1 973). It was
taken only from Miniopterus australis in this survey, an anticipated assoc-
iation since Miniopterus is the usual host (Wilson, 1970).

Ixodes (Lepixodes) kopsteini Oudemans is known to parasitize bats in
the Ethiopian, Oriental and Australasian Regions including the Moluc-
cas (Kadarsan. 1971; Clifford et al., 1973). During this study it was col-
lected from 3 host species belonging to 2 families. The molossids.
Chaerephon jobensis which ranges from India and Sri Lanka, east to
southern China and Vietnam, and southeast to the Philippines. Borneo.
Bali and the Moluccas, and Mormopterus beccarii which ranges from the
Moluccas east to New Guinea, northern and central Australia and adja-
cent small islands, appeared to be the main hosts but a single specimen
was collected from the pteropodid, Rousettus amplexicaudatus. Ixodes (L.)
kopsteini possesses certain morphological and reproductive features
unknown in other ticks ( Kadarsan, 1 97 1 ). These morphological traits led
Anastos (1950) to exclude this unusual tick from his monograph on
Indonesian Ixodidae because he considered it to be. "probably a special
type of mite."

52 ENTOMOLOGICAL NEWS

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54 ENTOMOLOGICAL NEWS

Ornithodoros sp. larvae could not be identified further so little can be
stated regarding host associations or geographical distribution. Never-
theless, this tick genus occurred almost exclusively on Dobsonia viridis
(70 specimens) although 2 specimens were taken from a female Rouset-
tus amplexicaudatus.

The laelapid mite, Neolaelaps spinosa (Berlese) is parasitic on a variety
of fruit bats in the genus Pteropus Erxleben. It ranges from Sri Lanka to

Australia so the present record from P. temmincki on Seram is expected.

This mite occasionally has been found attached phoretically to nyc-

teribiidbat flies (Maa, 197 la) but this phenomenon was not observed in

the Seram material.

All 4 macronyssid mites were damaged protonymphs and could not
be identified beyond genus. Two genera were collected: Macronyssus,
which is most often retrieved from vespertilionid bats, agreeing with the
Miniopterus australis record documented here, and Trichonyssus which
occurs principally on vespertilionids but also on some other bat families
including Molossidae from which the Seram collections were made.
While the genus Macronyssus is virtually cosmopolitan in distribution,
there does not appear to be any previous records for Trichonyssus outside
of Australia and Tasmania (Radovsky, 1979; Micherdzinski and Dom-
row, 1985; Domrow, 1987).

Seven species of spinturnicid mites were collected. Ancystropus tap-
robanius (Turk) and A zeleborii Kolenati were both taken from Rousettus
amplexicaudatus which concurs with previously documented host re-
cords especially those from southern Asian species of Rousettus (Prasad,
1969; Domrow, 1972; Hadi etal., 1980). Three species of Meristaspis were
identified from different bat hosts. Meristaspis lateralis (Kolenati), which
ranges from the Near East (Palestine and Yemen) southeast to New
Guinea, is principally parasitic on Rousettus species (Delfinado and
Baker, 1963; Prasad, 1969). The present records of Meristaspis lateralis
from Rousettus amplexicaudatus therefore conform to this trend although
the single specimen of Meristaspis lateralis from Dobsonia viridis is an
exception. Meristaspis jordani (Radford) was collected only from Dob-
sonia viridis. Previous records show this mite to be parasitic mainly on
bats of the genus Dobsonia, and to range from Sulawesi and the Philip-
pines to New Guinea and the Bismarck and Solomon Islands (Prasad,
1969; Domrow, 1972). Meristaspis calcarata (Hirst) typically parasitizes
numerous species of Pteropus bats and ranges from Madagascar to Aus-
tralia and a number of Pacific islands (Delfinado and Baker, 1963;
Prasad, 1 969). The present record from P. temmincki fits this distribution.
Paraperiglischrus rhinohphinus (Koch) is associated with numerous
species of Rhinolophus bats, and ranges from the British Isles southeast
to Africa, southern Asia and Australia. In this survey it was collected

Vol. 101, No. 1. January & February 1990 55

from R. euryotis. Spinturnix psi (Kolenati) is another ectoparasite with a
wide geographical distribution. There are records of this mite from
France, southern Europe, Madagascar, southern Asia and Australia. In
this survey, Spinturnix psi was collected only from Miniopterus australis
which concurs with its known host preference for species of Miniopterus
(Baker and Delfinado, 1964; Prasad, 1969; Domrow, 1972).

The above records extend the known geographical distributions of
most of the ectoparasites collected during this survey. Geographical dis-
tributions of the bat species and their associated ectoparasites are not
congruent in the majority of cases. Different bat species are parasitized
by these ectoparasites in other parts of their range. Conversely, a few of
the ectoparasites were more host specific and may be expected to overlap
more closely with the distributions of their hosts. The dispersive abilities
of their volant hosts and the facility of some bat ectoparasites to utilize a
spectrum of host species probably explains, at least in part, the large
geographical distributions of many bat-associated arthropods.

ACKNOWLEDGMENTS

Gratitude is extended to J.E. Hill, British Museum (Natural History) (retired), London.
U.K.. for identifying the bats, and to J.E. Keirans. U.S. National Tick Collection, National
Institute of Allergy and Infectious Diseases, National Institutes of Health, c/o Department
of Entomology, Museum Support Center, Smithsonian Institution. Washington, D.C.. for
identifying the ticks. The bat and ectoparasite collections were made during Operation
Raleigh (Headquarters in Chelsea, London, UK). We also are grateful to R.G. Robbins,
U.S. National Tick Collection. National Institute of Allergy and Infectious Diseases.
National Institutes of Health, c/o Department of Entomology. Museum Support Center
Smithsonian Institution, Washington, D.C., and T.C. Maa. 2145 Ridgewood Road. Medina.
Ohio, for reviewing our manuscript. We thank also R.D. Gordon, N.E. Woodley. and M.E.
Schauff, Systematic Entomology Laboratory. Washington, D.C., for reading and com-
menting on the manuscript.

LITERATURE CITED

Anastos, G. 1950. The scutate ticks, or Ixodidae. of Indonesia. Entomol. Americana (new

series) 30: 1-144.
Arthur, D.R. 1956. The Ixodes ticks of Chiroptera (Ixodoidea. Ixodidae). Jour. Parasitol.

42: 180-196.
Baker, E.W. and M.D. Delfinado. 1964. Spinturnicidae of South East Asia and the

Pacific Region. Pacif. Insects 6: 571-591.
Clifford, C.M., D.E. Sonenshine, J.E. Keirans, and G.M. Kohls. 1973. Systematics of

the subfamily Ixodinae (Acarina: Ixodidae). I. The subgenera of Ixodes. Ann. Entomol.

Soc. Amer. 66:489-500.
Delfinado, M.D. and E.W. Baker. 1963. Mites (if the family Spinturnicidae from the

Philippines (Acarina). Pacif. Insects 5: 905-920.
Domrow, R. 1972. Acari Spinturnicidae from Australia and New Guinea. Acarologia 13:

552-584.
Domrow, R. 1987. Acari Mesostigmata parasitic on Australian vertebrates: an annotated

checklist, keys and bibliography. Invert. Taxon. 1: 817-948.

56 ENTOMOLOGICAL NEWS

Hadi, T.R., S. Nalim, S. Sukaeri and D.T. Dennis. 1980. Scrub typhus survey of Biak
and Owi islands: ectoparasites of small mammals and rickettsial isolations. S.E. Asian
Jour. Trop. Med. Publ. Hlth. 11: 220-226.

Hiregaudar, L.S., and D.V. Bal. 1956. Some Ectoparasites of bats from India. Agra Univ.
Jour. Res. (Sci.) 5: 2-134.

Hopkins, G.H.E., and M. Rothschild. 1956. An illustrated catalogue of the Rothschild
collection of fleas (Siphonaptera) in the British Museum (Natural History). With keys
and short descriptions for the identification of families, genera, species and subspecies
of the order. Vol. II. Coptopsyllidae, Vermipsyllidae, Stephanocircidae, Macrop-
syllidae, Ischnopsyllidae, Chimaeropsyllidae, Xiphiopsyllidae. Brit. Mus. (Nat. Hist.),
London xi + 445 Pages, 32 plates.

Jobling, B. 1951. A record of the Streblidae from the Philippines and other Pacific islands,
including morphology of the abdomen, host-parasite relationship and geographical
distribution, and with descriptions of five new species (Diptera). Trans. Roy. Entomol.
Soc. Lond. 102:211-246.

Kadarsan, S. 1971. Larval ixodid ticks of Indonesia (Acarina: Ixodidae). Ph.D. Thesis,
University of Maryland, 182 p.

Maa, T.C. 1962. Records and descriptions of Nycteribiidae and Streblidae (Diptera).
Pacif. Insects 4: 417-436.

Maa, T.C. 1966. Partial revision of the Cyclopodiinae (Diptera: Nycteribiidae). Pacif.
Insects 8: 648-685.

Maa, T.C. 1968. Additions to the Cyclopodiinae. Part I (Diptera: Nycteribiidae). Pacif.
Insects 10: 1-23.

Maa, T.C. 1971a. Revision of the Australian batflies (Diptera: Streblidae and Nyc-
teribiidae). Pacif. Insects Monogr. 28: 1-118.

Maa, T.C. 197 Ib. Review of the Streblidae (Diptera) parasitic on Megachiropteran bats.
Pacif. Insects Monogr. 28: 213-243.

Maa, T.C. 1975. On new Diptera Pupipara from the Oriental Region. Pacif. Insects 16: 465^86.

Maa, T.C. 1977. Family Streblidae, Pages 432-435. In M.D. Delfinado and D.E. Hardy
(eds.). A catalog of the Diptera of the Oriental Region. Volume III. Suborder Cyclor-
rhapha (excluding Division Aschiza). University Press of Hawaii, Honolulu.

Maa, T.C. and A.G. Marshall. 1981. Diptera Pupipara of the New Herbrides: taxonomy,
zoogeography, host association and ecology. Quart. Jour. Taiwan Mus. 34: 213-232.

Micherdzinski, W. and R. Domrow. 1985. The genus Trichonyssus Domrow on western
Australian bats (Acarina: Macronyssidae). Intern. Jour. Acarol. 1 1: 55-65.

Prasad, V. 1969. Bat mites (Acarina: Spinturnicidae) mainly from South-East Asia and the
Pacific Region. Acarologia 1 1: 658-677.

Radovsky, F.J. 1967. The Macronyssidae and Laelapidae ( Acari: Mesostigmata) parasitic
on bats. University of California Press, Berkeley and Los Angeles, 288 pp.

Theodor, O. 1959. A revision of the genus Cvclopodia (Nycteribiidae, Diptera). Parasitol-
ogy 49: 242-308.

Theodor, O. 1967. An illustrated catalogue of the Rothschild Collection of Nycteribiidae
(Diptera) in the British Museum (Natural History), with keys and short descriptions for
the identification of subfamilies, genera, species, and subspecies. Brit. Mus. (Nat. Hist.)
Publ. 655: 506 p., 5 plates.

Whitmore, T.C. 1981. Wallace's line and plate tectonics. Clarendon Press, Oxford, xii + 91 p.

Whitmore, T.C. 1987. Biogeographical evolution of the Malay archipelago. Clarendon
Press, Oxford, x + 147 p.

Wilson, N. 1970. New distributional records of ticks from southeast Asia and the Pacific
(Metastigmata: Agrasidae, Ixodidae). Oriental Insects 4: 37-46.

Vol. 101, No. 1, January & February 1990

ANNOTATED CHECKLIST OF THE

RHYACOPHILOIDEA AND INTEGRIPALPIA

(TRICHOPTERA) OF ALABAMA1

S.C. Harris2, P.K. Lago3

ABSTRACT: Distributional records of 132 species of caddisflies in the suborder Annuli-
palpia, (superfamily Rhyacophiloidea) and the suborder Integripalpia (superfamilies
Limnephiloidea, Phryganeoidea, Leptoceroidea, and Sericostomatidea) from Alabama
are listed, along with information on seasonal occurrence, habitat and relative abundance.
This checklist brings the total number of caddisflies known from Alabama to 323 species.

The caddisfly fauna of the southeastern United States has received
considerable attention in recent years and species checklists are now
available for Tennessee (Etnier and Schuster, 1979), North and South
Carolina (Unzickere/ a/., 1982), Kentucky (Resh, 1975), Virginia (Parker
and Voshell, 1981), and Mississippi including southeastern Louisiana
(Holzenthal el al., 1982). This is the third and final contribution in a
series of papers (Harris, 1986a; Lago and Harris, 1987) on the caddisfly
fauna of Alabama. This checklist presents distributional records for 132
species in the families Glossosomatidae and Rhyacophilidae (infra-
order Spicipalpia, Weaver, 1984); Goeridae, Limnephilidae, Uenoidae,
Brachycentridae, Lepidostomatidae, Phyrganeidae (infraorder Pleni-
tentoria. Weaver, 1984); and Odontoceridae, Calamoceratidae, Lep-
toceridae, Molannidae, Helicopsychidae, and Sericostomatidae (infra-
order Brevitentoria, Weaver, 1984). The Hydroptilidae which are also in
the infraorder Spicipalpia (Weaver, 1984) are not included in this chec-
klist, but were compiled in Harris (1986a). Of the 132 species identified
from Alabama, ten were described during the course of the study (Harris,
1986b, 1987, 1989; Vineyard and Wiggins, 1987) and two others will be
described in the near future. The species richness is indicative of the
range of physiographic regions represented in the state. These include
the East Gulf Coastal Plain, Piedmont Plateau, Valley and Ridge, and
Appalachian Plateau, including the Highland Rim Plateau and Cum-
berland Plateau subregions (Sapp and Emplaincourt, 1975).

CHECKLIST OF SPECIES

In the following checklist each species is followed by numerically

^Received April 20, 1989. Accepted June 8, 1989

"Biological Resources Division, Geological Survey of Alabama. Tuscaloosa, Ala-
bama 35486
-'Department of Biology. University of Mississippi. University, Mississippi 38677

ENT. NEWS 10 1( 1 ): 57-66, January & February', 1990

58 ENTOMOLOGICAL NEWS

coded county records. These counties and their location, both within the
state and within a physiographic province, are depicted in Figure 1.
General information on distribution and abundance in the state is
included with each species, followed by collection months and numbers
of specimens examined ( ). In most cases only adult males were exam-
ined, except as noted. Detailed locality and collection information will
be provided in an overall summary of the caddisfly fauna of the state, to
be published later. This checklist represents over 600 collections made
primarily with black-light traps between 1981 and 1987. Voucher speci-
mens are maintained in the insect collections of the Geological Survey of
Alabama and the University of Alabama.

SPICIPALPIA
HYDROPTILOIDEA

Glossosomatidae

Agapetus alabamensis Harris. 25. Known only from two small streams of the Cumberland

Plateau. April, May. (30)
A. avitus Edwards. 1. 2, 4. 10. Locally common in northern Alabama, particularly the

Highland Rim. April-June. (94)
A. crasmus Ross. 3, 4. Locally common in lower Appalachians of northeastern Alabama.

June. (51)

A. gelbae Ross. 1. Uncommon in small Highland Rim streams. April-June. (20)
A. hessi Leonard and Leonard. 3, 8, 12, 34. Locally common; primarily in small streams of

northern Alabama. May, June. ( 1 1 5)
A. iridis Ross. 17. Record from small stream in lower Appalachians provided by D.A. Etnier.

April. (4)

A. pinatus Ross. 17. Only records from a small stream in the lower Appalachians. May. (2)
A. rossi Denning. 8, 12, 17, 26, 31, 34. Frequently collected at scattered localities in northern

half of state. April-June. (319)
A. spinosus Etnier and Way. 5. Rarely collected; in small streams of the lower Appalachians.

May. (4)
A. tomus Ross. 14, 17, 20, 34. Uncommon; in small streams of northcentral Alabama.

May. (12)
Glossosoma nigrior Banks. 1.2,5,6,8-10, 13, 14, 17-20, 25. 27, 29, 31. 32. 51. Widespread and

common; primarily in small streams of northern Alabama. April-September. (405)
Matrioptila jeanae (Ross). 5, 17, 26, 29, 31, 32, 34. Most frequent in small Piedmont streams,

never collected in large numbers. May, June. (135)
Protoptila cahabensis Harris. 19. Known only from a small section of the Cahaba River.

May, August. October. (70)
P. georgiana Denning. 31. Only records from Hillabee Creek, a large Piedmont stream.

May. (8)
P. maculata (Hagen). 16-18. Locally abundant in small streams of lower Appalachians and

upper Piedmont. May, June, September, October. (134)
P. palina Ross. 3, 12-14, 17-20, 25, 26, 29, 31, 32. 34, 54, 63-65. Widespread and common in

Alabama, but most frequently collected in northern half of state. April-October.

(2491)

Vol. 101, No. 1, January & February 1990 59

RHYACOPHILOIDEA

Rhyacophilidae

Rhyacophila carolae Harris. 8. Known only from type locality, a small first order stream

in Bankhead National Forest. May. (2)

R. Carolina Banks. 1, 5, 6, 8, 10, 11, 13, 14, 17-22, 25,27-29, 31, 32, 34,42, 51, 53, 66. Most com-
mon and widespread Rhyacophila in Alabama; most commonly collected in northern

half of state. April-October. (353)
R.fenestra Ross. 1,2. Only records from extreme northwestern Alabama on Highland Rim.

June. (6)
R.fuscula (Walker). 14, 17. 18, 25, 27-29, 31, 32. Occurring in northern half of Alabama,

primarily in Piedmont. April-June, September, October. (81)
R. glaberrima Ulmer. 8, 11, 13, 18, 25. Infrequently collected at scattered north Alabama

localities. May, June, September, October. (25)
R. ledra Ross. 1-3, 6, 14, 17, 20, 21, 25, 39, 51, 52, 58, 66, 67. Widespread, but infrequently

collected in state. May, June. (46)
R. lobifera Betten. 14, 21,25. Common in a few small streams of the Cumberland Plateau in

central Alabama. April, May. (86)
R. nigrita Banks. 17, 18, 25, 29. Uncommon; mainly in small Piedmont streams. May,

June. (13)

R. teddyi Ross. 17. Only records from small streams in lower Appalachians. May. (2)
R. torva Hagen. 1 7, 1 8, 3 1 . Infrequently collected in small Piedmont and lower Appalachian

streams. May, June. (15)
R. vuphipes Milne. 34. Larval records from Little Cahaba River provided by D.A. Etnier.

This record is tentative but the specimens key to/?, vulphipes in Unzicker^ra/. ( 1982) and

not to any of the recorded species from the Cahaba River system or from central

Alabama. April. (2)

PLENITENTORIA

LIMNEPHILOIDEA

Goeridae

Goera calcarata Banks. 1,5,6, 16- 19, 27, 28. 34. Common in northern Alabama, particularly

in small streams of lower Appalachians. April-October. (273)
G. townesi Morse. 8, 12, 17, 18, 25, 27, 29, 32. Similar in occurrence to G. calcarata, but more

frequently collected in Piedmont and Cumberland Plateau streams. May, June

(233)

Limnephilidae

Hydatophylax argus (Harris). 25. Larval records from several small streams of the Cum-
berland Plateau. April, May. (37)

Ironoquia kaskaskia (Ross). 22. Only record from a large, sand-bottom river. October. (1)
/. punctatissma (Walker). 1 8, 20, 25, 34, 35, 43. 45. Most often collected along large rivers, but

generally uncommon in our collections. September, October. (34)

Platycentropus radiatus '(Say). 8, 12. Rare in small streams of Cumberland Plateau. May. (3)
Pycnopsyche gentilis (MacLachlan). 18, 27. Rarely collected in small streams of lower
Appalachians. March (larva), October (adult). (2)

60 ENTOMOLOGICAL NEWS

P. Indiana (Ross). 18, 20, 22, 25, 34, 67. Uncommon in small streams, mainly in west central

Alabama. October, November. (30)
P. lepida (Hagen). 18, 22, 25. Uncommon in several small streams of northern Alabama.

September, October. (13)
P. luculenta (Betten). 18, 25, 34. Locally common in small streams of northern Alabama.

October. (65)
P. scabripennis (Rambur). 1, 16, 18, 25, 34,45. Uncommon at scattered localities in northern

half of Alabama. June, September, October. (27)
P. virginica (Banks). 18. Single record from Coldwater Spring provided by K.L. Manuel.

November. (1)

Uenoidae

Neophylax acutus Vineyard and Wiggins. 4. Records from small streams in southern
Appalachians (Vineyard and Wiggins, 1987). October. (10)

N. atlanta Ross. 8. Record from small stream on Cumberland Plateau. November. (1)

N. concinnus McLachlan. 8, 25. Larvae collected from several small streams of Cumberland
Plateau. March, May. (5)

N. oligius Ross. 25. Collected along several small streams of Cumberland Plateau. Novem-
ber. (5)

N. ornatus Banks. 8, 25. Rare along several small streams of Cumberland Plateau. April.

(2)

N. securis Vineyard and Wiggins. 4. Record from small stream in southern Appalachians
(Vineyard and Wiggins, 1987). October. (1)

Brachycentridae

Brachycentrus chelatus Ross. 67. Endemic to the Coastal Plain, collected by sweeping.

March. (6)
B. numerosus (Say). 25, 34, 39, 54. Collected in a few streams in western Alabama, mainly as

larvae. March, May. (14)
Micrasemacharonis Banks. 17. Record from a small stream in lower Appalachians provided

by D.A. Etnier. (23)
M. n. sp. 51, 53, 65-67. Endemic to the Coastal Plain and common in small streams; being

described by J. Chapin and J. Morse. March-August. (226)
M. rusticum (Hagen). 12, 17, 25, 26, 31, 34, 39, 45, 51, 54, 66, 67. Common and widespread

throughout Alabama. March-May. (697)
M. wataga Ross. 1, 3, 5, 6, 8, 10-12, 14, 17-21, 25-27, 29, 31, 32, 34, 43, 45, 51, 52, 64. Commonly

collected throughout state, but most abundant in northern portion. April-October.

(1007)

Lepidostomatidae

Lepidostoma latipenne (Banks). 10-12, 17-19, 25, 31, 34,45. Occurring at scattered localities in
northern half of state; never collected in large numbers. May, June, September,
October. (77)

L. tibiale (Carpenter). 21, 28, 32. Uncommon; primarily collected in small Piedmont
streams. May, June. (23)

L. togatum (Hagen). 17, 18, 23, 28, 29, 31, 32, 54. Locally common, especially in small Pied-
mont streams. May, June, September, October. (147)

L. weaveri Harris. 25. Only known from type locality, a small temporary stream of the Cum-
berland Plateau. March. (39)

Vol. 101, No. 1, January & February 1990 61

Theliopsyche melas Edwards. 5. Only record from a small temporary stream of the lower

Appalachians. June. (1)
T. tallapoosa Harris. 31. Known only from a small Piedmont stream. May. (3)

PHRYGANEOIDEA

Phryganeidae

Agrypnia improba (Hagen). 45. Records from vicinity of Auburn University October

(28)

A. vestita (Walker). 22, 25, 34. Rare in our collections. October. (3).
Banksiola concatenate! (Walker). 39, 45. 53. Only taken on Coastal Plain, but rare in our

collections. April, May. (3)
Ptilostomis ocelli/era (Walker). 22, 23. 25, 26, 33. 36, 42, 66, 67. Primarily collected on Coastal

Plain, particularly the northern portion. April-July. (32)
P. postica (Walker). 17. 18, 24, 27, 29. 35. 39.43,45, 51, 66. Widespread in state, but rare in our

collections. April-June. September. (16)

BREVTTENTORIA
LEPTOCEROIDEA

Odontoceridae

Psilotretafrontalis Banks. 11,17, 18, 25, 27, 53. Uncommon from scattered, primarily northern

portions of the state. April. May. October. (29)
P. labida Ross. 1. 19. 53. Rarely collected from scattered localities in Alabama. May.

June. (26)
P. rufa (Hagen). 17. Only record provided by D.A. Etnier from a tributary to Shoal Creek in

northeastern Alabama. April. (1)

Calamoceratidae

Anisocentropus pyraloides (Walker). l.X, 1 1. 12. 14, 17. 18.25,28,29,31,32,36,39,42,45,51-53,

58, 64-67. Widely distributed in state, but most abundant in small Costal Plain streams.

April-August. (561)
Heteroplectron americanum (Walker). 8. 1 7. 1 8, 25, 28, 29, 5 1 . Primarily occurring at scattered

localities in northern Alabama, but never collected in large numbers. April-June

(15)

Leptoceridae

Ceraclea alabamae Harris. 5. Only known in Alabama from the Little River. June. (248)
C. alces (Ross). 5. A north-central North American species, locally abundant in the Little

River. June. (158)
C. amylu.s (Vorhies). 3. 1 1. 12. 14. 17. 25. 28. 29. 31. 34. 45. Widespread, but uncommon in

northern half of state. May, June. (295)
C. cancellata(Betten). 1-6.8-10,13.14,16-18,20,24-26.28.29.31.32.35.39.41.43.45.49.51-54.

63-67. Widespread and common in state. May-September. (2651)
C. diluta (Hagen). 5. 66, 67. Locally common, both on the Coastal Plain and in the lower

Appalachians. March-May. (152)

62 ENTOMOLOGICAL NEWS

C.Jlava (Banks). 1-4,9-11,13, 16-18, 20,22,24-26, 29,32, 34,35,37-39,41, 45, 49,51, 53, 60,63-

67. Widespread and common in state. May, June. (2025)
C.maculata (Banks). 1-6, 8-18,20-26, 28,29,31-37,39-43,45, 49-55, 58, 60, 63-67. Abundant in

our collections from throughout the state. April-October. (8396)
C. mentiea (Walker). 24. Record from the Tombigbee River provided by D.A. Etnier.

July. (11)
C. neffi (Resh). 4, 5, 54. Rarely collected in lower Appalachians and from a single location

on the Coastal Plain. May, June. (5)

C. nepha (Ross). 1-3, 5. 6, 8, 9-14, 16-18, 20-22, 24, 25, 29, 31-34, 39, 45, 51. 58, 66, 67. Wide-
spread in state, but most frequently collected above fall line. April-June. (438)
C. ophioderus (Ross). 25, 26, 33-35, 41, 45, 49, 5 1 , 53. 54, 60, 63-67. Primarily Coastal Plain in

distribution. May-September. ( 1 192)
C. protonepha Morse and Ross. 1-3, 5, 6, 8-14, 17, 18, 20-23, 25, 26, 29, 31, 32, 34. 39,45, 51, 53,

54, 58, 64, 66, 67. Widespread, but most frequently collected in northern half of state.

March-June. (mO)
C. resurgens (Walker). 34, 35, 66, 67. Collected only from a few localities on the Coastal Plain.

March, April. (18)
C. tarsipunctata (Vorhies). 1-6, 8-10, 12-14, 16-21, 24-26, 29, 31, 32, 34, 35, 37, 39-45,49-54, 58,

63-67. Widespread and common in Alabama. April-June. (4075)
C. transversa (Hagen). 1-3, 5, 6, 12-18, 25, 28, 29, 31, 32, 34, 43, 45, 51, 64, 66. Occurring

throughout Alabama, but most abundant in northern counties. April-July. (283)
Leptocerus americanus (Banks). 1, 8, 1 1-13, 17. 21 24, 33, 38, 39, 41, 43, 51, 53. Infrequently

collected in small streams of Cumberland Plateau and western Coastal Plain. April-
June. (84)
Mystacidessepulchralis (Walker). 1 . 2, 4-6, 8-11,14-18, 20, 22, 25, 26, 28, 29, 3 1 , 34, 45. Restricted

to northern half of Alabama, primarily in small streams. May-October. (233)
Nectopsyche Candida (Hagen). 1,8-12, 16-18.20-25,29, 3 1, 33, 35, 36, 39-42,45,49, 51-55, 58,64-

67. Common throughout state except lower Appalachians. May-August. (943)
N. exquisita (Walker). 1 , 3, 4. 9- 14. 16, 1 7. 20-22. 25, 26, 28, 29, 3 1, 32, 34-36, 39-41 , 45, 49, 50, 52,

54, 60, 63-67. Widespread and common in state. May-September. (743)
N. paludicola Harris. 66, 67. Endemic to small streams of the Coastal Plain. May-
August. (82)
N. pavida (Hagen). 1-6. 9-26. 3 1-37, 39, 40, 42, 43, 45, 51-55, 58. 60, 63-67. Most common and

widespread Nectopsyche in state. May-October. (1275)
N. spiloma (Ross). 34, 35. 66. 67. Restricted to large Coastal plain rivers. May-October.

(844)
Oecetisavara (Banks). 1,10-12, 20, 23-26, 28, 32-36, 39-42, 45, 49, 50, 53-55.65. Most frequently

collected on Coastal Plain, particularly western portion. May-September. (2893)
O.cineruscens(Hagen). 1,2,4,6. 11.14-16, 18,20.22,24-26,32,34-39,43.45,50-53,58,60,64,66.

67. Widespread in state, but never collected in large numbers. April-October. (241 )
O. davtona Ross. 52. 65-67. Rare; in small Coastal Plain streams. April. May. August. (12)
O.ditissa Ross. 1,2.5.6.8-12, 14-26. 29, 31, 33-41, 43.45. 51-55, 58, 60, 63-67. Widespread and

common in state. April-October. (877)
O.georgia Ross. 12, 17.22,23,25,36,42,51-53.65-67. Primarily occurring in Coastal Plain

streams. May-August. (171)

O. inconspicua (Walker). 1-6. 8-26, 28, 29, 3 1 -43, 45, 49-55, 58, 60. 63-67. One of the most com-
monly collected caddisflies in the state. April-October. (6487)

O. morsel Bueno-Soria. 34. 35. Only collected from the Cahaba River. June-October. ( 14)
O. nocturna Ross. 1 -6. 9-26. 28, 29. 3 1 -36. 38-43, 45, 49-53, 55. 63-67. Widespread and common

in Alabama. April-October. (2658)
O.osteni 'Milne. 11, 17, 20, 23-25, 33. 34. 36, 40, 45, 5 1,54. 60, 64-67. Primarily Coastal Plain in

occurrence. March-October. (451)

Vol. 101. No. 1, January & February 1990 63

O.persimilis (Banks). 1-3, 5. 6, 8-26. 29, 3 1-34, 36, 37,39-41. 45,49-55, 58, 60. 63-67. Widespread
and common in state. April-October. (2144)

O. scala Milne. 5. 18, 34. Collected from several streams of the lower Appalachians and in
the Cahaba River system. June-October. (401 )

O. sphyra Ross. 10-12. 16-18. 22-26. 31-37. 39. 41. 42. 45. 51-55, 58. 60. 63-67. Most frequently
collected on Coastal Plain, particularly along sand-bottomed rivers. May-Septem-
ber. (7697)

Setodes dixiensis Holzenthal. 34, 35. Restricted to the Cahaba River system. May. June. Sep-
tember. October. (496)

S. guttatus (Banks). 34. 54. Only collected at two localities, locally common. May. August.
September. (48)

S. incertus (Walker). 32. Restricted to several small Piedmont streams. May. (5)

S. stehri (Ross). 54. Only records from the Sepulga River, where S. guttatus also occurs.
May. September. (62)

Triaenodes aba Milne. 20. 39. Rarely collected in Alabama. May. June. (2)

T. cumberlandensis Etnier and Way. 5, 8, 17, 21, 22, 25. Collected from scattered localities on
Cumberland Plateau and lower Appalachians. May, June, August. (63)

T. dipsia Ross. 25. Only record from a small headwater stream of the Cumberland Plateau.
July.(l)

T. flavescens Banks. 1, 17. Rarely collected, in northern Alabama. June. (8)

T. florida Ross. 64. Collected from Lake Jackson. June. (12)

T. helo Milne. 60, 66, 67. Rare in collections; restricted to Coastal Plain. April-June. August,
September. (11)

T: /^//a (Walker). 1-6,8-12, 14. 16-18,20,22-26,29,31,33-37.39,42.45.49,51-55.58,60,63-67.
Widespread and common in Alabama. April-October. (814)

T. injusta (Hagen). 1-4, 8, 9, 14. 17. 18, 20-22, 25. 26, 34. 64. Primarily occurring in northern
portion of state. April-September. (199)

T. marginata Sibley. 1, 14, 17. 18,20,22, 25,29, 31,45. 51. As with T. injusta. most frequently
collected in northern half of Alabama. April-August. (59)

T. melaca Ross. 39. Collected among small streams with sand-gravel substrates. April.
May. (4)

T. n. sp. 42, 52, 65, 66. Restricted to small Coastal Plain streams; being described by K.L.
Manuel. May, June, August. (29)

T. nox Ross. 29. 34. 45. Collected in Piedmont region and from a tributary to the Cahaba River.
May (4)

T. ochracea Betten and Mosely. 1 , 4. 6, 8, 1 7, 22, 23, 25, 29, 5 1, 52. 66. 67. Collected infrequent-
ly throughout the state. April-June. (64)

T.perna Ross. 5,9, 16, 18,20,22.25.26,34.52.66,67. Distribution and abundance similar to
T. ochracea. May-September. (58)

T. smithi Ross. 8, 25, 35, 52, 53, 64. Uncommon in collections, primarily from western por-
tion of Alabama. May. June. August. (16)

T. taenia Ross. 17. 18, 39, 45. Most specimens taken in small Piedmont streams. May.
June. (5)

T. tarda Milne. 6. 14. 18. 20. 24. 25. 32. 38. 41. 49. 50. 51. Collected at scattered localities
throughout the state. May. June. (68)

7! tridonta Ross. 51. Collected from several small Coastal Plain streams with substrates of
sand-gravel. April. (17)

Molannidae

Molanna blenda Sibley. 17. 18. 20. 25. 26. 29. 45. 51. 53. 66. Widespread, but infrequent!)
collected in state. April-June. (27)

ENTOMOLOGICAL NEWS

HIGHLAND RIM PLATEAU

CUMBERLAND PLATEAU

VALLEY AND RIDGE

PIEDMONT
PLATEAU

EAST GULF

COASTAL

PLAIN

Figure 1. Counties and physiographic provinces of Alabama.

Vol. 101. No. 1, January & February 1990 65

M. tryphena Betten. 1, 8, 12, 14, 17, 18, 25, 28, 36, 45, 51, 53, 64-67. Widespread in state, but

never collected in large numbers. April-September. (56)
M. ulmerina Navas. 12-14, 17, 18, 20, 21, 24, 25, 28, 31, 39, 52-55, 58, 64, 66, 67. Most widely

occurring and frequently collected Molanna in the state. April-September. (108)

SERICOSTOMATOIDEA

Helicopsychidae

Helicopsyche borealis (Hagen). 1, 8, 10. 12, 14, 17, 19, 20, 25, 27, 28, 32, 34. 39, 50, 51, 53, 54.
Widespread and common in state, particularly above the fall line. April-October.
(871)

Sericostomatidae

Agarodes alabamensis Harris. 51. Only known from the type locality, a small Coastal Plain
stream. April. (2)

A. crassicornis (Walker). 12, 21, 22, 45, 52, 53, 58, 63, 65-67. Most common in small sand-
bottom streams of the lower Coastal Plain. April-June. (211)

A. griseus Banks. 8, 10, 17, 31, 32, 45. Collected above the fall-line, most frequently in the
Piedmont region. April-June. (73)

A. libalis Ross and Scott. 22, 24, 36, 65-67. Commonly collected in small Coastal Plain
streams. April-August. (307)

A. stannardi (Ross). 10-12. Uncommonly collected in several small streams of northwestern
Alabama. May, June. (10)

SUMMARY

This checklist, combined with those previously published (Harris, 1986a. 1988; Lago
and Harris, 1987), brings the total number of caddisflies in Alabama to 323 species. This
total is comparable to the 298 species reported from Tennessee (Etnier and Schuster, 1979)
and the 330 species from North and South Carolina (Unzicker et ai, 1982). This species
total is probably a reflection of the range of physiographic regions represented in the state
(fig. 1) as well as the result of intensive collecting. Collections from neighboring Mis-
sissippi, for example, which lies almost entirely within the East Gulf Coastal Plain yielded
only 139 caddisfly species (Holzental et al., 1982).

The 323 species of caddisflies were distributed among 57 genera and 17 families. Hyd-
roptilidae were best represented (97 species), followed by Leptoceridae (56 species), Hyd-
ropsychidae (47 species), and Polycentropodidae (32 species). These four families which
accounted for 72% of the total fauna in Alabama are noted for their ability to exploit warm
water habitats typical of Alabama (Wiggins, 1977).

ACKNOWLEDGMENTS

In the course of compiling this checklist many caddisfly workers offered invaluable
assistance in identifying or verifying identifications of specimens. The help of O.S. Flint,
Jr., E.R. Fuller, R.W. Holzenthal. K.L. Manuel, J.C. Morse, R.N. Vineyard, J.S. Weaver, G.B.
Wiggins, J.A. Wojtowicz, and the late D.G. Denning in this effort is greatly appreciated.
Several individuals provided material or made specimens and records available to us,
including W.E. Clark, Auburn University Insect Collection; K.C. McGiffen and J.D.
Unzicker, Illinois Natural History Survey; O.S. Flint, Jr., National Musuem of Natural His-
tory; K.L. Manuel, Duke Power Company; and D.A. Etnier, University of Tennessee. P.E.

66 ENTOMOLOGICAL NEWS

O'Neil, M.F. Mettee, and B.J. Armitage assisted in field collections. P.E. O'Neil and K.L.
Manuel reviewed an early draft of the manuscript and offered suggestions for improve-
ment. Peggy Marsh typed the numerous drafts of the paper.

LITERATURE CITED

Etnier, D.A., and G.A. Schuster. 1979. An annotated list of Trichoptera (caddisflies) of

Tennessee. J. Tennessee Acad. Sci. 54:15-22.
Harris, S.C. 1986a. Hydroptilidae (Trichoptera) of Alabama with descriptions of three

new species. J. Kansas Entomol. Soc. 59:609-619.
. 1986b. New species of caddisflies (Trichoptera) from Alabama. Proc. Entomol.

Soc. Washington 88:30-41.

1987. A new species of Agarodes (Trichoptera: Sericostomatidae) from south-

eastern United States. Proc. Entomol. Soc. Washington 89:74-76.

_. 1989. New Trichoptera from Alabama. J. New York Entomol. Soc. 97:309-316.

Holzenthal, R.W., S.C. Harris, and P.K. Lago. 1982. An annotated checklist of the cad-
disflies (Trichoptera) of Mississippi and southeastern Louisiana Part III: Lim-
nephiloidea and conclusions. Proc. Entomol. Soc. Washington 84: 513-523.

Lago, P.K., and S.C. Harris. 1987. An annotated list of the Curvipalpia (Trichoptera) of
Alabama. Entomol. News 98:255-262.

Parker, C.R., and J.R. Voshell, Jr. 1981. A preliminary checklist of the caddisflies
(Trichoptera) of Virginia. J. Georgia Entomol. Soc. 16:1-7.

Resh, V.H. 1975. A distributional study of the caddisflies of Kentucky. Trans. Kentucky
Acad. Sci. 36:6-16.

Sapp, C.D., and J. Emplaincourt. 1975. Physiographic regions of Alabama. Alabama
Geol. Survey Map 168.

Unzicker, J.W., V.H. Resh, and J.C. Morse. 1982. Trichoptera, p. 9.1-9.138 in A.R.
Brigham, W.V. Brigham, and A Gnilka (eds.). Aquatic insects and oligochaetes of
North and South Carolina. Midwest Aquatic Enterprises, Mahomet, IL.

Vineyard, R.N., and G.B. Wiggins. 1987. Seven new species from North America in the
caddisfly genus Neophvlax (Trichoptera: Limnephilidae). Ann. Entomol. Soc. Amer.
80:62-73.

Weaver, J.S., III. 1984. The evolution and classification of Trichoptera, part 1: The
groundplan of Trichoptera. p. 413-419 in J.C. Morse (ed.). Proceedings of the fourth
international symposium on Trichoptera. Dr. W. Junk Publishers, Series Entomo-
logica 30.

Wiggins, G.B. 1977. Larvae of the North American caddisfly genera (Trichoptera). Univ.
Toronto Press, Toronto, 401 p.

Vol. 101, No. 1, January & February 1990 67

COLLECTIONS OF DUFOUREA VERSATILIS
(HYMENOPTERA: HALICTIDAE) FROM IDAHO1

Sharon Luce Walsh^

ABSTRACT: Dufourea versatilis was collected outside California for the first time on
Mimulus nanus (Scrophulariaceae) at two areas in central Idaho during the summers of
1987 and 1988.

Insect visitors oi Mimulus nanus (Scrophulariaceae) were collected in
the summers of the 1987 and 1988 at two locations in central Idaho: one
at Craters of the Moon National Monument, Butte County and the other
north of Ketchum, Elaine County. The vegetation in both areas consists
of rabbit-brush (Chrysothamnus nauseosus), sagebrush (Artemesia triden-
tata), antelope bitter-brush (Purshia tridentata), and many annual and
perennial herbs. The mean annual precipitation in each area is approx-
imately 38 cm. Insect visitors were collected twice weekly starting in late
May when the plants started blooming and continued until plants stopped
blooming. In 1987 insects were collected through July, whereas in 1988,
because of drought conditions, all M. nanus plants were finished bloom-
ing by the first of July.

One insect collected, Dufourea versatilis (Bridwell) (Halictidae), pre-
viously had been collected only in California (Hurd 1979). Both male
and female bees were collected, but during 1987 only 1 male was collec-
ted at the Craters site whereas 1 male and 3 females were collected from
the Ketchum site. In 1988 noD. versatilis were collected at the Craters site,
but 19 of the 25 insect visitors collected at the Ketchum site were D. ver-
satilis. Of these 19, 14 were females and 5 were males.

Dufourea versatilis is an oligolege of Mimulus species in California
(Hurd 1979). In all sampling periods during this study, D. versatilis was
observed foraging only on M. nanus at the two areas. It is likely that it has
never been collected outside California because insect pollinators of
most Mimulus species have not been collected elsewhere. I predict that D.
versatilis will be found in any location that has populations of M. nanus,
including eastern Oregon, eastern Washington, northern Nevada, wes-
tern Montana, central and western Idaho, and western Wyoming around
Yellowstone and Grand Teton National Parks.

ACKNOWLEDGMENTS

I would like to thank J. Anderson, R. Anderson, N. Huntly, K. Holte, and two anon-
ymous reviewers for comments on an earlier version of this manuscript. Additionally. I

1 Received July 5, 1989. Accepted November 9, 1989.

2Department of Biological Sciences, Idaho State University, Box 8007. Pocatello, ID 83209

ENT. NEWS 101(1): 67-68, January & February, 1990

68 ENTOMOLOGICAL NEWS

thank the personnel, especially T. Griswold, at the U. S. D. A. Bee Lab at Utah State Univer-
sity in Logan, Utah for identification of the collected bees and wasps and the National Park
Service at Craters of the Moon National Monument for their cooperation during the dura-
tion of this project. This research was funded by a Grant-in-Aid of Research from Sigma XI
and a grant from the Idaho State University Graduate Student Research Committee.

LITERATURE CITED

Hurd, P.O. 1979. Superfamily Apoidea. pp. 1741-2209. In: Krombein, K. V., Hurd, P. D.,
Smith, D. R., and Burks, B. D. 1979. Catalog of Hymenoptera in America North of Mex-
ico, Vol. 2. Smithsonian Institution Press, Washington, D.C.

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VOL. 101

USISSN0013-872X
MARCH & APRIL, 1990 NO. 2

ENTOMOLOGICAL NEWS

'O immigrant synanthropic flower flies (Diptera: Syrphidae)
new to North America

F.C. Thompson, F.D. Fee, L.G. Berzark 69

Redescription of males, biological notes, & identifica-
tion of American Acanthochalcis (Hymenoptera:
Chalcididae) Jeffrey A. Halstead

Lepidostoma (Nosopus) ozarkense (Trichoptera: Lepidosto-
matidae), a new species from Arkansas

O.S. Flint, Jr., P.A. Harp

A new species of Neotrichia (Trichoptera: Hydroptilidae)
from Colorado with additions & corrections to the
distribution & records of Colorado Trichoptera

D.E. Ruiter

A new species of Zonana (Homoptera: Cicadellidae)

from Venezuela Paul H. Freytag

Eurypauropus spinosus (Arthropoda: Pauropoda: Eurypauro-
podidae) from Arkansas & a key to the North American
Eurypauropus species Robert T. Allen

Dyscinetus morator (Coleoptera: Scarabaeidae) feeding on
roots of azaleas (Rhododendron spp.) - a scientific note

C.L. S fames. Jr.

Seasonal flight activity of male velvet ants (Hymenoptera:
Mutillidae) in south Florida M. Deyrup, D. Manley

Taxonomic distribution & phylogenetic significance of the

male hairpencil in the Tortricinae (Lepidoptera: Tortricidae)

John W. Brown 109

Acalyptrate Diptera reared from higher fungi in north-
eastern Ohio B. Bunyard, B.A. Foote 117

Improved culture techniques for mass rearing Galleria
mellonella (Lepidoptera: Pyralidae)

F.A. Eischen, A. Dietz 123

ANNOUNCEMENTS

BOOKS RECEIVED & BRIEFLY NOTED

SOCIETY MEETING OF FEBRUARY 23, 1990

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Vol. 101, No. 2, March & April 1990 69

TWO IMMIGRANT SYNANTHROPIC FLOWER

FLIES (DIPTERA: SYRPHIDAE) NEW TO

NORTH AMERICA1

F. Christian Thompson , Frank D. Fee-', Larry G. Berzark'*

ABSTRACT: Two flower flies are recorded from North America for the first time: Eris-
talinus(Eristalodes)taeniops -Florida; and Syrittaflaviventris- Texas and Mexico. Diagnostic
characters are given for these species, along with biological data.

Two flower flies are recorded from North America for the first time:
Eristalinus (Eristalodes) taeniops fWiedemann) - Florida; and Syritta
flaviventris Macquart - Texas and Mexico. Both are hemisynanthropes,
close associates of human ecosystems, and common filth flies in the Old
World. Diagnostic characters are given for these species, along with
biological data.

Two synanthropic flower flies, one presently unknown in the New
World and the other unknown from North America, were recently col-
lected for the first time in the United States. Syrittaflaviventris Macquart
was collected in southern Texas and Mexico, and Eristalinus taeniops
(Wiedemann) was collected in southern Florida. Both belong to pre-
dominantly Old World tropic groups and are properly hemisynanthropes
(Povolny, 1971).

Key to the New World species of Syritta

1. Face silvery white pollinose; antenna extensively dark, from all black in most males to
basoflagellomere more than 2/3 r dark brown in females and some males; fore and
midlegs entirely orange; wing without spurious vein (fig. 4) and with orange veins,
almost completely bare, only sparsely microtrichose on apical margins; male hind
femur with a large basoposterior ventral tubercle (fig. 5); male hind tibia expanded
apically; male abdomen with orange areas on 2nd and 3rd segments much more exten-
sive and only narrowly separated medially (female similar, figs. 6-7); male 4th sternum
deeply excavated (depth much greater than breadth) and with strong yellow
bristles flaviventris Macquart

— Face golden pollinose; antenna entirely or more than 2/3rd pale orange; fore and
midlegs not brownish black posteriorly on femora and apically on tibiae and on tarsi;
wing with spurious vein (fig. 3) and brownish black veins, more extensively micro-
trichose, microtrichose areas extending into apical cells; male hind femur without
tubercle; male hind tibia slender; male abdomen with orange areas on 2nd and 3rd
segments reduced and broadly separated medially (female similar, figs. 8-9); male 4th
sternum shallowly excavated (breadth much greater than depth ) and with only few fine
hairs pipicns Linnaeus

'Received April 28, 19X9. Accepted October 10, 1989.

•^Systematic Entomology Laboratory, ARS, USDA,c/o U. S. National Museum NHB-168,

Washington, DC 20560

351 1 West Ridge Avenue, State College, PA 16X03
4521 46th Street, Sacramento. CA 95X19

ENT. NEWS 101(2): 69-74, March & April. 1990

70 ENTOMOLOGICAL NEWS

Syritta flaviventris Macquart

Syritta flaviventris Macquart, 1842: 135. Type locality: Senegal. Type depository: Male,

MNHN, Paris.
Syritta spinigera Loew, 1848: 331. Type localities: Greek islands, Sicily & Turkey. Type

depository: synrypes, males & females, ZMHU, Berlin.

Syritta flaviventris andpipiens are so similar that no description is given.
The above key serves as a differential diagnosis for these species.

North American Records: MEXICO, NUEVO LEON, Apodaca, "E. L.
Mezquital", 26 May 1984 (C. Alvarez Pereyra; IIBIII lot # 84-07029.)
USA, TEXAS, Hidalgo County, Bentsen-Rio Grand Valley State Park, 2 1
October 1984 (F. D. Fee), 1 female, Relampago, 15- 19 October 1986 (F. D.
Fee), 6 males, 1 female. The material from Texas was collected in a
locality about 48 miles east on the Mexican specimen (initial record).
The specimens from Relampago were all collected in or along the banks
of an abandoned drainage ditch or canal. The males were patrolling and
visiting flowers of Schinus, Serjania and Polygonum. The female was
taken on Polygonum. Voucher specimens deposited in the National
Museum of Natural History, Smithsonian Institution, Washington; other
specimens retained in Fee Collection.

Syritta flaviventris is readily distinguished from pipiens, the only other
New World species of the genus, by the absence of a spurious vein (fig. 4)
and the presence of a strong basoposterior ventral spur on the male hind
femur (fig. 5); and from other Old World congeners by male genitalic
characters (Thompson, 1972: 170, fig. 69).

Syritta is not indigenous to the New World but has been introduced with
Man. The larvae of Syritta breed in almost any kind of waste, andpipiens
has been frequently reported to breed in human feces(Farrar 1987: 361,
Henning 1952: 189). Flaviventris was first recorded from the New World
by Fluke (1960), who recorded the species from Brazil, Sao Paulo, collec-
ted in 1954. Marnef (1967) recorded it from Chile as Austrosyritta cortesi
Marnef (synonymy by Thompson 1971), and Argentina was added to its
New World distribution by Thompson, et at. (1976: 119). In the Old
World, S. flaviventris ranges from the Mediterranean (Spain to Bulgaria
and Turkey), south to South Africa (Cape of Good Hope), and is found
on Saint Helena. Campos and Pena (1973: 225; Smith and Vockeroth
( 1980: 507)) recorded it from Easter Island. Syritta pipiens is found through-
out northern North America and is recorded as far south as central Mex-
ico (Durango and Chihuahua) (Thompson, et al. 1976: 119).

Vol. 101, No. 2, March & April 1990 71

Key to the New World Species of Eristalinus

1. Eye with large brown fasciae in addition to smaller brown puncta (fig. 1)

taeniop.s (Wiedemann)

— Eye with only small brown puncta (fig. 2) aeneus (Scopoli)

Eristalinus (Eristalodes) taeniops (Wiedemann)

Eristalis taeniops Wiedemann, 1818: 42. Type locality: South Africa, Cape of Good Hope.
Type depository: Males & females, NM, Vienna.

Head: black, extensively gray pollinose and white pilose; face with a medial and sub-
lateral shiny vittae; frontal triangle and front partially black pilose; vertex shiny; antenna
black, except basoflagellomere rarely orange basally and on ventral 1/4 or less, arista
orange becoming brown apically; eye yellow with brown fasciae, light brown pilose.

Thorax: black, gray pollinose and white pilose; mesonotum indistinctly vitiate;
scutellum yellow, white pilose except with some black pile medially; plumula. squama and
halter yellowish white. Wing hyaline, bare. Legs: femora reddish brown to black except
orange on apex; tibiae orange basally, brownish black apically; front tibia orange on basal
2/3, mid tibia on basal 3/4, rarely entirely orange, hind tibia on basal 1/4 or less; tarsi orange
on basal 2 tarsomeres, dark brownish on apical 3 tarsomeres; leg pilose yellowish white
except for a few black hairs apicoventrally on hind femur and extensively black pilose on
hind tibia.

Abdomen: first tergum orange on lateral quarter, black medially, gray pollinose, white
pilose; 2nd tergum orange except narrowly brownish black on basal and apical margins,
yellowish white pilose except black pilose on dark apical margin, slightly gray pollinose
medially; 3rd tergum orange on basal 1/4 to 1/2, apically brownish black, dull, gray
pollinose on medial 1/3 in males, basal 1/3 in females, yellow pilose except fora few apical
black hairs; 4th tergum black, rarely orange basomedially. dull black pollinose. with
grayish white sinuous fascia on basal 1/2 in male, grayish white pollinose on basal 1/3 in
female, shiny on apical margin, white pilose; 5th tergum black, black pollinose. white
pilose; venter white polose and sparsely white pollinose, basal sterna usually orange,
except rarely brownish black medially, 4th sternum brownish black. Male genitalia black.

New World record: USA, FLORIDA, Dade County, Florida City. 7
December 1985 (L. G. Bezark) 1 male (USNM). The specimen was swept
from Bidens, a common weedy composite, along a hedgerow of tamarisk
trees (Tamarix] adjacent to a field of cultivated tomatoes.

Eristalinus taeniops is easily distinguished from all other New World eris-
talines (species with sinuate R4+5 vein) by its fasciate eyes (fig. 1). The
only other species of Eristalinus known from the New World, aeneus
Scopoli, has punctate eyes (fig. 2), and all other New World eristalines
have no maculation on the eyes. Eristalinus taeniops belongs to a small
group of species of Eristalinus which have distinct fasciae on the eyes in
addition to puncta.

ENTOMOLOGICAL NEWS

-^^^^••M^^^^^MM^^^^^M*

Figs. 1-5. 1-2. Heads of Eristalinux species, lateral view. 1. E. taeniops (Wiedemann). 2. E.
aeneus Scopoli. 3-4. Wings ofSyritta. dorsal view. 3. S.pipiens Linnaeus. 4. S.flaviventris Mac-
quart. 5. Hind leg of Syritta flaviventris Macquart. lateral view.

Vol. 101, No. 2, March & April 1990

Eristalinus is not indigenous to the New World, but two species have now
been introduced. Eristalinus larvae, commonly called rat-tailed maggots,
have been recorded to breed in putrid waters associated with man, such
as sewers, privies, etc (Ferrar 1987: 359-360, Hennig 1952: 184-185). Eris-
talinustaeniops has never previously been recorded from the New World.
In the Old World, the species ranges from the Mediterranean (Spain to
Greece and Bulgaria), east to Pakistan, and south to South Africa (Cape
of Good Hope). Eristalinus aeneus Scopoli ranges from California to
Ontario and New Hampshire, south to Texas and Georgia; and in the
Old World aeneus ranges throughout the Palaearctic Region (Peck 1988:
182, Knutson el alia 1975:347) and has been introduced into Tanzania
in Africa (Smith and Vockeroth 1980: 501) and Wake Island, the Hawaiian
and Gilbert Islands in the Pacific Ocean.

6. Male 7. Female

flaviventris

8. Male 9. Female

pi pi ens

Figs. 6-9. Abdominal patterns of Syritta species, dorsal view. 6. S. flaviventris. male. 7. S.
flaviventris. female. 8. 5. pipiens. male. 9. S. pipienx. female.

74 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

We thank David A Nickle, Douglass R. Miller and Norman E. Woodley of the Sys-
tematic Entomology Laboratory, USDA Washington, D.C.; Amnon Freidberg of Zoology
Department, Tel-Aviv University, Israel; Wayne N. Mathis of the Smithsonian Institution
(USNM), Washington. D. C.; and J. R. Vockeroth of Biosystematics Research Centre,
Agriculture Canada. Ottawa, for their critical reviews of the manuscript.

LITERATURE CITED

Campos S., L. & L. E. Pena G. 1973. Los insectos de Isla de Pascua (Resultados de una

prospeccion entomologica). Revta Chil., Ent. 7: 217-229.
Ferrar, P. 1987,. A guide to the breeding habits and immature stages of Diptera Cyclor-

rhapha. Entomonograph 8, 2 vols., 907 pp. E. J. Brill / Scandinavain Science Press.

Leiden & Copenhagen
Fluke, C. L. 1960. Concerning the Catalogue of Neotropical Syrphidae. Revta Brasil. Ent.

9: 169.
Hennig, W. 1952. Die Larvenformen der Diptern. Eine Ubersicht uber die bisher

bekannten Jugendstadien der zweiflugeligen Insekten. 3. Teil. vii + 628 pp. Akademie-

Verlag GmbH., Berlin [syrphids, pp. 159-190]
Knutson, L. V., F. C. Thompson & J. R. Vockeroth 1975. Family Syrphidae. pp. 307-374.

In Delfinado, M. D. & D. E. Hardy (eds.), A Catalog of the Diptera of the Oriental

Region. Vol. 2, Suborder Brachycera through Division Aschiza, Suborder Cyclor-

rhapha. ix + 459 pp.. Univ. Press Hawaii, Honolulu.
Loew, H. 1848. Dipterologisches. Stettin, ent. Ztg 9: 329-332.
Macquart, J. 1842. Dipteres exotiques nouveaux ou peu connus. Mem. Soc. R. Sci. Agric.

Arts, Lille 1841: 65-200, 22 pis. Also, published separately as his "Dipteres exotiques

nouveaux ou peu connus," VOL. 2, Pt. 2, pp. 5-140, 22 pis. Paris 1842.
Peck, L. V. 1988. Syrphidae. Pp. 1 1-230 in Soos, A (ed.). Catalogue of Palaearctic Diptera.

Vol. 8, Syrphidae — Conopidae, 363 pp., Akademiai Kiado, Budapest.
Povolny, D. 1971.Synanthropy. Pp. 16-54./«Greenberg,B., Flies and Disease. Vol. 1, Ecol-
ogy, Classification and Biotic Associations, xii + 865 pp., Princeton Univ. Press.

Princeton, New Jersey.
Smith, K.G. V. & J. R. Vockeroth 1980. 38. Family Syrphidae. Pp. 487-510. In Crosskey,

R. W. (ed.), Catalogue of the Diptera of the Afrotropical Region. 1437 pp., British

Museum (Natural History), London.
Thompson, F. C. 1 971. The genus Nepenthosvrphus with a key to world genera ofTropidini.

J. Kansas Ent. Soc. 44: 523-534.
1972. Acontribution to a generic revision of the Neotropical Milesinae (Diptera:

Syrphidae). Arquivos Zool., Mus. Zool. Univ. Sao Paulo 23: 73-215.
Thompson, F. C., J. R. Vockeroth & Y. S. Sedman. 1976. Family Syrphidae. Cat. Dipt.

Amer. s. United States 46, 195 pp.
Wiedemann, C. R. W. 1818. Neue Insecten vom Vorgebirge derGuten Hoffnung. Zool.

Mag. (Wiedemann's) 1(2): 40-48.

Vol. 101, No. 2, March & April 1990 75

REDESCRIPTION OF MALES, BIOLOGICAL

NOTES, AND IDENTIFICATION OF AMERICAN

ACANTHOCHALCIS (HYMENOPTERA:

CHALCIDIDAE)i

Jeffrey A. Halstead^

ABSTRACT: Males of Acanthochalcis nigricans and /I. unispinosa are redescribed, voucher
specimens designated, diagnostic characters illustrated, and a key to the American species
presented. Characters to distinguish Acanthochalcis from other Nearctic Chalcididae are
presented. The taxonomic and biological information on Acanthochalcis is summarized
and new hosts, rearings, and floral visitations are recorded. A third species, A. gigas, is
known from Madagascar.

Acanthochalcis are the largest chalcidids in the Nearctic region with
males ranging in length from 6 to 1 1 mm and females from 7 to 24 mm.
These wasps are black with reddish brown to brown legs and smokey or
yellowish wings. Females are unique among Nearctic Chalcididae in
having a spear-like ovipositor which projects posteriorly a distance
equal to or greater than the length of the abdomen. Acanthochalcis are
uncommonly collected, though have been taken by sweeping flowering
vegetation and netting individuals around host inhabiting trees and
shrubs.

To improve the taxonomic information on American Acanthochalcis,
the males of A. nigricans Cameron and A. unispinosa Girault are rede-
scribed, the species distinguished in a key, the taxonomic history re-
viewed, and characters presented to distinguish this genus from other
Nearctic Chalcididae. This information improves the brief original
male descriptions and is needed to identify Acanthochalcis and its Amer-
ican species. Male voucher specimens are deposited with their female
holotype for use in future studies. The redescriptions will also be useful
to delineate the presently unknown male of A. gigas Steffan (from Mada-
gascar), and help establish the relationship between American and
Madagascaran Acanthochalcis.

Both American Acanthochalcis are parasitoids of buprestid beetle lar-
vae which inhabit oak trees (Quercus spp.) (Halstead and Haines 1985);
however, new information indicates buprestid hosts in saltbush (Atri-
plex sp.), ocotillo (Fouguieria splendens), and Baccharis sp. Overall, the
biological information on Acanthochalcis is very limited. This informa-
tion is summarized and new host, rearing, and floral visitation records

^Received March 23, 1989. Accepted June 30, 1989.
-California State University Fresno, Fresno, CA 93740.
Present address: 1 10 W. Barstow #112. Fresno, CA 93704.

ENT. NEWS 101(2): 75-80, March & April. 1990

76 ENTOMOLOGICAL NEWS

are presented to encourage and aid future research.

Steffan (1959) recognized \nAcanthochaIcis two American species A
nigricans (Cameron 1884) and A. unispinosa (Girault 1917, 1921; in Gordh,
etal. 1979) and A. gigas (Steffan 1950) from Madagascar. Drawings of the
apex of both the abdomen and scutellum for nigricans were presented.
Characters for the previously unknown males of A. nigricans and A. unis-
pinosa were presented in a key. The male of A. gigas is unknown. Steffan
also proposed subgenera, placing nigricans mAcanthochalcis, unispinosa
in Belochalcis, and gigas in Trypanochalcis, but the designation of sub-
genera for each species seems unwarranted on the basis of the few
species and at least for nigricans and unispinosa on morphological
grounds.

Acanthochalcis, the only member of the tribe Cratocentrini in the
Nearctic region, may be distinguished from other Nearctic Chalcididae
by the following characters: head without horns on dorsal area of frons,
antennae inserted in center of frons, abdomen sessile, hindtibiae with
one apical spur and the apex obliquely sinuate (Brachymerinae); post-
marginal vein much longer than marginal vein, scrobe cavity almost
touching vertex of clypeus, mesopleuron deeply excavated by a aceta-
bulum which receives the mesotibia and mesofemur in response, (9)
with 4 visible tergites, (cT) with 7 visible tergites (Cratocentrini); pro-
podeum produced laterally into a flattened triangular mass (dorsal
view).

Key to American species of Acanthochalcis Cameron

1. Propodeum dorsally with a median tooth (Fig. 9), in lateral view with posterior margin
truncate; hindfemur narrowly ovoid (Fig. 4); (9) epipygidium with ventral margin
straight: (c?) abdomen dorsoventrally flattened (Figs. 2. 6) unispinosa Girault

1.' Propodeum dorsally without a tooth, in lateral view projecting posteriorly as a triangu-
lar tooth which overhangs base of hindcoxae (Fig. 8); hindfemur broadly ovoid (Fig. 3);
(9) epipygidium with ventral margin toothed (Fig. 7); (cf) abdomen oval (Figs. 1.
5) nigricans Cameron

Acanthochalcis nigricans Cameron, Male Redescription

Figures 1,3,5,7,8

Type-species 9 (Cameron 1884). cT description (Steffan 1959).

Length: 1 1mm. Black with labrum clypeus, coxae, trochanters, femora (hindfemora
apically and basally). tibiae, tarsi, tergites 1-5 posteriorly and ventrally, sternites 1-6 pos-
teriorly, epipygidium, hypopygidium, and hindwing vention reddish brown. Pubescence
white, dense patches on axillae posteriorly, propodeum laterally, hindcoxae dorsally, and
subme Jian length of tergite 6.

Pronotum with anterior, dorsolateral margin rounded, posterior margin emarginate;
propodeum laterally produced as a flat triangular posterior projection, in lateral view pos-

Vol. 101, No. 2. March & April 1990

terior margin produced as a tooth, with a strong transverse basal carina and two lon-
gitudinal submedian carina, rectangular area between these carinae polished; hindfemur
broadly ovoid, 1 1 teeth along ventral margin, a sharp tooth on inside at base, outer and
inner surface polished, setal depressions separated by a distance equal to their own
diameter, inner surface with sparse setal depressions; hindcoxa length 3X its greatest
width, inner surface with sparse setal depressions; forewing infuscate, apical margin with a
darkened band; hindwing hyaline, slightly infuscate apically; stigma with a prominant
uncus; postmarginal vein 2.5X length of marginal vein.

Abdomen sessile, ovate, dorsal margin convex in lateral view; tergite 2 (T2) with 3-4
punctures at median length; tergites laterally perpendicular to body axis, with dense
setigerous punctures except for coriaceous band along posterior and ventral margins of Tl -
5; epipygidum triangular, matte, basal 1/6 punctate, with a flat median and carinate sub-
lateral ridge; sternites convex, stemites 1-5 (Sl-5) punctate medially. S6 punctate
throughout.

Types. — Holotype female collected from Sonora, Mexico; type in the British Museum
of Natural History (BMNH). I designate a male voucher specimen with a red label:
"Voucher Specimen. Male redescription./fra/7//)0c7itf/m nigricans Cameron, det. J.A. Hal-
stead 1987" with data: "Arizona. Pima Co., Madera Canyon, Vlll-24-\911,ProsopisjuIiflora.
R.W. Brooks." Voucher male deposited in BMNH.

Variation. — Males 8-1 1 mm, usually about 10mm. Females 7-24 mm, usually about 20
mm. Halstead ( 1987) noted three rubinistic colored (i.e., typically black areas are reddish
brown) males from El Salvador. A male from 23 mi S. Matias Romero, Oaxaca, Mexico, IV-
22-1962, F.D. Parker (University of California Davis collection) is also this color.

Host/rearings. — A probable host Chrysobothrisfemorata (Olivier) (Coleoptera: Bup-
restidae) was discussed by Halstead and Haines (1985) and additional probable hosts
Dicerca horni Crotch or Polycesta California (LeConte) (both Coleoptera: Buprestidae) in
interior live oak (Quercus Wisliienu) suggested. Label data on specimens denotes the
following host associations or rearings. Three females (Palm Springs, San Bernardino Co.,
California) were reared from Chrysobothris sp. in the roots of saltbush (Atriplex sp.). A
female (6 mi E. of Apache Junction on Hwy. 88, Maricopa Co., Arizona) was reared from
Baccharis sp. A female (Tucsun. Pima Co., Arizona) was associated with Chrysobothris
edwardsi Horn (Buprestidae) in a dead ocotillo (Fouquieria splendens).

Floral visitations/possible host substrates. — Atriplex. Baccharis glutinosa. B. ne-
glecta, Bebbia juncea. Encelia sp., Encelia farinosa, Flourensia cernua associated with Tachar-
della cornuta (Homoptera: Lacciferridae), Helianthits annuus, Koeberlinia, Larrea, Lupinus
inflatus. Placelia distorts. Prosopis chilensis, P. juliflora. Sphaeralcea. and Stanleya pinnata.

Distribution. — California east to Kansas, south through Mexico to Costa Rica
(Halstead 1987).

Acanthochalcis unispinosa Girault, Male Redescription

Figures 2, 4, 6, 9
Original description 9 (Girault 1917. 1921). tf description (Steffan 1959).

Length 1 1 mm. Black with tegulae, tibiae, tarsi, and posterior and ventral margins of
Tl-5 brown.

Pronotum with anterior, dorsolateral margin squared, posterior margin deeply emar-
ginate; propodeum laterally produced as a flat triangular posterior projection, though not
as acute posteriorly as in nigricans: in lateral view posterior margin squared, with a
transverse medially arching carina and a strong median carina which meet to torm a raised
acute toothlike process, area between these carina with a reticulation of carinae; hind-
femur narrowly ovoid, ventral margin with 12 teeth, without a tooth on inside at base, outer
and inner surface with dense setal depressions and densely setose; hindcoxa length 5X its

ENTOMOLOGICAL NEWS

Figs. l-6.Acanthochalcis spp. (males)./!, nigricans: 1, Abdomen, lateral view; 3, Hindfemur.
lateral view; 5, Abdomen, dorsal view. A. unispinosa: 2, Abdomen, lateral view; 4 Hind-
femur, lateral view; 6, Abdomen, dorsal view. Scale lines 2.0 mm.

Vol. 101, No. 2, March & April 1990

TOOTH

Figs. l-%.Acanthochalcisnigricans: V.epipygidium of female, lateral view; 8. propodeum and
metapleuron, lateral view. Fig. 9. Acanthochalcis unispinosa, propodeum, only carinae in
medical area drawn, dorsal view.

greatest width, inner surface with dense setal depressions and apical 1/3 with prominant
transverse striae; forewing infuscate, with a yellow tint; hindwing infuscate, darker from
under marginal vein to apex; stigma with a prominant uncus; postmarginal vein 2X length
of marginal vein.

Abdomen sessile, dorsoventrally compressed, dorsal margin flat in lateral view; T2
with 6-7 punctures at median length; tergites (especially T2-5) flaring outward from body
axis, with dense setigerous punctures except for a smooth band along posterior and ventral
margins of Tl-6; epipygidium triangular, coriaceous, basal 1/4 punctate, with a median
and submedian carina; sternites convex, punctures on S2-5 originating from anterior
margin and forming a half circle shaped area; SI and S6 punctate medially.

Types. — Holotype female collected from Belfrage, Texas; type in the United States
Museum of Natural History (USNM). I designate a male voucher specimen with red label:
"Voucher Specimen. Male Redescription./lraA7//!oc/!a/m unispinosa Girault, del. J.A, Hal-
stead 1987" with data: "CA. Tulare Co., Ash Mtn. Kaweah Powerhouse #3. Vl-19-1982, coll.
R.D. Haines." Voucher male deposited in USNM.

Variation. — Males 6-11 mm, usually about 10 mm. Females 17-24 mm, usually about
21 mm.

Host/rearings. — A probable host Polycesta California (Coleoptera: Buprestidae) in
Valley Oak (Quercus lobata) was suggested by Halstead and Haines (1985). Label data on
specimens denotes a rearing and a possible host substrate. A female (Pinon Flats, San
Diego Co. California) was reared from scrub oak (Quercus dumosa). A female (near
Eskimizin, Pima Co., Arizona) was flying about dead limbs of palo verde (Cercidium
sp.).

Distribution. — California, Arizona, Texas.

ACKNOWLEDGMENTS

I thank D.J. Burdick (California State University, Fresno), R.D. Haines (Tulare County
Agricultural Commissioner's Office, Visalia, California), N.J. Smith (Fresno County De-
partment of Agriculture, Fresno, California), S.R. Shaw (Harvard University, Cambridge.
Massachusetts), and an anonymous reviewer for editorial comments on earlier drafts of
this paper. I thank E.E. Grissell (Systematic Entomology Laboratory. ARS-USDA) for the
opportunity to examine types and Z. Boucek (CAB Institute of Entomology, London.
England) for comparing specimens of A. unispinosa. I lastly thank the Kings River Conser-
vation District. Fresno, California for the use of word processing equipment.

80 ENTOMOLOGICAL NEWS

LITERATURE CITED

Cameron, P. 1884. Biologia Centrali- Americana. Insecta. Hymenoptera: (Families Ten-

thredinidae-Chrysididae.). Vol. I. 487 pp.

Girault, A.A. 1917. NewChalcid Flies. Privately published, Glenn Dale, Maryland, 6pp.
1921. New serphidoid cynipoid, and chalcidoid Hymenoptera. Proc. U.S. Natl.

Mus. 58: 177-216.
Gordh, G., A.S. Menke, E.G. Dahma, and J.C. Hall. 1979. The privately reprinted

papers of A.A. Girault. Mem. Amer. Entomol. Instil. No. 28: 400 pp.
Ha 1st cad, J.A. 1987. New distribution records for Acanthochalcis nigricans Cameron-
including Central America (Hymenoptera: Chalcididae). Pan-Pac. Entomol. 63: 236.
. and R.D. Haines. 1985. On the biology of Acanthochalcis nigricans Cameron and

Acanthochalcis unispinosa Girault (Hymenoptera: Chalcididae). Pan-Pac. Entomol. 61:

227-228.
Peck, O.C. 1963. A catalog of theNearcticChalcidoidea (Insecta: Hymenoptera). Canad.

Entomol. Suppl. 30: 1092 pp.
Steffan, J.R. 1950. Note sur la tribu des Cratocentrini (Hym.: Chalcididae Brachy-

merinae); description de deux especes nouvelles. Bull. Natl. Mus. Nat. Hist. Paris 22:

596-602.
. 1959. Revision de la tribu des Cratocentrini (Hym.: Chalcididae). Acta Entomol.

Mus. Natl. Prague 33: 287-325.

INSECT FIELD DAY

The fourth annual Insect Field Day sponsored by the American Entomological Society
and the Young Entomologists' Society will be held Saturday June 23, 1990 from 9 to 5 at the
Fair Hill Nature Center in Fair Hill, Maryland. Entomologists, young and old. amateur or
professional and their friends are invited to join in a day of collecting, identifying, sharing,
and socializing. For registration forms, directions, program, and other information write
to: Insect Field Day, American Entomological Society. Academy of Natural Sciences. 19th
& Race Sts., Philadelphia, PA 19103.

Vol. 101, No. 2, March & April 1990 81

LEPIDOSTOMA (NOSOPUS) OZARKENSE

(TRICHOPTERA: LEPIDOSTOMATIDAE),

A NEW SPECIES FROM ARKANSAS1

Oliver S. Flint, Jr.2, Phoebe A. Harp3

ABSTRACT: Lepidostoma (Nosopus) ozarkense, a new species from northern Arkansas and
eastern Oklahoma belonging to the Modestum Group, is described. The distinctive male
and female genitalia are described and figured. The spring emerging adults were taken
near small streams which may dry up in summer. Details of the physicochemical and
biological characteristics of the topotypical stream are given.

While surveying benthic organisms of intermittent streams. Harp
collected examples, both adult and larvae, of a number of Trichoptera
that were sent to Flint for identification. In these collections a species of
Lepidostoma that appeared to be undescribed was discovered. Conse-
quently, correspondence with D.E. Bowles at the University of Arkansas
revealed that he had additional examples of this species from other
localities in Arkansas and Oklahoma. The recently published revision
of the New World species of Lepidostoma (Weaver 1988) confirms that
this species is heretofore unknown.

Lepidostoma (Nosopus) ozarkense Flint & Harp, new species

Adult: Length of forewing, male 6— 7mm, female 7— 7.5mm. color in alcohol uniformly
dark brown. Venation in both sexes typical of group (Weaver 1988, Figs. 46a. 46b). Male
with scape about 3 times as long as broad; maxillary palpus apparently one-segmented,
filled mesally with long spatulate hairs (essentially as in Weaver 1988, Fig. 32). Male
genitalia (Figs. 1-4): Ninth segment annular, widened ventrally, with a ventrolateral brace.
Tenth tergum elongate, blunt apically. with scattered setae arising from protuberant bases;
with a narrow, dorsomesal incision about a third length of segment, with a darkened, inter-
nal sclerotization around base of incision; with a basoventral process as long as dorsal lobe
and tapering to an apical point in both dorsal and lateral aspects. Clasper with basodorsal
process rather straight,apex truncate in dorsal and obliquely truncate in ventral aspect,
lying on segment one; segment one slightly inflated basally, hirsute ventrolaterally, mem-
branous mesoventrally, curved and bluntly pointed mesad; segment two a small subapical
lobe directed mesad. Phallus lacking sclerotized parameres, but with a bilobed, apicodor-
sal, membranous appendage; apex slightly decurved, with a darkened internal phallo-
tremal sclerite, beyftnd which it is membranous. Female genitalia (Figs. 5-6): Eighth
sternum with lateral fourths strongly sclerotized, almost square in shape, concave, connec-
ted mesally by a lightly sclerotized band along anterior margin. Ninth tergum with a long
anterolateral apodeme. ventral margin strongly concave; dorsally with a distinct projec-
tion subapically. Membrane ventrolaterally between eighth and ninth segments bearing
convoluted sclerotized and semisclerotized bands. Vaginal sclerite in ventral aspect with

1 Received June 26, 1989. Accepted July 20. 1989.

Department of Entomology, National Museum of Natural History. Smithsonian Institu-
tion, Washington, DC 20560.

•'Department of Biological Sciences, Arkansas State University, State University, AR

72467-0599.

ENT. NEWS 101(2): 81-87, March & April, 1990

ENTOMOLOGICAL NEWS

an arcuate posterior bridge, with lateral angles projecting, centrally with a keyhole-like
opening.

Material: Holotype, male: U.S.A., Arkansas. Independence Co., unnamed stream
(Tl IN, R6W, Sec. 2 SW1/4), 5.6km N Pleasant Plains on US Hwy 167, 18 April 1987, Phoebe
A Harp. NMNH Type. Paratypes: Same data, 2 males, 7 females; same, hut 21 Mar 1987, 1
male. Johnson Co.. spring-run. 7mi W Oark on Hwy 215, 16 Apr 1987. D.E. Bowles, UV
light, 1 male; Sulphur Creek, 89mi N Hagarville on Hwy 123, 29 Mar 1986, C.D. Row-
botham, lab reared, 1 male. Logan Co., Mt. Magazine, Gutter Rock Creek, 6 May 1988,
Bowles & Mathis, UV light, 1 male, 2 females; same, but 14 May 1988, D.E. Bowles, UV light.
1 male; same, but Dripping Springs, Gutter Rock Creek, 20 Apr 1987, sweeping. 19 males, 14
females. Pope Co., Petite Jean State Park, SW slope, stuck to fire pink near ephemeral
stream, 16 Apr 1988, R. Leschen, 1 male. Oklahoma, LatimerCo., Robbers Cave State Park,
Rough Canyon spring run, 2 Apr 1988. D.E. Bowles. 12 males, 2 females. Paratypes
deposited in collections at Arkansas State University (Department of Biological Sciences),
University of Arkansas (Department of Entomology), and National Museum of Natural
History.

Figs. 1-6. Lepidostoma (Nosopus) ozarkense n.sp. 1 , male genitalia, lateral. 2, male ninth and
tenth terga, dorsal. 3. male phallus, lateral. 4, male clasper, ventral. 5, female vaginal
sclerite, ventral. 6, female genitalia, lateral.

Vol. 101, No. 2. March & April 1990 83

This species is a member of the Modestum Group (Weaver 1988:35)
and is closely related to L. Ontario Ross. These are the only two species in
which the males possess a basoventral process from the tenth tergum
and the only apparently significant and consistent difference between
them lies in the length and shape of this process. In ozarkense the process
is only as long as the tenth tergum and is pointed in both lateral and dor-
sal aspects. However, in Ontario this process is almost 1 .5 times as long as
the tenth tergum, and, although it may appear pointed or furcate in
lateral aspect depending on the angle, in dorsal aspect it is flattened,
curved, widened and shallowly furcate apicad (Ross 1941, Fig. 98; Weaver
1988, Fig. 72). The females are quite similar in all species of the group,
and specific differences are less certain. However, the lightly sclerotized,
convoluted bands ventrad of the ninth tergum and the projecting lateral
angle of the vaginal sclerite in ozarkense appear to be diagnostic.

DISCUSSION

This species brings to seven the number of species described in the
Modestum Group, all of which are limited to eastern North America.
Four of the species, lydia Ross, modestum (Banks), Ontario Ross, and pic-
tile (Banks), have rather similar distributions, running in general from
Newfoundland or Quebec south along the Appalachian Mountains into
the western Carolinas or northwestern Georgia (cf. Fig. 7, the distribu-
tion of Ontario). Three species appear to have very restricted ranges: com-
pressum Etnier & Way in Cumberland Co., Tennessee, weaveri Harris in
Tuscaloosa Co., Alabama, and ozarkense n.sp. in Independence. Johnson.
Logan, and Pope Cos., Arkansas and Latimer Co., Oklahoma. All of
these species occupy areas outside the broad range of Ontario. This pat-
tern is suggestive of either relictual distributions resulting from expand-
ing and contracting ranges related to earlier advances and retreats of ice
sheets, or random dispersal events from the main range of the species
group.

ECOLOGICAL - BIOLOGICAL DATA

The topotypical stream, an intermittently flowing second-order tri-
butary of Salado Creek, is located in the Boston Mountains of the Ozark
Plateau. The sampling site (Fig. 8) was a 100m stretch of the stream
which parallels highway US 167, approximately 0.8km south of the

ENTOMOLOGICAL NEWS

Salado Creek bridge (Harp 1988). The stream margins are heavily tores-
ted, primarily with loblolly pine (Pinus taeda L.), shortleaf pine (Finns
echinata Mill.), eastern redcedar (Juniperus virginiana L.), southern red
oak (Quercus falcata Michx.), and white oak (Quercus alba L.).

Selected physicochemical parameters were measured from 27 Dec
1986 through 18Jul 1987(Table 1). This unnamed intermittent stream is a
slightly acidic, soft-water stream. Alkalinity ranged between 19 — 88ppm
with the highest values recorded during low flow. The pH ranged from
6.6 — 7.0, averaging 6.7. The low alkalinity and pH values reflect the soil
type, Enders stony fine sand loam (Ferguson et al. 1982).

Flow volume measured 120 — 300,000cc/sec. Temperature ranges in-
cluded: water 5.0 — 27.5°C, and air, -1.5 — 28°C. The first adult Lepidos-
toma ozarkense, a male, was taken on 2 1 March. Flow peaked on this date;
water temperature was 15°C, and air temperature 24°C. The remaining
specimens were collected on 18 April. On this date the flow measured

Ontario

weaver!
• compressum
D ozarkense

Fig. 7. Recorded distributions of Lepidostoma (Nosopus) Ontario Ross, L. weaver/ Harris,/,.
compressum, Etnier & Way, and ozarkense n.sp.

Vol. 101, No. 2, March & April 1990

Fig. 8. Toporypic habit of Lepidostoma (Nosopus) ozarkense n.sp.

19,000cc/sec; water temperature was 14.5°C, and the air was 25UC. Thus
this trichopteran was present during high flow and when water tempera-
ture was above 14°C.

Aquatic samples, totalling fifteen, were taken twice monthly, each the
result of a two hour collecting period. A Turtox Indestructable Dip Net
was the primary collecting instrument, but smaller samples were taken
from a Tyler Standard Screen (mesh size 0.295mm) after disturbing the
bottom. Adults, which were taken incidentally, were not included in the
sample statistics. The aquatic stages of a minimum of 54 taxa were collec-
ted in the samples. Of these taxa, 46 were insectan: 12 beetles, 10 stone-
flies, 8 caddisflies, 6 flies, 5 mayflies, 4 water bugs, and 1 fishfly. The
eleven most common insects (numbers 10 and 1 1 had identical counts),
in order of abundance, were: Amphinemura delosa (Ricker), Prosimulium
spp., Pseudocloeon spp.,Ameletus spp., hoperla spp.,Agapetus illini Ross,
Neophylax concinnus McLachlan, Clioperla sp.. Tipula spp.. Stenonema
femoratum (Say), and Perlesta spp. A paper presenting the full results of
the survey is being prepared by the junior author.

ENTOMOLOGICAL NEWS

Table 1. Selected physicochemical parameters in an unnamed intermittent stream, Independence County,
Arkansas, 27 December 1986 - 18 July 1987.

— i OO

3 1-H

I-S

in
f-

CM
CM

to
o

in
o

CM tO

3 •'T

r*-

t^-

oo to
oo

c o

3 CM

— ^

00
CM

O
Oi

m
o

* to

co co

0
CM

m to
m

cS*'^-*

o
oo

CM tO

CO CM

m
<n

„

li 00

oo to

CXT
-s:

u —

CO CM

co r-

.0 -H

.fi

o r—
u.

c •*?•

CO CM

i-a

CO —

••8

tj t—
V CM

1=1

m
o

CM tO

0
CM

o
o

O
O

t— o

r--

ro to

O5 tO
CM

o
m

o
oo

f- tO
CM

o
m

'30

to
co to

to
m

Oi to

4-9

-o

^«

o>
rt

E
U

a/
-o

t—

-o

CO
09

oo
co

—1 CJ
<-t-i Ci>
C/3

C E

cu u

U
3

Current factors 0.7

- 0
2 O>

o w

E O
3 O
—i O

O i—

to to

Q.
O.

a
oc =

—i Q

-C

4-1

'co
CO

Vol. 101. No. 2. March & April 1990 87

Although 37 Lepidostoma larvae (fifteenth in order of abundance)
were collected at the topotypical site, they were not reared, and we feel it
wisest not to risk an erroneous aasociation at this time. Both Bowles and
Harp are planning work on the immature stages of this species.

ACKNOWLEDGMENTS

We wish to extend thanks to George L. Harp for assisting in the field and lab. for iden-
tifying the beetles and confirming many other identifications, to Barry K. Poulton for sug-
gesting the study site, to Bill P. Stark. Richard W. Baumann and Barry K. Poulton for
identifying the stoneflies. to Rhonda K. Boyer for identifying the Prosimulium. to Peggy A.
McDaniel. Betty G. Cochran. Geoffrey R. Harp. Mitchell K. Marks. D. Cecilia Reiman.
and Tim W. Steward for providing additional valuable field assistance, and to D.E. Bowles.
W.N. Mathis. P J. Spangler and two anonymous individuals for reviewing this manuscript

LITERATURE CITED

Ferguson, D.V., J.S. Lawrence & C.E. McFadden. 1982. Soil Survey of Independence
County. .Arkansas. U.S. Dept. Agr. & Soil Conserv. Sen. 148 pp. U.S. Govt. Printing
Office:' 1982 0-373-040.

Harp, P.A. 1988. Aquatic Macroinvertebrates of an Ozark Intermittent Stream. Indepen-
dence County. Arkansas. Unpublished Master's Thesis. Arkansas State University.
State University. Arkansas. 25 pp.

Ross, H.H. 1941. Descriptions and Records of North American Trichoptera. Trans. Amer.
Entomol. Soc. 67: 35-126.

Weaver, J.S., III. 1988. A Synopsis of the North American Lepidostomatidae (Trichop-
tera). Contrib. Amer. Entomol. Inst. 24: iv + 141.

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88 ENTOMOLOGICAL NEWS

A NEW SPECIES OF NEOTRICHIA (TRICHOP-

TERA: HYDROPTILIDAE) FROM COLORADO
WITH ADDITIONS AND CORRECTIONS TO THE
DISTRIBUTION AND RECORDS OF COLORADO

TRICHOPTERA1

D.E. Ruiter2

ABSTRACT: Neotrichia downsi, new species, is described from Jackson County, Colorado.
Illustrations of the male and female genitalia are provided. Additions and corrections to
the list of Colorado Trichoptera are also included. Fifteen species are added to the Colorado
list, bringing the total number of species reported from Colorado to 188.

As part of continuing studies on the taxonomy and distribution of
Rocky Mountain Trichoptera, a new hydroptilid caddisfly was dis-
covered in material collected by Dr. Wilbur G. Downs from the North
Platte River. Notes on new distributional records of other Colorado
Trichoptera and corrections to the list by Herrmann et al. (1986) are
also provided.

Neotrichia downsi, new species
Figure 1

Male: Body and appendages typical for subgenus, light brown, total body length 2 mm.
Bracteole of IX segment constricted throughout basal third. In dorsal view, membranous X
tergite evenly tapered, a pair of stout setae located laterally, approximately one-third from
apex. Inferior appendages heavily sclerotized; ventrally triangular, with a minute, mesal
tubercle near midpoint; rapidly tapered in lateral view, apex slightly upturned, bearing
several minute tubercles. Subgenital plate heavily sclerotized, massive; in lateral view
hooked slightly ventrad apically, extending 0.75 length of inferior appendage. Phallus with
long, wide basal portion, tapering evenly to constriction at spiral process, apical portion
slightly expanded at base with apex incised; spiral process encircles phallus approx-
imately 1.5 times.

Female: Body and appendages typical for subgenus, light brown, no apparent sexual
dimorphism, total body length 2 mm. Sclerotized portion of VIII sternite resembling an
inverted champagne glass. Bursa with apex long and slender. Scale-like setae at distal
margin of VII sternite smoothly rounded at apex.

Etymology: Named for the collector, Wilbur G. Downs, in recognition of his extensive
collection of insects from the type locality.

Holotype male: Colorado, Jackson County, Ginger Quill Ranch at the North Platte
River, altitude 2,370 meters (7,700 feet) above mean sea level, 7 July 1986, W.G. Downs,
black light trap collection. Deposited in the Illinois Natural History Survey collection.

Paratypes: Same as above, except 4 July 1987, 1 male, 1 female. The female is in poor
condition, only the abdomen is relatively intact. Deposited in the United States National
Museum collection.

'Received July 17, 1989. Accepted September 1 1, 1989.
21588 South Clermont, Denver, Colorado 80222.

ENT. NEWS 101(2): 88-92, March & April, 1990

Vol. 101, No. 2, March & April 1990

Fig. \.Neotrichia downs/. A. male, apex of abdomen, ventral; B. male, apex of abdomen,
dorsal; C. male, apex of abdomen, lateral; D. male, phallus, ventral; E. female. VIII sternite;
F. female, bursa, ventral; G. female, scale setae of VII sternite.

90 ENTOMOLOGICAL NEWS

This species belongs to the collata species group, closely related to N.
okopa Ross, N. sonora Ross and N. osmena Ross. This species appears
most similar to N. okopa Ross from which it can be readily separated by
the dorso-ventrally flattened inferior appendages, constricted basal third
of the spatulate lateral appendages, and the evenly tapered X tergite. The
new species can be distinguished from N. sonora by the absence of the
long membranous IX tergite, and lack of long, sclerotized lateral projec-
tions on the tenth tergite. Downsi differs from osmena by the shape of the
inferior appendages and lack of the sclerotized, forceps-like structure of
the X tergite. However, the phallus ofN. downsi is indistinguishable from
that of N. osmena. The female association is inferred from the syn-
chronous collection with the male and the distinct difference from N.
halia Denning, the only other Neotrichia species collected at the type
locality during nearly ten years of collecting.

Corrections and New Distributional Records

Lepidostoma grisea (Banks) originally included by Herrmann et al.
(1986) is removed from the Colorado distribution list. This record was
questioned by John Weaver (personal communication) and after a diligent
search, this specimen could not be located. Weaver (1988), also syn-
onymized several species of Lepidostoma occurring in Colorado (L. mon-
eka Denning 1968 = L. ormea Ross 1946; L. mira Denning 1954 = L.
cascadense (Milne) 1936; L. strophis Ross 1938 = L. cinereum Banks 1899;
L. veleda Denning 1948 = L. pluviale (Milne) 1936).

Since publication of the Colorado list (Herrmann et al., 1986) several
new records have been published and additional species collected through
the efforts of W.G. Downs, B.C. Kondratieff and this author. The county,
known Colorado emergence period, and notes where appropriate are
listed below.

Culoptila cantha Ross, Jackson County, 2 August.

Cheumatopsyche speciosa (Banks), Moffatt County, 11 July. This

appears to be the first record of this species west of the continental divide.

Previously reported from locations in the east with the most western

records being from Montana and Oklahoma (Gordon, 1974; Nimmo,

1987).

Hydropsyche alhedra Ross, Grand and Jackson Counties, 23 July -

20 August.

Hydropsyche bidens Ross, Weld County, 3 August. Previously reported

from the eastern portion of the Great Plains (Hamilton and Schuster,

Vol. 101. No. 2. March & April 1990 91

1979; Nimmo. 1987). Its collection in eastern Colorado indicates it has
successfully crossed the prairie to the upper Platte River basin.
Hydropsyche slossonae Banks, Grand County, (Schefter and Wiggins.
1986). Jackson County. 15 July - 16 August.

Hydroptila angusta Ross. Moffatt County. 1 1 July. This appears to be
the first record for this species from west of the continental divide. Pre-
viously reported from the east with western records in New Mexico.
Texas and Kansas (Ross. 1944; Schuster and Hamilton. 1978).
Mayatrichia ayama Mosely. Mesa County'. 30 July.
Ceraclea arielles (Denning). Jackson County. 15 July - 9 August.
Ceraclea resurgens (Walker). Grand and Moffat Counties. (Herrmann.
1988) 25 July - 1 August.

Limnephilus apache Flint. Larimer and Pueblo Counties, 27 July - 2
August. This species has not been reported since Flint's 1965 description
from collections in Arizona and New Mexico. The collections in Larimer
and Pueblo Counties represent significant northern distributional
extensions.

Limnephilus selatus Denning, Grand County, 3 August.
Pycnopsyche guttifer (Walker), Jackson County, 31 July - 20 Sep-
tember.

Polycentropus aureolus (Banks). Jackson County, 1-9 August. Prev-
iously reported from the northern states and Canada (Nimmo. 1986)
with the nearest records from Idaho and North Dakota.
Agrypnia colorata Hagen. Larimer County, 28 June.A colomla is also a
seldom reported species. Most records are from Canada (Betten. 1934;
Ross and Spencer, 1952) with the nearest previous records from Mon-
tana (Roemhild, 1982).

Gumaga griseola (McLachlan). Yuma County, 25 June. (Kondratieff
and Ward. 1987). I have examined this specimen and while the specimen
was about half the typical size of specimens from California, the geni-
talia were nearly consistent with California specimens. Hamilton and
Schuster (1980) also noted structural differences in Kansas populations
of G. griseola. While it is clear the collection of Gumaga is a new record
(including a new familial record) for the state, further collections will be
necessary to determine the variation within the species.

Given the fairly detailed lists available for several of the adjacent
steles (Kansas: Schuster and Hamilton, 1978; Hamilton and Schuster.
1979, Hamilton and Schuster. 1980; Utah: Baumann and Unzicker.
1981; Wyoming: Ruiter and Lavigne. 1985). it appears many of the
species which may be on the fringe of their distributional areas are also
extremely localized in their distribution, having not been reported from
neighboring states.

92 ENTOMOLOGICAL NEWS

With the removal of four species, the description of one species, and
the addition of the 1 5 species just listed, the total number of Trichoptera
species reported from Colorado is 188.

ACKNOWLEDGMENTS

The author wishes to thank J. D. Unzicker. Boris Kondratieff and Steve Harris for pro-
viding critical comments on early versions this manuscript. J.D. Unzicker also provided
material of N. okopa Ross for comparison. O.S. Flint, Jr. identified the L. apache Flint
specimens. Wil Downs must he sincerely thanked for providing thousands of specimens
from the Ginger Quill locality.

LITERATURE CITED

Baumann, R.W. and J.D. Unzicker. 1981. Preliminary checklist of Utah caddisflies

(Trichoptera). Encyclia 58:25-29.
Flint, O.S. Jr. 1965. New species of Trichoptera from the United States. Proc. Ent. Soc.

Wash. 67(3): 168-176.
Gordon, A.E. 1974. A synopsis and phylogenetic outline of the nearctic members of

Cheumatopsyche. Proc. Acad. Nat. Sci. Phila. 126:1 17-160.
Hamilton, S.W. and G.A. Schuster. 1979. Records of Trichoptera from Kansas, II: The

families Glossosomatidae, Helicopsychidae, Hydropsychidae and Rhyacophilidae.

Tech. Publ. State Biol. Surv. Kansas 8:15-22.
Hamilton, S.W. and G.A. Schuster. 1980. Records of Trichoptera from Kansas. Ill: The

families Limnephilidae, Phryganeidae, Polycentropodidae, and Sericostomatidae.

Tech. Publ. State Biol. Surv. Kansas 9:20-29.
Herrmann, SJ. 1988. New record and range extension forCeraclea resurgens (Trichoptera:

Leptoceridae) from Colorado, with ecological notes. Ent. News 99(5):253-259.
Herrmann, S.J., D.E. Ruiter and J.D. Unzicker. 1986. Distribution and records of

Colorado Trichoptera. Southwestern Nat. 31:421-457.
Kondratieff, B.C. and J.V. Ward. 1987. Taeniopteryx burksi (Plecoptera: Taeniop-

terygidae) in Colorado, with notes on aquatic insects of plains streams. Ent. News 98: 13-

16.
Nimmo, A.P. 1986. The adult Polycentropodidae of Canada and adjacent United States.

Quaes. Ent. 22:143-252.
Nimmo, A.P. 1987. The adult Arctopsychidae and Hydropsychidae (Trichoptera) of

Canada and adjacent United States. Quaes. Ent. 23:1-189.
Roemhild, G. 1982. The Trichoptera of Montana with distributional and ecological

notes. Northwest Sci. 56:8-13.
Ross, H.H. 1944. The caddis flies, or Trichoptera. of Illinois. Bull. 111. Nat. Hist. Surv. 23: 1-

326.

Ross, H.H. and G.J. Spencer. 1952. A preliminary list of Trichoptera of British Colum-
bia. Proc. Ent. Soc. Brit. Col. 48:43-51.
Ruiter, D.E. and R.J. Lavigne. 1985. Distribution of Wyoming Trichoptera. University of

Wyoming. Agricultural Experiment Station Publication Number SM47. 102 pp.
Schefter, P.W. and G.B. Wiggins. 1986. A systematic study of the Nearctic larvae of the

Hydropsyche morosa group (Trichoptera: Hydropsychidae). Life Sciences Mis-
cellaneous Publication. Royal Ontario Museum. 94 pp.
Schuster, G.A. and S.W. Hamilton. 1978. Records of the trichopteran families Hydro-

ptilidae, Philopotamidae, and Psychomyiidae from Kansas. Tech. Publ. State Biol.

Surv. Kansas 6:36-47.
Weaver, o.S. III. 1988. A synopsis of the North American Lepidostomatidae (Trichoptera).

Cont. N. A. Ent. Inst. 24(2): 1-141.

Vol. 101, No. 2, March & April 1990 93

A NEW SPECIES OF ZONANA (HOMOPTERA:
CICADELLIDAE) FROM VENEZUELA1 2

Paul H. Freytag-'

ABSTRACT: A new species ofZonana is described from Venezuela. This fairly small genus
previously was known from only two species from Central America, so the known range of
the genus is extended to the upper part of South America.

New species ofZonana is described at this time from Venezuela. This
species is the third species placed in this genus; the other two were des-
cribed from Central America (DeLong and Freytag 1963).

Zonana bilobata sp. n.

(Figures 1-5)

Length of males 6.4-6.7 mm.; female unknown.

Resembling flamma in general size, shape and color, but differs in shape of aedeagal
shaft and processes.

Color: Similar to flamma, except with three additional dark spots along commissure of
each forewing. Dark punctures present on pronotum and along basal costal area of
forewing.

Male genitalia: Pygofer truncate and bilobed at apex, with small dorsal subapical
lobe. Genital plate short with truncate apex. Style short, sharply pointed at apex, with an
enlarged rounded caudal lobe. Aedeagus short, subapical ventral cleft forming lateral
hook-like lobes with several small spines along ventral margin; apex rounded, with two
darker sclerotized spine-line areas.

Holotype male: VENEZUELA. Aragua. El Limon, 450 m., 26- VI- 1974, En Luz de Mer-
curio, F. Fernandez Y. & F. Fernandez H., in the University of Central Venezuela collec-
tion. Paratypes: One male, VENEZUELA. Aragua, El Limon. 450 m.; 1-1-1981. F. Fer-
nandez Y. Coll.. in the University of Kentucky collection; one male. VENEZUELA.
Bolivar. Rio Surukum. Carretera, Sta. Elena Icabara. 850 m.. 19-31- 1-19895. F. Fernandez
Y., Anibal Chacon and Jurg Demarmels Colls.; one male. VENEZUELA. Miranda. Valle
Rio Cuira, 280m.. S. of Panaquire.approx. 10° 12' N; 66° 17' 0, 18-21 -IX- 1979. F. Fernandez
Y. & J. A. Clavijo Colls., both in the University of Central Venezuela Collection.

Note: This species can be separated from the closest species,/7awwa ,
on the basis of the distinct aedeagus and the additional punctures on the
pronotum and costal area of the forewing.

I wish to thank the late Dr. Fernandez Yepes who was instrumental in
collecting this most interesting species and making the specimens avail-
able for study.

'Received July 13, 1989. Accepted August 3, 1989.

2This paper is published with the approval of the Director of the Kentucky Agricultural

Experiment Station as journal article no. 89-7-147.
•^Department of Entomology, University of Kentucky, Lexington, KY 40546-0091.

ENT. NEWS 101(2): 93-94. March & April. 1990

ENTOMOLOGICAL NEWS

BILOBATA

0.5 mm

Figures 1-5, Zonana bilobata sp. n., male genitalia. Fig. 1, caudal view of aedeagus; fig. 2,
lateral view of aedeagus; fig. 3, lateral view of style; fig. 4, ventral view of genital plate; fig. 5,
lateral view of genital capsule and anal tube. All drawn to the same scale.

LITERATURE CITED

DeLong, D. M. and P. H. Freytag. 1963. Studies of the Gyponinae 2. A New Genus
Zonana (Homoptera: Cicadellidae). Ohio J. Sci. 63(6): 262-265.

Vol. 101, No. 2, March & April 1990 95

EURYPAUROPUS SPINOSUS (ARTHROPODA:
PAUROPODA: EURYPAUROPODIDAE) FROM

ARKANSAS AND A KEY TO THE NORTH
AMERICAN EURYPAUROPUS SPECIES1'2'3

Robert T. Allen4

ABSTRACT: Eurypauropus spinosus is reported for the first time in Arkansas. A key is given
to the three North American species of Eurypauropus.

In the closing section of his paper on the Pauropoda of the United
States, Scheller (1985) notes "that there are large lacunae in the knowl-
edge of the ranges" of most pauropod species. This paper serves to fill in
one of those lacunae for Eurypauropus spinosus Ryder, and to provide a
key, based on information from the literature whereby other workers can
identify Eurypauropus specimens.

The family Eurypauropodidae is easily recognized by the large,
sclerotized tergal plates on each body segment. These plates conceal the
head, terminal segments, and the legs when viewed from above. Brues, et
al. ( \ 954) present a key that will allow workers to place Pauropoda in the
correct families. As far as known, Eurypauropus is the only genus in the
family Eurypauropodidae occuring in North America.

In the United States four species of Eurypauropus have been des-
cribed, one of which has been synonymized. Eurypauropus spinosis ap-
pears to be the most common species and has been reported from eleven
states, Fig. 4. Recent collections, listed here, add two new localities from
Arkansas to the list of areas from which E. spinosus is known. The two
Arkansas localities are as follows: Pulaski County, Pinnacle Mountain
State Park, east summit trail, 25 January 1988, Robert T. Allen collector;
Yell County, entrance to Mount Nebo State Park, 15 April 1988. C. E.
Carlton collector.

Eurypauropus californicus MacSwain & Longham was described
from California, but was reduced to a synonym of E. spinosus by Remy
(1956). In the same paper Remy described a new species, E. uncigerhom
Kentucky. In the most recent and complete paper on the Pauropoda of
the United States, Scheller (1985) described E. washingtonensis from
Olympic National Park in Washington State. The following key, based
on characters extracted from the papers by Remy (1956) and Scheller

^Received October 25, 1988. Accepted July 27. 1989.

^Published with the approval. Director, Arkansas Agricultural Experiment Station,

University of Arkansas, Fayetteville. AR 72701.

This research supported, in part, by a grant from Arkansas Nature Conservancy.
^Professor, Entomology Department, University of Arkansas. Fayetteville. AR 72701

ENT. NEWS 101(2): 95-97, March & April, 1990

ENTOMOLOGICAL NEWS

Figures 1-3. The terminal segments of the three species of Eurypauropus from the United
States. Fig. \,E.spinosus( redrawn from Remy, 1956); Fig. 2,E. unciger( redrawn from Remy,
1956); Fig. 3. E. washingtonensis (redrawn from Scheller. 1985).

Vol. 101, No. 2, March & April 1990

(1985) will enable workers to identify Eurypauropus specimens from
North America.

Key to North American Species of Eurypauropus

1. Style (st) of the pygidial tergum elongate (Figs. 1 & 3) 2

Style of the pygidial tergum short, oblong-oval (Fig. 2) unciger Remy

2. Style elongate, straight, not bent (Fig. 1) spinosus Ryder

Style elongate, elbowed with a distinct bend (Fig. 3) woshingtonensis Scheller.

Figure 4. Map showing the distribution of Eurypauropus species in North America. The
crosshatching for E. spinosus represents one or more records from a state, not a general
distribution.

LITERATURE CITED

Brues, C. T., A. L. Melander, & F. M. Carpenter. 1954. Classification of insects. Cam-
bridge. Mass. U. S. A.

Remy, P. A. 1956. Pauropodes des Etats-Unis deAmerique. Mem. Soc. natn. Sci. nat.
math. Cherbourg 47:1-48.

Scheller, U. 1985. Taxonomic and distributional notes on pauropods from the United
States (Myriapoda. Pauropoda: Pauropodidae, Eurypauropodidae). Ent. scand.
16:237-257.

98 ENTOMOLOGICAL NEWS

SCIENTIFIC NOTE

DYSCINETUS MORATOR (COLEOPTERA: SCARABAEIDAE)

FEEDING ON ROOTS OF AZALEAS

(RHODODENDRON SPP.)1' 2

C. L. Staines, Jr.3

Dyscinetus morator (Fab.) (Coleoptera: Scarabaeidae), a common scarab distributed
from New York south to Florida and west to Texas and Mexico (Gordon & Anderson. 1981),
is widely distributed in Maryland (Staines, 1986). The adults are attracted to lights and
remain in the area of the lights during the day by hiding under available debris (Woodruff,
1970). Dyscinetus morator has been reported to attack rice, Oryza saliva L. (Anonymous,
1953); pangola grass pastures, Digitaria decumbens Stent. (Anonymous, 1956); caladium
bulbs, Caladium x hortulanum, cranberry, Vaccinium macrocarpon Ait. (Woodruff, 1970);
corn, Zea mays L. (Anonymous, 1980); carrot, Daucus carota L., radish, Raphanus sativus L.,
lettuce, Lactuca saliva L. (Foster et al.. \986); and waterhyacmth.Eichhorniacrassipes (Marl.)
Solms (Buckingham & Bennett, 1989). This species is associated with wet soils and marsh
areas (Anonymous, 1980; Buckingham & Bennett, 1989).

On 2 August 1982, during a routine inspection of a commercial nursery in Baltimore
County. Maryland there were discovered larvae, pupae, and teneral adults of an unknown
scarab around the root zone of container-grown azaleas (Rhododendron spp.). All life
stages were abundant and root feeding was obvious on the azaleas. E. J. Ford, USDA,
APHIS, PPQ, Baltimore (retired), identified the adults as Dyscinetus morator. the rice
beetle.

The azalea growing area of this nursery was bounded on three sides by salt marsh. The
azaleas were grown in containers and were maintained under damp conditions, thus pro-
viding a suitable habitat for the beetle. The production area was not lighted, so the infesta-
tion did not result from light-attracted adults. Adult beetles may have been attracted by the
moist, high organic matter content of the potting mixture used by the nursery.

This is the first report of Dyscinetus morator colonizing a woody ornamental. Subse-
quent surveys in this nursery have failed to yield further populations of this beetle in
azaleas, though the beetle is common outside of the growing area.

LITERATURE CITED

Anonymous. 1953. USDA Coop. Econ. Insect Report. 3:725.

Anonymous. 1956. USDA Coop. Econ. Insect Report. 6:1079.

Anonymous. 1980. USDA Coop. Plant Pest Report 5:66.

Buckingham, G. R.&C. A. Bennett. 1989. Dyscinetus morator ( Fab.) (Coleoptera: Scara-
baeidae) adults attack waterhyacinth, Eichhornia crassipes (Pontederiaceae). Coleopts.
Bull. 43:27-33.

Foster, R. E.etal. 1986. Dyscinetus morator (Coleoptera: Scarabaeidae) as a pest of carrots
and radishes in Florida. Florida Entomol. 69:431-432.

Gordon, R. D. & D. M. Anderson. 1981. The species of Scarabaeidae (Coleoptera)
associated with sugarcane in south Florida. Florida Entomol. 64:1 19-138.

Staines, C. L. 1986. An annotated checklist of the Scarabaeoidea (Coleoptera) of
Maryland. Maryland Entomol. 2:79-89.

Woodruff, R. E. 1970. The "rice beetle", Dyscinetus morator (Coleoptera: Scarabaeidae).
Florida Dept. Agric. Entomol. Cir. 103, 2pp.

^Received May 20. 1989. Accepted July 29, 1989

^Maryland Department of Agriculture Contribution Number CN 58-89
•^Maryland Department of Agriculture, Office of Plant Industries and Resource Conserva-
tion, Plant Protection Section, 50 Harry S Truman Parkway, Annapolis, Maryland
21401.

ENT. NEWS 101(2): 98, March & April, 1990

Vol. 101, No. 2, March & April 1990 99

SEASONAL FLIGHT ACTIVITY OF MALE
VELVET ANTS (HYMENOPTERA: MUTILLIDAE)

IN SOUTH FLORIDA1

Mark Deyrup , Donald Manley^

ABSTRACT: Seasonal male flight data from a four-year study are presented for 19 species
of mutillids belonging to the genera Ephuta, Photomorphus, Pseudomethoca. Sphaerop-
thalma, and Timulla. A total of 2,391 specimens was collected. At the study site in southern
Florida, most species flew during at least 8 months of the year. Two species flew only late in
the year. All species showed marked fluctuations in abundance from year to year.

Velvet ants (Hymenoptera: Mutillidae) are solitary parasitoid wasps,
most of them having as hosts pupae or mature larvae of bees and wasps
(Mickel, 1928). Male flight activity is particularly important for gene flow
in this group, as females are always apterous, and aerial dispersal is by
males. Males of a few species pick up and transport females, and might
cross barriers that would baffle an earth-bound female (Evans, 1969). We
have collected in Malaise traps copulating pairs of Timulla floridensis,
Ephuta floridana, E.pauxilla, and an associated pair of E. slossonae. Male
flight activity also offers the best methods for estimating relative num-
bers of adults of a species, as the males seem rather vulnerable to Townes
traps (modified Malaise traps). Females can be sampled with pitfall
traps, but we have found that the numbers of specimens collected in such
traps is relatively small, even when there are many traps and their
efficiency is enhanced with long metal barriers (drift fences) that direct
the mutillids into the traps. Nothing has been published on the seasonal
flight patterns of male mutillids. The purpose of this paper is to con-
tribute basic information on this little-known group of insects, and to
provide a data base that could be used in comparative studies of sea-
sonality elsewhere.

MATERIALS AND METHODS

The study site is on the Archbold Biological Station (Highlands
County), located at the southern end of the Lake Wales Ridge in south-
central Florida. The site is in a transitional zone between warm and sub-
tropical zones. Winters are mild and dry, with temperatures during some
years falling below 0°C for a few hours. Sheltered microhabitats are

^Received June 8, 1989. Accepted July 10, 1989

-Archbold Biological Station. P.O. Box 2057, Lake Placid, FL 33852
•'Department of Entomology, Clemson University. Pee Dee Resi
tion Center, Route 1, Box 531, Florence. SC 29501

ENT. NEWS 101(2): 99-108, March & April, 1990

100 ENTOMOLOGICAL NEWS

frost-free. Summers are warm and humid, with daytime temperatures
over 25°C. Mutillid species diversity is high, with over 30 species known.
The vegetation of the study site is a thin canopy of sand pine (Pinusclausa
Chapman), with a thick 1.5 - 3.5m understory of scrub oaks (Quercus
spp.), staggerbush (Lyonia spp.), silk bay (Persea humilis Nash), scrub
pawpaw (Asimina obovata Nash), and scrub hickory (Carya floridana
Sargent). The paths through this thick brush appeared to act as flight cor-
rdiors for mutillids.

Mutillids were collected in 2 small Townes traps (modified Malaise
traps) that were set up across 2 east-west paths. The traps were kept in
place and continuously monitored from May 1983 through December
1986. Each trap was annually replaced with an identical trap to forestall
the effects of wear. Specimens were collected 3 times a week. The mutil-
lids were identified by the authors. Specimens of all species are in the
collection of the Archbold Biological Station and in the collection of
Donald Manley. Synoptic collections of most species have also been
deposited in the Florida Collection of Arthropods (Gainesville) and the
collection of Denis Brothers (University of Natal, Pietermaritzburg,
South Africa).

RESULTS

Over the 4-year period, the traps collected 2,391 specimens belonging
to the genera Ephuta, Pseudomethoca, Timulla, Photomorphus, and
Sphaeropthalma. Many specimens of Dasymutilla were also collected;
these are not discussed here because of problems with the identification
of some species.

Ephuta floridana Schuster. 50 of 51 specimens taken May-December (Figure 1).
Annual numbers from 6 (1984) to 27 (1986).

E. stenognatha Schuster. 94 of 100 specimens taken May-November (Figure 2).
Annual numbers from 4 (1986) to 64 (1983).

E. slossonae (Fox). 132 of 134 specimens taken April-November (Figure 3). Annual
numbers from 20 (1984) to 46 (1986).

E. margueritae Schuster. All 118 specimens collected April-December (Figure 4).
Annual numbers from 8 (1984) to 44 (1986).

E. pauxilla Bradley. 40 of 42 specimens collected April-November (Figure 5). Annual
numbers from 3 (1093) to 15 (1985. 1986).

E.spinifera Schuster. No specimens collected 1983-1986. 8 specimens collected in July-
August 1987, in a temporary trapping site in mature sand pine scrub.

Photomorphus paulus (Bradley). 885 of 888 specimens taken April-November
(Figure 6). Annual numbers from 85 (1985) to 354 (1984).

P. alogus Viereck. All 1 8 specimens collected June-December. Annual numbers from 1
(1986) to 9 (1983).

P. archboldi Manley and Deyrup. All 12 specimens collected May-October. Annual
numbers from 1 (1986) to 5 (1984).

Vol. 101. No. 2. March & April 1990 101

Pseudomethoca oculata (Banks). All 129 specimens taken September-December
(Figure 7). Annual numbers from 13 (1985) to 59 (1983).

P. sanbomii (Blake). 69 of" 71 specimens collected May-November (Figure 8). Annual
numbers from 3 (1985) to 50 (1983).

P. simillima (Smith). 63 of 67 specimens collected October-November (Figure9). Annual
numbers from 12 (1983) to 32 (1986).

P.torrida Krombein. 41 8 of 425 specimens collected May-November (Figure 10). Annual
numbers from 58 (1985) to 158 (1983).

P. vanduzei Bradley. All 1 8 specimens collected April-October. Annual numbers from
2 (1984) to 6 (1985).

Sphaeropthalma pensylvanica (Lepeletier). All 17 specimens collected April-De-
cember. Annual numbers from 3 (1985, 1986) to 6 (1983).

Timulla dubitata (Smith). 25 of 26 specimens collected May-September (Figure 1 1 ).
Annual numbers from 1 (1985, 1986) to 12 (1983, 1984).

T. floridensis (Blake). 262 of 269 specimens collected March-December (Figure 12).
Annual numbers from 46 (1984) to 1 18 (1983).

T. ornatipennis (Bradley). 2 specimens. July 1983.

T. vagans (Fabricius). 1 specimen. May 1983.

DISCUSSION

At the Archbold Biological Station, males of most mutillid species
are active over a long period. Eight of the 12 species represented by at
least 25 specimens were active during 8 or more months of the year. The
prolonged flight season is likely to be related to the mild climate of the
study site, though complementary data from farther north are needed to
support this conclusion. As can be seen from the graphs (Figures 1-12),
most species are slow to begin flight activity in the spring. Flight activity
is seasonally less symmetrical than monthly average temperatures at the
station. Two species, P. oculata and P. simillima, have flight seasons res-
tricted to late in the year (September-December). There are no corre-
sponding spring-flying species. One species, P. paulus, shows a consis-
tently bimodal pattern reflecting a reduction in numbers in August
and September.

When species are active over a long period, a record of flight activity
does not provide good estimates of longevity. P. oculata and P. simillima,
with their short flight seasons, pro vide more useful information. Assum-
ing that large reductions from one month to the next reflect natural mor-
tality rather than a drastic effect of the traps on local populations, many
males of these species must live a month or less. The short life spans of
these species may not be typical if cooler temperatures in December and
late November are killing males. There is a case of an individual male
Dasymutilla foxi (Cockerell) that lived about 6 months in captivity (J.
Schmidt, personal communication).

The flight activity of males is presumably related to emergence of
females. Females apparently mate only once, immediately after emer-

102 ENTOMOLOGICAL NEWS

gence (Brothers, 1972; J. Schmidt, personal communication). Male flight
activity need not closely reflect total seasonal activity of females, as at
least some female mutillids appear to be long-lived insects (Schmidt,
1978). Female P. oculata and P. simillima, for example, are active in spring
and early summer, when there are no records of males. The fall and win-
ter emergence of these species may actually indicate a heavy dependence
on early spring hosts, rather than a dependence on fall or winter
hosts.

All species showed marked variation in annual abundance, with
maximum annual collections at least twice minimum collections. The
seven most frequently collected species showed no clear evidence that
their numbers were similarly and simultaneously affected by common
factors such as rainfall or temperature. The fluctuations seen in these
species may be unintelligible until we have identified hosts and know
something of the population dynamics of these hosts.

SUMMARY

At our study site in southern Florida, most species of mutillids fly
through much of the year. With the exception of 2 species of Pseudo-
methoca, there is no evidence of phenological niche partitioning or
dependence on hosts that are only available for a short season. All abun-
dant species showed large population fluctuations during the 4-year
study period. We hope that our information will encourage publication
of similar data sets from other areas. We would like to think thatour work
is the first of a series of studies of geographic phenological variation in
mutillids, that can be interpreted in terms of evolutionary pressures exer-
ted by different climatic regimes.

ACKNOWLEDGMENTS

We thank Nancy Deyrup for tabulating the data and creating and preparing the graphs.
Justin Schmidt (Carl Hayden Bee Research Center, Tucson, AZ), and Fred Lohrer (Arch-
bold Biological Station), and two anonymous specialists reviewed the manuscript and pro-
vided useful comments. The manuscript was typed by Patricia Bache.

LITERATURE CITED

Brothers, DJ. 1972. Biology and immature stages of Pseudomethoca F. frigida, with notes
on other species (Hymenoptera: Mutillidae). Univ. Kansas Sci. Bull. 50: 1-38.

Evans, H.E. 1969. Phoretic copulation in Hymenoptera. Entomol. News 80:113-124.

Mickel, C.E. 1928. Biological and taxonomic investigations on the mutillid wasps. Bull.
U.S. Nat. Mus. 143: ix + 351 pp.

Schmidt, J.0. 1978. Dasvmutilla occidentalis: a long-lived aposematic wasp (Hymenoptera:
Mutillidae). Entomol. News 89: 135-136.

Vol. 101, No. 2, March & April 1990

103

15 -i

10 -

5 -

cc.

0
30

25 -
20 -
15
10

5
0

EPHUTA FLORIDANA

I I I T

EPHUTA MARGUERITAE

1 2 3 4 5 6 7 8 9 10 11 12

1983

1984

1985

1986

Figures 1 and 2. Seasonal (light, Ephutafloridana and E. margueritai-

104

ENTOMOLOGICAL NEWS

9 -

7 -

5 -

3 -

OC
HI

EPHUTA PAUXILLA

EPHUTA SLOSSONAE

2 3 4 5 6 7 8 9 10 11 12
MONTH

1984 1985 1986

Y///1
1983

Figures 3 and 4. Seasonal flight, Ephuta pauxilla and E. xlossonae

Vol. 101. No. 2. March & April 1990

105

24 -

20 -

16 -

12 -

8 -

4 H

LLJ
CD

EPHUTA

STENOGNATHA

x/l L I I . I r. . I I . . - I iv,

I i I n«^.^rf i— _^_ ^

200

150 -

100

50 -

PHOTOMORPHUS

PAULUS

z xx/

1 2 3
1983

4 5 6 7 8 9 10 11 12
MONTH

1984 1985 1986

Figures 5 and 6. Seasonal flight, Ephuta stenognatha and Photomorphus • paulus

106

ENTOMOLOGICAL NEWS

QC
LU
CO

70
60

50
40
30
20

12 -
8 -
4 -

PSEUDOMETHOCA OCULATA

I I I i r

PSEUDOMETHOCA

SANBORNII

1 2 3 4 5 6 7 8 9 10 11 12

MONTH

1983 1984 1985 1986

Figures 7 and 8. Seasonal flight, Pseudomethoca oculata and P. sanbomii

Vol. 101, No. 2, March & April 1990

107

60 -
50 -
40 -
30 -

20 -
10 -

DC
LU
m

PSEUDOMETHOCA SIMILLIMA

I I

160 -i

120 -

80 -

40 -

0 -

PSEUDOMETHOCA

TORRIDA

1 2 3 4 5 6 7 8 9 10 11 12

MONTH

1983

1984

1985

1986

Figures 9 and l(). Seasonal flight, Pseudomethoca simillima and P. torrnla

108

ENTOMOLOGICAL NEWS

DC
111

7 -

TIMULLA DUBITATA

90 i

50 -

30 -

10 -

TIMULLA FLORIDENSIS

1 2 3 4 5 6 7 8 9 10 11 12

MONTH

1983 1984 1985 1986

Figures 11 and 12. Seasonal flight, Timullu dubitata and T.floridensis

Vol. 101, No. 2, March & April 1990 109

TAXONOMIC DISTRIBUTION AND PHYLO-

GENETIC SIGNIFICANCE OF THE MALE

FORELEG HAIRPENCIL IN THE TORTRICINAE

(LEPIDOPTERA: TORTRICIDAE)'

John W. Brown^

ABSTRACT: I report for the first time male foreleg hairpencils in the tortricine tribes
Euliini, Schoenotenini, and Atteriini. The structures are identical in Euliini and Schoeno-
tenini, and their shared possession is considered a putative synapomorphy for these two
tribes. Although similar in structure and point of origin, the hairpencil in Atteriini lies
along the anterior longitudinal axis of the femur rather than along the outer base of the
coxa as in Euliini and Schoenotenini. The atteriine hairpencil may represent an indepen-
dently derived structure, or, more likely, a modification of that found in the other two
tribes.

Male secondary structures (e.g., costal fold, corema, hairpencil) are
common and diverse among exoporian and ditrysian Lepidoptera
(Varley 1962; Birch 1972, 1985). Although these structures may be strik-
ingly unique and almost certainly homologous among taxa sharing
them, their value in elucidating phylogenetic relationships is dimin-
ished by the fact that they are evolutionarily more labile than many other
adult morphological features. Shared possession of uniquely derived
male secondary structures provides compelling evidence of common
ancestry (Birth 1972; Varley 1962), but their absence may be meaningless
in a phylogenetic context since the correct position in the transformation
series (i.e., primitively absent - present - secondarily lost) may be impos-
sible to determine.

Few characters of the legs of tortricids have been studied on a com-
parative basis. Examples include the work of Falkovitch (1962) on male
secondary structures in Olethreutinae, and that of Yasuda (1972) on tar-
sal setae of Japanese Tortricinae. Horak ( 1984) concluded that legs pro-
vide fewcharacters useful in resolving phylogenetic relationships within
Tortricinae. However, my recent discovery of male foreleg hairpencils
among the tortricid tribes Euliini, Schoenotenini, and Atteriini may be
useful in demonstrating the monophyly of these taxa and in elucidating
relationships among their members.

Hairpencil in Euliini and Schoenotenini
Structure. The foreleg hairpencil in Euliini and Schoenotenini con-

1 Received May 20, 1989. Accepted July 3, 1989.

Department of Entomology, NHB-127, Smithsonian Institution, Washington. D.C.,
20560.

ENT. NEWS 101(2): 109-1 16. March & April. 1990

1 10 ENTOMOLOGICAL NEWS

sists of a fascicle of 15-25 slender, pale yellow, hairlike scales, that arise
together from the proximal end of the femur and extend along the outer
surface of the coxa to near its base (Figs. 1-3). Scales on the trochanter
and the distal portion of the coxa form a broad groove that partially con-
ceals the basal portion of the hairpencil. On the thorax near the base of
the coxa, there is an invaginated region and associated cuticular flap
which receives the distal ends of the hairpencil (Fig. 4). The function of
the hairpencil is unknown; presumably it plays a role in dissemination
of short-range courtship pheromones. Baker and Carde (1979a, 1979b)
and Baker, Nishida, and Roelofs (1981) have shown that male abdom-
inal androconia ofGrapholita (Olethreutinae), consisting of an eversible
membranous sac, are used to release a close-range female attractant.

Taxonomic distribution. The euliine/schoenotenine hairpencil has
a sporadic distribution throughout the two tribes (Tables 1 and 2); i.e. the
structure is present in 48% of euliine and 43% of schoenotenine genera
examined. In addition to those genera listed in Table 1, several under-
scribed euliine genera and many unplaced species of ""Eulia" Hiibner
possess the hairpencil. In some genera all species have the hairpencil; in
others it is absent and presumably secondarily lost in one or more
species. Its presence in the presumably most primitive as well as the more
derived genera of Euliini (Brown and Powell, in prep.) suggests that it
represents a synapomorphy uniting the Euliini and Schoenotenini rather
than a character derived numerous times within the two tribes.

The Euliini, as recently redefined by Powell (1986), previously was
considered a para- or polyphyletic assemblage of primarily Neotropical
genera, characterized by symplesiomorphies of male and female geni-
talia (Horak and Brown 1990). Possession of the hairpencil demon-
strates the common ancestry of genera that share this unique structure;
however, absence of the hairpencil among several taxa does not neces-
sarily exclude them from the tribe since it may have been lost secon-
darily. The structure is absent in more than half of the genera included
by Powell (1986) in the Euliini; on the basis of this and other characters, a
few of these genera likely are assigned incorrectly to this tribe (e.g.,
Ecnomiomorpha Obraztsov, Deltobathra Meyrick).

Hairpencil in Atteriini

Structure. In Atteriini, the hairpencil typically consists of a dense
brush of > 50 fine, flattened, setalike scales, arising along a narrow patch
on the anterior edge of the femur immediately distad of the junction with
the trochanter. The scales are shorter in relation to coxa length, more
flattened, and greater in number than those in Euliini and Schoeno-
tenini. When not displayed, the hairpencil lies along the longitudinal

Vol. 101, No. 2, March & April 1990

• •• hairpencil

TROCHANTER

Fig. 1. Diagram of male foreleg illustrating position of hairpencil.

112

ENTOMOLOGICAL NEWS

Figs. 2-3. Scanning electron micrograph of hairpencil of Eulia ministrana L. (Euliini).

Fig. 4. Thoracic receptacle of hairpencil in Eulia ministrana.

Fig. 5. Scanning electron micrograph of hairpencil in Tinacrucis sp. (Atteriini).

Vol. 101, No. 2, March & April 1990 113

axis of the femur, extending approximately to the distal end of the femur
(Fig. 5).

Taxonomic distribution. The foreleg hairpencil is present in seven
of eight described atteriine genera; it is absent in the monotypic Holop-
tygma Powell which is the only atteriine with a costal fold. It is absent in
one or more species of six of the seven genera (Table 3).

The monophyly of the Atteriini is convincingly demonstrated by the
modified "corethrogyne" scales on sterna VI and VII of the female and
the correlated oviposition behavior (Powell 1986). The corethryogyne
scaling is evidently absent in Sisurcana Powell where no unequivocal
association of males and females is known (Powell 1986). The unique
hairpencil provides further evidence for the monophyly of the Atteriini,
as well as confirming the membership of Sisurcana.

CONCLUSIONS

Two types of male foreleg hairpencils are found in the Tortricinae,
one type in Euliini and Schoenotenini, and the other in Atteriini. The
hairpencils are similar in structure, configuration, and point of origin.
Consequently, the shared presence of these structures may be inter-
preted as evidence of the common ancestry of these tribes.

Males of an undescribed genus that possess convincing atteriine
genitalia [i.e. "Philedone" aluminias (Meyrick) and "P."" citrochyta (Mey-
rick)], have typical euliine/schoenotenine hairpencils. It is possible that
the genus represents an early branch of Atteriini (before the develop-
ment of the atteriine hairpencil), and that the atteriine hairpencil repre-
sents a modification of that found in the Euliini and Schoenotenini. If
this interpretation is correct, the character supports relationships unsus-
pected by previous workers attempting to reconstruct tortricoid phylogeny.

ACKNOWLEDGMENTS

I thank the following for providing comments and suggestions on various drafts of the
manuscript: R.L. Brown (Mississippi State University), J. F. G. Clarke (National Museum
of Natural History). M. E. Epstein (National Musuem of Natural History), J. A. Powell
(University of California, Berkeley), D. L. Wagner (University of Connecticut), and two
anonymous reviewers.

1 14 ENTOMOLOGICAL NEWS

Table 1. Distribution of male foreleg hairpencil in Euliini ("-" = structure not observable;
"F" = genera known only from females).

No. ofspp. % possession

examined of hairpencil

Acroplectis Meyrick 1 0

Anopina Obraztsov 30 100

Anopinella Powell 8 0

Apolychrosis Amsel 4 0

Apotomops Powell 2 0

Bicavernaria Razowski 1 100

Bonagota Razowski 6 0

Chicotortrix Razowski 1 100

Chileulia Powell 1 0

Chilips Razowski 2

Chrysoxena Meyrick 1 0

Clarkenia Razowski 5 0

Clarkeulia Razowski 18 83

Coryssovalva Razowski 1 0

Deltinea Pastrana 1 0

Dorithia Powell 17 0

Eriotortrix Razowski 2 100

Eniocornutia Razowski 2 100

Emocornutina Razowski 1 100

Eulia Hiibner 1 100

Gauruncus Razowski 3 100

Helicteulia Razowski 1 100

Hynhamia Razowski 3 67

Inape Razowski 10 90

Neoeulia Powell 1 0

Nesochoris Clarke 2 0

Oregocerata Razowski 1

Orgyuncus Razowski 1

Ortognathosia Razowski 1 0

Paraptila Meyrick 4 100

Popavanita Razowski 2 50

Proeulia Clarke 20 85

Pseudomeritastis Obraztsov 4 75

Ptyongnathosia Razowski 1 100

Quasieulia Powell 1 0

Rhythmologa Meyrick 1 F

Seticosta Razowski 15 100

Silenis Razowski 3 100

Telurips Razowski 1 0

Terinehrica Razowski 5 100

Uelia Razowski 1 0

Uncicida Razowski 1 0

Vol. 101, No. 2, March & April 1990 115

Table 2. Distribution of male foreleg hairpencil in Schoenotenini ("F" = genera known
only from females.

No. ofspp. % possession

examined of hairpencil

Antigraptis Meyrick 1 F

Archactenis Diakonoff 100

Barygnathella Diakonoff 9 0

Campotenes Diakonoff 4 50

Choristenes Diakonoff 1 F
Cornuticlava Diakonoff 3

Copidostoma Diakonoff 1 F

Diactenis Meyrick 1 0

Homalernis Meyrick 2 F

Litotenes Diakonoff 1 100

Metachorista Meyrick 6 0

Neotenes Diakonoff 1 100

Rhopalotenes Diakonoff 4 100

Rhabdotenes Diakonoff 6 100

Schoenotenes Meyrick 100

Stenotenes Diakonoff 2 0

Saetotenes Diakonoff 6 100

Oligotenes Diakonoff 2 0

Orthocomotis Dognin 25 0

Paracomotis Razowski 1 0
Protarchella Diakonoff 3

Proactenis Diakonoff 1 0

Zenotenes Diakonoff 1 0

Table 3. Distribution of male foreleg hairpencil in Atteriini.

No. ofspp. % possession

examined of hairpencil

Anacrusis Diakonoff 8 88

Archipimima Powell 3 67

Atteria Walker 4 100

Holoptygma Powell 1 0

Sisurcana Powell 4 25

Templemania Busck 4 25

Tina Powell 1 100

Tinacrucis Powell 3 67

LITERATURE CITED

Baker, T.C. and R. T. Carde. 1979a. Courtship behavior of the oriental fruit moth
(Grapholita molesta): experimental analysis and consideration of the role of sexual
selection in the evolution of courtship pheromones in the Lepidoptera. Ann. Entomol.
Soc. Amer. 72:173-188.

1979b. Analysis of pheromone-mediated behavior in Grapholita molesta. the

oriental fruit moth (Lepidoptera: Tortricidae). Environ. Entomol. 8:956-968.

1 16 ENTOMOLOGICAL NEWS

Baker, T.C., R. Nishida and W. L. Roelofs. 1981. Close-range attraction of female orien-
tal fruit moths to herbal scent of male hairpencils. Science 214:1359-1361.

Birch, M.C. 1972. Male abdominal brush-organs in British noctuid moths and their value
as a taxonomic character. Entomol. 105: 185-205.

. 1985. Eversible structures, in The Moths of Great Britain and Ireland, vol.

9:9-18.

Falkovitch, M. J. 1962. Use of secondary sexual characters in classification of the
Palearctic Olethreutinae (Lep., Tortricidae). Entomol. Rev. Washington 41:878-885.

Horak, M. 1984. Assessment of taxonomically significant structures in Tortricinae (Lep..
Tortricidae). Mitt. Schweiz. Entomol. Gesell. 57:3-64.

and R. L. Brown. 1990. Taxonomy and phylogeny,//; L. P. S. van derGeest (ed.).

Tortricoid Pests. Elsevier, Amsterdam (in press).

Powell, J. A. 1986. Synopsis of the classification of Neotropical Tortricinae, with descrip-
tions of new genera and species (Lepidoptera: Tortricidae). Pan-Pac. Entomol. 62:372-
398.

Varley, G. C. 1962. A plea for a new look at Lepidoptera with special reference to the scent
distributing organs of male moths. Trans. Soc. Brit. Entomol. 15:3-40.

Yasuda, T. 1972. The Tortricinae and Sparganothinae of Japan (Lep.: Tortricidae). Part 1.
Bull. Univ. Osaka Prefect. Ser. B 24:53-134.

BOOKS RECEIVED AND BRIEFLY NOTED

THE BUTTERFLIES OF HISPANIOLA. A. Schwartz. 1989. Univ. of Florida Press,
Gainesville, FL. 580 pp. 7 pi. $35

Each species account provides detailed collection and distribution data, accompanied
by a range map. Species accounts also contain substantial ecological information. One
new species is described. A key to the butterflies of Hispaniola is included.

ECOLOGY AND NATURAL HISTORY OF TROPICAL BEES. D.W. Roubik. 1989.
Cambridge University Press. 514 pp. 22 pi. $69.50

This book summarizes and interprets worldwide research on the diversity of bees,
emphasizing their function within the tropical biota and including their interaction with
human populations. The book draws together several major themes of ecology, natural his-
tory, and evolution and is intended to be a reference for research workers in the field.

Vol. 101, No. 2. March & April 1990 117

ACALYPTRATE DIPTERA REARED FROM
HIGHER FUNGI IN NORTHEASTERN OHIO1

Britt Bunyard, B.A. Foote2

ABSTRACT: Living fungi were collected from two sites in northeastern Ohio between the
fall of 1987 and fall of 1988. Flies reared from this material in the laboratory were deter-
mined to species.

Fourteen species of acalyptrate Diptera, comprising five families and eight genera, were
reared from 44 species (14 families) of higher fungi. Among flies reared was Leiomyza
laevigata [Asteiidae]. No descriptions of the immature stages of any of the over 100 species
of Asteiidae worldwide are available. Also reared was Drosophilaguttifera [Drosophilidae],
another species whose biology is poorly known.

Most mycetophagous Diptera appeared to be generalists with respect to utilization of
fungal species. Larvae of several species were probably scavengers, utilizing decaying
fungal material.

Aside from a few publications (Buxton, 1960; Pielou. 1966; Pielou
andMathewman, 1966; Pielou and Verma, 1968;Shorrocks. 1973; Valley.
el « /., 1 969), the study of mycetophagous Diptera associated with mush-
rooms remains in a pioneer stage (Graves and Graves, 1985).

The purpose of this study was to determine the species of acalyptrate
Diptera associated with higher fungi in northeastern Ohio (Portage Co.).
Generalizations about the trophic relationships of certain Diptera to
their fungal hosts (strict mycophagy, polyphagy, saprophagy) are also
given.

MATERIALS AND METHODS

Two sites in Portage Co. were chosen for the collection of fungi: Towner's
Woods near Kent and West Branch State Park near Ravenna.

Material was collected between September of 1987 and September of
1988. Each collected fungus was placed in a plastic bag or wrapped in
wax paper to prevent larvae of one mushroom from entering another.
Fungi were then identified using various sources (Graham, 1944; Lin-
coff, 1 98 1 ). To avoid incidental occurrences of Diptera with the fungi (e.g.
resting or hiding in crevices), only adults which actually emerged from
larvae occurring within the fungus were counted.

Upon emergence, adult Diptera were retained alive for at least 24
hours to allow the exoskeleton to harden and then killed and preserved.
Adults were either pinned or placed directly into 70% ethanol. Larvae

1 Received July 10, 1989. Accepted September 2, 1989.

-Department of Biological Sciences, Kent State University, Kent, Ohio 44242

ENT. NEWS 101(2): 117-121, March & April. 1990

118

ENTOMOLOGICAL NEWS

were killed in boiling water and then preserved in 70% ethanol. Preser-
ved specimens were placed in the Kent State University collection of
Diptera.

Rearing required the construction of special containers which allowed
the fungi to remain in a somewhat natural condition. The bottom of
these rearing chambers consisted of the bottom of petri dishes (10 x
100mm) to which had been added moistened pulverized peat moss. It
was necessary to keep the peat moss substrate in contact with the fungus
initially and to moisten it frequently to prevent desiccation. As the
fungus decayed, the substrate absorbed moisture produced by the fungus.
The remainder of the rearing container consisted of clear plastic tubing
having a diameter of 90mm and cut to various lengths. To the top of this
was glued fine polyester mesh material. The rearing chamber was placed
over the fungus in the petri dish. The fungal material was retained in the
rearing containers for at least three months.

RESULTS AND DISCUSSION

Species of acalyptrate Diptera reared from the fungal material are
given in table 1.

Table 1. Associations between acalyptrate Diptera species and their fungal hosts.

Acalyptrate Species

1234567

Fungal Host Species

8 9 10 11 12 13 14 15 16 17 18 19 20

Asteiidae
Leiomyza laevigata

Chloropidae

Gaurax atripalpus

Tricimba lineella

Tricimba sp.

Nartshukialla sp. poss. melancholica

Drosophilidae
Drosophila duncani
Drosophila fatleni
Drosophila gutlifera
Drosophila putrida
Drosophila testacea
Drosophila tripunctata
Mycodrosophila dimidiala

Ephydridae
Athryroglossa granulosa

Lonchaeidae
unknown spp.

1 Agaricus arvensis

2 Lepiota rachodes

3 Amanita muscaria

4 Boletus chrysenteron

5 Pluteus cervinus

6 Laetiporus sulfureus

7 Polyporus pubescens

8 Russula compacta

9 Russula vesca

10 Pholiola mutabilis

1 1 Pholiola squarrosa

12 Psilocybe polytrichophila

13 Collybia dryophila

14 Marasmius oreades

15 Mycena galericulala

16 Omphalotus olearius

1 7 Oudemansiella radicala

18 Pleurotus ostreatus

19 Pleurotus pulmonaria

20 Tricholomopsis platyphyla

Vol. 101, No. 2, March & April 1990 119

Leiomyza laevigata Meigen [Asteiidae] has been reared from fungi
previously (Buxton, 1960; Sahrosky, 1957). The family Asteiidae is a
small one of some 100 species of which little is known. No descriptions of
the immature stages of any species of Asteiidae are available.

Gaurax atripalpus Sabrosky [Chloropidae] has been reared pre-
viously from Fames sp., [Polyporaceae] (Valley, etal., 1969). It is apparent
from this and our more recent rearings that G. atripalpus utilizes the fungi
as both a food source and site of overwintering.

The rearing of Athyroglossa granulosa Cresson [Ephydridae] from

fungi is surprising, because members of this family usually have larvae

that are aquatic or semiaquatic. Grimaldi and Jaenike (1983) reared

adults from larvae feeding in decaying skunk cabbage, Symplocarpus

foetidus (L.) Nutt.

There are many known species of fungivorous Drosophilidae. Droso-
phila falleni Wheeler, D. putrida Sturtevant, D. testacea von Roser, D.
tripunctata Loew, and Mycodrosophila dimidiata Loew, which were all
reared in this study, are all well known fungal feeders (Jaenike. 1977.
1978; Jaenike, et al., 1983; Patterson and Stone, 1952). However, none is
known to be monophagous (Jaenike, 1978; Lacy, 1984). Drosophila dun-
cani Sturtevant and D. guttifera Walker are also known to be fungal
feeders but little is known about the life history of either species (Patter-
son and Stone, 1952).

Two drosophilids, D. falleni and D. putrida, were reared from toxic
species ofAmanita mushrooms. These two as well as a few other species
have been reared in the past (Jaenike, 1977; Jaenike, et at., 1983) from
mushrooms containing toxic amanitins. The amanitins are alkaloid
compounds which are potent inhibitors of RNA polymerase II, the
enzyme which transcribes genes that encode messenger RNA's ( Wieland.
1968). Therefore, these compounds are potentially toxic to all eukar-
yotes. How these drosophilids manage to avoid being affected deleter-
iously is not known (Jaenike et al., 1983).

An interesting question raised by this study concerns how so many
different acalyptrate species, sometimes of the same genus, can coexist in
the same substrate (Table 1 )? As many as six species were found in the
same fungal sporophore at the same time. When examined more closely
it was apparent that in most cases larvae of the different species were
feeding on the same material, at the same time, and in a similar manner
(i.e. burrowing through the fungal sporophore). Apparently, competitive
exclusion is not a factor here, as the fungal food probably was not a limit-
ing resource. However, Grimaldi and Jaenike (1984), demonstrated that
mycophagous larvae frequently do exhaust the food available in indi-
vidual mushrooms. It is probable that predators or parasites of these lar-

120

ENTOMOLOGICAL NEWS

vae functioned to reduce competition (both inter - and intraspecific)
between larvae. Many parasitic wasps were obtained from many of the
fungi surveyed.

Another point made evident by this study as well as earlier one (Bux-
ton, 1960) is that certain fungi are more attractive to species of Diptera
than other fungi. For example, Pluteus cervinus Fr. [Pluteaceae] pos-
sessed the greatest diversity of acalyptrate species (Table. 1 ). In contrast,
several species of fungi were repeatedly examined, but no species of
acalyptrate Diptera were obtained. The reasons for this are unclear and
further research in this area is necessary. A final point to be made by this
study concerns the mechanisms utilized by the many species of fungi-
vorous Diptera to cope with the fact that fungal sporophores represent
an ephemeral and unpredictable food source. The months of May and
September produced the highest number of species of emerging adults
(Fig. 1). Many species of Diptera probably initiate a reproductive
diapause during times of no larval food sources, such as the dry summer
months. Other species may utilize other food sources, including live,
injured, and decaying vegetation. An example is Drosophila gutlifera
[Drosophilidae], a rare species that was believed to be strictly myco-
phagous (Patterson and Stone, 1952). However, we discovered that this
species, at least in laboratory rearings, readily accepted other substrates
(tomato juice, agar, bananas, commercial Drosophila medium) as an
ovipositional site and larval food source (Bunyard and Foote, 1990).
Similar results have been obtained in laboratory rearings of other
mycetophagous species of the guinaria group of Drosophila (Grimaldi,

Fig. 1 Number of rearings per month for
five families of acalyptrate Diptera.

N
o
s

r
e
a
r

9
s

Lonchaeldae
Astelldae

Chloropldae
Phorldae

I 1 Drosophilidae

Vol. 101, No. 2. March & April 1990 121

pers. comm.). However, no field-based records of rearings from non-
fungi sources have been reported for these species, and the laboratory
results thus may not reflect reality.

ACKNOWLEDGMENTS

We wish to thank D. Grimaldi. American Museum of Natural History, New York City,
and K. Valley. Pennsylvania Department of Agriculture. Harrisburg, for their assistance in
identification of Diptera.

We are also grateful to S. Mazzer. Kent State University, for identification of fungal
specimens.

LITERATURE CITED

Bunyard, B. and B. A. Foote. 1990. Life history and host selection of Drosophila guttifera

[Diptera: Drosophilidae], a consumer of mushrooms. Entomol. News. In Press.
Buxton, P. A. 1960. British Diptera associated with fungi. III. Flies of all families reared

from about 150 species of fungi. Entomol. Mon. Mag. 96: 61-94.
Graham, V.O. 1944. Mushrooms of the Great Lakes Region. Chicago Nat. Hist. Museum.

Chicago, p. vii-390.
Graves, R. C. and C. F. Graves. 1985. Diptera associated with shelf fungi and certain

other micro-habitats in the highlands area of western North Carolina. Entomol. News

96: 87-92.
Grimaldi, D. and J. Jaenike. 1983. The Diptera breeding on skunk cabbage. Symplocar-

pusfoetidus (Araceae). J. N.Y. Entomol. Soc. 91: 83-89.
1984. Competition in natural populations of mycophagous Drosophila. Ecology

65: 1113-1120.
Jaenike, J. 1977. Resource predictability and niche breadth in the Drosophilu guinaria

species group. Evolution 32: 676-678.

. 1979. Host selection by mycophagous Drosophila. Ecology 59: 1286-1288.

Jaenike, J., D. A. Grimaldi, A. E. Sluder, and A. L. Greenleaf. 1983. oc-Amanitin

tolerance in mycophagous Drosophila. Science 221: 165-167.

Lacy, R. C. 1984. Ecological and genetic responses to mycophagy in Drosophilidae (Dip-
tera). pp. 286-301. In Q. Wheeler and M. Blackwell. eds. Fungus/Insect Relationships:

Perspectives in Ecology and Evolution. Columbia Univ. Press. NY. NY.
Lincoff, G. H. 1981. The Audubon Society Field Guide to North American Mushrooms.

Albert Knopf, Inc., NY. 926pp.
Patterson, J.T. and W. S. Stone. 1952. Evolution in the genus Drosophila. The Mactnillan

Company, NY 610pp.
Pielou, D. P. 1966. The fauna of Polyporus betulinus (Bulliard) Fries [Basidiomycetes:

Polyporaceae] in Gatineau Park. Quebec. Can. Entomol. 98: 1233-1237.
Pielou, D. P. and W. G. Matthewman. 1966. The fauna of Fomes fomeniarius : (Linnaeus

ex. Fries) Kickx growing on dead birch in Gatineau Park. Quebec. Can. Entomol.. 98:

1308-1312.

Pielou, D. P. and A. N. Verma. 1968. The arthropod fauna associated with the birch brac-
ket fungus. Polyporus betulinus. in eastern Canada. Can. Entomol. 100: 1 179-1 199.
Sabrosky, C. W. 1957. Synopsis of the New World species of the dipterous family Astei-

idae. Ann. Entomol. So. 50:43-61.
Shorrocks, B. and A. M. Wood. 1973. A preliminary note on the fungus feeding species

of Drosophila. J. Nat. Hist. 7:551-556.
Valley, K., T. Wearsch and B. A. Foote. 1969. Larval feeding habits of certain Chloro-

pidae (Diptera). Proc. Entomol. Soc. Wash. 71: 29-34.
Wieland, T. 1%X. Poisonous principles of mushrooms of the genus. Amanita. Science 159:

946-952.

122 ENTOMOLOGICAL NEWS

SOCIETY MEETING OF FEBRUARY 28, 1990

EVOLUTION OF EUPHILOTES BIOTYPES: TRIALS AND TRIBULATIONS

OF THE BUTTERFLY BLUES

By Dr. Gordon F. Pratt

Darwin's book. The Origin of Species, changed the way we view ourselves and the world
around us; nevertheless, the origin of species remains a central concern of evolutionary
biologists. Because insects represent the great majority of known species, they are often the
subject of studies on speciation. Dr. Gordon F. Pratt, now a post doctoral associate at the
University of Delaware, is interested in insect speciation. While he was a graduate student
in California he studied a fascinating complex of blue butterflies of the genus Euphilotes
whose larvae feed exclusively on the blossoms and seeds of one or another of the species of
wild buckwheat, Eriogonum.

Variation in altitude, soil type, and climate in the southwestern United States have
created habitat islands suitable for the growth of Eriogonum and in turn Euphilotes. The
tight coupling between the flowering of particular host species and the butterfly's life cycle
has resulted in many local races that are behaviorly distinct. For example, some emerge in
the spring and others in the fall. Some adults fly near the ground where the flowers of their
food plant are, while others fly higher for flowers of a different species. The sister species
Euphilotes enoptes and E. battoides have sharp differences in male and female genitalia;
however, electrophoretic analysis of allozymes in various host races show they are closely
related. These studies also indicate that host races are evolving rapidly and suggest that E.
battoides evolved from the E. enoptes host race that feeds on Eriogonum umbellatum.

The adults of Euphilotes obtain nectar from the same plants on which they lay their eggs.
The larvae are cryptic and assimilate pigments from the flowers on which they feed. They
are attended by ants. Pupation occurs in the ground beneath the host plant. Presumably, in
desert environments where moisture triggers seed germination or growth of perennial
species, moisture also serves to synchronize adult emergence with blossoming. The fact
that pupae can survive up to five years in the laboratory helps to explain the absence of
adults in dry years and their reappearance several years later. The ability of the butterfly to
evolve different local populations adapted to specific host plants has resulted in sympatric
populations of a single species that do not interbreed.

Dr. Pratt's talk at the Philadelphia Academy of Natural Sciences drew one of the largest
audiences for a membership meeting in five years. Eighteen members and twelve guests
were present. Like Dr. Pratt's talk, the notes of local entomological interest focused on the
Lepidoptera and speciation. Dr. Chuck Mason reported that both the E and Z pheromone
races of the European corn borer, Ostrinia nubilalis, are common in Delaware. The E-type
emerges early and produces offspring that feed on a variety of early host plants other than
corn. The other type produces the Z isomer of the pheromone, emerges later, and attacks
corn in its first generation. The second generation of both types reproduce on corn. Hybrid
individuals produce a mixture of the pheromones. Dr. Duke Eisner noted a possibly
related situation of cranberry truilworm.Acrobasis vaccinii. Attempts to attract males with a
pheromone extracted from theconspecific race feeding on blueberry have been unsuccess-
ful. Dr. Ken Frank noted that female cecropia moths, Hyalophora cecropia, that emerge in
captivity later than their wild counterparts will sometimes attract males of the introduced
cynthia silkmoth, Sam/a ov7f/;/a, which normally emerge later than cecropia in the wild. At
the end of the meeting Dr. Frank distributed a number of cecropia coccoons he had res-
cued from a city site likely to be destroyed before summer.

Harold B. White
Corresponding Secretary

ENT. NEWS 101(2): 122, March & April. 1990

Vol. 101, No. 2. March & April 1990 123

IMPROVED CULTURE TECHNIQUES FOR MASS
REARING GALLERIA MELLONELLA (LEPIDOP-

TERA: PYRALIDAE)1

j •}

Frank A. Eischen , Alfred DietzJ

ABSTRACT: Supplementing an artificial diet with 5% pollen, honey, or beeswax signifi-
cantly increased survival of adult Galleria mellonella. Survival on the basic diet averaged
27.4%. When honey, pollen, or wax was added, survival was 44.7%. 80.8%. and 89.6%. respec-
tively. Developmental time for moths fed diets containing 5% pollen or a combination of
pollen, honey, and wax was shorter by approximately 2-5 days. Newly emerged virgin
females which had fed as larvae on a 100% pollen, honey, and wax diet weighed 36% more
than females derived from larvae fed the artifical diet. A phagostimulatory hypothesis
is suggested.

Additionally, 78% of prepupae accepted cut plastic soda straws as puparial sites and
spun their cocoons in them. This greatly facilitated the handling of individual prepupae
and pupae and should be a convenience to both research programs and commercial wax
moth producers.

The greater wax moth, Galleria mellonella(\~.), is easily reared and is
often used as a laboratory animal. However, our initial attempts to
establish a culture from wild stock resulted in low survival rates (ca. 30%)
when an artificial diet [Stoneville( SV) developed by King etal. 1979] was
used. However, pilot studies indicated that survival was good (ca. 95%)
on a diet composed wholly of their naturally occurring larval food, i.e.
pollen, honey, and beeswax. Inspection of the artificial diet (King el al.
1979) did not reveal nutritional shortcomings. Previous studies have
shown that small amounts of beeswax improved growth rates (Beck
1960, Dadd 1966, Young 1964). Further, Dadd (1966) observed higher
survival rates during portions of larval development when beeswax was
added to an artificial diet. These findings suggested that small amounts
of natural food could play an important nutritional role in adult sur-
vival. We report here the results of a study done to determine why sur-
vival differed so strikingly on artificial diet compared with a diet com-
posed of natural ingredients.

MATERIALS AND METHODS

Six diets were tested. Their composition by weight was as follows: 1 )
100% Stoneville(SV), 2) 95% SV + 5% pollen(P), 3) 95% SV + 5% hon-
ey(H),4) 95% SV + 5% beeswax(W), 5) 95% SV + 5% PHW, 6) 100% PHW.

^Received June 19. 1989. Accepted September 23. 1989.

"Dept. Of Entomology. Washington State University, Pullman, WA 99164

- Dept. of Entomology. University of Georgia. Athens. GA 30602

ENT. NEWS 101(2): 123-128. March & April. 1990

124 ENTOMOLOGICAL NEWS

The Stoneville diet consists largely of baby foods (Gerber mixed and
high protein cereals), gycerol, wheat germ, water, sucrose, and vitamins
(see Kingetat. 1 979 for details). Its protein composition is about 13%. The
pollen-honey-wax diet was made by mixing bee-collected pollen (63%;
see Dietz 1982 for plant species represented) with chipped honey comb
(37%). The honeycomb (cappings) contained approximately 50% honey.
The protein content of this diet is estimated to be about 13%. Twenty
grams of diet were placed in 100ml glass rearing jars, sealed and kept
frozen until used. Fortified diets were prepared by placing 19gofSVdiet
in the rearing jar, and then Ig of either pollen, honey, or wax was sprink-
led or dripped onto its surface.

Moths were obtained by removing overwintering prepupae from a
dead honey bee hive in Athens, Georgia. They were incubated at 30°C.
Adults emerged in about two weeks and mated inter se. Eggs were collected
between strips of pleated wax paper. Three days after laying, egg masses
were divided into small clusters of 1 0- 1 5 eggs. These clusters were placed
on filter paper in petri dishes and incubated at 30°C + 2°C. Just before
hatching, eggs were examined and clusters containing at least 10 larvae
that appeared normal were selected for testing. Excess larvae were des-
troyed by puncturing the eggs with a needle. Eggs were then placed in
small aluminum foil baskets ( 16mm x 10mm) and randomly assigned to
diets. Twenty replications of each diet were performed. After addition of
the eggs, the rearing jars were sealed with a solid metal screw lid contain-
ing a rubber interseal. Rearing jars were held at 32°C + 1 °C, 40 + 10%
RH, and 12:12LD. Seven days after infestation, egg clusters were re-
moved and percent hatch determined. Also, the solid lids were replaced
with a similar lid containing a 9mm hole. This hole was covered with a
piece of transparent tape, sticky side out. Several small holes were
punched in the tape with a small pin to allow air exchange. When the
first prepupae were observed, this tape was replaced with a circular piece
of aluminum screen. At this stage, rearing jars were inspected daily. As
adults appeared they were removed, sexed, and their emergence date
noted. The first twenty adult virgin females that emerged from the SV
and the pollen-honey-wax diets were weighed. Sixty days from the time
of egg laying, the contents of the rearing jars were examined. Cocoons
were counted and pupal mortality noted.

Survival and emergence data were analyzed with a 1-way ANOVA,
and differences among treatments evaluated with Duncan's multiple
range test. Student's t-test was used to determine differences in adult
female weights.

To confirm that the supplemented Stoneville diet was suitable for a
mass rearing program, we fed the Stoneville+5% fresh frozen pollen to
ca. 6,000 hatching G. mellonella larvae (eggs were weighed, not counted).

Vol. 101, No. 2. March & April 1990 125

As prepupae emerged from the diet and began spinning light cocoons,
they were removed and placed in 3.81 glass jars containing cut plastic
soda straws (2 X 0.5cm; SweetheartR). About 70 prepupae were added to
jars containing 200-300% excess of cut straws (scattered horizonally on
the bottom of the jar), which were covered with a single circular piece of
toweling. Jars were kept in a darkened incubator (same conditions as in
above dietary trials).

RESULTS

The addition of 5% of either pollen, honey, or beeswax (PHW) to the
Stoneville(SV) diet resulted in a significant increase in adult survival
(Table 1 ). Adult survival on the basic diet averaged 27.4%. The addition of
honey increased survival to 44.7% (P < 0.05). Fortification with either
pollen, wax, or a combination of pollen, honey, and wax produced sur-
vival rates that did not differ significantly from the control diet (89.2%).

Males reared on the SV+5% pollen, SV+PHW, and control diet
eclosed in a significantly shorter time (ca. 3-5 days, P < 0.05) than moths
on the other three diets (Table 1). Average time to first emergence was
slightly shorter for males (0.2-1 .3 days) than that required by females, but
this difference was not significant. Newly emerged virgin females weighed
on average less when reared on the SV diet, than on the control diet ( 1 14
and 156mg, respectively; P < 0.001). During the mass rearing trial 78% of
prepupae spun their cocoon inside the cut soda straws. The adult sur-
vival rate for larvae fed the Stoneville+5% pollen diet during this trial
was estimated to be about 84%.

DISCUSSION

The striking increases in survival caused by small amounts of pollen,
honey, or beeswax was not the result of an altered physical consistency,
nor the addition of essential nutrients (Haydak 1936. 1940. Allegret 1964;
Dadd 1966; Marston and Campbell 1973). Dadd (1966) suggested that
the inclusion of beeswax served as a source of metabolic water. However,
the basic Stoneville diet contains about 17% water and supports good lar-
val growth (King etal. 1979). The 300% increase in survival with the addi-
tion of 5% beeswax to this diet suggests that beeswax performed an
additional function.

Eischen et al. (unpublished) found in a preliminary test that newly
hatched larvae preferentially chose artificial diets to which alcoholic
extract of pollen was added. This suggests that pollen provided phago-
stimuli attractive to G. mellonella. Other observations support this view.
Balazs (1958) reported that newly hatched larvae preferred honeycomb

126 ENTOMOLOGICAL NEWS

to an artificial diet, even though this diet contained honey and beeswax.
He also noted that older larvae fed the same artificial diet chose hon-
eycomb when given the opportunity. Haydak (1936) reported that larvae
attacked old combs in those areas that contained pollen. We have on
numerous occasions observed newly hatched larvae that have been
given the Stoneville diet wander about the inner walls of their rearing
container when food was nearby. Newly hatched larvae given a honey-
pollen-wax diet were rarely seen away from the food. The natural history
of this moth also lends support to a phagostimulatory hypothesis. Adult
females oviposit in and around bee colonies; generally in crevices where
eggs are protected from the bees (Paddock 1918, Nielsen and Brister
1977). Once hatched, the larvae must search for food. Though distances
may be short, it would be adaptive to be able to locate food (which also
serves as shelter) rapidly in the hostile interior of a honey bee colony.

The shorter times to eclosion of moths fed diets containing pollen
indicate that even small amounts of pollen are effective in promoting
development. Since pollen was localized on the upper surface of the diet
mass, it seems unlikely that its consumption would have been uniformly
distributed during larval growth. If the phagostimulatory hypothesis is
true, then consumption and consequent beneficial effects occured dur-
ing the earliest instars. From this it follows that an early pollen meal may
supply a nutrient, perhaps protein, that allows faster development, while
apparently wax and honey do not.

Prepupal acceptance of soda straws greatly facilitated the handling
of prepupae and pupae. Large numbers of known age groups can be held
in relatively small containers. This is convenient when storing prepupae
under refrigerated conditions (ca. 15.5°C). We find that chilled, lightly
spun prepupae are easy to use by fishermen, a slight squeeze on the straw
prompts the prepupae to crawl out. Because male prepupae emerged
from the diet slightly before females, the first harvest of prepupae results
in a high percentage of males. This is an added benefit when separation
of sexes is desired. Since these tests, we have successfully reared many
thousand G. mellonella using this technique.

These findings should be of value to those who wish to preserve the
genetic variation observed in wild stock or to increase the yield of G.
mellonella in newly-established cultures (Bush 1975). However, cultures
that have been reared for many generations on a particular artificial diet
and consequently undergone selection for it may not show greatly im-
proved survival. Nevertheless, under some conditions it could be pru-
dent as well as cost effective to supplement artificial wax moth diets with
small amounts of properly stored pollen or unprocessed honeycomb.

Vol. 101, No. 2, March & April 1990

127

Table 1 . Developmental characteristics ofGalleria mellonella reared on an artificial diet for-
tified with honey, pollen, or beeswax

% adult

days to
Istcf

days to adult
1st 9 9

Diet

survival
(X±SE)

emergence
(X±SE)

emergence weight
(X±SE) (X±SE)

Stoneville

27.4 ±

4.7 a1

42.7

± 1.1 a

43.9

± 0.8 a 1 14.2 ± 5.3 mg2

Stoneville +

44.7 ±

6.0 b

43.3

± 1.3 a

43.5

± 0.8 a

5% honey

Stoneville +

89.6 ±

3.8 c

42.8

± 0.7 a

43.7

± 1.0 a

5% wax

Stoneville +

80.8 ±

4.1 c

39.7

± 0.8 b

41.0

± 0.9 b

5% pollen

Stoneville + 5%

82.5 ±

4.7 c

39.2

± 0.8 b

39.4

± 0.8 c

honey, pollen, wax

Honey, pollen, wax

89.2 ±

2.6 c

37.6

± 0.7 b

37.8

±0.7 cd 156.0 ±5.8

Means in the same column followed by the same letter do not differ significantly at the 5%
level according to Duncan's multiple range test.

-Means weights are significantly different (P < 0.001). Female weights on other diets were
not recorded.

ACKNOWLEDGMENTS

We thank R.D. Akre and C.W. Bersiford for reviewing the manuscript. J. Fields helped
with the statistical analysis. This investigation was supported in part by Cooperative
Agreement 25-2 1 -RC293-078 between the University of Georgia (A. Dietz. principle inves-
tigator) and the Honey Bee Breeding. Genetics, and Physiology Laboratory, USDA-ARS.
Baton Rouge. LA.

LITERATURE CITED

Allergret, P. 1964. Interrelationship of larval development, metamorphosis and age in a
pyralid lepidopteran, Galleria mellonella(L.), under the influence of dietetic factors.
Experimental Gerontology 1: 49-66.

Balazs A. 1958. Nutritional and nervous factors in the adaption of Galleria mi'llomJIti to
artifical diet. Acta Biol. Acad. Sci. Hung. 9: 47-69.

Beck, S.D.1960. Growth and development of the greater wax moth Galleria mellonella(L.)
(Lepidoptera: Galleriidae). Trans. Wis. Acad Sci.. Arts Lett. 49: 137-148.

Bush, G.L. 1975. Genetic variation in natural insect populations and its bearing on mass-
rearing programmes. In pp. 9-17: IAEA/FAO Panel Proc. Sen. Controlling Fruit Flies
by the Sterile Insect Technique. Vienna, 1973.

128 ENTOMOLOGICAL NEWS

Dietz, A., R. Krell, and M.S. Brower. 1982. Pollination and our seashores, pp. 57-66. In:
Proc. 10th Pollination Conference, Southern Illinois University, July 1982.

Dadd, R.H. 1966. Beeswax in the nutrition of the wax moth, Galleria mellonella (L.). J.
Insect Physiol. 12: 1479-1492.

Haydak, M.H. 1936. Is wax a necessary constituent of the diet of wax moth larvae? Ann.
Entomol. Soc. Am. 29: 581-588.

Haydak, M.H. 1940. The length of development of the greater wax moth. Science. 91:
525.

King, E.G., G.G. Hartley, D.F. Martin, J.W. Smith, T.E. Summers, and R.D. Jack-
son. 1979. Production of the tachinid Lixophaga diatraeae on its natural host, the sugar-
cane borer, and on its unnatural host, the greater wax moth. U.S.D.A.. S.E.A. Advances in
Agricultural Technology Southern Series No. 3.

Marston, N. and B. Campbell. 1973. Comparison of nine diets for rearing Galleria
mellonella. Ann. Entomol. Soc. Am. 66: 132-136.

Nielsen, R. A. and D. Brister. 1977. The greater wax moth: Adult behavior. Ann. Entomol.
Soc. Am. 70: 101-103.

Paddock, F.B. 1918. The beemoth or waxworm. Texas Agric. Exp. Stat. Bull. 231: 3-38.

Young, R.G. 1964. Digestion of wax by the greater wax moth, Galleria mellonella(L.). Ann.
Entomol. Soc. Am. 57: 325-327.

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ENTOM

MAY & JUNE, 1990

LNEWS

' host record for An iso torn a basalis (Coleoptera: Leiodidae)
CT<3 ^ I reeding in sporocarps of slime mold Lycogala flavofuscum
~; Myxomycetes: Liceales) Q.D. Wheeler, E.R.Hoebeke 129

r7 J\^"

Antennal anomaly in Oxypoda opaca (Coleoptera: Staphylinidae)
from New York E. Richard Hoebeke 133

A new species and new record of water-penny genus Psephenops
(Coleoptera: Psephenidae) from Costa Rica

Pa ulJ. Spangler 137

New record and range extension for Rhyacophila wallowa
(Trichoptera: Rhyacophilidae) from Rocky Mountain
National Park, Colorado Scott J. Herrmann 141

HPLABEL: a program and microfont for the generation of
date/locality labels using a laser printer

D.C. Darling, R.C. Plowright 143

Microcylloepus formicoideus (Coleoptera: Elmidae), a new
riffle beetle from Death Valley National Monument,
California William D. Shepard 147

The status of Pison doggonum (Hymenoptera: Sphecidae)

Arnold S. Menke 154

Distributional notes on North and Central American

Dilaridae (Neuroptera) Kevin M. Hoffman 155

First United States record of Ascalobyas (Neuroptera:
Ascalaphidae), a range extension from northeastern
Mexico to Texas Roy C. Vogtsberger 158

Biological notes on Drosophila guttifera (Diptera:
Drosophilidae), a consumer of mushrooms

B. Bunyard, B.A. Foote 161

Propylea quatuordecimpunctata: additional U.S. records of
an adventive lady beetle (Coleoptera: Coccinellidae)

A.G. Wheeler, Jr.

SOCIETY MEETING OF MARCH 28, 1990
SOCIETY MEETING OF APRIL 25, 1990
BOOK REVIEWS

164

Additions to the Papilionoidea (Lepidoptera) of the

Revillagigedo Islands, Mexico John W. Brown 167

N«

Invertebrate populations in the nests of a screech owl
(Otus asi'o) and an American kestrel (Falco sparverius)
in central New York J.I?. Philips, D.L. Dindal 170

153
160
136, 146, 157, 169

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Vol. 101, No. 3, May & June 1990 129

NEW HOST RECORD FOR ANISOTOMA BASALIS

(COLEOPTERA: LEIODIDAE) BREEDING IN
SPOROCARPS OF THE SLIME MOLD LYCOGALA
FLAVOFUSCUM (MYXOMYCETES: LICEALES)1

Quentin D. Wheeler, E. Richard Hoebeke2

ABSTRACT: Two breeding populations of the primitive staphylinoid beetle Anisotoma
basalis (Leiodidae: Agathidiini) were found feeding within mature aethalial sporocarps of
the uncommon slime mold Lycogala jlavofuscum (Myxomycetes: Liceales) on the campus
of Cornell University, Ithaca, New York, in 1983 and 1989. These discoveries represent the
first documented insect association for this myxomycete.

An emerging pattern of host utilization in the agathidiine Leiodidae
suggests that many species of these beetles are able to breed on a diverse
range of host Myxomycetes (slime molds). Few instances of host specificity
are suspected to date. The majority of species have only been associated
with mature sporocarps (Blackwell 1984; Lawence, 1989; Lawrence &
Newton, 1980; Newton, 1984; Wheeler, 1979), although a few records
exist for both Anisotoma (Russell, 1979; Wheeler, 1980) and Agathidium
(Newton, 1984; Wheeler, 1984a, 1984b, 1987) on plasmodia of host slime
molds. Many records involve common myxomycetes that produce large
fruiting bodies [e.g., Fuligo septica (L.) Wiggers] or dense masses of
smaller fruiting bodies [e.g., Stemonitis fusca Roth]. The slime mold
Lycogala epidendrum (L.) Fries, a small, puffball-like species, has also
been recorded as an agathidiine host (Lawrence & Newton, 1 980; Black-
well, 1984).

Myxomycete host records have been reported by Lawrence & New-
ton (1980), based on label data, field observations and from published
accounts in the North American literature, for eight North American
Anisotoma, including slime mold species of the genera Comatrichia,
Fuligo, Lycogala, Metatrichia, Stemonitis, Trichia and Tubifera. Precise
information on feeding habits and preferences, and microhabitats of the
majority of leiodid beetles is scarce.

In this paper we provide evidence for a new adult and larval host
record for Anisotoma basalis (LeConte) (Leiodidae: Agathidiini). Two
breeding populations of this primitive staphylinoid beetle were found
feeding within mature aethalial sporocarps of the uncommon slime
mold Lycogala Jlavofuscum (Ehrenb.) Rost. These populations were dis-
covered by one of us ( ERH) on two separate occasions, one on a prostrate
log and the other on a standing dead tree, along a wooded hillside on the

1 Received September 16, 1989. Accepted November 14, 1989.
2Department of Entomology. Cornell University, Ithaca, New York 14853

ENT. NEWS 101(3): 129-132, May & June, 1990

130

ENTOMOLOGICAL NEWS

Fig. 1. Mature aethalial sporocarp of Lycogalaflavofuscum. Beetle exit holes through the
peridium of the fruiting body are readily visible. Dime indicates scale.

Cornell University campus, Ithaca, New York (Tompkins Co.). These
discoveries represent the first documented insect association for this
myxomycete.

On 4 June 1983, a large sporocarp of L.flavofuscum (approx. 5.1 x 4.6
cm.) (Fig. 1) was found on the surface of a prostrate log (Fig. 2) along a
wooded slope on the campus of Cornell University. Within this mature
fruiting body a massive population of both larvae and adults ofAnisotoma
basalis was discovered. This population was observed and sampled for
more than a week. Because we only partially disturbed the fruiting body,
no complete census of the beetles was made. However, more than 200 lar-
vae were collected and slide-mounted, and more than that number were
collected into ethanol. We estimate that the total number of adult A.
basalis was at least 100-200 and that the number of larvae probably
exceeded 400-500.

Fig. 2. Microhabitat of the slime mold Lycogalaflavofuscum. Arrow indicates position of
fruiting body (of Fig. 1) on prostrate log.

Fig. 3. Mature aethalial sporocarp of Lycogalaflavofuscum. An adult Anisotoma basalis is
visible near exit hole of fruiting body. Scale line = 1.0 cm.

Vol. 101, No. 3, May & June 1990

131

132 ENTOMOLOGICAL NEWS

Again on 21 June 1989, several meters from the original 1983 collec-
tion site, another smaller sporocarp of L.flavofuscum (approx. 2.1 x 2.5
cm.) was discovered on the trunk of a standing, dead basswood tree
(Tilia), approximately 5 ft. above the ground. It, like the previous sporo-
carp, had also been penetrated by adult specimens of A. basalts (Fig. 3),
but no larval specimens were observed.

Both sporocarps are deposited in the Cornell University Insect Col-
lection as voucher specimens of the slime mold.

All three larval instars associated with adult A. basalts from this
myxomycete host have been described by Wheeler (1990a), as the basis
for a study on ontogeny (Wheeler, 1990b).

ACKNOWLEDGMENTS

We thank M. Blackwell (Louisiana State University, Baton Rouge) for verification of
the host identification, and J. K. Liebherr and J. V. McHugh (Cornell University) and two
anonymous reviewers for providing comments on an early draft of this paper.

LITERATURE CITED

Blackwell, M. 1984. Myxomycetes and their arthropod associates, pp. 67-90. In Q. Wheeler
and M. Blackwell (eds.), Fungus-Insect Relationships. New York: Columbia University
Press.

Lawrence, J. F. 1 989. Mycophagy in the Coleoptera : feeding strategies and morphological
adaptations, pp. 1-23. In Wilding, N., Collins, N. M., Hammond, P. M., and J. F. Webber
(eds.), Insect-Fungus Interactions. London: Academic Press.

Lawrence, J. F. and A. F. Newton, Jr. 1980. Coleoptera associated with fruiting bodies of
slime molds (Myxomycetes). Coleopt. Bull. 34:129-143.

Newton, A. F., Jr. 1984. Mycophagy in Staphylinoidea (Coleoptera), pp. 302-353. In Q.
Wheeler and M. Blackwell (eds.), Fungus-Insect Relationships. New York: Columbia
University Press.

Russell, L. K. 1979. Beetles associated with slime molds (Mycetozoa) in Oregon and
California (Coleoptera: Leiodidae, Sphindidae, Lathridiidae). Pan-Pac. Entomol. 55:1-9.

Wheeler, Q. D. 1979. Slime mold beetles of the genus Anisotoma (Leiodidae): classifica-
tion and evolution. Syst Entomol. 4:251-309.

Wheeler, Q. D. 1980. Studies on Neotropical slime mold/beetle relationships. Part I.
Natural history and description of a new species of Anisotoma from Panama (Coleop-
tera: Leiodidae). Proc. Entomol. Soc. Wash. 82:493-498.

Wheeler, Q. D. 1984a. Evolution of slime mold feeding in leiodid beetles, pp. 446-479. In
Q. Wheeler and M. Blackwell (eds.), Fungus-Insect Relationships. New York: Colum-
bia University Press.

Wheeler, Q. D. 1984b. Associations of beetles with slime molds: ecological patterns in the
Anisotomini (Leiodidae). Bull. Entomol. Soc. Amer. 30:14-18.

Wheeler, Q. D. 1987. A new species ofAgathidium associated with an "epimycetic" slime
mold plasmodium on Pleurotus fungi (Coleoptera: Leiodidae); Myxomycetes:
Physarales; Basidiomycetes: Tricholomataceae). Coleopt. Bull. 41:395-403.

Wheeler, Q. D. 1990a. Morphology and ontogeny of postembryonic larval Agathidium
and Anisotoma (Coleoptera: Leiodidae). Amer. Mus. Novitates (In Press).

Wheeler, Q. D. 1990b. Ontogeny and character phylogeny. Cladistics (In Press).

Vol. 101, No. 3, May & June 1990 133

ANTENNAL ANOMALY IN OXYPODA OPACA

(COLEOPTERA: STAPHYLINIDAE)

FROM NEW YORK1

E. Richard Hoebeke2

ABSTRACT: A specimen ot Oxypoda opaca, collected in Tompkins Co., NY, in June 1988,
was found to exhibit a teratological aberration of its left antenna, whereby the distal anten-
nomeres (articles III-XI) are compressed into a short club-like structure. This aberration is
illustrated by scanning electron microscopy, and compared with appendage anomalies
previously recorded in other Oxypoda species.

Instances of teratology (i.e., individuals exhibiting structural abnor-
malities) are not uncommon in many species of beetles (Coleoptera),
especially among artificially reared specimens (Crowson, 198 1). Teratol-
ogical malformations have been studied and categorized for the Coleop-
tera in the monographic works of Balazuc (1948, 1969).

Among members of the family Staphylinidae, the phenomenon of
teratology was reviewed in some detail by Frank (1981) who chronicled
all recorded morphological aberrations, including the presence of
supernumerary appendages, fusion or loss of appendages, anomalies of
body segmentation and malformations of the thorax and male genitalia.
Recently, Segers (1987) reported a case of triophthalmy and other teratol-
ogical aberrations in the Staphylinidae, anomalies previously unreported
for the family.

Because of a paucity of published information on teratological speci-
mens in the largest of the staphylinid subfamilies, the Aleocharinae, it
seems advisable to report on morphological abnormalities exhibited in
specimens taken under natural conditions. Therefore, in this note I
document a teratological specimen of the aleocharine Oxypoda opaca
(Gravenhorst), a Palearctic species recently reported for the first time
from North America (Hoebeke, 1989). This teratology, an antennal
malformation, is thoroughly described and further illustrated with scan-
ning electron photomicrographs. Additional literature on teratologies in
members of the genus Oxypoda is summarized.

A single New York specimen was found to have an extreme aberra-
tion of the left antenna, a type of meiomely (following the classification
and terminology of Balazuc, 1948). The antenna is 3-segmented; anten-
nomeres III-XI are apparently compactly fused, resulting in a clubbed
structure with no apparent segmentation (Figs. 1-3). The basal article
(scape) is somewhat stouter than that of the normal antenna, but article

Deceived August 7, 1989. Accepted December 14, 1989.

Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853

ENT. NEWS 101(3): 133-136, May & June, 1990

134

ENTOMOLOGICAL NEWS

II appears to be of nearly normal dimensions. Visible at high magnifica-
tion (540x), there are two large, craterlike structures, perhaps thermo- or
hygroreceptors, in juxtaposition on the dorsal surface (Fig. 3). In addi-
tion to being strongly setose, the surface of the "club" is covered with mic-
rosculpture consisting of an imbricate network of distinct microlines.
The overall length of the abnormal left antenna is 0.48 mm, compared to
1.17 mm for the normal right antenna.

The normal 1 1-segmented, right antenna is characterized as follows:
basal article (scape) and articles II and III elongated; articles IV-X sub-
quadrate to feebly transverse and not increasing much in length; and ter-
minal article (XI) elongate, pointed apically and equal to or slightly
longer than articles IX+X combined. The terminal article of the normal
antenna bears a pair of rather small, inconspicuous coeloconica-type
sensilla, probably thermo- or hygrosensitive in function. One sensillum
is located on the dorsal surface (Fig. 4, arrow) while the other occurs on
the opposite (ventral) side.

Figs. 1-4. Scanning electron photomicrographs of adult Oxypoda opaca. 1 , Frontal aspectof
head showing abnormal left antenna, 125x. 2, Closeup of abnormal left antenna, 208x. 3,
Enlargement of "club" of abnormal left antenna, 540x. 4, Terminal article (XI) of normal
antenna showing coeloconica-type sensillum (arrow) (dorsal surface), 350x.

Vol. 101, No. 3, May & June 1990 135

It is not possible to ascribe the antennal aberration described above
to any genetic mechanism (mutational) or to any other external factor
(i.e., extreme environmental conditions, injury, etc.) acting on an earlier
developmental stage such as the pupa.

Although the pair of sensilla on the apical segment of the normal
antenna (Fig. 4, minute, coeloconica-type, dorsal/ventral in position) are
very different from the pair of "sensilla-like" structures on the deformed
antenna (Fig. 3, extremely large, crater-like, dorsal/dorsal in position), I
surmise that these structures are probably homologous. Thus, it is the
author's opinion that the deformed antenna does indeed represent an
entire antenna and not a partial one, with the loss of intermediate or
apical segments.

The coated specimen is deposited in the Cornell University Insect
Collection. Complete collection data are as follows: NY: Tompkins Co.,
Town of Ulysses, N. of Jacksonville, 22 June 1988, E. R. Hoebeke. (male).

A search of the literature reveals 3 other references to teratology of
appendages in Oxypoda. Uhmann (1919) reported a specimen of O.
opaca, found near Dresden (E. Germany), with an abnormal right an-
tenna. The distal antennomeres, beyond article V, were shorter in length
and more compressed than the analogous articles of the normal left
antenna. Segmentation, however, was still evident in the abnormal an-
tenna. Keys (1936) documented a specimen of 0. opaca from New Forest
(S. England) with "four segments only to each of its anterior tarsi, where-
by its tarsal formula was 4-5-5 instead of the 5-5-5 which is proper to the
genus." Segers (1987) presented a case of symphysomely (the left antenna
showing 2 partly fused articles) in a normal female of O. brachyptera
(Stephens) collected from a pasture at Poeke (Belgium, 6-5-1982, trapped
in pitfalls).

ACKNOWLEDGMENTS

I am grateful to J. Howard Frank (University of Florida, Gainesville), James K. Liebherr
(Cornell University), and two anonymous reviewers for reading a draft of this paper and
providing helpful comments.

LITERATURE CITED

Balazuc, J. 1948. La teratologie des coleopteres et experiences de transplantation sur

Tenebrio molitor L. Mem. Mus. Nat. Hist. Natur., Paris (n.s.) 25:1-293.
Balazuc, J. 1969. Supplement a la teratologie des coleopteres. Redia 51:39-1 1 1.
Crowson, R. A. 1981. The biology of the Coleoptera. Academic Press Inc., London.

802 pp.
Frank, J. H. 198 1 . A re .dew of teratology in Staphylinidae, with description of a teratologi-

cal specimen ofTachinusaxillaris Erichson (Coleoptera, Staphylinidae, Tachyporinae)

from Florida. Florida Entomol. 64:337-340.

136 ENTOMOLOGICAL NEWS

Hoebeke, E. R., 1989. First record of the Palearctic species Oxypoda opaca (Gravenhorst)
from North America (Coleoptera: Staphylinidae: Aleocharinae). Jour. N.Y.
Entomol. Soc. 97:448-454.

Keys, J. H. 1936. Notes on variations in the tarsal segmentation of two staphylinid beetles.
Entomol. Mon. Mag. 72:151-153.

Segers, R. 1987. A case of triophthalmy and other teratological aberrations in Staphy-
linidae (Coleoptera). Bull. Ann. Soc. Roy. Beige Entomol. 123:179-184.

Uhmann, E. 1919. Zwei Staphyliniden mit abnormalen Bildunen (Col.). Entomol. Mitt.
8:214-216.

BOOK REVIEW

WORLD CROP PESTS. W. Helle, Editor in chief. 1989. Elsevier Publ.
Co., Amsterdam, The Netherlands, and P.O. Box 1663, Grand Central
Station, New York, NY 10163

VOL. 3A FRUIT FLIES: THEIR BIOLOGY, NATURAL ENEMIES,
AND CONTROL. AS. Robinson & G. Hooper, eds 1989. $161.

This volume was not reviewed. The four major parts deal with Taxonomy and Zoogeo-
graphy; Pest Status; Biology and Physiology; and Behavior.

VOL. 3B. FRUIT FLIES: THEIR BIOLOGY, NATURAL ENEMIES,
AND CONTROL. AS. Robinson & G. Hooper, eds. 1989.

This is the second of two volumes dealing with the frugivorous Tephritidae. It is divided
into 5 sections concerned with Genetics, Rearing, Population detection. Ecology and Con-
trol. Under these basic headings the 43 contributing authors address topics from the
specific ("ie. Cytoplasmic Incompatibility in Rhagoletis Cerasi") to the general (ie. "Life
History Strategies of Tephritid Fruit Flies"). The graphics are well presented and the book
attractively designed.

In addition to scope this volume has depth. The abundance of organized references
alone is invaluable to anyone used to dealing with the cosmopolitan and often scattered
fruit fly literature. Within minutes of opening the book, I had found interesting but
unfamiliar data and papers. However, the reader should be warned that there was con-
siderable delay between the writing of many of these chapters and publication so that there
are fewer recent citations than the 1989 publication date would suggest.

With that caveat excepted, I believe this book would be an important reference to any-
one with an interest in fruit flies. This includes not only those who are directly involved
with fruit fly control but also those interested in the more "basic" endeavors of Ecology,
Behavior, etc. I, for one, already find myself referring to one or the other of these volumes
several times a day.

JOHN M. SIVINSKI

USDA/ARS

Insect Attractants, Behavior, and

Basic Biology Laboratory

Gainesville, Florida

Vol. 101, No. 3, May & June 1990 137

A NEW SPECIES AND NEW RECORD OF THE

WATER-PENNY GENUS PSEPHENOPS

(COLEOPTERA: PSEPHENIDAE)

FROM COSTA RICA1

Paul J. Spangler2

ABSTRACT: Members of the genus Psephenops have not been recorded previously from
Costa Rica. A new species, Psephenops prestonae. from Costa Rica is described and com-
pared to Psephenops maculicollis Darlington (1936) described from Colombia and here
reported from Costa Rica. A diagnosis of Psephenops maculicollis is given and the distinc-
tive aedeagus of each species is illustrated.

The aquatic larvae of water-penny beetles are usually common
throughout the year in shallow streams with sand, gravel, and rocky sub-
strates and a moderately fast velocity. The riparian adult psephenids,
however, are short lived, elusive, and not commonly collected, especially
not in series. Consequently, a number of species of water-penny beetles
have been described from one or only a few adult specimens, as is this
new species. The single male of Psephenops prestonae, n. sp., was collected
by Warren E. Steiner and companions J.M. Hill & S.E. Frye.

I collected seven specimens of Psephenops maculicollis Darlington
( 1 936) in San Isidro, Costa Rica, in 1 967 but have not previously reported
on them.

These are the first species of water-penny beetles known from Costa
Rica and bring the number of water-pennies known from middle America
to eight.

Psephenops prestonae, new species

Figures 1. 2

Holotype cf — Form and size: Body flattened, oblong; thorax narrowed anteriorly;
elytra almost parallel sided, apices rounded. Length, 2.46 mm; greatest width, 1.26 mm.

Color: Covered with golden pubescence. Head black except genae behind eyes yellow
brown. Maxillary palpal segments dark reddish brown. Labial palpal segments lighter red-
dish brown than maxillary palpus. Ventral surface of head dark reddish brown. Antennal
segments all dark reddish brown. Pronotum black. Scutellum black. Elytron uniformly
reddish brown. Prosternum dark reddish brown; mesosternum. metasternum, and abdomen
blackish brown; hypomeron and epipleuron reddish brown. Coxae of all legs reddish
brown; tibia and tarsi of all legs dark reddish brown.

Head: Frontal area shallowly depressed; punctures coarse, separated by Vz to 1 times
puncture diameter. Clypeus almost on same plane as head. Labrum short, broad; shallowly
emarginate apically. Eyes prominent, hemispherical. Antenna densely pubescent; extend-

iReceived March 7. 1990. Accepted March 8. 1990.

"Department of Entomology, National Museum of Natural History, Smithsonian Institu-
tion. Washington, D.C. 20560

ENT. NEWS 101(3): 137-140. May & June, 1990

138 ENTOMOLOGICAL NEWS

ing almost to base of pronotum; basal segment enlarged, 1/3 wider than and twice as long as
segment 2; segment 3 about 1/3 longer than segment 4; segments 4 — lOsubequal; segment
11 slightly longer than segment 10. Maxillary palpus about 1/3 as long as antenna, 4
segmented; segment 1 shortest, about V* as long as segment 2; segment 2, a third longer than
segment 3; segment 3 about 3/4 as long as segment 4; segment 4 moderately flattened,
swollen, with an elongate sensory area on apex. Labial palpus very small, 3 segmented;
segments 1 and 2 short, stout, subequal in length; segment 3 much narrower and about \Vi
times as long as segment 2.

Thorax: Pronotum widest at base and about a third wider than long; discal area with
moderately coarse and fine punctures intermixed; coarse punctures separated by Vz to 2
times puncture diameter; fine punctures separated by 1 to 3 times puncture diameter; disc
moderately carinate on meson on posterior half and concave on each side of carina; base
strongly bisinuate; anterolateral angles moderately rounded; posterolateral angles sub-
acute; all borders narrowly rimmed. Scutellum subtriangular. Elytron widest slightly
behind midlength; with fine and coarse punctures intermixed; fine punctures separated by
Vz to 1 times puncture diameter; coarse punctures separated by 2 to 4 times puncture
diameter. Prosternum short in front of procoxae. Prosternal process long and narrow,
extending to midlength of mesocoxae, carinate on apex. Mesosternum with narrow, deep,
longitudinal sulcus. Metasternum swollen, incised and concave between metacoxae; mid-
line with narrow longitudinal sulcus on posterior two-thirds; shallowly and narrowly de-
pressed on each side of midline. Leg with femur robust; tibia slender, slightly arcuate and
slightly bent subapically. Protibia without distinct posterolateral denticle at apex. Tarsi
with segments 1—4 broad, with dorsal surface pubescent and ventral surface densely
pubescent. All legs with tarsal segments 1, 2, and 3 with large fiat pubescent lobes. Tarsal
claw moderately robust.

Abdomen: Sterna 5, 6, and 7 broadly subtruncate along posterior margins.

Genitalia: Trilobate and as illustrated (Figures 1 & 2).

Female. — Unknown.

Type data. — Holotype cT: COSTA RICA: CARTAGO PROVINCE: Pejibaye, 24 Mar
1987, W. E. Steiner, J.M. Hill, S.E. Frye; deposited in the U.S. National Museum of Natural
History, Smithsonian Institution.

Etymology: The specific epithet prestonae is named for Emily D. Preston, presently
Director of the Stony Brook Nature Center and Sanctuary in Norfolk, Massachusetts, in
appreciation of her strong interest in conservation activities.

The completely rimmed pronotum, the protibia without an apical
posterolateral denticle, and the distinctive aedeagus of Psephenops pres-
tonae (Figures 1, 2) will distinguish it from the similar P. maculicollis.

Psephenops maculicollis Darlington, 1936

Figures 3, 4

Diagnosis: Head black. Pronotum reddish yellow, darker across base; with large, round
shining, piceous mark on disc extending from apex to a point 2/5 from base and extending
as a fine line on carina on meson almost to base; narrowly rimmed laterally and pos-
teriorly. Scutellum reddish brown. Protibia with distinct, apical, posterolateral denticle.
Aedeagus as illustrated (Figures 3, 4).

Specimens examined: COSTARICA: SAN JOSE: San Isidro(l mi S), Rio General, 1
July 1967, Paul J. Spangler, 6 tf, 1 9; deposited in the National Museum of Natural History,
Smithsonian Institution.

Vol. 101, No. 3, May & June 1990

139

/ •> *••. .< v.

. ifjt {-,'«/ *» -—

"*• X S ."'A '"'» ~

'^ ^CY^"'

o.lmm

0 imm

Figures 1 and 2. PSephenops prestonae, new species, holotype. aedeagus. 1. ventral view: 2.
lateral view.

Figures 3 and 4. Psephenops maculicollis Darlington, aedeagus. 3. ventral view; 4, lateral
view.

140 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

I thank the following individuals for their help in making this species known: Warren
E. Steiner, for collecting and making available this new species and many other aquatic
beetles for my research; Young T. Sohn, Smithsonian Institution Biological Illustrator, for
the line drawings; and Phyllis M. Spangler, for typing the manuscript

LITERATURE CITED

Darlington, PJ. 1936. A List of the West Indian Dryopidae (Coleoptera), with a New
Genus and Eight New Species, Including one from Colombia. Psyche 43(2-3):65-83, 1 1
figures.

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American Entomological Society
c/oAcademy of Natural Sciences
19th & Race Sts.
Philadelphia, PA 19103 (215)561-3978

Vol. 101, No. 3, May & June 1990 141

NEW RECORD AND RANGE EXTENSION FOR

RHYACOPHILA WALLOWA (TRICHOPTERA:

RHYACOPHILIDAE) FROM ROCKY MOUNTAIN

NATIONAL PARK, COLORADO1

Scott J. Herrmann

ABSTRACT: Rhyacophila wallowa is reported for the first time from Colorado. This record
represents a southeastern extension of the known range of the species. Current state dis-
tribution records of/?, wallowa include OR, CA, WA, MT, ID, WY, UT and CO.

Three adult males of Rhyacophila wallowa Denning were collected in
Rocky Mountain National Park (RMNP), Larimer County, Colorado,
on 3 August 1984. The specific collection site in RMNP was at the con-
fluence of Fall River and Chiquita Creek at an altitude of 2640 m amsl
(8660 ft amsl) in T5N, R74W, Sll on the east side of the Continental
Divide. The three adults were collected by the author with a standard
sweep net at sunset. This is the first reported record of R. wallowa
from Colorado.

Denning's (1956) holotype male was collected 15 July 1949 from the
Wallowa River, Wallowa County, Oregon. Anderson (1976) and Wold
(1974) reported several other Oregon sites in addition to some from
Washington, Idaho, northern California and northern Wyoming.
Wold's (1974) Wyoming record is a single metamorphotype male collected
from the Gardner River, Yellowstone National Park, 12 July 1961.
Baumann and Unzicker ( 198 1 ) included R. wallowa in their checklist of
Trichoptera of Utah, as did Newell and Potter (1973) and Roemhild
(1982) of Montana. In Montana Roemhild (1982) commonly found this
species in cold alpine streams east and west of the Continental Divide.
The first published record from Colorado represents a southeastern
extension of the known range of R. wallowa: it also represents the six-
teenth species of Rhyacophila to be reported from Colorado (Herrmann.
Ruiter and Unzicker 1986). Other species of Trichoptera collected simul-
taneously with R. wallowa at the Fall River (RMNP) site included: Rhya-
cophila hyalinata Banks, Rhyacophila harmstoni Ross, Ecclisomyia
maculosa Banks, Oligophlebodes minuta (Banks), Onocosmoecus unicolor
(Banks), Agrypnia deflata (Milne) and Hydropsyche oslari Banks. Ruiter
and Lavigne (1985) stated that little is known about the habits of this
species. At the RMNP collection site Fall River is classified as a cold,
lower montane, second-order stream and Chiquita Creek as a cold.

'Received November 13, 1989. Accepted January 2, 1990

-Department of Life Sciences, University of Southern Colorado. Pueblo, CO 81001-4901.

ENT. NEWS 101(3): 141-142. May & June. 1990

142 ENTOMOLOGICAL NEWS

lower montane, first-order stream.

ACKNOWLEDGMENTS

I am grateful to Jay H. Linam and James E. Sublette for prepublication reviews; to
David R. Stevens for authority to collect Trichoptera in Rocky Mountain National Park
and to Joan M. Herrmann for assistance and patience with field collections.

LITERATURE CITED

Anderson, N.H. 1976. The distribution and biology of Oregon Trichoptera. Agr. Exp. Sta.

Tech. Bull. 134. Oreg. St Univ., Corvallis.
Baumann, R.W. and J.D. Unzicker. 1981. Preliminary checklist of Utah caddisflies

(Trichoptera). Encyclia 58: 25-29.
Denning, D.G. 1956. Several new species of western Trichoptera. Pan-Pac. Entomol.

32: 73-80.
Herrmann, SJ., D.E. Ruiter and J.D. Unzicker. 1986. Distribution and records of

Colorado Trichoptera. Southwest. Natur. 31: 421-457.

Newell, R.L. and D.S. Potter. 1973. Distribution of some Montana caddisflies (Trichop-
tera). Proc. Mont. Acad. Sci. 33: 12-21.
Roemhi Id, G. 1982. The Trichoptera of Montana with distributional and ecological notes.

Northwest Sci. 56: 8-13.
Ruiter, D.E. and R J. Lavigne. 1985. Distribution of Wyoming Trichoptera. Agr. Exp. Sta.

Publ. SM 47. Univ. Wyo., Laramie.
Wold, J.L. 1974. Systematics of the genus Rhyacophila (Trichoptera: Rhyacophilidae) in

western North America with special reference to the immature stages. M.S. Thesis,

Oreg. St. Univ., Corvallis.

Vol. 101, No. 3, May & June 1990 143

HPLABEL: A PROGRAM AND MICROFONT FOR

THE GENERATION OF DATE/LOCALITY LABELS

USING A LASER PRINTER1

D. Christopher Darling2, R. C. Plowright3

ABSTRACT: A computer program for the direct generation of date/locality labels for
pinned insect specimens is described. These labels are compared with those produced by
offset printing and by reduced photocopying.

One of the major bottlenecks in the curation of insect collections is
the generation of date/locality labels for specimens. For pinned and
pointed specimens, these labels have to be small (approximately 25
characters per inch), permanent, and legible. Most insect collections
now use a two-step process to produce labels. First, labels are printed
either with a typewriter or a printer in a standard font (usually 10-12
characters/inch) and a paste-up board is assembled. A "stat" is prepared
by photographically reducing the board to produce a master of the
required size when the paste-up board is filled. Labels are then offset
printed onto high quality paper (i.e. high rag content, acid neutralized).
These labels are of excellent quality (Fig. Id) but labels cannot be
efficiently generated until a paste-up board is filled, which can result in a
delay of many months in the preparation of the final labels. Some collec-
tions produce labels only 2 or 3 times a year. In addition, offset printing
usually has a minimum run of at least 50 copies per original, resulting in
many wasted labels for small lots of specimens.

These constraints on the efficient generation of date/locality labels
have no doubt resulted in the loss of a great deal of valuable ecological
and distribution data. Very general labels are often used, giving only a
minimum of information, e.g. U.S.A., OR: Benton Co., June 1988, D.C.
Darling. Often much more detailed information is contained in field
notes, but these data often fail to become associated with specimens
because of inefficiencies in label production, and are ultimately lost.
One solution is to produce "generic" labels for collecting localities and to
add a second label with more specific ecological information, such as
floral associations, hosts, or collecting method. There are at least two
problems with this approach: double labelling is time consuming and
affords an additional opportunity for mislabelling; and insect speci-

^Received August 14, 1989. Accepted November 25, 1989.

~ Department of Entomology, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario.

Canada, M5S 2C6 and Department of Zoology, University of Toronto, Toronto, Ontario.

Canada, M5S 1A1
^Department of Zoology, University of Toronto, Toronto, Ontario. Canada. MS5 1A1

ENT. NEWS 101(3): 143-146, May & June, 1990

144

ENTOMOLOGICAL NEWS

mens can quickly become cluttered with labels.

Many collections and collectors have experimented with producing
date/locality labels with reducing photocopy machines. When printed
on high quality paper these labels have the required permanence and
can be produced quickly, but usually with considerable wastage of
paper. Two reductions of 64% are necessary to produce labels of the cor-
rect size and the final labels lack the clarity of offset printed labels (Fig.
lb,c). These labels are generally regarded as unacceptable by major
museums as a standard curatorial procedure.

Computer programs that have been published for generating insect
labels on a microcomputer are either very basic (Kissinger 1982) or
primarly concerned with the mechanics of generating multiple copies of
labels and storing and managing files of labels (Ellis et al. 1985). These
programs streamline the production of output suitable for assembling
paste-up boards but do not generate labels directly. Labels must still be
reduced for offset printing or with a photocopy machine.

Laser printing technology is ideally suited to the production of labels

INDIA: Tamil Nadu, 29

km S Ootacamund

Nilqiris, 1 100 m.

Oct. 7, 1985

DC Darling, NF Johnson

INDIA: Tamil Nadu, 29

km S Ootacamuod

Nilgim, 1100 m

Od 7, 1985

DC Darling, NF Johnson

MCM

INDIA: Tamil Nadu, 29

km S Ootacamund

Nl lair Is, 11 00 m.

Oct. 7, 198^

DC Darling VF Johnson

INDIA: Tawi I Nadu, 29

km S Ootacamund

Ni 1911- is, 1 100 m.

Oct. 7, 1985

DC Dar I ing, NF Johnson

COSTA RICA, Puntarenas:
ca.8 km NW San Vito
u.vJight. 11 FEB 1988
O8 50'Nx82 58'W. 2307 m
ROM#880007. B.Hubley

COSTA RICA, Punlarcnas
ca.8 km NW San Vito
uv.lighi 11 FEB 1988
O8*50iNx82*58'W. 2307 m
ROM#880007. B.Hubley

OOn» WCA. PWI
«.« m NMr Bar >
u.,fff 11 rFP KM
CM fc-voi M-w no? >

COSTA RICA, Puntarenas

ca.8 km NW San Vito
u.v.l ight . 11 FEB 1988
08"50'Nx82058'W. 2307 m
ROM»880007. B.Hubley

COSIft RICft, Puntarcnat:
ca.8 km NU San Vito
u. v. I ight . 1 1 FEB 1988
Oe°f.O'N«8?°58'U. 2307 m
ROn*88OOO7. B.Hubley

Figure 1 : A comparison of date/locality labels, a, laser printer with smallest commercially-
available font, 6 point Helvetica, b, HP laser printer (10 point Times Roman) and two
photocopy reductions of 64% (40% of original) with Kodak Ektaprint 1 50. c, laser printer (6
point Helvetica) and two photocopy reductions of 64% (40% of original) with Kodak Ekta-
print 150. d, offset printing, e, HPLABEL. Scale line, 1 cm.

Vol. 101, No. 3, May & June 1990 145

for pinned or pointed specimens. The printing process results in extremely
high quality labels which should have excellent long term permanence
under normal storage conditions. There is, however, some concern about
the permanence of laser printed labels for specimens stored in alcohol or
when labels are exposed to various reagents and high temperatures [see
Insect Collection News, 2(2):26-27 (1989)]. Microcomputers and laser
printers (LaserJet series II) are quickly becoming a standard feature of
most museum and university departments. Unfortunately, the smallest
available fonts (6 point) are too large to directly generate date/locality
labels (Fig. la). Alternatively, commercially-available desktop publish-
ing programs are available to generate labels directly. These tend to be
rather cumbersome, requiring considerable word processing skills, and
cannot be incorporated easily into programs for generating multiple
copies of individual labels. For example, Fancy Font can generate high
quality 3 point labels. We present here an edited microfont and computer
program (HPLABEL) for the direct production of date/locality labels
using an IBM-PC (or compatible) microcomputer and Hewlett-Packard
LaserJet Series II printer.

The microfont is an edited version of a public domain font supplied
to us by Henry Spencer (Department of Zoology, University of Toronto).
A BASIC program was written by CP to streamline the generation of
labels. The program uses either BASICA or GWBASIC and prompts the
user for up to 5 lines of label data. After previewing the label, the user is
asked how many copies of the label are required. The program then
prompts for additional labels. The labels are stored in a buffer and printed
to efficiently use label paper.

Figure 1 compares labels produced by HPLABEL (Fig. le) with stan-
dard labels generated by offset printing (Fig. Id) and with labels generated
by a reducing photocopy machine (Fig. le). Although not as sharp as
offset printing, we think that the HPLABELS are acceptable for the
routine labelling of specimens. These labels are clearly superior to those
produced by a 40% reduction of either 10 point (Fig. Ib) or 6-point fonts
(Fig. le). With these reduced labels the letters run together making the
labels difficult to read and the labels are either too large (Fig. Ib) or too
small and illegible (Fig. le).

This program is currently being used for label production in the
Department of Entomology, Royal Ontario Museum. A copy of the font,
program and operating instructions is available by sending a formatted
5.25 inch diskette to DCD.

ACKNOWLEDGMENTS

We thank Henry Spencer (Department of Zoology, University of Toronto) for supply-

146 ENTOMOLOGICAL NEWS

ing the initial font, Allma Edwards (Department of Entomology, American Museum of
Natural History) for bringing Fancy Font to our attention, and David Grimaldi and James
Woolley for comments on the manuscript.

LITERATURE CITED

Ellis, K. A., Surgeoner, G. A. and Ellis, C. R. 1985. Versatile program for generating
insect labels on an IBM-PC microcomputer. Can. Ent. 117: 1447-1448.

Kissinger, D. G. 1982. Insect label production using a personal computer. Proc. Ent. Soc.
Wash. 84: 855-857.

BOOK REVIEW

WORLD CROP PESTS. W. Helle, Editor in chief. 1989. Elsevier Publ.
Co., Amsterdam, The Netherlands, and P.O. Box 1663, Grand Central
Station, New York, NY 10163

VOL. 2. APHIDS: THEIR BIOLOGY, NATURAL ENEMIES, AND
CONTROL. AIC Minks & P. Harrewijn, eds. 1 989. Vol. 2A 450 pp., $ 1 76;
Vol. 2B 382 pp., $169; Vol 2C 322 pp., $145.

Vol. 2A contains chapters on morphology and systematics; anatomy and physiology;
reproduction, cytogenics, and development; biology; aphids and their environment; evol-
ution; and structures of population and species.

2B includes two long chapters on techniques and natural enemies.

2C also contains just two chapters: Damage by aphids and control of aphids. The latter
includes, among the nine subjects, biological control, host plant resistance, and behavioral
modification. The black & white photographs, tables and graphs are not numerous, but are
of high quality. Comprehensive lists of references follow each chapter. Slight deficiencies
in coverage of North American papers are more than compensated for by the inclusion of
European papers that are rarely seen here.

The series title implies that these large-format (8 1/2" x 12") books may only be of
interest to economic entomologists. However, a broad range of subjects, by a large number
of world authorities, is included. The series will be useful reference to workers in many sub-
disciplines of entomology, in addition to those concerned with arthropod control.

The sample (Vol. 2C) that I received contains up-to-date reviews and syntheses of the
vast world literature. The scope and depth are most ambitious. The cost is not as high as it
appears because the large format includes approximately 30% more information per page
than the average (6" x 9") book.

Volumes 1 (Spider mites) has also been published, and several others are in prep-
aration.

- William H. Day,
Beneficial Insects/Lab., USDA
Newark, DE

Vol. 101, No. 3, May & June 1990 147

MICROCYLLOEPUS FORMICOIDEUS

(COLEOPTERA: ELMIDAE), A NEW RIFFLE

BEETLE FROM DEATH VALLEY NATIONAL

MONUMENT, CALIFORNIA*

William D. Shepard2

ABSTRACT: Microcylloepus formicoideus sp. nov. is described from Travertine Springs,
Inyo Co., California. A survey of local permanent springs indicates that the species only
exists at the type locality. Relationships with other congeners is discussed.

Microcylloepus was erected in 1935 by Hinton for the sole species
Stenelmis pusillus LeConte. Since then, 24 other species have been added
to the genus (Brown 198 1 ). Microcylloepus is essentially a Neotropical ele-
ment that has invaded the Nearctic. Within the US occur five described
species and perhaps several undescribed species. The eastern M. pusillus
has had four subspecies described but they represent only color morphs.
Populations have been found that have all the morphs co-occuring. In
the western Nearctic several species have been taken, particularly from
warm springs in the Basin and Range desert. Also in the western US
occurs M. similis Hinton which has long been considered to be a sub-
species of M pusillus. Although it resembles M. pusillus in many aspects,
it is a valid species, as is seen with a side-to-side comparison and when
examining the genitalia.

In the course of a survey of the riffle beetles of Death Valley two new
species were discovered. One is described here; the second presents a
more complicated problem requiring more analysis and will be des-
cribed later.

Microcylloepus formicoideus new species

Body: Body elongate, parallel-sided (Fig. 1); 1.29-1.53 mm in length. 0.50-0.56 mm in
width. Surface coarsely punctate and coarsely asperate; sculpturing very pronounced.
Dorsum, head and hypomera black; sterna and legs rufous; antennae and palpi testaceous.
Legs projecting well beyond sides of body.

Head: Head withdrawn into prothorax up to eyes. Dorsum densely granulate and
punctate; setae thick, arcuate, decumbent and separated by approximately half their
length. Genae and postocular areas smooth; setation fine, dense and decumbent. Frons
with fronto-clypeal suture almost straight; angles slightly obtuse, raised, and continuing
around base of antennae. Clypeus with apex broadly arcuate; angles broadly rounded.
Labrum with apex straight, sides diverging slightly to base; surface shiny and alutaceous;
setation sparse, fine and straight on disc and apically dense, coarse and curved. Mandibles

'Received October 30, 1989. Accepted March 7, 1990.

2Department of Biological Sciences, California State University at Sacramento.
Sacramento. CA 958 19.

ENT. NEWS 101(3): 147-153, May & June, 1990

148 ENTOMOLOGICAL NEWS

with 3 short teeth; prostheca present; lateral lobe large. Maxillae with 4-segmented palp,
last segment 2.5 times as long as wide, segments 1-3 subequal and half the length of the last;
galea digitiform, apically with several stout curved setae; lacinia elongate-rectangular,
fused to stipes, medially with several stout setae, apically with many stout setae with curved
tips forming a multitiered brush; cardo subrectangular and divided by an oblique suture.
Labium with 3-segmented palpi borne on short palpigers; mentum and submentum with
numerous setae; prementum reduced to palpiger; ligula apically with 20-25 cultriform
setae. Antennae 11 -segmented with pedical and scape longer and wider than following
segments except for eleventh which is twice as long as penultimate and cultriform.

Pronotum: Surface densely granulate and punctate, granules separated by own width,
punctae smaller and contiguous. Length slightly greater than greatest width. Sides bi-
sinuate, explanate and convergent both apically and basally; margins dentate; apices pro-
jecting beyond anterior margin. Disc in basal half with Y-shaped carina connecting to
sublateral carinae; moderately convex in apical half; median longitudinal depression in
middle half with length 3 times width. Sublateral carinae broken at one-half; basal pieces
apically swollen.

Scutellum: Shape ovoid, widest near base; surface flat and granulate.

Elytra: Surface densely granulate and punctate; setation as on dorsum of head. Striae
with punctae basally very large and nearly confluent, becoming smaller apically but
extending almost to apex. Second, fourth and sixth intervals flat, almost obliterated basally
by large punctae. First intervals slightly raised. Third, fifth and seventh intervals carinate,
seventh interval carinae reaching almost to apex, third interval carinae reaching just past
one-half, fifth interval carinae intermediate in length. Fifth and seventh intervals joined at
humeri which are only slightly wider than base of pronotum. Epipleura extending almost
to apex.

Wings: Only short basal portions remain.

Prosternum: Surface granulate with widely spaced setae. Anterior margin straight
across middle with sides broadly rounded dorsally; margin projecting under head and
covering most of mouthparts. Anterior half of prosternum strongly directed ventrally. Pros-
ternal process depressed in middle and apically broadly rounded. Prosternal carinae prom-
inent, broadly V-shaped and forming raised margins of prosternal process. Hypomera
densely granulate and coarsely asperate. Episterna strongly directed dorsally.

Mesosternum: Surface as on prosternum. Disc depressed and pentagonal; median
longitudinal sulcus present on apical half; posterior margin raised and slightly arcuate.
Mesosternal carinae prominent and bordering mesocoxal cavities.

Metasternum: Surface and sculpturing as in mesosternum. Median longitudinal sul-
cus deeper than mesosternal sulcus, cleft-like and in apical half of metasternum. Posterior
margin arcuate. Metasternal carinae broadly divergent from base.

Legs: Legs long and narrow. Pro- and mesocoxae globular; metacoxae transverse. All
coxal cavities open. Femora and tibiae long, subequal to pronotal length. All surfaces
granulate, except for tarsomeres which have only scattered granules. Tarsi with two ventral
rows of short, coarse setae. Last tarsomere with ventral apex prolonged into a tooth about as
long as broad.

Abdomen: First sternite with prominent carinae strongly divergent in anterior half,
then parallel or slightly convergent to posterior margin; disc anteriorly depressed and slop-
ing up to posterior margin of sternite, asperate and with scattered large granules; sides
directed dorsally. Second, third and fourth sternites similar; medially asperate with scat-
tered large granules, number of granules declining posteriorly. Fifth sternite with only a
few granules medially; lateral margins produced into teeth which clasp epipleura.

Genitalia: Male with median lobe constricted medially and broadly attenuated
apically; parameres almost parallel-sided, apices curving under median lobe (Fig. 2).
Female with sinuate baculi; hemisternites as long as broad; styli relatively short and curved
(Fig. 3).

Vol. 101, No. 3, May & June 1990

149

Figs. 1-3. Microcylloepus formicoideus sp. nov. 1. adult, dorsal view. 2, male genitalia. 3.
female genitalia.

150 ENTOMOLOGICAL NEWS

Tomentum: Occurs on genae, femora, medial surface of distal half of tibiae, pro- and
mesepimera, lateral portions of meso- and metacoxae, and lateral portions of all ab-
dominal sternites.

DIAGNOSIS

M. formicoideus is readily distinguished from all other North and
Central American congeners by its small size and relatively long legs
(these characters give the ant-like appearance that suggested its name).
Also very diagnostic is the lack of distinct elytral humeri, a result of
brachyptery (H. P. Brown, pers. comm.). This character is shared with M.
angustus Hinton from Mexico (Hinton 1940). The latter is somewhat
larger and possesses a distinctly different aedeagus. The male genitalia
of M formicoideus have parameres resembling those of M thermarum
Darlington while the median lobe is similar to that of M. inaequalis
(Sharp) and M. moapus La Rivers. However, the genital characters in-
volve subtle differences and Hinton (1940) notes that " a number of
species may have the structure of the male genitalia identical."

TYPE LOCALITY

The type locality is: CA, Inyo Co., Death Valley National Monument
(DVNM), 2.5 miles (4 km) east of Death Valley (along US Highway 190),
Travertine Spring. The spring heads are numerous and their outflows
combine while flowing southward toward the highway. Upon reaching
the highway all flow is channeled into a concrete-lined canal that crosses
under the highway and parallels it westward toward Furnace Creek Inn,
the local hotel. M. formicoideus occurs in the spring heads and along the
stream course almost to the road. Initially M. formicoideus occurs alone in
the springhead but it is eventually replaced along the stream course by
another congener.

An extensive survey of other DVNM water sources showed this to be
the only locality where M. formicoideus occurs. The benthic community
along the spring outflows is remarkably diverse and may represent the
most diverse aquatic insect fauna in DVNM. This area and community
certainly deserve protection and preservation. Texas Spring (which is
closer to the hotel, campgrounds, date-palm orchard and visitors' center)
has been completely diverted to human water uses. Whatever com-
munity was there is now lost. Because of the proximity of the two springs,
and the great likelihood that their outflows were contiguous, their com-
munities were probably the same. Thus the benthic community in Traver-
tine Spring probably represents the only remaining portion of a much
larger community.

Vol. 101, No. 3, May & June 1990 151

TYPES

Holotype male, allotype female and 37 para types collected 23/1/1984
by WDS from Furnace Creek canal, a concrete canal diverting water to
Furnance Creek Wash from Travertine Springs. Additional paratypes
from the type locality include 53 collected by WDS on 16/IV/1984, 134
collected by Hugh Leech on 25/XII/1962, and 26 collected by Raymond
Bandar on 25/XII/1962. The holotype, allotype and several paratypes
will be deposited in the National Museum of Natural History at the
Smithsonian Institution. Additional paratypes will be deposited in the
collections of Harley P. Brown (Norman, OK), Louisiana State Univer-
sity (Baton Rouge, LA), William D. Shepard (Sacramento, CA), Death
Valley National Monument museum (Death Valley, CA), the California
Department of Food and Agriculture (Sacramento, CA) and Monte L.
Bean Museum at Brigham Young University (Provo, UT). The Leech
and Bander paratypes are in the collection of the California Academy of
Sciences (San Francisco, CA).

ETYMOLOGY

The name M. formicoideus is chosen to note the slender body and long
legs which give individuals an ant-like appearance.

DISCUSSION

In Brown (1972) this species keys to couplet #70 which separates M.
browni (Hatch) from M. moapus. M. formicoideus can be distinguished
from M. browni by its smaller length and width. M. formicoideus can be
distinguished from M. moapus by several characters: smaller size; black
versus brown dorsal color; elytra with more pronounced carinae and
punctae, surface more asperate and less shiny; anterior portion of pros-
ternum more strongly directed ventrally. In Hinton's (1940) work on
Mexican e\mids,M. formicoideus keys to M angustus from which it can be
distinguished by several characters: general surface densely granulate;
longer carina on third intervals; sides of prosternal process not parallel;
metasternal carinae strongly divergent; first abdominal sternite with
carinae parallel posteriorly.

M. thermarum is a closely related species from which Microcylloepus
formicoideus varies in its more pronounced elytral sculpturing. In A/, ther-
marum the third elytral intervals are only slightly prominent, the fifth
intervals are flat Lnd the seventh intervals have "fine, inconspicuous cos-
tae" (Darlington 1928).

152 ENTOMOLOGICAL NEWS

This new species, with its putative close relationship with M. moapus,
M. thermarum, and M. angustus, recalls La Rivers' (1949) prediction of
intervening forms indicating all to be just intergrading populations of
one species. While close examination reveals character differences equi-
valent to those between other species oiMicrocylloepus, one is led to won-
der about the effects of the similar habitats in which these species live.
Hinton (1940) doesn't mention the habitat forM angustus, but all the rest
come from warm springs. The relatively uniform warm temperatures
may well alter developmental pathways leaving morphological varia-
tion canalized. Sweeney (1984) indicates that higher tempertures may
maximize larval developmental rates leading to smaller than normal
adults. Another species inhabiting warm springs in the Owens Valley of
California has lost temporal synchronization of pupation (WDS, un-
published data). Alternatively, one may assume that this species group
represents a different lineage from that including M. pusillus, which
exists throughout the eastern half of the US. The genus is sorely in need
of revision. Only then can questions of the source of variation in Microcyl-
loepus be properly answered.

ACKNOWLEDGMENTS

Hugh Leech kindly loaned specimens from his collection and copies of portions of his
personal field notes. Both Hugh Leech and Harley P. Brown provided copies of their cor-
respondence regarding this species. The California Academy of Sciences provided space
and equipment to study their specimens. The staff of Death Valley National Monument
provided access to collecting sites, much information and other valuable assistance with-
out which this work could not have been accomplished. Part of this study was accom-
plished during a summer research position at the University of Oklahoma. I thank H. P.
Brown and C. B. Barr for reviewing the manuscript.

LITERATURE CITED

Brown, H. P. 1972. Aquatic Dryopoid Beetles (Coleoptera) of the United States. Biota of

Freshwater Ecosystems Identification Manual No. 6. U. S. Environmental Protection

Agency. U. S. Government Printing Office, Washington. 82 pp.
1981. A distributional survey of the world genera of aquatic dryopoid beetles

(Coleoptera: Dryopidae, Elmidae, and Psephenidae sens. lot.). Pan-Pac. Entomol.,

57(1): 1-6.

Darlington, P. J. 1928. New Coleoptera from western hot springs. Psyche, 35(1): 1-6.
Hinton, H. E. 1935. Notes on the Dryopoidea (Coleoptera). Stylops, 4(8): 169-179.
. 1940. A monographic revision of the Mexican water beetles of the family

Elmidae. Novit. zoolog., 42:217-396.
Horn, G. H. 1870. Synopsis of the Parnidae of the United States. Trans. Amer. Entomol.

Soc., 3:29-42.
La Rivers, 1. 1949. A new species ofMicrocylloepus from Nevada (Coleoptera: Dryopidae).

Entomol. News, 60(8):205-209.
Le Conte, J. L. 1852. Synopsis of the Parnidae of the United States. Proc. Acad. Nat. Sci.

Phil., 6:41-45.

Vol. 101, No. 3, May & June W90 153

Sharp, D. 1882. Insecta, Coleoptera, Haliplidae, Dytiscidae, Gyrinidae, Hydrophilidae.

Heteroceridae, Parnidae,Georissidae,Cyathoceridae. Biol.centrali-americana, 1(2): 1-

144.
Sweeney, B. W. 1984. Factors influencing life-history patterns of aquatic insects, pp 56-

100. In V. H. Resh and D. M Rosenberg (eds.) The Ecology of Aquatic Insects. Praeger.

New York. NY. 625 pp.

SOCIETY MEETING OF MARCH 28, 1990

EVOLUTION AND HYBRIDIZATION OF ADMIRAL BUTTERFLIES

Dr. Austin P. Platt, Speaker

Insect taxonomists beware! If all groups are as interesting and complex as the butterfly
genus Limenitis, most disputes between lumpers and splitters may never be resolved. Only
in recent years, more than a century after the principal morphological types were de-
scribed, has a reasonable understanding of the relationships among admiral butterflies
emerged. As Nobelist Arthur Kornberg once said, "I have yet to see a complicated problem
which, when looked at in the right way, doesn't become more complicated."

Dr. Austin P. Platt of the University of Maryland, Baltimore County, has studied the
genetics and speciation of admiral butterflies for more than two decades. These well known
and widely distributed butterflies include the viceroy, Limenitis archippus; the banded pur-
ple, L. arthemis arthemis; the red-spotted purple, L. a. astyanax; Weidemeyer's admiral, L.
weidemeyerii; and Lorquin's admiral, L. lorquini. The latter four are really allopatric races of
a single "super species" which, except for arthemis and styanax, rarely hybridize in the wild.
Subspecies of the viceroy, L. a. archippus and L. a.floridensis, are mimics respectively of the
monarch, Danaus plexippus, and the queen, D. gilippus. Similarly the red-spotted purple
and Lorquin's admiral are thought to be mimics of the pipevine swallowtail. Bonus
philenor, and the California sister, Adelpha bredowii. Based on cladistic analyses and con-
siderations of geographical distribution, all mimetic forms of Limenitis are probably
derived from an ancestral form resembling the northern banded purple.

The banded phenotype is widespread in related genera and is controlled by a single
autosomal gene. Genetic analyses of natural and laboratory hybrids between various
species and subspecies of Limenitis indicate that several genes modify the banding pattern.
Dr. Platt offered the intriguing hypothesis that the distinctive nonmimetic transvese black
band on the hind wing of the monarch-like viceroy represents the vestige of a dark-
margined white band that has collapsed as the result of modifying genes. The plausibility
of this hypothesis was strengthened by comparing the wing patterns of hybrids between the
viceroy and each of the members of the L. arthemis super species group that were displayed
by Dr. Platt (See Bull. Ent. Soc. Am. (1983) 29(3): 10 - 20).

In addition to a discussion of the evolutionary relationships within Limenitis, Dr. Platt
discussed their interesting life cycle. The eggs are commonly laid on willow or aspen leaves.
They look like minature geodesic domes. The larvae that hatch from them- establish
characteristic feeding stations. In response to photoperiod, halfgrown 3rd instar larvae
accumulate glycerol as a natural antifreeze and retreat to hibernacula in which they over-
winter. These leaf-enclosed structures are easy to recognize and can be collected for pop-
ulation studies.

Dr. Plan's talk at the University of Delaware was attended by seventeen members and
four guests.

- Harold B. White
Corresponding Secretary

154 ENTOMOLOGICAL NEWS

THE STATUS OF PISON DOGGONUM
(HYMENOPTERA: SPHECIDAE)1

Arnold S. Menke2

ABSTRACT: Pison doggonum, originally described from Mexico, is a member of the wes-
tern Pacific island fauna and may be a synonym of P. iridipenne.

Pison doggonum Menke was described in 1988 from a single female
specimen labelled simply "Mejico, Mus. Drews." It now appears that the
holotype had erroneous locality data. Recently while examining some
Pison from islands in the western Pacific Ocean I noted that specimens of
P. iridipenne Smith looked similar to doggonum. Pison iridipenne occurs in
Hawaii, Fiji, Samoa, Society Islands, Tuamotu Archipelago, Bolabola
Is., Marquesas, Bismarck Archipelago, and islands in Micronesia
(Krombein, 1949, Bohart and Menke, 1976, Tsuneki, 1982). I borrowed
the holotype of doggonum from the Zoologisk Museum in Copenhagen,
and compared it with material of iridipenne identified by Karl Krombein.
The two taxa appear identical and doggonum may be a junior synonym,
although Krombein (1949) suggests that iridipenne is possibly a complex
of species in which the males offer the best differences. For that reason I
think it would be premature to synonymize doggonum with iridipenne. In
any event, diggonum should be considered as a member of the western
Pacific insular fauna, and not the Neotropical Region.

The discovery just related underscores the fact that describing new
species from single, ancient specimens with meagre locality data is a
risky, unwise business.

LITERATURE CITED

Bohart, R. M. and A. S. Menke. 1976. Sphecid wasps of the world. Univ. Calif. Press,

Berkeley. 695 p.
Krombein, K. V. 1949. the Aculeate Hymenoptera of Micronesia. I. Scoliidae, Mutillidae,

Pompilidae and Sphecidae. Proc. Hawaiian Ent. Soc. 13:367-409.
Menke, A. S. 1988. Pison in the New World: a revision (Hymenoptera: Sphecidae: Trypox-

ylini). Contrib. Amer. Ent. Inst. 24(3):1-171.
Tsuneki K. 1982. Sphecidae collected by the Noona Dan Expedition to the Bismark and

Solomon Archipelagoes (Hymenoptera). Spec. Publ. Japan Hymen. Assoc. (19): 1-58.

'Received December 4, 1989. Accepted December 4, 1989.

Systematic Entomology Laboratory, Agricultural Research Service, USDA, % U.S.
National Museum, Washington DC 20560

ENT. NEWS 101(3): 154, May & June, 1990

Vol. 101, No. 3, May & June 1990 155

DISTRIBUTIONAL NOTES ON NORTH AND

CENTRAL AMERICAN DILARIDAE

(NEUROPTERA)1'2

Kevin M. Hoffman^

ABSTRACT: Nallachius americanus is recorded from South Carolina and central Georgia
and a male of Nallachius pulchellus is reported from Costa Rica. An emendation is made to
the existing key for New World Dilaridae to accomodate the presence of a forked costal
crossvein in each forewing of the N. pulchellus specimen.

The sixteen species of New World Dilaridae are rarely-collected
neuropterans of which only three are known from North and Central
America (Adams 1979, Penny \98\). Nallachius americanus (McLachlan)
has been recorded from Puerto Rico, Venezuela, and the eastern United
States (the District of Columbia, Florida, Georgia, Indiana, Kentucky,
Maryland, Michigan, Pennsylvania, Texas and Virginia) (Gurney 1947,
MacLeod and Spiegler 1961, Adams 1970, Lawson and McCafferty
1984); Nallachius championi (Navas) is known from only one locality in
Guatemala (Adams 1970); and Nallachius pulchellus (Banks) has been
recorded from Cuba and the southwestern United States (Arizona)
(Alayo 1968, Adams 1970). The present paper records N. americanus from
South Carolina and central Georgia, N. pulchellus from Costa Rica, and
provides an emendation to the existing key for New World Dilaridae to
accomodate the presence of a forked costal crossvein in each forewing of
the Costa Rican specimen of N. pulchellus. All specimens are deposited in
the Clemson University Arthropod Collection (CUAC), Department
of Entomology.

New United States records for Nallachius americanus

GEORGIA: Crawford County, approximately 5 miles SSE of Roberta at Spring Creek,
8.1X1983, UV light trap, S.W. Hamilton and R.W. Holzenthal, Icf; SOUTH CAROLINA:
Aiken County, Aiken State Park. 7.VII.1988, UV light trap, K. M. Hoffman. Icf; Kershaw
County, Spears Creek at U.S. Route 601, 2.VI.1988, UV light trap, K. M. Hoffman and J. D.
Spooner, Icf; Pickens County, Clemson University Experimental Forest surrounding Lake
Issaqueena, Wildcat Creek, elevation 235 m., 12-20. VI. 1988, Malaise trap. K. M. Hoffman,
Icf; same collection data except 27.VI.-4. VII. 1988, Icf.

The only previous record for Georgia was from Decatur County in
the extreme southwestern corner of the state, well within the Upper Coastal
Plain. The Crawford County locality extends the range of this species in

^Received September 21, 1989. Accepted October 23. 1989.

^Technical Contribution No. 2993 of the South Carolina Agricultural Experiment Station.

Clemson University.

Department of Entomology, Clemson University, Clemson. SC 29634-0365.

ENT. NEWS 101(3): 155-157. May & June. 1990

156 ENTOMOLOGICAL NEWS

Georgia to the Sandhills region near the center of the state. The South
Carolina localities in Aiken and Kershaw Counties are likewise within
the Sandhills, whereas the Pickens County locality is in the upper
Piedmont.

New record for Nallachius pulchellus

COSTA RICA: Guanacaste, Parque Nacional Guanacaste, Maritza, Rio Tempisquito,
10.958 N, 85.497 W. 19-20. VII.1987, elevation 550 m., at light. R. W. Holzenthal, J. C.
Morse, P. J. Clausen, Itf.

This record represents the first dilarid reported from Costa Rica and
confirms the prediction of Adams (1970) that this species would event-
ually be found in Central America. The specimen was identified by com-
paring the genitalia both with the description and figures of Adams
(1970) and with the cleared genitalia of the holotype in the Museum of
Comparative Zoology (MCZ), Harvard University. However, a problem
was encountered when using the key of Adams (1970) because the second
character used to distinguish N. pulchellus in couplet 6 is "costal veinlets
simple," and each forewing of the Costa Rican specimen has a distinctly
forked costal crossvein at half length. Furthermore, this crossvein is in
approximately the same location as a forked costal crossvein figured by
Alayo(1968, Fig. 10B) fora male TV. pulchellus from Cuba. In view of the
variation in this character, the phrase "costal veinlets simple" should be
deleted from couplet 6 in the key by Adams ( 1 970) and from couplet 8a in
the key by Penny (1981), which was modified from the key by Adams.
These deletions will not affect the second halves of these couplets and
will actually alleviate some confusion, because costal veinlets were not
used as characters in the second halves.

ACNOWLEDGMENTS

Gratitude is extended to Scott R. Shaw, MCZ, Harvard University, for the opportunity
to examine the holotype of A', pulchellus. I thank Albert W. Johnson, J. Ben Kissam, and
John C. Morse, Clemson University, for their helpful comments on this manuscript.
Examination of the holotype of N. americanus was made possible by a grant from the Ernst
Mayr Fund of the MCZ, and the South Carolina dilarids were collected during surveys fun-
ded by the South Carolina Heritage Trust Program and the U.S. Fish and Wildlife Service.
This support is gratefully acknowledged. The collection of N. pulchellus in Costa Rica was
conducted on National Science Foundation Grant BSR-8512368.

LITERATURE CITED

Adams, P.A. 1970. A review of the New World Dilaridae. Postilla 148: 1-30.
Alayo, D. 1968. Los Neuropteros de Cuba. Poeyana (Ser. B) 2: 5-127.
Gurney, A. B. 1947. Notes on Dilaridae and Berothidae, with special reference to the
immature stages of the Nearctic genera (Neuroptera). Psyche 54: 145-169.

Vol. 101. No. 3, May & June 1990 157

Lawson, H. R. and W. P. McCafferty. 1984. A checklist of Megaloptera and Neuroptera

(Planipennia) of Indiana. Great Lakes Entomol. 17: 129-131.
MacLeod, E. G. and P. E. Spiegler. 1961. Notes on the larval habitat and developmental

peculiarities of Nallachius americanus (McLachlan) (Neuroptera: Dilaridae). Proc.

Entomol. Soc. Wash. 63: 281-286.
Penny, N. D. 1981. Neuroptera of the Amazon Basin. Part 2. Dilaridae. Acta Amazonica

11: 383-390.

BOOK REVIEW

THE ANTS. Bert Holldobler and Edward O. Wilson. 1990. Belknap
Press of Harvard University. 732 pp. $65.

CONTENTS: Introduction; Classification (139 pp.) including keys, illustrations of world
genera; Colony formation and structure (36 pp.); Behavior (s.l.) (338 pp.); Communities ( 1 7
pp.); Symbiosis (120 pp.); Specialized predators, fungus growers, and harvesters (50 pp.);
Weaver ants (1 1 pp.); Study methods (4 pp.); Glossary, bibliography, index (83 pp.)

This is an overwhelming narrative of the lives of ants, crammed with facts and ideas
and glorious illustrations. There is no way that I can comment on specific facts and theories
in this book; there are simply too many of them, and one cannot review an elephant by
critically examining a few hairs. Suffice it to say that if you are a biologist you must have
this book for the sheer excitement of its brilliantly detailed revelation of a bizarre and
somehow symbolic world, like one of the busier visions of Hieronymus Bosch. If you are a
myrmecologist you will just have to get used to lugging around a 7-lb. volume, because it
will be difficult to study, or even think about ants, without this book at hand.

In an age of books that are edited hodgepodges of disparate authors, THE ANTS is uni-
que in both its exhaustive coverage and its cohesion. I wish I could say it was a practical
model for scientific writing, but the fact is that few of us are up to such a tour de force. Any-
body who has attempted to meld information from a dozen references into a succinct and
literate paragraph knows the difficulty of such a task; Holldobler and Wilson do this,
apparently effortlessly, for hundreds of pages, utilizing thousands of different references.
Enormous numbers of studies are briefly described, with the inclusion of exactly the
primary data one might need to draw one's own conclusion, even though the authors have
supplied their personal interpretations. In other words, the authors, experts though they
are, write for an audience presumed to have its own initiative and powers of deduction In
another eschewal of arrogance, the authors avoid sarcastic treatment or unceremonious
dumping of the less plausible theories that circulate through myrmecology. Throughout
the book there is an emphasis on fascinating unanswered questions, thereby providing
both foundation and direction for future work.

To write such a book, one must, at least temporarily, deliver oneself over to obsession.
Thus, though "communites" of ants and ant-plant symbioses are carefully analyzed, there
is relatively little coverage of the role of ants in entire ecological communities, and almost
an avoidance of the dreadful topic of ants as the principle fodder of a host of other animals,
including many vertebrates. The limitations of sociality are hardly explored, so after read-
ing the book one might wonder how there can be more than a million species of arthropods
not apotheosized into sociality. Still, it is an obsession that is never tedious, always creative.
Even the statement (Chapt. ) ) that humans and ants represent the summits of evolution in
vertebrates and arthropods respectively, manages to create a rather endearing new hybrid
bias: antopocentrism.

To see this book is to covet it, and the price is modest. THE ANTS is destined to become as
widely distributed as its extraordinary subjects.

158 ENTOMOLOGICAL NEWS

FIRST UNITED STATES RECORD OF
ASCALOBYAS (NEUROPTERA: ASCALAPHIDAE),
A RANGE EXTENSION FROM NORTHEASTERN

MEXICO TO TEXAS1

Roy C. Vogtsberger

ABSTRACT: A female specimen of Ascalobyas albistigma (Neuroptera: Ascalaphidae),
collected west of Juno in August, 1 973 from Val Verde Co., Texas, was discovered in the Mid-
western State University (Texas) insect collection. This is the first record of this genus in the
United States and extends its known range by approximately 575 miles (925 km.).

While identifying the Neuroptera in the Midwestern State University
insect collection, the author discovered a very large and peculiar owlfly
in the collection. The specimen was collected west of Juno in Val Verde
Co., Texas in August, 1973 by H. L. Horry, graduate biology student. The
specimen was probably caught with an aerial net, since the habitat on the
collection label read "air".

The Texas specimen was readily assigned to the ascalaphid sub-
family Haplogleniinae, since the eyes were entire and not divided by a
transverse sulcus. According to the literature, Ascaloptynx (or Neurop-
teryx) had previously been the only members of this subfamily known to
occur in the United States. The distinguishing feature of this genus in the
New World, besides the entire eyes, is the petiolate wings, with forewings
having a prominent thumb-like projection on the posterior margin near
the base of each wing. Penny (1981) placed Ascaloptynx in the tribe
Verticillecerini.

Since the specimen in question lacks petiolate wings and the promi-
nent thumb-like projections, it is assigned to the tribe Haplogleniini.
Penny ( 1 98 1 ) recognized three genera of American Haplogleniini: Amoea,
Ascalobyas, and Haploglenius. The Texas specimen was assigned io Asca-
lobyas on the basis of the antennae being shorter than half the length of
the forewing, costal field of forewing and apical fourth of all wings being
dark brown, and the yellowish-white pterostigmas. Penny ( 1 98 1 ) presented
Ascalobyas as a new genus because the old genus name, Byas Rambur,
1842 was preoccupied.

Weele's (1908) monograph with descriptions of the two known species
of Byas (= Ascalobyas) mentions the type species, microcerus Rambur, as
an eastern and Antillean species and a western species, albistigma
(Walker). He suggested they could be combined under one species,

^Received July 6, 1989. Accepted January 2, 1990.

2Department of Biology, Midwestern State University, Wichita Falls, Texas 76308.

ENT. NEWS 101(3): 158-160, May & June, 1990

Vol. 101, No. 3, May & June 1990 159

because the terminalia are superficially quite similar, and the extremely
small differences are never constant. However, he chose to maintain
them as two distinct species due to lack of transitional forms from the
intermediate region of the two species' distributions. Measurements of
the Texas specimen (Table 1) and color descriptions most nearly coin-
cide with those given by Weele (1908) for the western albistigma, which
would be logical for a Texas locality.

The northernmost published record for Ascalobyas albistigma
(Walker), 1853 is San Pedro Sula, Honduras (Weele, 1908). Penny (1981)
reported that "Ascalobyas is presently known from two species ranging
from northern Brazil to Panama," but mapped the distribution of Ascalo-
byas to extend northward to San Pedro Sula, Honduras. The statement,
therefore, was probably an error and should have read, ". . .from northern
Brazil to Honduras". This makes the Texas specimen a 1343 mile (2161
km.) range extension from the northernmost published record for the
species.

Oliver Flint (pers. comm., 1989), curator of Neuropteroids at the
National Museum of Natural History, Smithsonian Institution, reports
several examples of the species from well into northeastern Mexico in
the NMNH collection. The northernmost specimens are from El Salto
Falls on the San Luis Potosi-Tamaulipas border near El Naranjo and,
previous to the Texas specimen, were considered to represent the northern
range extent for the genus. Therefore, the Val Verde record is a northern
range extension of 'Ascalobyas albistigma by approximately 575 miles (925
km.) from El Salto Falls in northeastern Mexico to Val Verde Co., Texas, a
new record for the United States. The owlfly, damaged in transit to the
Smithsonian Institution as a dried specimen to be compared with the
Mexican specimens, has been relaxed and is nowpreserved in alcohol in
the Midwestern State University insect collection.

TABLE 1 - Length measurements (in mm.) of body, wings, and antennae of Ascalobyas
females.

Total body Forewing Hindwing Abdomen Antennae -Reference

microcerus
albistigma
Texas specimen

Weele 1908

36-38

42-44

38-43.5

19-20

Weele 1908

160 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

I wish to thank Oliver S. Flint, Jr. for his taxonomic expertise and valuable comments,
Elsa Galbraith for her excellent translation of Week's German monograph of the Ascala-
phidae (1908), Norman Horner, Fred Stangl, Jr., and anonymous reviewers for critically
reviewing earlier drafts of this manuscript, and Horner for making the Texas specimen of
Ascalobyas available for study.

LITERATURE CITED

Penny, N.D. 1981. Review of the generic level classification of the New World Ascal-

aphidae (Neuroptera). Acta Amazonica 11 (2):391-406.
Weele, H.W. van der. 1908. Ascalaphiden. Monographisch Bearbeitet. Coll. zool. Edm.

Selys Longchamps. Fasc. VIII. 326 pp., 254 figs., 2 pis.

SOCIETY MEETING OF APRIL 25, 1990

PATTERNS IN NATURE
Dr. Richard Weber, Speaker

Dr. Richard Weber, entomologist from the University of Delaware and eminent insect
photographer, treated his audience of 21 members and 30 guests to a unique experience.
Not only did he show them remarkable pictures of insects but he also regaled them with his
world view, quick wit, and playful humor. While many seek to learn and emulate Dick
Weber's technical skills, it was evident that technique, though important, was peripheral to
the real essence of his photography. Pretty pictures come to life when they tell a story or pro-
voke a question. As Albert Szent-Gygorgyi said, "See what others have seen, but think what
no one has ever thought." Dick Weber's success comes from photographing what he
thinks.

It was not possible to be passive in the audience because questions, banter, and repartee
were part of the show. "What's going on here? What good are cryptic or aposematic colora-
tion against night predators? Does anyone know the name of this plant? Has anybody seen
this? Can you imagine the smell I put up with photographing maggots on that rabbit car-
cass? Did you ever see a bird dropping that looked like an insect? Look at that, a fly eating a
wasp!" These were part of the barrage of illustrated, thought-provoking comments and
anecdotes from an evening that was memorable for other reasons as well.

Prior to Dr. Weber's talk. President Roger Fuester presented the annual Calvert Prize to
Jennifer Reynolds for her science fair project, "Does the color of light affect the reproduc-
tion of Drosophila melanogasterl" She and the first and second runners up, Kimberly
Wallace and Chaitanya Rao, displayed their insect-related projects. Margot Livingston,
the first Calvert Prize recipient in 1987, was present for the ceremony.

In addition to the Calvert Prize, special certificates of appreciation were presented to
unsuspecting members of the Society for their great commitment and continuing con-
tributions to the Society. Howard P. Boyd, president from 1977-1981 and editor of Entomo-
logical News since 1974, described his association with AES as one of the most rewarding
and enjoyable activities of his life. Mildred Morgan, office secretary since 1979, also
received a certificate of appreciation. She confided that she almost quit after three weeks
but that Howard Boyd convinced her to stick it out a "bit longer." Both expressed their
heartfelt thanks to the Society for the honor. Jesse Freese, treasurer since 1969, was also
honored but was unable to attend the meeting.

(Continued on page 163)

Vol. 101, No. 3, May & June 1990 161

BIOLOGICAL NOTES ON DROSOPHILA

GUTTIFERA (DIPTERA: DROSOPHILIDAE),

A CONSUMER OF MUSHROOMS1

Britt Bunyard, B. A. Foote2

ABSTRACT: Information is given on the life history and larval feeding habits ofDrosophila
guttifera, a species previously considered to be strictly mycophagous. Laboratory rearings
indicate that larvae of this species can be polyphagous.

Patterson and Stone (1952) listed Drosophila guttifera Walker, a wide
ranging species in the eastern and midwestern states, as "being so rare as
to have nothing known about its life cycle." However, they felt that it was
probably mycophagous.

The purposes of the present study were to elucidate the life history of
D. guttifera, describe its larval feeding habits, and to determine whether
the species is truly mycophagous.

MATERIALS AND METHODS

Adult flies were reared from larvae occurring in sporophores of gill-
bearing mushrooms collected in an old field that is part of the Kent State
University campus in Portage County, Ohio. Sporophores were placed
in breeding containers containing a layer of moistened peat moss to
allow emergence of adults. To establish a laboratory colony, adults
emerging from the field-collected mushrooms were transferred to breed-
ing jars containing pieces of the commercial mushroom Agaricus bis-
porus Singer. Adults emerging from commercial mushrooms were then
placed in vials containing Instant Drosophila Medium (Carolina Bio-
logical Supply) and allowed to cycle. Subsequent rearings utilized such
"unnatural" foods as decaying lettuce, bananas, and tomato juice.

We determined oviposition preference by offering females different
food substrates and recording the number of eggs deposited on each sub-
strate. For this study, an isoline was developed by allowing a virgin
female to mate with only one male. All adults used in the preference tests
were of this isoline reared on Instant Drosophila Medium.

The substrate was nutrient agar to which was added (while still
liquid) homogenized aliquots of four different food sources: commercial
mushroom, iceberg lettuce, tomato juice, and banana. Each of the liquid
mixtures of food and agar was poured into a sterile petri plate and

Received November 1, 1989. Accepted January 5, 1990.
"Department of Biological Sciences, Kent State University, Kent, Ohio 44242

ENT. NEWS 101(3): 161-163. May & June. 1990

162 ENTOMOLOGICAL NEWS

allowed to solidify. The ratio of sustrate to agar was 10: 1 as measured by
volume. Plates of pure agar were also poured to the same heights as those
with food. Once solidified, four 2.0 cm diameter discs were cut from each
of the agar plates. A disc taken from each of the food/agar plates were
placed in holes of corresponding diameter and depth in the plain agar
plates. As a control, discs of pure agar were used. Ten gravid, non-
anesthetized females were then added to all test plates and allowed to
oviposit for 24 hrs. at 27° C. Adults were then removed, and the number
of eggs occuring on each substrate counted. Each test was replicated five
times. Females were not used in subsequent tests.
Voucher specimens have been deposited in AMNH and USNM.

RESULTS AND DISCUSSION

Adults of D. guttifera, were reared only from sporophores oiCollybia
dryophila (Bull, ex Fr.) Kummer (Tricholomataceae) and Psilocybe poly-
trichophila Pk. (Strophariaceae) out of 44 different species of field-collected
mushrooms held in emergence chambers (Bunyard and Foote, 1990).
Life history data obtained for flies reared on Instant Drosophila Medium
are given in Table 1.

Adult flies were successfully reared from all four food substrates as
well as from plain agar on which a culture of baker's yeast was actively
growing. In contrast, all larvae placed on pure agar died before form-
ing puparia.

Tests to determine preference of oviposition site (Table 2) showed
mushrooms to be the preferred food substrate. The lettuce substrate was
also frequently selected, but the banana and tomato juice substrates were
poorly utilized.

The laboratory rearings, particularly the frequent use of rotting let-
tuce as an ovipositional site, lead us to suggest that D. guttifera, may be
capable of utilizing decaying plant material in nature and thus is not
necessarily restricted to a fungal diet. Of course, countering that sugges-
tion is the fact that adults have only been reared under natural con-
ditions from larvae feeding in mushrooms.

Vol. 101, No. 3, May & June 1990 163

Table 1. Life history data from Drosophilaguttifera reared on Instant Drosophila Medium at
27° C. (Mean and 1 S. D.)

Incubation Period 3-4 days (x'= 3.2 ± 0.5, n = 30)

Larval Period 4-5 days (x = 4.4 ± 0.4, n = 40)

Prepupal Period 2.0 days (n = 4)

Pupal Period 5-8 days (x = 6.0 ± 1.0, n = 28)

Adult Longevity 28-45 days (x = ± 3.8, n = 23)

Table 2. Results of oviposition preference tests. Tests were conducted at 27° C for 24 hrs. and
involved 50 females.

Number of eggs laid on each substrate

mushroom banana tomato lettuce agar
185 13 15 125 12

LITERATURE CITED

Bunyard, B., and B. A. Foote. 1990. Acalyptrate Diptera reared from higher fungi in

northeastern Ohio. Entomol. News. In Press.
Patterson, J. T., and W. S. Stone. 1952. Evolution in the Genus Drosophila. The Mac-

Millan Co., NY. 610p.

(Continued from page 160)

In notes of local entomological interest Dr. Paul Schaefer followed up on a discussion
at the November meeting. He showed a number of photographs by Dwight Kuhn of Dex-
ter, Maine showing Polistes wasps that had been parasitized by the strepsipteran, Xenospec-
kii. Few in the audience had ever seen a strepsipteran, let alone pictures of the fleeting,
free-living adult male mating with the grub-like parasitic female still within the abdomen
of the host wasp! We all came away realizing that this odd insect is more common than first
thought and no doubt paper wasps will be examined more closely this summer as a
result.

Harold B. White,
Corresponding Secretary

164 ENTOMOLOGICAL NEWS

PROPYLEA QUATUORDECIMPUNCTATA:
ADDITIONAL U. S. RECORDS OF AN

ADVENTIVE LADY BEETLE
(COLEOPTERA: COCCINELLIDAE)1

A. G. Wheeler, Jr.2

ABSTRACT: The distribution of Propylea quatuordecimpunctata, an Old World aphido-
phagous coccinellid known in North America from Quebec, New York, and Vermont, is
extended to Maine. Occurrence of this adventive predator is attributed to spread of pop-
ulations along the St. Lawrence River in Quebec rather than to intentional releases for
biocontrol purposes. The native and exotic plants on which P. quatuordecimpunctata was
collected in Maine and at an additional site in Vermont are listed; probable aphid prey
are noted.

The Palearctic coccinellid Propylea quatuordecimpunctata (L.) feeds
on numerous European aphid species associated with diverse crops and
habitats. Evaluated in Oklahoma for its biocontrol potential against the
greenbug, Schizaphis graminum (Rondani) (Rogers et al. 1972 a,b), it was
released in Delaware, New Jersey, and Oklahoma beginning in 1970.
Attempts to establish this predator were unsuccessful (Gordon 1985,
Dysart 1988).

At about the time that P. quatuordecimpunctata was imported from
France by the U. S. Department of Agriculture (Rogers et al. 1971), an
established population was detected in North America during 1968 near
Ste. Foy, Quebec (Chantal 1972). Its fortuitous introduction into eastern
Canada thus predated intentional releases in the United States.

Subsequent collections in Quebec, mostly along the St. Lawrence
River, led Larochelle and Lariviere (1980) to suggest that it had been
introduced with ship traffic associated with the St. Lawrence Seaway.
Dysart (1988) and Schaefer and Dysart (1988) concurred that this coc-
cinellid probably was accidentally introduced into the New World. By
1987, P. quatuordecimpunctata had been collected in 14 counties in Quebec,
Chittenden and Grand Isle counties in northern Vermont, and at Platts-
burgh (Clinton County) in northeastern New York (Dysart 1988). Gor-
don's (1985) diagnosis and illustration of the adult facilitate recognition
of this distinctive species in the Nearctic fauna.

Larochelle and Lariviere (1980) and Dysart ( 1 988) suggested that this
aphidophagous coccinellid, having adapted to Quebec's harsh climate,
probably will spread throughout most of North America. Herein, I
extend the known Nearctic distribution to Maine, cite an additional Ver-

1 Received September 9, 1989. Accepted October 21, 1989.

^Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA

17110-9408.

ENT. NEWS 101(3): 164-166, May & June, 1990

Vol. 101, No. 3, May & June 1990 165

mont record, and give ecological notes on these collections. Voucher
specimens have been deposited in the insect collections of Cornell
University, Ithaca, New York (CUIC), and Pennsylvania Department of
Agriculture, Harrisburg (PDA).

Propylea quatuordecimpunctata: Additional U.S. Records

On 11 June 1988, 1 collected seven adults on the University of Ver-
mont campus at Burlington, Chittenden County. Two were beaten from
terminals of tatarian honeysuckle, Lonicem tatarica L., infested with the
aphid Hyadaphis tataricae (Aizenberg); five were collected from an orna-
mental cinquefoil, Potentilla parvifolia Lehm.

In 1989, 1 swept four adults on 3 July from hairy vetch, Vicia villosa
Roth, along Old Coldbrook road near exit 44 of 1-95 at Hampden
(Penobscot County), Maine. The most likely prey species on vetch was
the pea aphid, Acyrthosiphon pisum (Harris). Two days later an adult was
collected in Houlton (Aroostook County), Maine, by beating branches
of speckled alder, Alnusrugosa (DuRoi) Spreng., infested with the woolly
alder aphid, Paraprociphilus tessellatus (Fitch).

R. V. Flanders called my attention to a previous Maine record, one
not formally published but cited in the National Pest Happenings news-
letter [5 (3): 4, 1989], issued by the Texas Agricultural Extension Service.
One adult was collected 7 June 1988 on raspberry at Monmouth (Ken-
nebec County), by M. P. Tully.

DISCUSSION

Schaefer and Dysart (1988) noted that P. quatuordecimpunctata is
being propagated and redistributed by the USDA APHIS Biological
Control Program. No recoveries, however, have been made from any of
the release sites, including those in Maryland and Pennsylvania (P. W.
Schaefer, personal communication) and North Falmouth, Massachu-
setts (R. V. Flanders, personal communication). The Maine records
probably can be attributed to natural dispersal of the well-established
Quebec populations rather than to intentional releases or to a separate
introduction with European commerce. The Penobscot County collec-
tion near Bangor extends the known U. S. distribution about 350 km east
of the Plattsburgh, New York - Burlington, Vermont area. The Houlton
site in Aroostook County lies about 240 km from the nearest population
recorded in Quebec; Monmouth (Kennebec County) is about 230 km
from the nearest Quebec population.

166 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

I thank P. W. Schaefer (USDA, Beneficial Insects Research Lab., Newark. Delaware)
for allowing me to use recent information relating to releases of P. quatuordecimpunctata in
eastern North America, R. V. Flanders (USDA Biological Control Lab., Niles, Michigan)
for informing me of the Cape Cod releases and the previous Maine record, R. D. Lehman
(PDA, BPI) for confirming my identification, and Lehman and K. Valley (PDA, BPI) for
reviewing the manuscript.

LITERATURE CITED

Chantal, C. 1972. Additions a la faune coleopterique du Quebec. Nat. Can. 99:243-244.

Dysart, R. J. 1988. The European lady beetle Propylea quatuordecimpunctata: new locality
records for North America (Coleoptera: Coccinellidae). J. New York Entomol. Soc.
96:119-121.

Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. J. New
York Entomol. Soc. 93:1-912.

Larochelle, A. and M.-C. Lariviere. 1980. Propylea quatuordecimpunctata L. (Coleoptera:
Coccinellidae) en Amerique du Nord: etablissement, habitat et biologic. Bull. Invent.
Insectes Quebec 2:1-9.

Rogers, C. E., H. B. Jackson, G. W. Angalet, and R. D. Eikenbary. 1972. Biology and
life history of Propylea 14 - punctata (Coleoptera: Coccinellidae), an exotic predator of
aphids. Ann. Entomol. Soc. Am. 65:648-650.

Rogers, C. E., H. B. Jackson, and R. D. Eikenbary. 1972. Voracity and survival of Pro-
pylea 14 -punctata preying upon greenbugs. J. Econ. Entomol. 65:1313-1316.

Rogers, C. E., H. B. Jackson, R. D. Eikenbary, and K. J. Starks. 1971. Sex determina-
tion in Propylea 14 - punctata (Coleoptera: Coccinellidae), an imported predator of
aphids. Ann. Entomol. Soc. Am. 64:957-959.

Schaefer, P. W. and R. J. Dysart. 1988. Palearctic aphidophagous coccinellids in North
America. Pages 99-103 in E. Niemczyk and A. F. G. Dixon, eds. Ecology and effective-
ness of Aphidophaga. SPB Academic Publishing, The Hague, Netherlands.

Vol. 101, No. 3, May & June 1990 167

ADDITIONS TO THE PAPILIONOIDEA

(LEPIDOPTERA) OF THE REVILLAGIGEDO

ISLANDS, MEXICO1

John W. Brown2

ABSTRACT: Five species of butterflies are reported from the Revillagigedo Islands, Mex-
ico, for the first time: Phoebis agarithe (Pieridae); Chlorostrymon simaethis, Leptotes marinus,
and Hemiargus ceraunus (Lycaenidae); and Danaus gilippus (Nymphalidae). These new
records increase to ten the number of Papilionoidea recorded from this archipelago.

The Revillagigedo Archipelago is comprised of four islands located
approximately 500 km SSWof Cabo San Lucas, Baja California Sur, and
about 600 km W of Cabo Corrientes, Jalisco. Politically, the islands
belong to the state of Colima, Mexico. The islands are oceanic and
volcanic in origin. Socorro is the largest and most diverse topographi-
cally; Clarion and San Benedicto are considerably smaller; and Roca
Partida is little more than a rock jutting from the sea (Palacios-Vargas,
Llampallas, and Hogue 1982).

Vazquez (1958, 1959, 1960) cited five butterfly taxa from the Revillag-
igedos, four of which represent endemic subspecies: Battus philenor
insularis (Vazq., \956)(Papi\ionidae), Strymon melinusclarionensis(Heid.,
1933) (Lycaenidae), Strymon columella clarionica (Vazq., 1958) (Lycae-
nidae), and Erynnisfuneralis(ScuddeT& Burgess, 1870)(Hesperiidae), all
from Clarion; and Strymon columella socorroica (Vazq., 1958) (Lycae-
nidae) from Socorro. Palacios-Vargas et al. (1982) added a sight record of
the widespread Neotropical Phoebis sennae (Linnaeus, 1758) (Pieridae)
from Socorro. The purpose of this note is to present the first records of
five additional species of Papilionoidea from Socorro Island, and the
first capture record of Phoebis sennae. All of these are widespread Neo-
tropical species that occur commonly on the Mexican mainland. Depos-
itories are abbreviated as follows: LACM, Natural History Museum of
Los Angeles County, Los Angeles, California; SDNHM, San Diego
Natural History Museum, San Diego, California. All specimens were
determined by the author.

PIERIDAE

Phoebis sennae (Linnaeus): Socorro Island, Bahia Braithwaite, 1M, 16 April 1987, D. K.
Faulkner (SDNHM). The specimen is best referred to P. sennae marcellina (Cramer), the
common mainland subspecies.

'Received December 4, 1989. Accepted January 11, 1990.

-Entomology Section, Natural History Museum of Los Angeles County, 900 Exposition
Blvd., Los Angeles, CA, 90007

ENT. NEWS 101(3): 167-169, May & June, 1990

168 ENTOMOLOGICAL NEWS

Phoebis agarithe (Boisduval): Socorro Island, Bahia Braithwaite, 1M, 16 April 1987, D.
K. Faulkner (SDNHM). Identification of the specimen is based on the straight, unbroken
discal line of the ventral forewing (Brown 1929; Howe 1975). The single specimen is in poor
condition inhibiting subspecific assignment.

LYCAENIDAE

Chlorostrymon simaeihis (Drury): Socorro Island, 3200', 12F, 15 April 1955, E. Y. Dawson
(LACM). The specimens are phenotypically indistinguishable from C. simaethis sarita
(Reakirt), the common Mexican mainland subspecies (Nicolay 1980; Johnson 1989).

Leptotes marinus (Reakirt): Socorro Island. Bahia Braithwaite, 2M, IF, 16 April 1987, D.
K. Faulkner (SDNHM).

Hemiargus ceraunus (Fabricius): Socorro Island, 3200', IF, 18 April 1955, E. Y. Dawson
(LACM). I follow Nabokov (1945) and Clench (1965), treating H. ceraunus as distinct from
H. hanno (Stoll). Intra-population variability in H. ceraunus inhibits meaningful use of
trinomials in this species.

NYMPHALIDAE

Danausgilippus (Cramer): Socorro Island, naval base at Cornwallis Bay, 1 F, 27 Novem-
ber 1988, K. Kaiser & J. Johnston (LACM).

DISCUSSION

No butterfly species has been recorded from either San Benedicto or
Roca Partida. Of the ten species recorded from Socorro (n = 7) and
Clarion (n=4), it is surprising that only one is known from both islands,
i.e. Strymon columella, represented by an endemic subspecies on each
island. It is likely that the absence of additional taxa in common to both
islands reflects insufficient sampling rather than ecological equilibria.

ACKNOWLEDGMENTS

I thank David K. Faulkner (SDNMH) and Charles L. Hogue (LACM) for allowing me
to examine specimens in their care and for providing helpful comments on the brief
manuscript. I also thank Robert K. Robbins and Lee D. Miller for critically reviewing
the paper.

LITERATURE CITED

Brown, F. M. 1929. A review of the genus Phoebis (Lepidoptera). Amer. Mus. Novit. 368: 1-22.
Clench, H. K. 1965. Variation and distribution of Hemiargus huntingtoni (Lepidoptera:

Lycaenidae). J. New York Entomol. Soc 73:41-45.
Howe, W. H. 1975. The butterflies of North America. Doubleday and Co., Inc., Garden

City, New York. 633 pp.
Johnson, K. 1989. Revision of Chlorostrymon Clench and description of two new austral

neotropical species (Lycaenidae). J. Lepid. Soc. 43:120-146.
Nabokov, V. 1945. Notes on neotropical Plebejinae (Lepidoptera: Lycaenidae). Psyche

52:1-61.

Vol. 101, No. 3, May & June 1990 169

Nicolay, S. S. 1980. The genus Chlorostrymon and a new subspecies of C.simaethis.J. Lepid.
Soc. 34:253-256.

Palacios-Vargas, J. G., J. Llampallas, and C. L. Hogue. 1982. Preliminary list of the
insects and related terrestrial Arthropoda of Socorro Island, Islas Revillagigedo, Mex-
ico. Bull. Southern California Acad. Sci. 81:138-147.

Vazquez G., L. 1958. Notas sobre Lepidopteros de las Islas Revillagigedo. I. Ann. Inst.
Biol. Mexico 28:301-307.

Vazquez G., L. 1959. Notas sobre Lepidopteros de las Islas Revillagigedo. II. Ann. Inst.
Biol. Mexico 29:349-353.

Vazquez G., L. 1960. X. Observaciones sobre los artropodos. Pp. 217-234, in Adem, J., E.
Cabo, L. Blasquez, F. Miranda, A. Villalobos. T. Herrera, B. Villa, and L. Vazquez. La
Isla Socorro, Archipelago de Las Revillagigedo. Monogr. Inst. Geofisica, Univ. Nac.
Autonoma Mexico 2:1-234.

BOOK REVIEW

BIRD BLOW FLIES (PROTOCALLIPHORA) IN NORTH AMERICA
(DIPTERA: CALLIPHORIDAE) WITH NOTES ON THE PALE-
ARCTIC SPECIES. 1989. C.W. Sabrosky, G.F. Bennett, T.L. Whitworth.
Smithsonian Institution Press, Washington, D.C. and London. 312
pp. $16.95.

This monographic revision of a blowfly genus whose larvae feed upon the blood of nest-
lings of many passerine birds in both enthralling and very informative. This is the third
revision of the genus in the last 60 years. In the last previous treatment of its included
species. Hall (1948. The Blowflies of North America. Thos. Say Foundation, p 179-201)
flatly declared that " Protocalliphora . . . does not occur in North America." The current
authors have painstakingly settled this matter. Although misidentified type specimens and
designated lectotypes have led to some confusion concerning the proper generic name.
North American usage and the current work will certainly establish Protocalliphora as the
correct generic name of the bird blowflies. Species in the genus possess a fascinating
biological association with birds, and their larvae have unique anatomical attributes to
facilitate their 'parasitic' mode of feeding. While Hall (1948) recognized 10 species, this
work recognizes 26 species, 15 of which are described as new, and 2 species that are now
known to be Holarctic in distribution, P. chrysorrhoea and P. braueri. Each species treated
here is given a complete description, including a diagnosis, description of male and female,
descriptions of the immature stages, particularly the puparium, the material examined,
with the type series for new species, distribution informatioin, and ecology and biology,
along with appropriate remarks. These descriptions present admirable models of what
every entomologist would desire to have in a revision. Unfortunately, larval cephalo-
pharyngeal skeletons and the prothoracic fringe of the mature larvae are not illustrated;
one must refer to Hall (1948, PL 38) for these details. Also, the details of the posterior
spiracles and the integumentary armature from the puparia are not easily resolved in the
halftone illustrations. Certainly, this work will be useful to entomologists, ornithologists,
ecologists, and field workers. After over 30 years of work, study, and compilation, the
authors have presented an excellent book.

Stuart E. Neff, Professor

Department of Biology, Temple University

Philadelphia, PA 19122

170 ENTOMOLOGICAL NEWS

INVERTEBRATE POPULATIONS IN THE NESTS

OF A SCREECH OWL (OTUS ASIO) AND AN

AMERICAN KESTREL (FALCO SPARVERIUS)

IN CENTRAL NEW YORK1

James R. Philips , Daniel L. DindaP

ABSTRACT: Screech owl (Otus asio) nest material from a tree hole in Syracuse, N.Y., con-
tained 22,991 arthropods of 61 species. Arthropod density was 131/g dry weight of nest
material. An American Kestrel (Falco sparverius) nest in a nest box in Jamesville, N.Y.,
yielded 26,553 invertebrates of 93 species. Arthropod density was 38/g dry weight of nest
material. Lists of the species found and their populations are presented, and their trophic
and symbiotic relationships are discussed. Bird parasite levels were extremely low. Litter
fauna was dominant in the screech owl nest, while stored products fauna was dominant in
the Kestrel nest.

Nests of birds harbor a wide variety of invertebrates,, including soil
and litter, parasitic, predatory and coprophilic organisms. Numerous
studies have demonstrated that birds' nests are reservoirs of domestic
and stored products pests as well, containing populations of carpet
beetles (Dermestidae), clothes moths (Tineidae), house dust mites (Pyro-
glyphidae), stored products mites (Glycyphagidae), and poultry mites
(Macronyssidae) (Woodroffe and Southgate, 1951; Woodroffe, 1953,
1954; Baker et #/., 1956). Nests of birds of prey (Falconiformes and Strigi-
formes) serve as a habitat for necrophilic arthropods as well as other
nidicoles, since they contain carrion and regurgitated pellet remnants of
their prey (Philips and Dindal, 1977).

The check-lists of Hicks (1959, 1962, 1971) serve as excellent guides to
the literature on insects in birds nests, and they demonstrate how poorly
raptor nest fauna is known. Prior to this study, only six species of inver-
tebrates were known from eastern screech owl (Otus asio (L.)) nests
(Baker, 1904, Bequaert, 1955, Gehlbach and Baldridge, 1987; Linsley,
1944; Robinson, 1941; Vaurie 1955) and only eight species were known
from American kestrel (Falco sparverius L.) nests (Balgooyen, 1976,
Bequaert, 1955; Capelle and Whitworth, 1973; Hill and Work, 1947;
Roest, 1957; Williams, 1947). The objective of this study was to inves-
tigate the invertebrate community of a screech owl and a kestrel nest, and
to determine the levels of parasites, pest, and other species infestation
in them.

Deceived September 27, 1989. Accepted December 30, 1989
2Babson College, Wellesley, MA 02157

•%tate University of New York, College of Environmental Science and Forestry, Syracuse,
NY 13210

ENT. NEWS 101(3): 170-192, May & June, 1990

Vol. 101, No. 3, May & June 1990 171

METHODS

Eastern screech owls occur in mixed woodlands east of the Rocky
Mountains, feeding on small vertebrates and invertebrates and roosting
and nesting in hardwood tree hollows or occasionally in nest boxes
(Hekstra, 1973; Karalus and Eckert, 1974). In 1975, screech owls were
observed nesting and roosting in a tree hole 9m high in a white oak
(Quercus alba L.) in Oakwood Cemetery, Syracuse, New York. On 12
March 1976, a 1 75 g dry wt. sample of nest material was collected from the
tree hole. At this time the hole was being used by the owls, but egg-laying
had not yet taken place. Most of the sample material was placed in mod-
ified Tullgren funnels (Murphy, 1962) for extraction of microarthropods,
and weighed when dry. The rest was placed in culture dishes so that
nidicolous insect larvae could be raised to adults for identification. On
16 June 1976, after nesting was over, a second small sample of nest
material was collected and used for invertebrate extraction. Although
the March sample removed most of the litter from the tree hole, it did not
ruin the hole as a nest site, since the owls nested there in 1976 and 1977 as
they had several years previously.

American kestrels inhabit open country in North and South America,
like the screech owl feeding on small vertebrates and invertebrates and
nesting in tree holes or nest boxes (Brown and Amadon, 1968). On 24
June 1976, the day after the nestlings fledged, we collected an American
kestrel nest (705 g dry wt.) in a nest box 5m high in a dead tree in James-
ville, New York. The nest material was processed in the same way as the
screech owl nest.

RESULTS

The March screech owl nest sample included as much of the nest
debris as could be obtained through the tree hole. The material consisted
mainly of bits of oak leaves and small twigs. Small amounts of grass,
woodchips, dirt, eggshell, excreta, feathers and pellet material (chitin,
hair and bones) were also present. Prey remains included a skull and jaw
of Microtus pennsylvanicus (Ord.). Funnel extraction yielded 22,991 mic-
roarthropods of 61 species (Table 1). Fifteen species of insects were
fqund, but 99% of the individuals were mites. Arthropod density was 1317
g dry wt. of nest; individual species and their populations are given in
Table 2.

The June screech owl postnesting sample containing basal material
from the nest hole was not analyzed in detail. A survey of the June nest
fauna revealed that almost all of the species present were also present in
the March sample. Relative populations were quite different, but that is

172 ENTOMOLOGICAL NEWS

probably more the result of the site disturbance in March - removing so
much of the litter in the hole - than the result of changes associated with
the nesting cycle. Two species were present in June that were not observed
in March - Dendrophaonia scabra G. -T. (Diptera: Muscidae) and Trox
variolatus Mels. (Coleoptera: Trogidae).

The American kestrel nest was composed mainly of sawdust with bits
of sticks, leaves, moss, grass, wood, excreta, chitinous prey remains, pel-
lets of hair, bone and chitin and several prey feet. Identifiable prey
remains included two skulls of Microtus pennsylvanicus (Ord.) and many
fragments of the carabid beetle Calosoma frigidum Kirby. One avian
skull was found, probably that of a sparrow. The nest sample contained
26,553 individuals of 93 species (Table 1). Thirty-one species of insects
were found, but 90% of the individuals were mites. Arthropod density
was 38/g dry wt; individual species and their populations are given in
Table 3.

BIOLOGICAL ANNOTATIONS ON COLLECTED TAXA

Screech Owl Nest Fauna

Ramusella clavipectinata (Mich.), the numerically dominant species
in this nest community, is a widespread oribatid mite fungivore which
Mahunka (1986) found to be very abundant in Hungary. Conditions in
the tree-hole full of highly decomposed leaf litter were near optimal for
this mite, judging by its tremendous population. The nest oribatids are
part of the humus fauna involved in the decomposition of the nest
material.

The astigmatic mites found included nest humus and animal remains
fauna, with several new genera and species (Fain and Philips, 1977a,
1977b, 1978a, 1979, 1981). Fuscarus was previously known only from
rodent nests, and nothing was known of its food habits (Krantz, 1978).
We observed fungal spores as gut contents of all stages of Fusacarus
tenuipes (Fain and Philips), and the fungus species was the same as that
eaten by Oppia and Acotyledon .

Acotyledon paradoxa Ouds. was previously known only from phoretic
deutonymphs and a protonymph. The discovery of all stages of this
species has enabled taxonomists to clarify the systematic position of the
genus. This species has been found on a bat and in granaries in the Soviet
Union, and in mouse nests in the U.S.A. (Fain and Philips, 1978c).

Histiogaster robustus Wdrg. is known from trees (Woodring, 1966b).
Other species in the genus have been found associated with bark beetles.

The histiostomatids collected were represented mainly by the ento-
mophilic nonfeeding hypopus stage. Development of this stage from
protonymphs may have been triggered by the extraction process. His-

Vol. 101, No. 3, May & June 1990 173

tiostomatids were not common in the nest leaf litter, but were abundant
in the basal hole debris of the postnesting sample. One species of Myi-
anoetus is known to be phoretic on Muscina stabulans (Fall.) ( Hughes and
Jackson, 1958). In the June sample we observed 20 Histiostoma paucipu-
teolum Fain and Philips hypopodes on the head, legs, and abdomen of a
Trox variolatus Mels. beetle larva. An adult T. variolatus carried 53 of these
hypopodes, almost all on the legs. In the March sample, one histerid bee-
tle adult oWendrophilusxavieri Mars, bore 20 of these hypopodes, mainly
on the elytra and abdomen. Feeding stages of histiostomatid mites
generally strain microorganisms from wet substrates (Krantz, 1978).

Besides Histiostoma hypopodes, the adult trogid beetle carried 137
hypopodes ofEuglycyphagus, on the pronotum, metasternum, elytra and
abdomen. Euglycyphagus is fungivorous to some extent and, like the his-
tiostomatids, was most abundant in the basal debris. Histiostomatid and
glycyphagoid mites have not previously been found on trogid beetles,
although Fain and Philips ( 1978b) have described a new species of win-
terschmidtiid mite from Australian trogid beetles.

Sapracarus tuberculatus Fain and Philips, known only from this nest,
may be a strict nidicole or general tree-hole dweller. Its entomophilic
hypopus indicates a phoretic relationship with some insect. This mite is
presumably detritivore, but no gut contents were visible in the speci-
mens examined.

Echimyopus is a mite genus known as deutonymphs from hair follicles
and skin galls of Central American rodents and marsupials (Fain, 1969;
Fain etal., 1973; Fain and Lukoschus, 1976). One previous record of the
genus exists for North America, from squirrel hosts (Whitaker et al.
1975). Adults and nymphs of this genus had not previously been found
(Fain and Philips, 1981).

Orycteroxenus hypopodes occur on mammal hair, and the occurrence
of one hypopus of O. soricis (Ouds.) can be correlated with our finding of
prey remains of one of its hosts, Microtuspennsylvanicus (Fain, 1969; Fain
and Whitaker, 1973). Neoxenoryctes hypopodes also are pilicolous, and
the nymphs and adults are at least partly fungivorous.

The chigger Miyatrombicula cynos Ewing is known from raccoons
and squirrels (Ewing, 1937; Whitaker^ al., 1975). All stages of this mite
were found in the nest, and over half of the larvae were engorged. Since
many chiggers show little host specificity, it is possible tht the screech
owl could be an acceptable host for this species, but more likely the hole
had been used by squirrels previously, and the owls had only recently
taken possession again for nesting.

Tarsocheylus paradoxus Berl. is known from tree-holes and rotting
bark (Atyeo and Baker, 1964). Pyemotids, scutacarids, tarsonemids, and
tydeids are all soil and litter groups, which pierce and suck fluids from

174 ENTOMOLOGICAL NEWS

fungi and/or nematodes (Moore et al., 1988). Proctotydaeus is an insect
associate (Andre, 1980) and may have reached the nest on moths, since
Treat ( 1 96 1 ) found one species phoretic on noctuid moths. Cheyletids are
common nest predators, and Woodroffe (1953) observed that the acarid
mite Tyrophagus is preferred prey for Cheyletus eruditus (Schr.); perhaps
in this nest it was mainly preying on Acotyledon, the most abundant
acarid mite present.

The mesostigmatic mites found included gamasine parasites of birds
and mammals, gamasine predators, and uropodine fungivores. Species
of Laelaps, Haemogamasus, Hyperlaelaps, and Androlaelaps fahrenholzi
(Bed.) are small mammal parasites with many hosts (Whitaker and
Wilson, 19"y '4). Androlaelaps casalis (Berl.) has been described as a mam-
mal and bird ectoparasite common in bird nests (Wilson and Bull, 1977.,
Rosen et al. 1985), but McKinley (1963) concluded that this species did
not pierce mammal or avian skin, and actually fed on other mites, such
as acaridids, and their eggs.

Hypoaspis, Blattisocius, and Proctolaelaps are common nest predators.
The size difference in the three Dendrolaelaps species suggests possible
differences in prey size classes. Many ameroseiids feed on fungi or
pollen, but can also be predators (Krantz, 1978). They occur in litter as
well as mammal nests. Among the uropodines, Metagynella parvula
Camin is known only from tree-holes (Camin, 1953). Its phoretic relation-
ships, if any, are unknown. One deutonymph of Trichouropoda martini
Hirschmann was phoretic on the histerid beetle Dendrophilus xavieri
Marseul. Trichouropoda martini is known from a red squirrel (Tamias-
ciurus) nest (W. Hirschmann, pers. comm.). Uroseius deutonymphs have
been found in passerine nests phoretic on trogid beetles (Sixl, 1971).

The pseudoscorpion Acuminochernes crassopalpus (Hoff) is a pred-
ator that has been previously recorded from treeholes (Park and Auer-
back, 1954) and mammal nests (Muchmore, pers. comm.). Chernetid
pseudoscorpions posses several adaptations for colonizing nests and
treeholes, including phoresy and the ability to store sperm for prolonged
periods (Jones, 1975).

Most of the fly larvae are known scavengers on animal remains, but
scenopinid larvae prey mainly on tineid moth larvae. The cecidomyid
larvae were probably saprovores associated with the decaying nest
plant matter.

Beetles associated with treeholes rather than birds' nests included the
curculionid adult and the melandryid, tenebrionid and staphylinid lar-
vae (Borroretal., 1981). Xylodromus is a new staphylinid record for New
York (Moore and Legner, 1975). The histerid beetles were the largest
predators in the nest.

Only a few fleas were found, and both species are mammal parasites.

Vol. 101, No. 3. May & June 1990 175

Orchopeas howardi (Baker) is most commonly found on tree squirrels
and O. leucopus (Baker) prefers Peromyscus (Main, 1970). Ixodes marxi
Banks, the single tick found, is also a tree squirrel parasite (Bequaert,
1945).

The single psocopteran appears to be an accident. It may have been
windborne or reached the nest on a bird or mammal, as phoresy occas-
ionally occurs in psocopterans (Mockford, 1967). The only wasp found.
Synergus, was extracted from a gall on one of the nest leaves.

The ectoparasites found included the squirrel tick, the squirrel flea, a
squirrel chigger, three parasitic mesostigmatic mite species known from
squirrels, and an astigmatic mite genus known from squirrels. Also, the
nest contained many mammal nest associates such as Fusacarus and
Neoxenoryctes. Oakwood Cemetry abounds in gray squirrels (Sciurus car-
olinensis Gmelin), and it seems inevitable that this tree-hole would be
occasionally visited by squirrels when not used by the owls. In 1977, the
owls again nested in this hole, as they had for several previous years, but
in 1978 squirrels took over the hole and the owls did not nest there.

Squirrel use of the hole explains the many squirrel parasites found.
We believe that the hosts o{ Echimyopus orphanus Fain and Philips and
Neoxenoryctes is the gray squirrel, and that these other genera like
Fusacarus will also be found in gray squirrel nests.

This nest community was characterized by very high species richness
and very lowequitability. The variety of food sources in the tree-hole nest
provide a habitat for many species which colonize the hole deliberately
or accidentally, and the protected microenvironment seems to provide
nearly optimal conditions for some species.

American Kestrel Nest Fauna

The kestrel nest invertebrate community was very different from the
screech owl nest community. Of 93 species in the kestrel nest, only 12
were also present in the screech owl nest. These species - 9 mites, a his-
terid beetle, a sphaerocerid fly, and a psocopteran - included no avian
parasites and were small components of both nests, comprising only 8%
of the individuals in the screech owl nest, and 4% of those in the kestrel
nest. Four of the mites are parasitic or phoretic on rodents. Three addi-
tional species may be common to both nests. Lack of adults of the
scenopinid fly, the tineid moth in the kestrel nest, and the uroseiid mite
in the screech owl nest, prevented complete identification.

Diversity in the kestrel nest was lower than in the screech owl nest.
Although species richness was higher than that of the screech owl nest,
equitability was lower, as was arthropod density. In this nest, a new
species ofLardoglyphus, a genus known from dried fish, butchers' offal.

176 ENTOMOLOGICAL NEWS

and hides (Hughes, 1976) was highly numerically dominant. This species
was fungivorous to some extent, and the hypopodes were phoretic on
Dermestes pulcher LeConte (Philips and Norton, 1978). It seems to have
some degree of host specificity. A nest box in a tree 18m from this kestrel
nest was examined and preliminary inspection has revealed neither Lar-
doglyphus falconidus Philips and Norton nor its dermestid host species.
Another genus of dermestid was present, as were the species ofLepido-
glyphus and Cheyletus which occurred in this kestrel nest.

Other genera of acarid mites in this nest with entomophilic hypo-
podes were Histiogaster, Michaelopus, Sancassania, and Schwiebea.
Michaelopus corticalis (Michael) has been found under dead tree bark
and in nests of several avian orders (Fain, 1982), and is probably sap-
rophagous like Sancassania and Schwiebea. Schwiebea terrana Jacot was
originally collected from pine litter and inhabits dead wood (Wood-
ring, 1966a).

Histiostomatid mite hypopodes were found phoretic on Trox seaber
(L.) in this nest, as they were on the other species of trogid beetle in the
screech owl nest. Thirteen hypopodes were found under the wings of the
trogid, in crevices at the border of the thoracic and abdominal regions of
the body. Twelve hypopodes were Histiostoma species number three, and
one was Hexanoetus conoidalis Fain and Philips. Hexanoetus conoidalis
was present in adult and immature stages in the screech owl nest. Histios-
toma species number 3 is identical to the Histiostoma species B found by
Philips etal. (1983) under the elytra of four specimens of Troxaequalis Say
in a saw-whet owl (Aegolius acadicus (Gmelin)) nest in Connecticut.

Two species of feather mites were found in the nest. The specimens of
Dubininia in the pellet sample may have been regurgitated with a pellet
or may have dropped off from a kestrel in the nest, as this genus is a
parasite of the Falconiformes as well as other birds. Analges probably
was derived from prey.

Lepidoglyphusfustifer(Ouds.) is a European stored products species.
It has not previously been recorded from North America (Hughes, 1976)
and the hypopus stage was not previously known from this species. We
have observed fungal spores in the gut of many of our specimens.

As with the screech owl nest, a few Orycteroxenus hypopodes were
found, as well as evidence of predation on Microtus. Glycyphagus
hypudaei (Koch) hypopodes are phoretic on Microtus and many other
small mammals. Myocoptesjaponensis Radford is a parasite whose type
host is Microtus pennsylvanicus and type locality is Ithaca, N.Y. (Fain and
Hyland, 1 970). Dermacarus sciurinus ( Koch), another mammal associate,
is known from a Peromyscus nest (Fain and Whitaker, 1973).

Tytodectes cerchneis Fain was previously known only from one badly
damaged specimen found on Falco tinnunculus L. in Rwanda, Africa

Vol. 101, No. 3, May & June 1990 177

(Fain 1967). The hypoderid mites are a poorly known group, and the life
cycle of very few species is completely known. The hypopus is a sub-
cutaneous parasite of birds, and the other immature stages are not active.
The adults are very short-lived and nonfeeding nidicoles (Fain and
Bafort, 1967).

The finding of Tytodectes represents a new U.S. record. Both Falco
sparverius and F. tinnunculus also have a nasal mite species in common,
and it would be interesting to survey falcon nasal, subcutaneous, and
other mites and compare that data with concepts in falcon taxonomic
relationships.

Nanacarus is a new New York record. This genus contains a number
of fungivorous species which inhabit polypore fungi (O'Connor, 1984).

Among the prostigmatic mites present were two mammal parasites -a
nymphal myobiid mite and a Demodex, as well as tarsonemid and tydeid
soil and litter species. There was no mammal carrion in the nest, so we
believe the parasites either came off prey in the nest before the prey
was eaten, or survived the bird's digestive tract and came out in regurgitated
pellet or fecal material.

Cheyletus trouessarti Oudemans was the most abundant predatory
mite in the nest. Cheyletids are often associated with infestations of
acarid mites (Baker, 1949) and we believe this species was probably feed-
ing mainly on the abundant Lardoglyphus in this nest. Anystids are very
mobile predatory mites which often feed on phytophagous insects and
mites. The one larvae found may have wandered into the nest box from
the tree the box was on, as these mites are long-legged rapid movers
(Krantz, 1978).

As with the screech owl nest, the mesostigmatic mites included gama-
sine mammal parasites, gamasine predators, and uropodine fungivores.
but there were no gamasine avian parasites. Poecilochirus necrophori
Vitzth. is known to be phoretic on the silphid beetle Nicrophorus
(Chmielewski, 1977) and the presence of four on Nicrophorus pustulatus
Hers, explains how this mite reached the nest. This mite probably preys
on nest fly larvae. Macrocheles muscaedomesticae, another immature dip-
teran predator (Peck and Anderson, 1969), is known to be phoretic on
muscid flies (Evans and Browning, 1956; Chmielewski, 1977). Copriphis
is a predatory gamasine which is phoretic on scarabaeid dung beetles.
Two females were found under the wings otTroxscaber near the thorax/
abdomen border. This species is identical to the species of "Eviphis"
found by Philips el al. ( 1983) under the elytra of Trox aequalis in a saw-
whet owl nest in Connecticut. Copriphis has been considered by some
taxonomists to be a synonym ofEviphis, but we agree with those regard-
ing it as a distinct genus. Ameroseiusapodius Karg was discovered by Karg
( 1 97 1 ) in compost in Germany, and has not previously been found in the

178 ENTOMOLOGICAL NEWS

U.S.A. Trichouropoda falconis was recently described as a new species
from this nest by Hirschmann and Wisniewski (1988). Trichouropoda
species number one was found as a phoretic deutonymph attached by
anal stalk to tibia IV of an adult histerid beetle, Carcinops pumilio
Erichson.

Very feworibatid mites were present in this nest, which was dominated
by animal remains fauna. Other humus fauna species were equally
scarce - one collembolan, one enchytraeid, two earthworms, and a few
Psocoptera. To our knowledge, earthworms have only been previously
recorded from nests of one bird - the rook (Corvusfrugilegus L.) in Britain
(Coombs, 1960). Earthworm presence in a kestrel nest may be due to prey
escape, since kestrels prey on earthworms (Balgooyen, 1976). Clubionid
spiders hunt both on the ground and in foliage (Kaston, 1972), and prob-
ably entered the nest while foraging on the tree.

Histerids again were the largest predators in the nest. While the food
of many species of staphylinids is not known, Koskel and Hanski (1977)
classify all aleocharines and staphylinines (like Philonthus) as
carnivores. Nicrophorus may prey on maggots, as well as eating carrion
(Steele, 1927). Lathridiid and tenebrionid fungivores are very common
in birds' nests and Tenebrio molitor L. is a stored products pest (Woodroffe
and Southgate, 1951).

Calosoma frigidum Kirby is a new prey record for the American
kestrel (Philips, 1977). This carabid beetle was known to be preyed upon
by only one bird - the ruffed grouse (Bonasa umbellus (L.)) (Larochelle,
1975), but this demonstrates the paucity of knowledge on avian insect
predation.

Allen (1973) has noted that this carabid preys upon several important
insect pests, and its habit of tree foraging may make this species more
susceptible to kestrel predation than cryptozoic ground carabids, par-
ticularly when, as Allen observed, large numbers of this carabid beetle
occur during major infestations of forest insect pests.

The trogid and dermestid beetles eat hair and feathers in animal
remains like carrion and raptor pellets. Trox scaber (L.) is known from
mammal nests and nests of several other birds, including the screech owl
(Vaurie, 1955). Unidentified species of Dermestes were recorded from
kestrel nests by Balgooyen (1976). Nicrophorus feeds on fresher carrion
than Dermestes.

Scavenging dipteran larvae occurred as in the screech owl nest, but in
much greater numbers and in greater variety, with muscid and scatopsid
flies present. Two families of avian parasitic flies inhabited this nest. The
biting midges (ceratopogonids) suck blood, but the food habits of the
nidicolous milichiid Camus hemapterus Nit. have been a subject of
debate. Bequaert (1942) has quoted the observations of earlier workers

Vol. 101, No. 3, May & June 1990 179

who discovered that it fed on blood and that it fed on skin and feather
secretions. Borror et al. ( 1981) consider it to be blood-sucking. Lloyd and
Philip (1966) agreed with Bequaert that the mouthparts are not adapted
for piercing host skin.

One parasitoid cynipid wasp inhabited this nest, but the hosts of
many species of Pseu deucoila are not known (Muesebeck et al., 1 95 1 ). The
mymarid wasp is a parasite of insect eggs (Borror et al. 1981). Many
braconid wasps are parasites of lepidopteran larvae (BorroTetal. 1981).
Lasius ants nest in rotting logs and stones and cultivate aphids (Arnett,
1985). The three nest anoplurans can be attributed to the birds' mam-
malian prey. The two flea larvae could not be specifically identified.

DISCUSSION

These raptor nest data present a considerably different picture of the
nest community than previous studies. Although Nordberg ( 1936) found
oribatid mites of many species in his European raptor nests, very few
species occurred in our nests. Nordberg found mainly feather mites
among the astigmatic mites in his nests, but our nests contained a wide
variety of astigmatic mites. The prostigmatic and mesostigmatic mite
components of raptor nests are also much more diverse than were prev-
iously realized. In particular, no mammalian-associated mites had been
reported from raptor nests. Our expectations that they would be found
still did not encompass the actual extent of the mammal nest and para-
site fauna in raptor nests.

The mite family Pyroglyphidae is very common in birds' nests
(Woodroffe, 1953; Krantz, 1978), but has never been previously found in
raptor nests, and only three specimens were found in our nest samples.
Its scarcity in the kestrel nest is even more surprising considering its
abundance in a tree swallow (Iridoprocne bicolor (Vieillot)) we examined
which was in a nestbox only 200 m away. More samples are needed to
confirm this apparent preference for nonraptor nests - perhaps the rap-
tor nest moisture and carrion create an unattractive environment. Wood-
roffe (1953) noted that these mites are tolerant of low humidities and
occur in very dry nests. Some nest pyroglyphids also occur in house dust
and cause house dust allergy (Krantz, 1978).

Referring to mites in bird nestbox debris, Herman (1936) stated that
"an estimate of billions in each nest seems conservative". He correctly
judged the importance of the mites, if not their actual density. Neverthe-
less, the densities of arthropods in the screech owl and kestrel nests are
high. Park et al. (1950) studied the tree-hole fauna of elm, sugar maple,
and beech trees, and arthropod densities ranged from 4.4-5.5/g. Ryder
and Freitag (1974) examined ring-billed gull (Lams delawarensis Ord)

180 ENTOMOLOGICAL NEWS

nests of weights similar to the screech owl and kestrel nests, and densities
averaged 1.5-1.7 arthropods/g.

The oribatid mite density in the screech owl nest appears to be
unmatched in any other reported nest. Gembestky and Andrechikova
(1969) examined 32 nests of 7 passeriform species and collected only 98
oribatid specimens. Bukva etal. (1976) found 979 oribatids in 278 small
mammal nests, and Kramarova and Mrciak( 1971) collected 13,364 from
303 small mammal nests.

Kramarova and Mrciak studied the invertebrate groups in small
mammal nests as we did with the raptor nest groups. Some additional
numbers from their 303 nests provide a useful comparison: 20,556 astig-
matic mites; 110,575 mesostigmatic mites; 679 prostigmatic mites; 71
pseudoscorpions; 22,2 1 3 Collembola. Vysotskaya and Nosek ( 1 974) studied
43,796 Collembola from 464 small mammal nests.

The dominance of gamasine mites, especially parasites, in small
mammal nests has also been shown by Drummond (1957) and Allred
and Roscoe (1957). Judging from our data, astigmatic or oribatid mites
can be dominant in raptor nests, and although a variety of mesostigmatic
mites occur, they are not present in tremendous numbers.

Park and Auerbach (1954) found that collembolans made up 27% of
the tree-hole fauna they studied. Considering their abundance in small
mammal nests and treeholes, the scarcity of collembolans in the raptor
nests examined is surprising. Further study is needed to determine the
frequency of Collembola in many raptor nests.

Woodroffe and Sou thgate (1951) studied passerine nests and observed
a succession of invertebrates. Avian ectoparasites dominated during ini-
tial nest construction and occupation. After birds left the nest,
scavenging invertebrates were dominant as feather debris and excreta
decomposed. The final stage of decomposition of the nest material was
dominated by humus fauna. If the nest was used as a winter roost and
reused the next year, the nest acted as a refugium and the scavenging
fauna persisted. Open, exposed nests decomposed faster and the scaveng-
ing stage was reduced or absent. Differences in the fauna of nests of dif-
ferent bird species were correlated with differences in the composition of
the nests.

This pattern is not consistent for all birds' nests. Freitag and Ryder
(1973) found almost no ectoparasites in their ring-billed gull nests. Sap-
rophagous mite populations peaked after gull egg-laying, while pred-
atory mite populations peaked at or after egg-hatching (Freitag et al,
1974).

Trophic category estimates of the raptor nest fauna are given in Table
4. There is no evidence that the raptor nests followed Woodroffe's

Vol. 101, No. 3. May & June 1990 181

scheme, as very fewavian ectoparasites were found. However, the scaveng-
ing animal remains fauna was dominant in our samples taken im-
mediately after falconiform nesting. The effects of screech owl nesting
upon the nest humus fauna were not observed because of our distur-
bance of the site. It is possible that this fauna could have remained domi-
nant throughout the entire nesting period, but an increase in the animal
remains fauna would still be expected as that food source became
more abundant.

Hagvar (1975) described coleopteran succession in raptor nests.
Nests used every year grow larger, older material decomposes, and the
number of niches increases. Old nests in use have the largest species
richness, and large old nests can remain humid in the center even
during droughts.

Succession in raptor nests is complicated by the many variables in
nest location and use. Nests may be used in other seasons and years by
other roosting species, and raptors may share their nests with a variety of
other vertebrates (Wilson, 1925; Sumner, 1933; Zarn, 1974).

Detailed investigations of raptor nest fauna are needed to elucidate
the intricate interrelationships in this microcommunity. Many raptors
are marginal or endangered species, and we need to know what, if any,
invertebrates may be a source of mortality and how frequently this
occurs. When nest trophic interactions are well known, biological con-
trol measures against undesirable invertebrates may be possible.

ACKNOWLEDGMENTS

We are very grateful to S. Allen and D. Crumb for locating the nests for us. Specific tax-
onomic identifications in many cases would have been impossible without the assistance
of many specialists. R. A Norton assisted us with many groups, especially oribatid mites,
and A. Fain expended a great deal of effort in collaborating with us in identifying and de-
scribing many of the astigmatic mites we found. Additional help was received from the
following invertebrate taxonomists: F. G. Andrews (Lathridiidae); F. Athias-Binche
(Uropodina); E. F. Cook (Scatopsidae); S. Cover (Formicidae); D. R. Davis (Tineidae): N.
M. Downie (Lathridiidae); R. J. Gagne (Cedidomyiidae, Muscidae); J Gaud (Analgidae);
R. W. Hamilton (Curculionidae); W. Hirschmann (Uropodina); W. I. Knausenberger
(Ceratopogonidae); J. F. Lawrence (Coleoptera larvae); E. E. Lindquist (Ascidae, Amero-
seiidae, Digamasellidae); A. J. Main, Jr. (Siphonaptera); J. F. McAlpine (Cecidomyiidae,
Milichiidae, Muscidae); A. S. Menke (Cynipidae); A. F. Newtown, Jr. (Histeridae, Staphy-
linidae); D. P. Schwert (Carabidae, Lumbricidae); G. Steyskal (Sphaeroceridae); H. J.
Teskey (Milichiidae, Muscidae, Sphaerceridae); M. K. Thayer (Staphylinidae); E. L. Todd
(Noctuidae); and N. Wilson (Laelapidae). R. Norton and F. Kurczewski reviewed the
manuscript. Preparation of this manuscript was supported by a grant from the Babson
College Board of Research.

182

ENTOMOLOGICAL NEWS

Table 1. Invertebrates in the nests of a screech owl and an American kestrel.

Group

Screech Owl % of American Kestrel % of
species total species total
individuals individuals individuals individuals

Arachnida

Acari

Acaridida

5,720

24.88

19,692

74.16

Actinedida

4,168

18.13

4,015

15.12

Gamasida

625

2.71

238

.90

Ixodida

.004

Oribatida

12,214

53.13

.18

Araneida

.005

Pseudoscorpionida

119

.52

Insecta

Anoplura

.01

Coleoptera

.12

485

1.83

Collembola

.005

Diptera

.40

2,007

7.56

Hymenoptera

.003

.03

Lepidoptera

.08

.01

Psocoptera

.003

.18

Siphonaptera

.02

.01

Oligochaeta

.01

Total

22,991

26,553

Table 2. Arthropod species populations in a screech owl nest.

Class Arachnida
Order Acarina

Suborder Acaridida

Family Acaridae

Acotyledon paradoxa Oudemans
Histiogaster robustus Woodring

Family Euglycyphagidae

Euglycyphagus intercalates Fain and Philips

Family Glycyphagidae

Echimyopus orphanus Fain and Philips
Fusacarus tenuipes Fain and Philips
Neoxenoryctes reticulatus Fain and Philips
Orycteroxenus soricis soricis (Oudemans)

Family Hemisarcoptidae

Sapracarus tuberculatus Fain and Philips

3.746
9

150

614

1,053

Vol. 101, No. 3, May & June 1990

183

Family Histiostomatidae

Comyianoetus denticulatus Fain and Philips

Histiostoma pauciputeolum Fain and Philips

Miyanoetus micromaculatus Fain and Philips

Hexanoetus conoidalis Fain and Philips
Suborder Actinedida
Family Cheyletidae

Cheyletus eruditus (Schrank)
Family Pyemotidae

Bakerdania sp. 1

Bakerdania sp. 2

Brennandania sp.
Family Scutacaridae

Imparipes sp.
Family Tarsocheylidae

Tarsocheylus paradoxus Berlese
Family Trombiculidae

Miyatrombicula cynos Ewing
Family Tarsonemidae

Tarsonemus sp.
Family Tydeidae

Proctotydaeus sp.

Microtydeus sp.

Tydeus sp. 1
Suborder Gamasida
Family Ameroseiidae

Kleemannia sn. sp.
Family Ascidae

Blattisocius dentriticus (Berlese)

Proctolaelaps pomorum (Oudemans)
Family Digamasellidae

Dendrolaelaps sp. nr. marylandae (Hurlbutt)

Dendrolaelaps sp. nr. pint Hirschmann

Dendrolaelaps sp. nr. presepum (Berlese)
Family Laelapidae

Androlaelaps casalis casalis Berlese

Androlaelaps fahrenholzi (Berlese)

Haemogamasus reidi Ewing

Hyperlaelaps micron (Ewing)

Hypoaspis lubrica Voigts and Oudemans

Laelaps alaskensis Grant
Family Metagynuridae

Metagynella parvula Camin
Family Polyaspinidae

Uroseius sp.
Family Trematuridae

Trichouropoda martini Hirschmann

species 1

species 2

Suborder Ixodida
Family Ixodidae

Ixodes marxi Banks

166

252
28
20

13
4

869
1,287

1,508

15
33

11
24
13

8
3
2
1

109
2

2
26

318
12

184 ENTOMOLOGICAL NEWS

Suborder Oribatida
Family Damaeidae

Epidameus canadensis (Banks) 1

Family Oppiidae

Ramusella clavipectinata (Michael) 12,056

Family Oribatulidae

Scheloribates sp. 1 53

Family Parakalummidae

Protokalumma n. sp. 4

Order Pseudoscorpionida
Family Chernetidae

Acuminochernes crassopalpus (Hoff) 1 19

Class Insecta

Order Coleoptera

Family Curculionidae

Phloeophagus variolatus Drury 1

Family Histeridae

Dendrophilus xavieri Marseul 1 1

Family Melandryidae

Scraptinae sp. 8

Family Staphylinidae

Xylodromus sp. 4

Family Tenebrionidae

Neatus tenebrioides (Palisot) 3

Order Diptera

Family Cecidomyiidae

Lestodiplosis sp. 19

Family Milichiidae

Leptometopa sp. 34

Family Scenopinidae

Scenopinus sp. 1

Family Sphaeroceridae

Leptocera (Coproica) n. sp. 39

Order Hymenoptera

Family Cynipidae

Synergus sp. 1

Order Lepidoptera

Family Noctuidae

Epizeuxis americalis (Guenee) 2

Family Tineidae

Tinea sp. 16

Order Psocoptera

Family Liposcelidae

Liposcelis bostrychophihts Badonnel 1

Order Siphonaptera

Family Ceratophyllidae

Orchopeas howardi howardi (Baker) 3

Orchopeas leucopus (Baker) 1 s

Vol. 101. No. 3, May & June 1990

185

Table 3. Invertebrate species populations in a American kestrel nest.

Phylum Annelida
Class Chaetopoda
Order Oligochaeta

Family Enchytraeidae

species 1 1

Family Lumbricidae

Dendrobaena rubida (Savigny) 1

Lumbricus sp. 1

Phylum Arthropoda
Class Arachnida
Order Acarina

Suborder Acaridida

Family Acaridae

Histiogaster carpio ( Kra mer) 1

Lardoglyphus falconidus Fain and Philips 18,581

Michaelopus corticalis (Michael) 1

Sancassania sp. 1

Schwiebea terrana Jacot 8

Family Analgidae

Analges sp. 4

Dubininia sp. 2

Family Glycyphagidae

Dermacarus sciurinus ( Koch ) 1

Glycyphagus hvpudaei (Koch) 1

Lepidoglyphus fustifer Oudemans 870

Orycteroxenus soricis soricis (Oudemans) 2

Family Hemisarcoptidae

Nanacarus n. sp. 3

Family Histiostomatidae

Hexanoetus conoidalis Fain and Philips 1

Histiostoma sp. 1 157

Histiostoma sp. 2 13

Histiostoma sp. 3 12

Miyanoetus n. sp. 16

Family Hypoderidae

Tytodectes cerechneis Fain 12

Family Myocoptidae

Myocoptes japonensis Radford 1

Family Pyroglyphidae

Dermatophagoides sp. 6

Suborder Actinedida

Family Anystidae

species 1 1

Family Cheyletidae

Cheyletus trouessarti Oudemans 1.859

Family Demodicidae

Demodex n. sp. 1

Family Myobiidae

species 1 1

186

ENTOMOLOGICAL NEWS

Family Tarsonemidae

Tarsonemus sp. 1
Family Tydeidae

Coccotydaeolus sp.

Tydeus sp. 1

Tydeus sp. 2
Suborder Gamasida
Family Ameroseiidae

Ameroseius apodius Karg
Family Ascidae

Blattisocius tarsalis (Berlese)

Proctolaelaps sp.
Family Digamasellidae

Dendrolaelaps sp. nr. presepum (Berlese)
Family Eviphididae

Copriphis sp
Family Laelapidae

Androlaelaps fahrenholzi (Berlese)

Hyperlaelaps microti (Ewing)

Laelaps alaskensis Grant
Family Macrochelidae

Macrocheles muscaedomesticae (Scopoli)
Family Parasitidae

Poecilochirus necrophori Vitzthum
Family Polyaspinidae

Uroseius lagenaeformis (Berlese)
Family Trematuridae

Trichouropoda martini Hirschmann

Trichouropoda falconis Hirschmann and Wisniewski

Trchouropoda sp. 1
Suborder Oribatida
Family Achipteriidae

Anachipteria sp.
Family Cymbaeremaeidae

Scapheremaeus sp.
Family Eremaeidae

Eremaeus sp.
Family Oppiidae

Oppiella nova (Oudemans)
Family Oribatellidae

Oribatella sp.
Family Oribatulidae

Eporibatula sp.

Oribatula tibialis (Nicolet)

Phauloppa pilosa (Banks)

Scheloribates sp. 1

Zygoribatula frisae (Oudemans)

Family Oripodidae

Oripoda sp.
Family Parakajummidae

Protokalumma depressa (Banks)

1,232

703

217

2
2

3
5
9

119
6

15
10

2
2
1
5
1

1
1
6
9

2
3

Vol. 101, No. 3, May & June 1990

187

Family Phthiracaridae

Phthiracarus setosellum Jacot
Family Tectocepheidae

Tectocepheus velatus (Michael)
Order Araneida

Family Clubionidae
species 1
Class Insecta

Order Anoplura

Family Hoplopleuridae
species 1
Order Coleoptera

Family Dermestidae

Dermestes pulcher LeConte
Family Histeridae

Carcinops pumilio Erichson

Dendrophilus punctatus (Herbst)

Dendrophilus xavieri Marseul

Euspilotus sp.

Gnathoncus sp.

Margarinotus merdarius (Hoffman)

larvae
Family Lathridiidae

Lthridim minutus (L.)
Family Silphidae

Nicrophorus pustulatus Herschel
Family Staphylinidae

Aleocharinae sp.

Philonthus sp.
Family Tenebrionidae

Tenebrio molitor L.
Family Trogidae

Trox scaber (L.)
Order Collembola

Family Entomobryidae

Willowsia buskii Lubbock
Order Diptera

Family Ceratopogonidae

Culicoides sp. nr. piliferus Root and Hoffman

Culicoides sp. nr. travisis Vargas
Family Milichiidae

Camus hemapterus Nitzsch

Leptometopa latipes (Meigen)
Family Muscidae

Fannia sp.

Muscina stabulans Fallen
Family Scatopsidae

Coboldia fuscipes (Meigen)
Family Scenopinidae

Scenopinus sp.
Family Sphaeroceridae

Leptocera (Coproica) n. sp.

7
6

2
9
4

278

17
1

7
1

38
1,509

5
434

188 ENTOMOLOGICAL NEWS

Order Hymenoptera

Family Braconidae

species 1 1

Family Cynipidae

Pseudeucoila sp.
Family Formicidae

Lasius alienus (Foerster)
Family Mymaridae

species 1 4

Order Lepidoptera

Family Tineidae
species 1
Order Psocoptera

Family Liposcelidae

Liposcelis bostrychophilus Badonnel 48

Order Siphonaptera

species 1

Table 4. Trophic classification of raptor nest fauna.

% individuals per nest
Screech American

Category Owl Kestrel

Animal remains saprovores

27.65

82.19

Nest material saprovores

60.01

8.35

Predators

8.47

9.11

Parasites of vertebrates

3.87

.33

Parasites of invertebrates

.02

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192 ENTOMOLOGICAL NEWS

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SEPTEMBER & OCTOBER, 1990 NO. 4

OMOLOGICAL NEWS

Biogeographic affinities of Ephemeroptera of Black

Hills, South Dakota W.P. McCafferty 193

Anthopotamus, new genus for No. American species
previously known as Potamanthus (Ephemeroptera:
Potamanthidae) W.P. McCafferty, Y.J. Bae 200

Cytology of Leptoglossus zonatus (Hemiptera:

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New lassinae (Homoptera: Cicadellidae) from

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Notiophilus palustris (Coleoptera: Carabidae), a
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Marking Cassidinae (Coleoptera: Chrysomelidae) larvae in
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New distribution record for sand-burrowing mayfly
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William L. Jacobs 219

New method for collecting adult phryganeid caddis-
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D.E. Bowles, K. Stephan, M.L. Mathis 222

Sucking lice (Anoplura) from Pakistan mammals, with
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L.A. Durden, R. Traub, K.C. Emerson 225

Checklist of caddisflies (Trichoptera) from West

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Vol. 101, No. 4, September & October 1990 193

BIOGEOGRAPHIC AFFINITIES OF THE

EPHEMEROPTERA OF THE BLACK HILLS,

SOUTH DAKOTA1' 2

W. P. McCafferty3

ABSTRACT: Records of mayfly species in South Dakota are few in number. Most records
are from the Black Hills region of southwestern South Dakota. Twelve new species records
(genera Caenis. Callibaetis, Dactylobaetis, Epeorus, Nixe. Paraleptophlebia, Siphlonurus.
Tricorythodes) are also based on collections from the Black Hills. The diversity of the mayfly
fauna in the Black Hills is relatively low, with only 19 species in seven families known. The
species mix consists of some widespread North American species, but also western con-
tinental species and eastern continental species whose respective easternmost and
westernmost range limits meet in the Black Hills. The somewhat insular nature of this
small montane region may explain low numbers of species, and its proximity to the main
body of Rocky Mountains may explain the presence of its western component. Eastern
species represented are primarily northeastern North American species with disjunct, pro-
bably relict populations now isolated in the lower Appalachian Mountains, the Ozark-
Ouachita Mountains, and the Black Hills.

Very few published records of Ephemeroptera species in South
Dakota have been established. Species previously reported from the
Black Hills region of southwest South Dakota are included in Table 1.
The Black Hills region referred to herein includes Custer. Fall River,
Lawrence, and Pennington Counties (Fig. 1). The previous records were
given by Morihara and McCafferty (1979) for the Baetidae, and by Allen
and Edmunds (1965) for Ephemerella inermis. Examination of addi-
tional materials of all of these previously reported species has substan-
tiated their presence.

Even fewer records have been established for areas of South Dakota
outside of the Black Hills region. In recent revisions, Bednarik and
McCafferty (1979) recorded Stenonema mediopunctatum arwini from the
Yellowbank River in extreme northeastern South Dakota, and Kon-
dratieff and Voshell (1984) recorded Isonychia rufa from Sioux Falls in
the southeast corner of the state. There are new state records of mayflies
based on species I have examined from eastern South Dakota, but these
will be published as part of revisionary studies now in progress.

Two primary factors have prompted the present report. First, it has
become increasingly important to survey the North American fauna
with respect to documenting biodiversity. Second, the Black Hills region
is important biogeographically since, as will be discussed below, it con-

^Received April 20, 1990. Accepted May 22. 1990.
"Published as Purdue Experiment Station Journal No. 12466.
•'Department of Entomology, Purdue University. West Lafayette, IN 47907.

ENT. NEWS 101(4):/r93-199. September &.Qctobe1| 1990

194 ENTOMOLOGICAL NEWS

tains one of the most unusual mixes of mayfly faunal elements found in
North America.

All species records for the Black Hills region reported by Morihara
and McCafferty (1979) and all but one reported here for the first time
(Table 1) have been based on collections made by me and by A. V. Pro-
vonsha and B. L. Heath in June, 1975 (Sites 2-17 in Table 2 and Fig. 1).
Site 1 (Table 2 and Fig. 1) was collected in November, 1977, by an
unknown collector. All larvae upon which data in Table 1 are based are
deposited in the Purdue Entomological Research Collection, West
Lafayette, Indiana.

In addition to a list of species taken in the Black Hills region, Table 1
indicates numbered collection sites where each species was found. These
sites are plotted on a map of the region (Fig. 1 ) and are detailed in Table 2.
Also included in Table 1 is an indication of the general North American
distribution of each of the Black Hills species.

A rather striking pattern of biogeographic affinity is found for a large
proportion of Black Hills species. Black Hills records represent the
extreme range margin for 13 of the 19 species listed. Ten of the species are
western continental or eastern continental species whose respective
easternmost or westernmost fringes of their ranges apparently include
the Black Hills. For the three other species, the Black Hills distribution is
apparently along their northern or northeastern range margin.

Among the Baetidae, the species listed as widespread are generally
distributed with a broad east-west transcontinental pattern. In the case
of Callibaetis fluctuans and Diphetor hageni, these transcontinental pat-
terns are somewhat restricted north and south, with either a mid-latitude
or more northern pattern, respectively. Fallceon quilleri is widespread but
only as far east as Illinois (Burks 1953, as B. cleptis) and Louisiana (Lager
1985) and is mainly southwestern and south-central in distribution. Fall-
ceon quilleri may actually represent a complex of more than one species,
but requires further systematic study (Waltz and McCafferty 1987a). In
the case of Callibaetis pictus, although the range is signified as western in
Table 1 , it also extends into some south-central states as far east as Texas.
Those species listed as eastern generally have midwestern as well as
more eastern distributions.

Baetis brunneicolor, B.flavisriga, and B. intercalaris are eastern species
whose westernmost distribution limits are represented by disjunct pop-
ulations in the Black Hills. Acentrella insignificans and Dactylobaetis
cepheus are western species whose easternmost limits appear to be in the
Black Hills. With respect to D. cepheus, this may have been the species
that Lehmkuhl (1976) reported as Dactylobaetis sp. from Saskatchewan.
Whereas biogeographic affinities of Dactylobaetis are Neotropical

Vol. 101, No. 4, September & October 1990 195

(Traver and Edmunds 1968), those ofAcentrella are Holarctic (Waltz and
McCafferty 1987b). The Black Hills are along the northern fringe of the
known ranges of Callibaetis fluctuans and C. pictus. Fallceon quilleri has
been reported from Saskatchewan (Lehmkuhl 1976), but the Black Hills
distribution may be disjunct and along its northeastern range margin.
Eventually, the Callibaetis species probably will be found in Saskat-
chewan.

The Heptageniidae is the only other family with more than one
species listed for the region. Epeorus grandis and Nixe criddlei are western
species whose eastern limits are apparently the Black Hills. Records of
the latter species from Fort Collins, Colorado (Traver 1935) are nearly as
far east.

Of the remaining species, Caenis arnica is widespread and ubiquitous
(Provonsha 1990), and Tricorythodes minutus is also widespread except
for its absence from the Southeast (Berner 1977). Siphlonurus colum-
bianus and Ephemerella inermis are western species whose eastern limits
are apparently the Black Hills, although E. inermis is reported from
Saskatchewan (Lehmkuhl 1976), and I have seen S. columbianus from
north-central Colorado. Hilsenhoff (1987) reported that either E. inermis
or a sibling of it occurs in Wisconsin. I would not expect this species in
Wisconsin, given its absence in places such as Manitoba and Minnesota
(Flannagan and Flannagan 1982, Lager et al. 1982), but I have not
studied the Wisconsin material. Paraleptophlebia mollis is a mainly
northeastern species whose previous westernmost known distribution
included Manitoba and Minnestoa (Flannagan and Flannagan 1982.
Lager et al. 1982).

In conclusion, there are at least three aspects of the Black Hills
mayfly fauna that are noteworthy. First, the diversity of species is rela-
tively small. This perhaps could be explained by the fact that the region is
a somewhat insular montane area. Its isolation from other montane
areas may be an obstacle to dispersal for many mayfly species.

Second, the Black Hills represent the easternmost limits for certain
western species. The montane nature of the region would perhaps explain
the presence of such species that have populations located proximally in
the main body of the Rocky Mountains and that are relatively good
dispersers.

Third, and what might appear to be the most difficult to explain, four
eastern species have westernmost limits in the Black Hills. Baetis bnm-
neicolor and B. flavistriga are, however, mainly northeastern and upper
midwestern in distribution (Morihara and McCafferty 1979) as is the
Paraleptophlebia mollis (Traver 1935). Although mainly lowland species
in their primary range, their range extensions outside of this area are dis-

196 ENTOMOLOGICAL NEWS

junct in montane areas: Unzicker and Carlson (1982) reported B. brun-
neicolor from North Carolina; Peters and Warren (1966) reported B.
flavistriga (as B. levitans) from the Ozark Mountains of Arkansas, and
Berner ( 1977) reported this same species from Tennessee and North Car-
olina; and Traver (1937) confirmed P. mollis from North Carolina, and
McCafferry and Provonsha (1978) found it in the Ouachita Mountains
of Arkansas. This type of pattern in North America probably indicates a
relict distribution, where certain cold-water species have become restric-
ted to northeastern regions and isolated in certain disjunct montane
areas outside the Northeast as a result of glacial events during the
Pleistocene.

Although the distribution oi Baetis intercalaris is similar to the three
other eastern, Black Hills species, this species is found in more divese
lowland and upland locations in the East, including areas such as
Florida and Alabama (Morihara and McCafferry 1979). The western dis-
junction in this instance is therefore more difficult to interpret. The
species does occur in the lower Appalachians and the Ozarks (Morihara
and McCafferty 1979), but these are not disjunctions. Perhaps the species
has been a better post-glacial disperser, and since it appears more ubi-
quitous, perhaps will be found in lowland areas adjacent to, and east of,
the Black Hills. This apparent anomaly, however, may not actually exist
if it is an artifact of our inexact taxonomy. Baetis flavistriga and B. inter-
calaris are closely related and morphologically cryptic as larvae, being
separable only by color pattern (Morihara and McCafferty 1979). There-
fore, it is possible that what we have determined to be B. intercalaris in the
Black Hills is a color variation of B. flavistriga that resembles B. inter-
calaris in other areas. Baetis ochris is another closely related and similar
species that may be involved.

ACKNOWLEDGMENTS

I thank A. V. Provonsha, Purdue University, and B. L. Heath, Lafayette, IN, for their
participation in the collecting phase of this research, R. D. Waltz, Indiana Division of
Entomology and Plant Pathology, Indianapolis, for his aid in identifying certain
specimens, and D. W. Bloodgood, Purdue University, for preparation of the regional map.

Vol. 101, No. 4, September & October 1990

197

Spearfish
LA WRENCE

PENNINGTON

Keystone
15

NEBRASKA

Fig. 1. Black Hills region of southwest South Dakota, with plotted mayfly collection
sites.

198 ENTOMOLOGICAL NEWS

Table 1. Distribution of the Ephemeroptera species of the Black Hills region of South
Dakota. Asterisked species are newly reported for South Dakota. Numbers refer to sites
plotted in Fig. 1. and detailed in Table 2.

SPECIES SITES N.A DISTRIBUTION
Baetidae

Acentrella insignificans 6 west

Baetis brunneicolor 5,15 east

Baetis flavistriga 5,11,15,16,17 east

* Baetis intercalaris 11,16,17 east

Baetis tricaudatus 6, 11, 12, 13, 14, 16 wide

* Callibaetis ferrugineus 2 wide

* Callibaetis fluctuans 10 wide

* Callibaetis pallidus 1 wide

* Callibaetis pictus 3. 7 west

* Dactylobaetis cepheus 6 west

Diphetor hageni 12,16 wide

Fallceon quilleri 6, 8 wide
Caenidae

* Caen is arnica 17 wide
Ephemerellidae

Ephemerella inermis 4, 11, 12, 16 west
Heptageniidae

* Epeorus grandis 13,14,15,16 west

* Nixe criddlei 5 west
Leptophlebiidae

* Paraleptophlebia mollis 11,12,15,16 east
Siphlonuridae

* Siphlonurus columbianus 15 west
Tricorythidae

* Tricorythodes minutus 5, 8, 9, 11. 15 wide

Table 2. Details of Black Hills mayfly collection sites.

1. Custer Co, Paulson's Pond, 8 mi SW Pringle.

2. Custer Co, Bismark Lake Campground.

3. Custer Co, Stockade Lake at US 16A.

4. Custer Co, Beaver Cr 1 mi E Rifle Pit Historical Site.

5. Custer Co, French Cr at Hazelrodt Picnic Grounds.

6. Fall River Co, Hot Brook at Chataqua Park, Hot Springs.

7. Fall River Co, Hawkwright Cr at Red Canyon.

8. Fall River Co, Fall R at US 385 nr confluence of Cheyenne R.

9. Fall River Co, Cheyenne R at US 385.

10. Fall River Co. Fall Creek Reservoir 2 mi N Hot Springs.

11. Lawrence Co, Boxelder Cr at Boxelder Forks Campground 1 mi W Nemo.

12. Lawrence Co, Jim Cr 3 mi S Nemo.

13. Lawrence Co, Spearfish Cr 1 mi N Cheyenne Crossing at US 14A.

14. Lawrence Co, Iron Cr at US 14A 12 mi S Spearfish.

15. Pennington Co, Grizzly Bear Cr at Keystone.

16. Pennington Co, Rapid Cr at mouth of Dark Canyon.

17. Pennington Co, Rapid City at light.

Vol. 101, No. 4. September & October 1990 199

LITERATURE CITED

Allen, R. K. and G. F. Edmunds, Jr. 1965. A revision of the genus Ephemerella (Ephemer-

optera: Ephemerellidae) VIII. the subgenus Ephemerella in North America. Misc. Publ.

Entomol. Soc. Am. 4: 243-282.
Bednarik, A. F. and W. P. McCafferty. 1979. Biosystematic revision of the genus

Stenonema (Ephemeroptera: Heptageniidae). Canad. Bull. Fish. Aq. Sci. 201: 1-73.
Berner, L. 1977. Distributional patterns of southeastern mayflies (Ephemeroptera). Bull.

Fla. St. Mus.. Biol. Sci. 22: 1-55.
Burks, B. D. 1953. The mayflies, or Ephemeroptera, of Illinois. Bull. 111. Nat. Hist. Surv. 26:

1-216.
Flannagan, P. M. and J. F. Flannagan. 1982. Present distribution and post-glacial origin

of the Ephemeroptera, Plecoptera. and Trichoptera of Manitoba. Man. Dept. Nat. Res.,

Fish. Tech. Rep. 82-1: 1-79.
Hilsenhoff, W. L. 1987. An improved biotic index of organic stream pollution. Great

Lakes Entomol. 20: 31-39.
Kondratieff, B. C. and J. R. Voshell, Jr. 1984. The North and Central American species of

Isonychia (Ephemeroptera: Oligoneuriidae). Trans. Am. Entomol. Soc. 110: 129-244.
Lager, T. M. 1985. Range extensions and ecological data for southern United States

Ephemeroptera. Proc. Entomol. Soc. Wash. 87: 255-256.
Lager, T. M., M. D.Johnson and W. P. McCafferty. 1982. The mayflies of northeastern

Minnesota (Ephemeroptera). Proc. Entomol. Soc. Wash. 84: 729-741.
Lehmkuhl, D. M. 1976. Mayflies. Blue Jay. 34: 70-81.
McCafferty, W. P. and A. V. Provonsha. 1978. The Ephemeroptera of mountainous

Arkansas. J. Kans. Entomol. Soc. 51: 360-379.
Morihara, D. K. and W. P. McCafferty. 1979. The Baetis larvae of North America

(Ephemeroptera: Baetidae). Trans. Am. Entomol. Soc. 105: 139-221.
Peters, W. L. and L. O. Warren. 1966. Seasonal distribution of adult Ephemeroptera in

northwestern Arkansas. J. Kans. Entomol. Soc. 39: 396-401.
Provonsha, A. V. 1990. A revision of the genus Caenis in North America (Ephemeroptera:

Caenidae). Trans. Am. Entomol. Soc. in press.
Traver, J. R. 1935. Systematic, pp. 237-739. In The biology of mayflies, by J. G. Needham. J.

R. Traver and Y-C. Hsu. Comstock Publ. Co., Ithaca, N. Y.
Traver, J. R. 1937. Notes on the mayflies of the southeastern states (Ephemeroptera). J.

Elisha Mitchell Sci. Soc. 53: 27-86.
Traver, J. R. and G. F. Edmunds, Jr. 1968. A revision of the Baetidae with spatulate-

clawed nymphs (Ephemeroptera). Pac. Ins. 10: 629-677.
Unzicker, J. D. and P. H. Carlson. 1982. Ephemeroptera. pp. 3.1-3.97. In A. R. Brigham,

W. U. Brigham and A. Gnilka (eds.). Aquatic insects and oligochaetes of North and

South Carolina. Midwest Aq. Enterprises, Mahomet, 111.
Waltz, R. D. and W. P. McCafferty. 1987a. New genera of Baetidae for some Nearctic

species previously included in Baetis Leach (Ephemeroptera). Ann. Entomol. Soc.- Am.

80: 667-670.
Waltz, R. D. and W. P. McCafferty. 1987b. Systematics of Pseudocloeon , Acentrella,

Baetiella, and Liebebiella, new genus (Ephemeroptera: Baetidae). J. New York Entomol.

Soc. 95: 557-568.

200 ENTOMOLOGICAL NEWS

ANTHOPOTAMUS, A NEW GENUS FOR

NORTH AMERICAN SPECIES

PREVIOUSLY KNOWN AS POTAMANTHUS

(EPHEMEROPTERA: POTAMANTHIDAE)1' 2

W.P. McCafferty, YJ. Bae3

ABSTRACT: Comparative studies of all life stages of potamanthid mayflies worldwide
have indicated that North American species historically placed in the genus Potamanthus
are generically distinct from all Palearctic Potamanthus, including the type of the genus, the
European species P. luteus (Linn.). Close phylogenetic relationships of the Nearctic species
of Potamanthidae are with East Asian genera, but not Palearctic Potamanthus. The new
genus Anthopotamus [type species = A. verticis (Say), new combination] is therefore estab-
lished for the Nearctic fauna of Potamanthidae. A brief synopsis of the new genus is
provided, and a biogeographic origin of the new genus is suggested.

The genus Potamanthus Pictet in North America was most recently
reviewed by McCafferty (1975). Although the classification (McCafferty
and Edmunds 1979) and the phylogeny (McCafferty 1979) of Potamanthidae
within the higher groups of Ephemeroptera have been recently pre-
sented, the systematics within the family have not been studied
comprehensively. Members of this clearly monophyletic family, com-
monly known as hacklegills (McCafferty 1981), are distributed in the
Holarctic and Oriental Realms, with several genera occurring in East
Asia, but only Potamanthus occurring in North America. We are
currently conducting a revisionary study of both species and genera of
the Potamanthidae. This has been possible only by bringing together
material from throughout the world for comparative study.

Potamanthus, as it has been constituted, is known from North
American and Eurasia. The type of the genus is P. luteus (Linn.), a
common European species that was first described in Ephemera in 1767.
We have studied adults and larvae of Nearctic and Palearctic Potamanthus
in detail, including/1, luteus. Our comprehensive comparisons with other
potamanthids from the entire range of the family clearly show that
species in North American are distinct from P. luteus. The degree of
demarcation is equivalent to, or more than, that found in well-established,
valid genera in the family. Asian species of Potamanthus do, however,
agree with the type concept of the genus. Thus, the North American
species constitute a separate and distinct grouping that require classification

Deceived December 22, 1989. Accepted March 27, 1990.
Purdue Experiment Station Journal No. 12335.
•^Department of Entomology, Purdue University, West Lafayette, IN 47907.

ENT. NEWS 101(4): 200-202, September & October, 1990

Vol. 101, No. 4, September & October 1990 201

in a new genus. The most compelling reason for removing the Nearctic
species from Potamanthus is our finding that they are phylogenetically
removed from Palearctic Potamanthus and are instead more closely
related to other East Asian genera.

Because ecological and behavioral data dealing with various North
American potamanthid species are about to be made available for publi-
cation (Bae and McCafferty unpublished, D.C. Tarter, pers. comm.) and
publication of the family revision will not be forthcoming for at least
another two years, it is appropriate to report this systematic finding and
make a new generic name available at this time. Therefore, a new genus
is established herein and a brief synopsis and discussion of the taxon is
given.

Anthopotamus, new genus

Type Species. Potamanthus verticis(Say), 1839:42, orginally as Baetis.
Species Included. Anthopotamus diaphanus (Needham), A. distinctus (Traver), A.
inequalis (Needham), A. mvops (Walsh), A. neglectus (Traver), A. rufous (Argo), A. verticis
(Say), and A. walkeri (Ide),' all NEW COMBINATIONS.
Distribution. Eastern half of North America.

Etymology. The name Anthopotamus is a masculine name taken from the Greekanthos
(flower) and potamos (river). The allusion to a "river flower" is analogous to the other
generic names in Potamathidae, which also denote river flowers.

Diagnosis. Larvae of Anthopotamus have long, well-developed
mandibular tusks, highly developed mouthpart setation, and well-
developed rows of long setae on the foretibiae (see Fig. 3 in McCafferty
1975 and Fig. 7.42 in McCafferty 1981). Potamanthus larvae, on the other
hand, have rudimentary tusks and poorly developed mouthpart and
foretibiae setation in comparison (e.g. see Fig. Ib in Macan 1979). Adults
of Anthopotamus can be distinguished from those of Potamanthus by the
generally smaller compound eyes of the males (ratio of eye diameter to
dorsal distance between compound eyes is less than 0.2), as well as a
much longer prothorax, and generally lighter coloration.

Relationships. Our cladistic analysis, details of which will be pub-
lished as part of the revisionary monograph, indicate that the Old World
genus Potamanthus (sensu novum) represents an ancestral, early branching
lineage within the Potamanthidae. Its sister lineage contains all other
Potamanthidae, a grouping consisting of East Asian genera as well as
Anthopotamus. Within this later Ymeage, Anthopotamus is intermediate in
position between the more ancestral Potamanthodes and the more derived
Rhoenanthus. Anthopotamus is not closely related phylogenetically to
Potamanthus, wherein its species were previously included.

Most probably, an Anthopotamus-Mke ancestral lineage spread to
North America via Beringia in pre-Pleistocene times. Vicariance of this

202 ENTOMOLOGICAL NEWS

Holarctic, Arcto-Tertiary distribution would have probably paralleled
the phyletic split into a Nearctic "Anthopotamus" lineage and the
Palearctic "Rhoenanthus" lineage. If more derived elements of
Potamanthidae were very widespread in the Holarctic during the
Tertiary, then perhaps glacial events during the Pleistocene would ex-
plain the present absence of this element in Europe and western North
America. Such Arcto-Tertiary and subsequent patterns are well documented
in plant and animal genera, e.g. see review of Pielou (1979).

ACKNOWLEDGMENTS

We thank numerous colleagues for providing fresh study material of Potamanthus
luteus: J. Alba-Tercedor, Granada, Spain; M.T. Gillies, Sussex, U.K.; N.J. Kluge, Leningrad,
USSR; R. Sowa and M. Klonowska-Olejnik, Krakow, Poland; and AG.B. Thomas,
Toulouse, France.

LITERATURE CITED

Macan, T.T. 1979. A key to the nymphs of the British species of Ephemeroptera with notes

on their ecology. Freshwat. Biol. Assoc. 20: 1-80.
McCafferty, W.P. 1975. The burrowing mayflies (Ephemeroptera: Ephemeroidea) of the

United States. Trans. Amer. Entomol. Soc. 101: 447-504.
McCafferty, W.P. 1979. Evolutionary trends among the families of Ephemeroidea. Proc.

2nd Internal. Conf. Ephem., Krakow, 1975: 45-50.

McCafferty, W.P. 1981. Aquatic entomology. Jones and Bartlett Publ., Boston. 448 pp.
McCafferty, W.P. and G. F. Edmunds, Jr. 1979. The higher classification of the

Ephemeroptera and its evolutionary basis. Ann. Entomol. Soc. Amer. 72: 5-12.
Pielou, E.G. 1979. Biogeography. John Wiley & Sons, New York. 351 pp.

Vol. 101, No. 4, September & October 1990 203

CYTOLOGY OF LEPTOGLOSSUS ZONATUS
(HEMIPTERA: COREIDAE)1

Richard J. Packauskas''

ABSTRACT: The diploid chromosome complement of Leptoglossus zonatus (Dallas)
males is shown to consist of 21 chromosomes: 1 8 autosomes, 2 microchromosomes, and an
X chromosome. There are two large homologous pairs (4-5 microns in length). The
remaining autosomes and the X chromosome range from 2-3 microns in length, and the
microchromosomes (which are round) are 1 micron in diameter. Major meiotic events of
the species are briefly characterized.

The Coreidae is a very large and diverse family, and many species
within it have been examined cytologically. Most coreids investigated
have a pair of microchromosomes and an XO sex mechanism. The
chromosomes are holokinetic (diffuse centromeric), as are those of all
Hemiptera (Thomas 1987). Coreid chromosome numbers range from 2n
= 13 to 2n = 28; the most common number is 2n = 21 (Ueshima 1979).
The genus Leptoglossus Gue'rin belongs in the coreine tribe Anisoscelidini.
some of whose members have been investigated cytologically (Piza 1 945,
1956; Wilson 1907, 1909).

Here I examine the cytology of Leptoglossus zonatus (Dallas) and
relate this to previous studies. °The chromosome number of this species
has never been published, nor have meiotic events in this genus been
charactetized.

MATERIALS AND METHODS

The figures and observations were based upon preparations made
from 3 male fifth instars collected in Panama and alcohol preserved
(70% ethyl) in August of 1987. A modification (Jane O'Donnell, unpubl.)
of a technique (Ueshima 1963) for preparation of alcohol-preserved
specimens was used with good results even after a year of storage: 1)
testes were dissected out in alcohol and placed for 1 2 hours in the fixative
isopropyl carnoy (3 parts pure isopropyl alcohol, 1 part glacial acetic
acid); 2) they were transferred to glacial acetic acid and heated gently
over an alcohol lamp for 5-10 minutes; 3) the preparation was allowed to
cool to room temperature, and then placed in a drop of aceto-carmine; 4)

^Received January 20, 1990. Accepted April 2. 1990.

-Department of Ecology and Evolutionary Biology. Box U-43, The University of
Connecticut, Storrs, Ct 06268

ENT. NEWS 101(4): 203-206. September & October. 1990

204 ENTOMOLOGICAL NEWS

a cover slip was applied and the material squashed with a few hard taps
of a pencil eraser. All preparations were subsequently mounted in
Diaphane to make them permanent (Sharma and Sharma 1972). Drawings
were made with the aid of an ocular grid, and measurements with an
ocular micrometer.

OBSERVATIONS

Observations were recorded as drawings; only brief explanatory
comments are offered.

Spermatogonial phases are easily seen in L. zonatus, but these are not
always easily analyzed. The spermatogonial chromosomes do not spread
well in a squash and can only be counted in broken cells, but they are
nevertheless useful in establishing a diploid count. Figure 1 is of a
spermatogonial phase, and shows 21 chromosomes.

Prophase events are also not easily analyzed in the Heteroptera
(Ueshima 1979), but are useful in establishing the size and shape of the
sex chromosomes. Figures 2 and 3 clearly show a single heteropyconotic
chromosome (the X chromosome).

Cells which are found in early diakinesis show chiasmata formation
among the autosomes. Figure 4 shows terminalization of chiasmata
among homologs of each of the autosomes. The microchrosomes (m) are
plainly visible as the two smallest chromosomes. The X chromosome (x)
can now be determined by comparison with Figures 2 and 3. Nine
homolog pairs can be seen, as can the X chromosome and the two
microchromosomes. Figure 5 shows late diakinesis, when terminalization
of chiasmata is complete and the autosome homologs have reached their
maximal lengths for this stage. This stage was the most useful in estab-
lishing a count. Thus, in L. zonatus we clearly see 9 pairs of autosomes,
the X chromosome, and two microchromosomes, for a total of 21.
Chromosomes were measured in 7 cells in late diakinesis. Two of the
autosome homolog pairs were consistently larger than the rest, 4-5 microns
in length. The remaining autosome pairs and the X chromosome were 2-
3 microns in length, and the microchromosomes (which are round) were
1 micron in diameter.

In metaphase I (Figures 6 and 7), the autosome pairs and the X
chromosome formed a ring around the two closely appressed micro-
chromosomes. Finally, in a metaphase II (Figure 8), the same type of ring
formed around the single diffuse microchromosome.

Vol. 101, No. 4. September & October 1990

205

-•*

13.

^ / m

, $r*

Figures: 1. spermatogonial phase. 2. prophase, zygotene. 1. prophase. leptotene
= diplotene, 4. early diakinesis, 5. late diakinesis. 6. metaphase I. polar view.
7. metaphase I. lateral view. 8. metaphase III. polar view, m == michrochromosome.
X = X chromosome. Scale bar = 5 microns.

206 ENTOMOLOGICAL NEWS

DISCUSSION

Other species of Leptoglossus have been looked at cytologically,
including L. dilaticollis Guerin (Piza 1956), L. gonagra (Fabricius) (Piza
1945),L. phyllopus (Linnaeus) (Wilson 1 909), and L. stigma (Herbst) (Piza
1956). All have a diploid complement of 21 chromosomes in the male
(ISA + 2m + X). Leptoglossus zonatus, therefore, falls into this already
established pattern.

These observations do not depart significantly from the more detailed
description of meiotic events in Coreus marginatus L. (a member of the
tribe Coreini) (Nokkala 1986); but the procedure and drawings herein
should provide a novice entry into this neglected field of cytology. Only
chromosome numbers were reported in all previous studies of this genus.
It is hoped that the brief characterization of meiotic events within this
species and the finding of disparate chromosome sizes will prove to be
of value in further analysis of the cytology and the complex evolutionary
history of this diverse and cytologically poorly known family.

ACKNOWLEDGMENTS

The author thanks Roberto Ibanez for collecting the specimens. Carl W. Schaefer for
reviewing a draft of this manuscript. Jane E. O'Donnell for much discussion, and Gregory
J. Anderson without whom this study would not have happened.

LITERATURE CITED

Nokkala, S. 1986. The mechanisms behind the regular segregation of the m-chromosomes

in Coreus marginatus L. (Coreidae. Hemiptera). Hereditas 105: 73-85.
Piza, S. deT. 1945. Estudoscitoldgosem Hemipterisda familiaCorediae. LuizdeQueiroz

2: 119-148.
Piza, S. de T. 1956. A note on chromosomes in three coreid bugs. Rev. Agric. (Lima).

31:32.
Sharma, A.K. and A. Sharma. 1972. Chromosome techniques. Butterworths & Co. Ltd..

London, and University Park Press. Baltimore. 575 pp.
Thomas, D.B. 1987. Chromosome evolution in the Heteroptera (Hemiptera):

Agmatoploidy versus aneuploidy. Ann. Entomol. Soc. Am. 80: 720-730.
Ueshima, N. 1963. New techniques in cytotaxonomy. Chromosome Inf. Service 4: 17-18.

Ueshima, N. 1979. Hemiptera II: Heteroptera. Animal Cytogenetics. Vol. 3: Insecta 6.

Gebruder Borntraeger. Berlin.

Wilson, E.B. 1907. The case ofAnasa tristis. Science 25: 191-193.
Wilson, E.B. 1909. Studies on chromosomes. IV. The accessory chromosome in

Syromastes and Pyrrhocoris, with a comparative review of the types of sexual difference

of the chromosome groups. J. Exp. Zool. 6:69-99.

Vol. 101, No. 4. September & October 1990 207

NEW IASSINAE (HOMOPTERA: CICADELLIDAE)
FROM MEXICO AND PERU1'2

H. Derrick Blocker3

ABSTRACT: Two new species of lassinae, Gargaropsis (Curistuva) poravis and Baldriga deor-
nitha, are described from Mexico. These were taken from foodballs regurgitated from the
crops of swifts that were captured and released unharmed in Chiapas, Mexico. Another
new species of lassinae, Garlica freytagi, is described from Peru.

Two new species of lassinae from foodballs regurgitated from the
crops of white-collared swifts, Streptoprocne zonaris (Apodiformes:
Apodidae) that were captured and released unharmed are described
from Chiapas, Mexico. This population of swifts was located at Cueva el
Chorreadero, 12km N.E. of Chiapa de Corzo, Chiapas. Three species
were represented by the three specimens of lassinaer recovered. These
were: Gargaropsis (G.) confinis Blocker, G. (Curistuva) poravis, n. sp., and
Baldriga deomitha, n. sp. Specimens were in excellent condition, although
some appendages were missing and the body was often coated with
saliva or mucus which often coats these foodballs. In addition, a new
species, Garlica Freytagi, is described from specimens collected in Peru.

Gargaropsis subgenus Curistuva Blocker

Gargaropsis (Curistuva) Blocker, 1979:14. Type species: Gargaropsis adibilis Blocker by

original designation.

Four species are presently assigned to this subgenus:
adibilis Blocker, 1975:561. Mexico.
filialis Blocker, 1975:563. Mexico.
poravis Blocker, new species. Mexico.
wilburi Blocker, 1975:564. Mexico.

Gargaropsis (Curistuva) poravis, n. sp.

(Figs 1-3)

Length of male 4.8 mm; head width 1.5mm; pronotal width 1.9mm. Female unknown.

Color stramineous, body and wings covered with small dark brown spots, larger dark
brown areas on scutellum and claval area at apex of forewings; face, legs, and venter of
thorax with some fuscous marking.

Vertex length slightly longest medially; anteclypeus not attaining margins of genae;
vein in appendix of forewing evanescent apically; hind femoral chaetotaxy 2-2-1 with one
of second pair reduced.

1 Received February 28, 1990. Accepted March 27, 1990.

2 Contribution 90-382-J from the Kansas Agricultural Experiment Station.
Department of Entomology, Kansas State University, Manhattan, KS 66506.

ENT. NEWS 101(4): 207-210, September & October, 1990

208 ENTOMOLOGICAL NEWS

Pygofer process acute apically; style bifid apically (fig. 1); connective straplike; aedeagus
with shaft tapered apically (fig. 2), bifid apically in dorsal view with apical processes slen-
der, curved cephalad (fig. 3), dorsal apodeme with arms broadly U-shaped.

Holotype male, from Mexico, Chiapas, Chiapa de Corzo, 3 vi- 16 vii 1985 (D. F. Whitacre).
food of (Aves) Streptoprocne zonaris, in The University of Kansas collection.

Gargaropsis (Curistuva) poravis is related to adibilis and keys to this
species in Blocker (1975). It can be distinguished by the wider, apically
tapered shaft of the aedeagus and by the shape of the apex of the style.

Baldriga Blocker

Baldriga Blocker, 1979:15. Type species: Baldriga knutsoni Blocker by original designation.

Six species are presently assigned to this genus:
deornitha Blocker. new species. Mexico.
galida Blocker, 1982:641. Mexico.
knutsoni Blocker, 1979:15. Mexico.
nielsoni Blocker. 1982:640. Panama.
painteri (Blocker). 1975:563. Mexico.
naasaca Blocker, 1979:16. Brazil

Baldriga deornitha, n. sp.

(Figs. 4-6)

Length of male 6.6 mm; head width 2.2 mm; pronotal width 3.0 mm. Female unknown.

Color stramineous dorsally; body and forewings with numerous fuscous dots; venter of
thorax fuscous.

Vertex parallel-margined; anteclypeus attaining margin of gena; hind femoral
chaetotaxy 2-1-1 or 2-1 (in same specimen).

Pygofer processes acute apically; style abruptly narrowed near apex, unique (fig. 4);
connective straplike. slightly arched in lateral view; aedeagus with shaft excavated in
lateral view (fig. 5), apex not bifid in dorsal view (fig. 6), apical processes exceeding
width of shaft.

Holotype male, from Mexico, Chiapas, Chiapa de Corzo, 3 vi-16 vii 1985. (D. F.
Whitacre), food of (Aves), Streptoprocne zonaris, in The University of Kansas collection.

Baldriga deornitha is related to painteri but can be distinguished by its
larger size and the unique shape of the apex of the style.

Garlica Blocker

Garlica Blocker, 1976:5 19. Type species: Garlica hepneri Blocker by original designation and

monotypy.

Five species are presently assigned to this genus:
exila Blocker, 1979:25. Brazil
freytagi Blocker, new species. Peru.
hepneri Blocker, 1976:519. Panama.
margina Blocker. 1982:645. Panama.
punctulata (Linnavuori), 1956:20. Brazil.

Vol. 101. No. 4. September & October 1990

209

Figs. 1-9. 1-3. Gargaropsis (Curistuva) poravis. \. Style, broad aspect. 2. Aedeagus, lateral
view. 3. Aedeagus. dorsal view. 4-6. Baldriga deorniiha. 4. Style, broad aspect. 5. Aedeagus,
lateral view. 6. Shaft of aedeagus, dorsal view. 7-9. Garlicafreytagi. 1. Style, broad aspect.
8. Aedeagus. lateral view. 9. Plate, ventral view.

210 ENTOMOLOGICAL NEWS

Garlica freytagi, n. sp.

(Figs. 7-9)

Length of male 4.5 mm; head width 1.9mm; pronotal width 2.0 mm. Female unknown.

Color stramineous with darker brown pattern on vertex, pronotum, and scutellum;
numerous dark setae on forewing, appendix and 1st apical cell bordered anteriorly by
fuscous veins; face with fuscous lines on either side of midline.

Vertex parallel-margined, not as wide as pronotum; ocelli more than 5X their diameter
from eye; hind femoral chaetotaxy 2:2:1 with one of second pair reduced; sternum VIII
elongate, extending beneath genital segment.

Pygofer with short fringe of microsetae on caudoventral margin and small tuft of micro-
setae on inner surface at same location; plates with two tufts of microsetae (fig. 9); pseudo-
styles inconspicuous, membranous; (styles with apex elongate, tapered, with two small
spine-like processes (fig. 7); connective rectangular; aedeagus with shaft bulbous in apical
1/2 (fig. 8).

Holotype male, PERU, Madre de Dios, along Manu de dios River, 12 ° S, 25 Aug 1988,
Manu Nat. Park, Cocha Salvador, at light (T. Myers), in The California Academy of Sciences
collection. Three male paratypes, same data, at The University of Kansas, The American
Museum of Natural History, and The University of Kentucky.

Garlicafreytagi, n. sp. is related to exila but can be distinguished by the
shape of the plates that are widest at the base and the presence of a small
tooth on the foot-shaped style apex. I take pleasure in naming this
species for my longtime and trusted colleague, Paul Freytag.

ACKNOWLEDGMENTS

I acknowledge the assistance of David Whitacre, University of California at Davis, who
furnished interesting material from the crops of Mexican swifts and Paul Freytag, Univer-
sity of Kentucy, who lent the specimens of Garlica.

LITERATURE CITED

Blocker, H. D. 1975. The Mexican genus Gargaropsis (Homoptera: Cicadellidae: las-

sinae). Ann. Entomol. Soc. America 68:561-564.
1976. Three new genera of Neotropical laassinae (Homoptera: Cicadellidae).

Ann. Entomol. Soc. America 69:519-522.

1979. The lassinae (Homoptera: Cicadellidae) of the Western Hemisphere. J.

Kansas Entomol. Soc. 52:1-70.

_. 1982. New species of lassinae from the Neotropics (Homoptera: Cicadellidae). J.

Kansas Entomol. Soc. 55:639-650.
Linnavuori, R. 1956. Neotropical Homoptera of the Hungarian National Museum and
some other European museums. Ann. Entomol. Fennici 23:144-150.

Vol. 101, No. 4, September & October 1990 211

NOTIOPHILUS PALUSTRIS

(COLEOPTERA: CARABIDAE), A EURASIAN

CARABID BEETLE NEW TO NORTH AMERICA1

A. Larochelle , M.-C. Lariviere^

ABSTRACT: The Eurasian carabid beetle Notiophilus palustris is recorded for the first time
for North America. Forty-four specimens have been found in Nova Scotia and Prince
Edward Island, Canada, in 1967 and 1987. While macropterous forms are rare in Europe,
individuals found in North America seem all to be macropterous, suggesting a recent
immigration.

The insect collection of the Nova Scotia Museum of Science
(Halifax) contains one specimen of the Eurasian carabid beetle
Notiophilus palustris (Duftschmid, 1812), collected by Ken Neil from
Nova Scotia: Halifax Co., Armdale, 1. VII. 1967. Forty-three individuals
of this species have also been discovered from the following localities:
Nova Scotia: Pictou Co.: Merigomish Island, 20.VII.1987 (25 exx.).
Prince Edward Island: Kings Co.: Basin Head, 13.VII.1988 (one ex.);
Panmure Island Provincial Park, 15.VII.1987 (one ex.); Red Point Provincial
Park, 15.VII.1987 (one ex.); Souris Beach Provincial Park, 14.VII.1987
(one ex.). Queens Co.: Lord Selkirk Provincial Park, 14.VII.1987 (one
teneral ex.); Wood Island Provincial Park, 13.VII.1987 (13 exx.). This is
the first record of this Eurasian carabid for North America.

All specimens have been found on sandy sea beaches, under vegetal
debris. According to Lindroth (1985), the species is hygrophilous and
usually occurs in rather shady places on humus-rich ground; it is found
in deciduous woods among the litter and in mosses as well as in open
habitats such as meadows and marshes, with high and dense vegetation.
In Europe, it is most common in spring when reproduction takes place,
but also in fall when the new adult generation emerges.

Notiophilus palustris has apparently been accidentally introduced
into North America, perhaps transported by ship. The species seems to
be well established in the Atlantic Provinces of Canada as demonstrated
by the large number of individuals found in numerous collecting local-
ities and the presence of a teneral individual which indicates that
breeding occurs locally. While macropterous forms are rare in Europe

,! Received February 4, 1990. Accepted March 31, 1990

2Lyman Entomological Museum and Research Laboratory, Macdonald College of
McGill University, Ste-Anne-de-Bellevue, Quebec H9X ICO, Canada.
Department of Entomology, Macdonald College of McGill University, Ste-Anne-de-
Bellevue, Quebec H9X ICO, Canada.

ENT. NEWS 101(4): 211-212, September & October. 1990

212 ENTOMOLOGICAL NEWS

(Lindroth, loco citato), all specimens examined (at least 20) have fully-
developed inner wings; this condition suggests a recent immigration into
North America. Moreover, the area has been well surveyed in the past
and while the beetle is conspicuous, it has never been recorded. This
carabid should spread rapidly in northeastern North America due to its
fully-developed, probably functional, wings and its eurytopic
characteristics.

The species is characterized as follows: Broad, black with shiny
cupreous luster; entire tibiae, base of palpi and four basal segments of
antennae pale; anterior diverging frontal furrows; eyes very large; head
wider than prothorax; pronotal sides strongly constricted basally and
rounded. Second elytral interval just behind the middle, more than three
times as wide as third interval; each elytron with two dorsal punctures
and two preapical punctures; intervals not reticulated; striae densely and
deeply striated, especially behind shoulder. Body length: 5-6.1 mm.

In Lindrotrfs key (1961) to North American Notiophilus, the species
will trace to Couplet 8 modified in the following manner:

8. Tibiae black. Elytra with 2. interval hardly broader than the following two together
JV. simulator Fall

- Tibiae yellowish brown. Elytra with 2. interval at least as broad as the three following
together 9

9. Second and third elytral striae obliterated before apex. Elytra quite dark or with ill-
defined pale apical vitta 9.1

- Second and third elytral striae evident to apex. Elytra always with well defined pale apical
vitta N. biguttatus (Fabricius)

9.1 Each elytron with 2 dorsal punctures. Head much wider than prothorax which is
strongly constricted basally. 5-6.1 mm. Nova Scotia and Prince Edward Island
N. palustris (Duftschmid)

- Each elytron with a single dorsal puncture. Head as wide as prothorax which is devoid of
basal contriction. Smaller: 3.4-4.7 mm. Eastern United States. .../V. novemstriatus LeConte

ACKNOWLEDGMENTS

We wish to thank Y. Bousquet (Canadian National Collection, Ottawa) for
confirming our identification of the species and for sending information concerning the
specimen contained in the Nova Scotia Museum. Sincere thanks also to V.R. Vickery
(Macdonald College of McGill University. Ste-Anne-de-Bellevue, Quebec) for revising the
manuscript.

LITERATURE CITED

Lindroth, C.H. 1961. The ground-beetles of Canada and Alaska. Opusc. Entomol.,

Suppl. 20: 200 pp.
Lindroth, C.H. 1985. The Carabidae (Coleoptera) of Fennoscandia and Denmark.

Fauna Entomol. Scand.. 15(1): 255 pp.

Vol. 101, No. 4, September & October 1990

A NEW MASTOGENIUS
(COLEOPTERA:BUPRESTIDAE) FROM JAMAICA1

Gary V. Manley^

ABSTRACT: A new species of Mastogenius (M. howdenontm) is described from Jamaica,
West Indies. The genus is widely scattered in the West Indies, but this is the first species
from Jamaica.

A series of Mastogenius specimens given to me recently by
Dr. H. Howden (Carleton University) proved to be undescribed. The
species is similar to M. coyolensis Manley from Honduras.

Mastogenius howdenorum

Manley, new species
(Figs. 1, 2 & 3)

HOLOTYPE, male: Elongate oval, shining, pronotum and elytra equally clothed with
dense long whitish recumbent pubescence, dorsal surface uniformly aeneus, pronotum
slightly brighter than elytra, ventral surface uniformly black, legs slightly reddish laterally,
tarsi brownish-red.

HEAD slightly convex, faint shallow round depression on frons, surface coarsely punctate,
clothed with recumbent hairs; eyes with inner margins parallel, not converging toward
each other at apex; antenna serrate from fourth segment, extending beyond the pronotum,
clothed with scattered hairs, segment four more than twice the length of segment three.
PRONOTUM uniformly convex, wider than long, narrower at apex than at base, widest
near middle; sides broadly arcuately diverging from apical angles to near middle, then
obliquely converging to posterior humeral angles; anterior margin slightly broadly
emarginate; posterior margin truncate, straight; marginal and submarginal carina more
widely separated near middle of pronotum, marginal carina not reaching anterior border
of pronotum, sloping almost to submarginal carina and ending just prior to anterior
margin of pronotum; surface uniformly, densely punctate; densely clothed with moderately
long white setae.

SCUTELLUM black, triangular, and smooth.

ELYTRA convex, slightly flattened along median suture, as wide as pronotum at base;
sides nearly parallel from humeral angles to near middle, then arcuately converging to tips,
which are truncate; oblique transverse depression near bases; surface irregularly punctate,
crenulate, densely clothed with relatively long white setae.

UNDERSIDES uniformly black, convex, punctate; abdomen clothed with scattered recum-
bent white setae; last sternite punctate, truncate at apices, clothed with scattered setae of
various lengths; prosternum punctate, scattered hairs most common near the middle,
without antennal grooves; posterior margin of hind coxal plates strongly sinuate, inner
margin wider than external margin.

'Received March 29, 1990. Accepted April 18, 1990.
214174 Hoffman Road, Three Rivers, Michigan 49093

ENT. NEWS 101(4): 213-215, September & October. 1990

214

ENTOMOLOGICAL NEWS

SIZE: length, 2.6 mm; width, 1.1 mm (humeri).

ALLOTYPE, female: no significant differences were observed except the female has

shorter antennae, not extending beyond the pronotum.

TYPES: Holotype male, Jamaica, Try. Duncans, VII-19-1966, AT. Howden (HAHC,

Ottawa). Allotype female and 3 male paratypes collected at the same locality and date as

the holotype. Paratypes in Howden and Manley collections.

This species is similar to M. coyolensis Manley from Honduras and
keys to the species in Manley, 1987 '. Mastogenius howdenorum differs from
M. coyolensis in the following ways: M. howdenorum is smaller and less
elongate, the pronotum is more uniformly rounded on lateral margins
and widest at or just behind the middle, the disk of the pronotum is more
uniformly convex, and the species is more densely pubescent on both
the pronotum and elytra.

Figure 1-2. Dorsal view and pronotum, Mastogenius howdenorum.

Vol. 101. No. 4. September & October 1990

215

Figure 3. Dorsal view of male genitalia, Mastogenius howdenorum.

ACKNOWLEDGMENTS

The author wishes to express thanks to the following persons and institutions for their
contribution to this work: Henry F. Howden, Carleton University, for providing the
specimens of this interesting new species and for making available the scanning electron
microscope; Lewis Ling for taking the photographs; Peter H. Carrington for doing the
drawing of the genitalia; Elinar A. Manley for suggestions during manuscript preparation.

LITERATURE CITED

Manley, G.V. 1987. New species of Mastogenius (Buprestidae:Coleoptera) from Central
America with notes and a key to species of Central and South America. Ent. News.
98(1): 1-9.

216 ENTOMOLOGICAL NEWS

MARKING CASSIDINAE

(COLEOPTERA: CRYSOMELIDAE) LARVAE IN
THE FIELD FOR POPULATION DYNAMICS

STUDIES1

M.A. Garcia, L.M. Paleari2

ABSTRACT: Marks on the exuviae held by caudal fork, a permanent structure in
Cassidinae larvae, can provide a simple method for the study of larval population
dynamics in this group.

Population-dynamics studies of insects are usually restricted to adults,
which can be easily individualized by some kind of mark. Many mortality
factors act on juvenile stages, however, and need to be studied. Since any
mark directly placed on larvae or nymphs can only be seen until ecdysis
occurs to the following instar, various indirect methods have been des-
cribed to study the population-dynamics of the immature insects.
Approximate correlation or indices of equivalence with population sizes
at different times during insect development can be obtained by mea-
sures of damage, counts of exuviae or collecting feces of the immature
insects (see Southwood 1978). These indirect methods can supply good
correlations for some species. Another method developed by Kiritani
and Nakasuji (1967) improved estimates of staged specific mortality
rates for a population developing through various life stages. A draw-
back of this method is the fixed sampling pattern that modifies the
population size and makes it difficult to obtain good estimates.

Cassidinae larvae offer a special opportunity for studies on popu-
lation dynamics of juvenile insects. One of their characteristics is the
accumulation of the exuviae and some feces attached to the caudal fork,
constituting a permanent structure called the annex by Buzzi (1988).
This structure can be present even on pupae and is suitable for marking.

The mobility of the caudal fork permits the annex to be adjusted over
the insect's body, giving a camouflaged aspect. The side of the annex that
is in contact with the larva's body is smooth and it is possible to count the
series of exuviae on it. Marks can be placed on this side of the annex,
without any modification of the larval aspect, giving little or no effect on
the probability of attack by natural enemies or survival of the young
insects in the field.

Deceived January 16, 1990. Accepted April 21, 1990.

2Laboratorio de Interacoes Inseto-Planta, Departamento de Zoologia, Institute de
Biologia, Universidade Estadual de Campinas, Campinas, SP. Brasil 13081.

ENT. NEWS 101(4): 216-218, September & October, 1990

Vol. 101. No. 4. September & October 1990

217

A population of Charidotis punctatostriata larvae was marked in
Campinas, Sao Paulo, Brazil, in a fallow field where the host plant
Pyrostegia venusta (Bignoniaceae) was abundant.

Marks were made using white nail polish, but a special marking pen
could just as well have been used, with each larva receiving its own
number.

Larvae of C. punctatostriata from the same egg mass show a tendency
to remain together. This made it easy to mark groups of different instars,
placing the white spot over the more recent exuviae attached to the
annex. The small white spot could only be seen when the larva spon-
taneously raised the annex or when this movement was provoked touching
it with a brush.

The five instars of C. punctatostriata larvae found in the fallow field
were observed and counted daily until they moulted to the next instar,
when they were collected and reared further in the laboratory, while the
observations continued.

None of the marks was lost in a month of observations. During this
time all the marked larvae completed their development in the labor-
atory, disappeared by predation in the field, were mummified by
parasitoids or died by disease (table 1). These data will be analysed and
discussed in a separate paper about the field biology of C. punctatostriata.

Table 1. Survival of Charidotis punctatostriata (Coleoptera: Chrysomelidae: Cassidinae)
larvae marked in the field, (number of individuals)

Instar
marked

1
2
3
4
5

IN THE FIELD

Number
marked

60
50
50
51
51

LATER IN LABORATORY

Surviving
to next
instar

14
12
38
34
34

Mumified

by
parasites

23
9

Died

diseases

8
4
5
2
2

Surviving

to
adults

6
6

25
9

ACKNOWLEDGMENTS

We are grateful to Keith S. Brown, Ivan Sazima and Joao Vasconcellos Neto for their
review of the manuscript and for their valuable suggestions.

218 ENTOMOLOGICAL NEWS

LITERATURE CITED

Buzzi, ZJ. 1988. Biology of neotropical Cassidinae, p. 559-580. In P. Jolivet, E. Petitpierre
& T.H. Hsiao (eds). Biology of Chrysomelidae Kluwer Acad. Publishers.

Kiritani, K. and F. Nakasuji 1967. Estimation of stage specific survival rate in insect
population with overlapping stages. Res. Popul. Ecol, 9: 143-152

Southwood, T.R.E. 1978. Ecological Methods, with particular reference to the study of insect
populations. Second edition. Chapman and Hall, London and New York, p. 288-301.

Vol. 101, No. 4, September & October 1990 219

A NEW DISTRIBUTIONAL RECORD FOR THE
SAND-BURROWING MAYFLY

DOLANIA AMERICANA
(EPHEMEROPTERA: BEHNINGIIDAE)1

William L. Jacobs^

ABSTRACT: The sand-burrowing mayfly, Dolania americana is reported for the first time
from the St. Croix River at the Wisconsin/Minnesota border. This record extends the range
of this unique species far northward from its known typical southeastern blackwater
habitats. A possible dispersal route for this species from the southeastern Coastal Plain
into the St. Croix River system is briefly discussed.

The only North American member of the sand-burrowing mayfly
family Behningiidae, Dolania americana Edmunds and Traver has been
previously reported from Alabama (Wester and Folkerts 1987), Florida
(Schneider 1966), Georgia (Benke et al. 1984), Louisiana (Dakin and
Felder 1981), North Carolina (Finn and Herlong 1980), and South
Carolina (Edmunds and Traver 1959, Harvey et al. 1980). The biology of
this mayfly is relatively well-known. Its preferred habitat is clean, shift-
ing sand rivers and large streams (Peters and Peters 1977, Tsui and
Hubbard 1979, Fink 1986, and Peters et al. 1987).

On July 24, 1989, one nymph was collected from the St. Croix River.
The collection site was near Soderbeck Landing, St. Croix National
Scenic Riverway (Burnett County, Wisconsin). This site was approxi-
mately 70m upstream from the confluence of the Snake River, which
originates in Minnesota. This nymph was collected with a Surber square
foot sampler at a depth of 70-85cm. Due to drought conditions, the river
was unseasonably low. The primary substrate consisted of fine grain
sand (0.0625-0. 125mm size range) and water pH was 7.4. Another speci-
men was collected August 21, 1989 at the same site. The two nymphs
measured 5mm and 7mm body length respectively, indicating a 2 year
life cycle.

Previous macroinvertebrate studies of the St. Croix National Scenic
Riverway have yielded no Dolania. Additionally, no nymphs were col-
lected during studies of a major Wisconsin tributary, the Namekagon
River. The current benthic macroinvertebrate study of the St. Croix
River involved collections every two weeks from late May until the end of
September. Samples were taken upstream and downstream of the fol-
lowing tributaries: Cranberry bog (a bog which flows into the

Deceived February 7. 1990. Accepted March 27. 1990.

-Colorado State University. Department of Entomology, Fort Collins, Colorado 80523

ENT. NEWS 101(4): 219-221, September & October. 1990

220 ENTOMOLOGICAL NEWS

Namekagon River just below Pacwawong Dam, Sawer County,
Wisconsin), Yellow River, Clam River (Burnett County, Wisconsin),
Kettle River, Snake River (Pine County, Minnesota), Sunrise River
(Chisago County, Minnesota), and Apple River (St. Croix County,
Wisconsin). It is interesting to note that Dolania has not been previously
reported from Wisconsin, although many streams have been intensively
surveyed by William L. Hilsenhoff and co-workers (for example
Hilsenhoff 1977).

Other mayflies collected with the Dolania specimens include Baetis
spp. (Baetidae); Isonychia sp. (Oligoneuriidae); Leucrocuta hebe
(McDunnough), Stenacron interpunctatum (Say), Stenonema spp.,
Rhithrogena pellucida Daggy, Epeorus vitreus (Walker) (Heptageniidae);
Choroterpes basalis (Banks)?, Leptophlebia cupida (Say), Paraleptophlebia
sp. (Leptophlebiidae); Potamanthus sp. (Potamanthidae); Ephemera
simulans Walker, Hexagenia limbata (Serville) (Ephemeridae); Ephoron
leukon Williamson (Polymitarcyidae); Serratella spp., Ephemerella spp.,
Eurylophella spp.,Attenella attenuata (McDunnough) (Ephemerellidae);
Tricorythodes sp. (Tricorythidae); Caenis spp.,Brachycercus sp. (Caenidae),
Baetisca obesa (Say) and B. lacustris McDunnough (Baetiscidae).

All of Minnesota and much of Wisconsin were covered with glacier
during the last glacial stage, the Wisconsinan (Flint 1971), indicating
post-glacial migration by the present day Ephemeroptera. This last
glacial stage began retreating about 12-1 5 x 10^ B.P. Aquatic dispersal by
nymphs opposed to aerial dispersal by adults appears to be the likely
mechanism in Dolania. The duration of the adult stage is less than two
hours (Peters and Peters 1977). Flannagan and Flannagan (1984) outline
post-glacial waterways from glacial Lake Agassiz (centered in Manitoba)
and present likely migration routes. Dolania probably would be included
in the South Agassiz group, inhabitants of the Gulf of Mexico watershed.
Migration northward likely occurred through the Mississippi-Missouri
River system when Lake Agassiz drained southwards. Contemporary
watershed barriers probably prevented recent northward migration of
Dolania and other mayflies from the southeastern Coastal Plain, indi-
cating that Dolania has probably been present in the St. Croix River
system before recent times.

ACKNOWLEDGMENTS

I thank William L. Peters, Florida A & M University for verifying the identification of
Dolania, and Boris C. Kondratieff. Terence P. Boyle and Mitch A Harris. Colorado State
University, for prepublication reviews. William L. Hilsenhoff. University of Wisconsin
and William L. Peters provided valuable comments which also improved the manuscript.
This study is part of the National Park Service supported research on resource inventory of
St. Croix National Scenic Riverway.

Vol. 101, No. 4, September & October 1990 221

LTERATURE CITED

Benke, A.C.,T.C. Van Arsdall, Jr., D.M.Gillespie, and F.K.Parrish. 1984. Invertebrate

productivity in a subtropical blackwater river: the importance of habitat and life

history. Ecol. Monogr. 54: 25-63.
Dakin, M.E., Jr. and Felder. 1981. A record of the mayfly Dolania americana in Louisiana

(Ephemeroptera: Behningiidae). Florida Entomol. 74: 454-455.
Edmunds, G.F. and J.R. Traver. 1959. The Classification of the Ephemeroptera I.

Ephemeroidea: Behningiidae. Ann. Entomol. Soc. Amer. 52: 43-51.
Fink, TJ. 1986. The reproductive life history of the predaceous, sand-burrowing mayfly

Dolania americana (Ephemeroptera: Behningiidae). PhD dissertation. Florida St.

Univ., Tallahassee.
Finn, P.L. and D.D. Herlong. 1980. New distributional record of Dolania americana

(Ephemeroptera: Behningiidae). Entomol. News 91: 102-104.
Flannagan, P.M. and J.F. Flannagan. 1984. The post-glacial origin and present

distribution of the mayflies (Ephemeroptera) of Manitoba, Canada, Pp. 149-169. In

Landa, V., T. Soldan. and M. Tonner (eds.). Proc. IVth Intern. Conf. Ephemeroptera.

Czechoslovak Acad. Sci.

Flint, R.F. 1971. Glacial and quaternary geology. J. Wiley and Sons;, Inc., NY. 892 pp.
Harvey, R.S., R.L. Vannote, and B.W. Sweeney. 1980. Life history, developmental

processes, and energetics of the burrowing mayfly Dolania americana. Pp. 21 1-230. In

Flannagan, J.F. and K.E. Marshall (eds). Advances in Ephemeroptera Biology. Plenum

Press, New York.
Hilsenhoff, W.L. 1977. Use of arthropods to evaluate water quality of streams. Wisconsin

Dept. Nat. Res. Tech. Bull. 100: 1-15.
Peters, J.G., W.L. Peters, and T.J. Fink. 1987. Seasonal synchronization of emergence in

Dolania americana (Ephemeroptera: Behningiidae). Can. J. Zool. 65: 3177-3185.
Peters, W.L. and J.G. Peters. 1977. Adult life and emergence of Dolania americana in

northwestern Florida (Ephemeroptera: Behningiidae). Int. Rev. Ges. Hydrobiol. 62:

409-438.
Schneider, R.F. 1966. Mayfly nymphs from northwestern Florida. Quart. J. Florida Acad.

Sci. 29: 203-206.
Tsui, P.T.P. and M.D. Hubbard. 1979. Feeding habits of the predaceous nymphs of

Dolania americana in northwestern Florida (Ephemeroptera: Behningiidae).

Hydrobiol. 67: 119-123.
Wester, E.E. and G.W. Folkerts. 1987. The may fly, Dolania americana (Ephemeroptera:

Behningiidae) in Alabama. Entomol. News 98: 253-254.

222 ENTOMOLOGICAL NEWS

A NEW METHOD FOR COLLECTING ADULT

PHRYGANEID CADDISFLIES
(TRICHOPTERA: PHRYGANEIDAE) 1

David E. Bowles2' 3, Karl Stephan4, Michael L. Mathis5

ABSTRACT: A fermented molasses trap was used to collect several specimens of the
caddisfly genus Ptilostomis (Trichoptera: Phryganeidae). Molasses bait traps are inex-
pensive, relatively maintenance free, and can be used as an effective collecting technique.
Molasses and other sugar-based materials are suggested to be a possible source of nutrition
for these caddisflies.

Representatives of the trichopteran family Phryganeidae are large,
attractive caddisflies often characterized by having distinctly patterned
wings. Ten genera and 27 species of phryganeids are represented in
North America (Wiggins 1984, Wiggins and Larson 1989). Depending on
geographical location phryganeids can be common and adults may be
obtained through a variety of techniques including fluorescent and UV-
light traps, Malaise and flight-intercept traps, sweeping, and laboratory
rearings of immature stages. Recently, one of the authors (KS) collected
several specimens of the phryganeid genus Ptilostomis from an eastern
Oklahoma woodland (Latimer County; near Red Oak) using a fer-
mented molasses trap.

A mixture of molasses and water (3:1 ratio) was placed into a 355 ml
plastic cup, approximately one-half full, and positioned approximately
2 m above ground level. The trap was being used to collect saprophagous
Coleoptera that are attracted to the fermenting molasses.

During June 1989, several specimens of Ptilostomis postica (Walker)
and two individuals of P. ocelli/era (Walker) were collected with the
molasses trap. Although specimens were removed from the trap daily,
the numbers collected on a particular day were not recorded. The molasses

Deceived December 15, 1989. Accepted February 12, 1990.

^United States Air Force School of Aerospace Medicine, Epidemiology Division, Brooks
AFB,TX 7823 5-5301.

This paper reports the results of research completed while a graduate student in the
Department of Entomology, University of Arkansas, Fayetteville, AR 72701. Opinions
and assertions contained herein are those of the authors and are not to be regarded as
official or as reflecting the views of the United States Air Force.

4Rt. 1, Box 913, Red Oak, OK 74563.
Department of Zoology, University of Arkansas, Fayetteville, AR 72701.

ENT. NEWS 101(4): 222-224, September & October, 1990

Vol. 101, No. 4, September & October 1990 223

trap was situated on a heavily forested hillside approximately 200 m
from the nearest permanent water sources including a spring, small
stream, and farm pond. All three sources of water could potentially serve
as the larval habitat (Bowles and Mathis 1 989). The molasses bait appar-
ently attracted the caddisflies during nocturnal periods, and the
specimens were found during daylight hours drowned in the fermented
mixture. No specimens were collected alive; no additional caddisfly
species were collected from the trap.

The total number of specimens collected for each species was similar
for both the molasses bait and UV-light trap samples that were taken
nightly during the same time period. The molasses bait yielded five
female and seven male specimens of/3, postica and two female specimens
of P. ocelli/era. UV-light trap collections yielded five females and six
males of P. postica and one female of P. ocelli/era. Males of the latter
species have been collected previously from the same area. Voucher
specimens of P. ocelli/era and P. postica are deposited in the University of
Arkansas Insect Collection.

The reason why the phryganeids were attracted to the molasses bait is
not known. However, some species of Trichoptera have been reported to
feed as adults, particularly on sweet materials such a flower nectar
(Crichton 1957), sugar baits, honeydew, honey, and fruit preserves
(Malicky 1989). The phryganeids probably were attempting to obtain
nourishment when they became trapped in the molasses mixture.
Crichton (1957) provided a detailed discussion of the structure and
imbibing function of phryganeid mouthparts and compared them with
those of representatives of 12 other trichopteran families. The protrusible
haustellum used in imbibing fluids was found to be most developed in
the Phryganeidae and Limnephilidae (Crichton 1957).

The molasses bait trap provides a relatively maintenance free and
inexpensive means for collecting phryganeid caddisflies. Molasses bait
traps could prove advantageous in situations where the collector has
time constraints such as simultaneously collecting from several local-
ities. A series of several bait traps placed in a given area could be used as
an effective collecting tool. Possibly, other sugar based materials also
might be effective attractants. Additional testing will be necessary to
evaluate the usefulness of molasses bait traps for collecting other
caddisfly species.

ACKNOWLEDGMENTS

Dr. Chad P. McHugh. Lt. Col. Dennis D. Pinkovsky and Capt. Terry L. Carpenter of the
USAF School of Aerospace Medicine kindly reviewed this manuscript. The comments of
two anonymous reviewers also enhanced this manuscript.

224 ENTOMOLOGICAL NEWS

LITERATURE CITED

Bowles, D.E., and M.L. Mathis. 1989. Caddisflies (Insecta: Trichoptera) of mountainous

regions in Arkansas, with new state records for the order. J. Kansas Entomol. Soc.

62:234-244.
Crichton, M.I. 1957. The structure and function of the mouthparts of adult caddisflies

(Trichoptera). Phil. Trans. Royal Soc. London, Ser. B. Biol. Sci. 241:45-91.
Malicky, H. 1989. Feeding of adult caddisflies. Trichopterist Newsletter 16:18.
Wiggins, G.B. 1984. Trichoptera, pp. 271-31 1. In, R.W. Merritt and C.W. Cummins [eds.],

An introduction to the aquatic insects of North America, 2nd ed. Kendall/Hunt:

Dubuque, Iowa.
Wiggins, G.B., and D J. Larson. 1989. Systematics and biology for a new Nearctic genus

in the caddisfly family Phryganeidae (Trichoptera). Can. J. Zool. 67:1550-1556.

Vol. 101, No. 4, September & October 1990 225

SUCKING LICE (ANOPLURA) FROM PAKISTAN

MAMMALS, WITH NOTES ON

ZOOGEOGRAPHY*

L.A. Durden , R. Traub , K.C. Emerson^

ABSTRACT: Anoplura collected from wild rodents, insectivores and canids in Pakistan
are documented. Twenty-two species of lice were recovered as follows: Hoplopleura (8),
Linognathoides (1), Neohaematopinus (1), Polyplax (12). At least 14 of these species are re-
corded from Pakistan for the first time and some significant range extensions are estab-
lished. Some of the records pertain to species previously known only from the type series or
from other limited collections. Zoogeographically, Pakistan's anopluran fauna has strong
Palearctic affinities although elements of this fauna showclear Oriental, Ethiopian or cos-
mopolitan associations.

Except for records of widespread species or descriptions of new taxa,
the sucking lice (Anoplura) of Pakistan are poorly known. This is sur-
prising because contributions documenting the anopluran fauna of
adjacent regions in Afghanistan (Smetana and Daniel, 1970), India
(numerous papers), Iran (Kim and Emerson, 1971), southern USSR
(numerous papers) and the People's Republic of China (numerous
papers) are available. Mishra (1981) produced a monograph of the
hoplopleurid sucking lice of the Indian subcontinent and included the
Pakistan fauna only from the Indus valley eastwards to the Indian bor-
der; however, that work principally addressed the Indian fauna and no
specific records for Pakistan are given.

This paper provides records of sucking lice from wild land mammals
(mainly rodents) obtained in Pakistan from 1962-1979. Most specimens
were collected by field teams of the Department of Microbiology, Uni-
versity of Maryland School of Medicine, Baltimore under the direction
of Robert Traub. Additional material was collected by Robert G. Tuck,
then with the Division of Mammals, National Museum of Natural His-
tory (NMNH), Washington, D.C. Collection data for the 22 species of
sucking lice recovered during these surveys includes hosts, collection
localities, altitude (if available) and dates, followed by remarks. Louse
synonymies listed are not new but are important to this study. Host
mammal names follow Honacki etal. (1982) and Anoplura classification
follows Kim and Ludwig (1978). Louse and host mammal material
documented here is deposited in the collections of the NMNH, Washing-
ton, D.C.

^Received 25 November, 1989. Accepted 30 March, 1990.

Department of Entomology, Museum Support Center. Smithsonian Institution, Wash-
ington, D.C. 20560

3560 Boulder Drive, Sanibel, Florida 33957

ENT. NEWS 101(4): 225-235, September & October. 1990

226 ENTOMOLOGICAL NEWS

Family Hoplopleuridae Ferris

1) Hoplopleura affinis (Burmeister, 1838)

Specimens examined: (44 collections (39cT,2539) ex Apodemus sp. (Rodentia,
Muridae)) HAZARA DIST.: Kaghan valley: Soch, Naran, 2430 m and 2615 m,
IXX1962, VII.1964, VII.1965 - 29 colls.; Shogran, 2385 m and 2770 m, VIII.1963,
VIII. 1964 - 3 colls.; Burawai, 3080 m, IX. 1963 - 5 colls.; Battakundi, 2740 m, VIII. 1963 - 1
coll.; 6.4 km E. of Lalazar, 2430 m, X. 1962 - 1 coll.; GILGIT AGENCY: Naltar, 2985 m.
VIII. 1964 - 4 colls.; Gupis, Kohighizar, 2360 m, VIII. 1964 - 1 coll.

Hoplopleura affinis is a common ectoparasite of Eurasian field mice
of the genus Apodemus. There is some confusion regarding the status of
Apodemus in Pakistan (Roberts, 1977); A. sylvaticus (Linn.) definitely
occurs in that country but A. flavicollis (Melchior) may also be present.
Hosts for the above-mentioned collections were listed as Apodemus sp.,
A. flavicollis or A. sylvaticus. To avoid erroneous records, a conservative
approach is taken here and all hosts are listed as Apodemus sp. Hoplo-
pleura affinis has previously been recorded from both A. flavicollis and
A. sylvaticus.

2) Hoplopleura alticola Mishra and Bhat, 1972.

Specimens examined: (1 collection (29) ex Alticola rovlei Gray (Rodentia, Arvico-
lidae)) HAZARA DIST.: Kaghan valley, 4155 m, VIII.1963.

Until now, this louse was known only from the type series taken from
the vole, A. roylei, in montane regions in the Indian states of Uttar
Pradesh and Himachal Pradesh (Mishra and Bhat, 1972; Mishra, 1981).

3) Hoplopleura captiosa Johnson, 1960 (synonym: musculi Wegner,

1961).

Specimens examined: (8 collections (6cT,179) ex Mus musculus Linn. (Rodentia,
Muridae)) HAZARA DIST.: Murree Hills, Dunga Gali, 2495 m, 1X1962 - 3 colls.;
Balakot,IX.1962-2colls.; LAHORE DIST.: Near Ravi River bridge, 215m, 1X1963-1
coll.; Changa Manga Forest, 11.1965 - 1 coll.; PARACHINAR AGENCY: Parachinar.
1540 m,III. 1964-1 coll. (5 collections (5cT,l 49) ex A/wssp. (Rodentia, Muridae)) CHIT-
RAL DIST.: Chitral, 1490 m, VIII. 1964 - 3 colls.; HAZARA DIST.: Kaghan valley,
Shogran, 2385 m, VIII. 1964 - 1 coll.; MALAKAND AGENCY: Amandarra, 845 m,
VIII.1964- 1 coll.

Hoplopleura captiosa is principally parasitic on the house mouse, M.
musculus and probably occurs in most areas of the world where this
largely peridomestic mammal is found. It would therefore be expected to
occur throughout most of Pakistan, particularly in association with
human settlements.

4) Hoplopleura maniculata (Neumann, 1909) (synonym: mitsuii

Kaneko, 1963).

Specimens examined: (6 collections (33cf,379,4 nymphs) ex Funambulus pennanti
Wroughton (Rodentia, Sciuridae)) LAHORE DIST.: Near Ravi River bridge, 215 m,
1X1963 - 2 colls.; Model Town, XI. 1962 - 1 coll.; unspecified locality, XI. 1962 - 1 coll.;
RAWALPINDI DIST.: Ayub Natl. Park, XI. 1962 - 1 coll.; SIND PROV.: Karachi,
University campus, VII. 1976 - 1 coll.

Vol. 101, No. 4. September & October 1990 227

This louse was originally described from palm squirrels. Funambulus
palmarum (Linn.), collected in India. It appears to be a widespread
parasite of squirrels of the genus Funambulus (F. palmarum, F. pennanti,
F. tristriatus (Waterhouse)) in India and Sri Lanka (Pratt and Stojano-
vich, 1961; Kaneko, 1963; Mishraetal., 1974;Mishra. 1981). However, the
specimens recorded here appear to be the first noted for Pakistan.

5) Hoplopleura merionidis Ferris. 1921.

Specimens examined: (1 collection (3cT.19)ex Meriones libvcus Lichtenstein (Roden-
tia. Cricetidae)) KALAT DIST.: unspecified locality. 2320 m, X.1963.

Hoplopleura merionidis has not previously been reported from Pakis-
tan although it is known from neighboring regions in Iran (Kim and
Emerson, 1971), the People's Republic of China (Ferris, 1921) and the
USSR (Sosnina, 1982). The type series from Shaanxi (as Shensi) Pro-
vince, China, was recovered from the gerbil. Meriones meridianus
(Pallas), while the Iran material was collected from both M. libvcus and
M. crassus Sundevall, and the USSR specimens were from both M. libvcus
(as M. erythrourus) and M. meridianus.

6) Hoplopleura pacifica Ewing, 1924.

Specimens examined: (8 collections (2cf ,99,17 nymphs) ex Rattus norvegicus (Berken-
hout) (Rodentia, Muridae)) SIND PROV.: Karachi: town area. 1.1977 - 3 colls.; Em-
press market 1.1977 - 3 colls.: East wharf, 1. 1977 -2 colls. (2 collections ( lcc,29) ex Rattus
rattus (Linn.) (Rodentia. Muridae)) GILGIT AGENCY: Kohighizar, 2360 m. IX. 1963.
(2 collections (2cT,l 19) ex Rattus sp. (Rodentia, Muridae)) HAZARA DIST.: Kaghan
valley. Shogran, 2385 m, VIII. 1964 - 1 coll.: RAWALPINDI DIST.: Ayub Natl. Park.
XII. 1962- 1 coll.

Hoplopleura pacifica has an almost global distribution along with that
of its domestic Rattus hosts.

7) Hoplopleura pavlovskyi Sosnina, 1951.

Specimens examined: (1 collection (29) ex Rattus turkestanicus
(Satunin) (Rodentia, Muridae)) GILGIT AGENCY: Gilgit. Chinar
Bagh, 1490m, VIII. 1964.

This species is similar to H. pacifica but whereas the dorsal apical
angle of the paratergal plate of abdominal segment seven is extended in
H. pacifica, no such lobe is present in H. pavlovskyi. Although H. pavlov-
skyi is well documented from numerous collections in Turkestan. USSR
(Sosnina, 1951, 1967, 1982), this species does not appear to have been
reported from other regions. The host rat, R. turkestanicus, ranges from
southern Turkestan, northeastern Iran and Afghanistan to northern
India and southwest China (Roberts, 1977; Honacki et al.. 1982).

8) Hoplopleura ramgarh Mishra. Bhat and Kulkarni, 1972.
Specimen examined: (1 collection (19) ex Mus soxicola Elliot (Rodentia. Murdiae)
SIND PROV.: Rani Kot, 1.1977.

This louse was previously documented only from several states in
India as an ectoparasite of the mice. A/, saxicola and M. platythnx Bennett

228 ENTOMOLOGICAL NEWS

(Mishra et a/., 1972, 1974; Mishra, 1980, 1981; Saxena, 1987). Mishra
(1980) pointed out that although mice belonging to the M. platythrix com-
plex are morphologically similar, their sucking lice are quite different
and could be utilized to aid studies of host systematics.

Family Polyplacidae Fahrenholz
9) Linognathoides palaearctus (Olsoufjev, 1938.)

Specimens examined: (2 collections (2<J,19, 7 nymphs) ex Marmora caudata (Geof-
froy) (Rodentia, Sciuridae)) HAZARA DIST.: Kaghan valley: Saif Ul Maluke, 3200 m,
X.1962 - 1 coll.; Besal, 3260 m. IX. 1963 - 1 coll.)

This louse is a specific parasite of M caudata (the long-tailed mar-
mot) and was originally described from Turkestan, USSR. It has also
been reported from Afghanistan (Smetana and Daniel, 1970), Jammu
and Kashmir state, India (Mishra et a/., 1974; Mishra, 1981), Pakistan
(Kim and Adler, 1982) and several provinces in the People's Republic of
China (Chin, 1985).

10) Neohaematopinus echinatus (Neumann, 1909.)

Specimens examined: (11 collections (53cf,609, 9 nymphs) ex Funambulus pennanti
Wroughton (Rodentia, Sciuridae)) LAHORE DIST.: Jahangir's tomb, IX. 1962 - 2
colls.; Near Ravi River bridge, 215 m, 1X1963 - 1 coll.; Model Town, XI. 1962 - 1 coll.;
Luliani, X.1963 - 1 coll.; unspecified localities, XI. 1962 - 2 colls; BAHAWALPUR
DIST.: Uch-Sharif, XII. 1963 - 2 colls; SIND PROV.: Rani Kot, LVI.1977 - 2 colls.

Neohaematopinus echinatus has previously been reported only from
India where it is a widely distributed parasite of the squirrels, Funam-
bulus pennanti, F. palmarum and F. tristriatus (Pratt and Stojanovich,
1961; Kaneko, 1963; Mishra et a/., 1974; Mishra, 1981; Kim and Adler,
1982). Funambulus pennanti is widely distributed in Pakistan but is the
only squirrel of this genus occurring in that country (Roberts, 1977).

\\)Polyplax asiatica Ferris, 1923 (synonym: turkestanica Blagovesht-
chensky, 1950).

Specimens examined: (17 collections (18<J,389, 9 nymphs) ex Nesokia indica (Gray
and Hardwicke) (Rodentia, Muridae)) LAHORE DIST.: Near Ravi River bridge, 215
m, IX. 1962, IX. 1963, I.IV.1964 - 6 colls.; Lahore, IX. 1962 - 4 colls.; 6.4 km NW of
Luliani, 1.1964 - 1 coll.; DERA ISMAIL KHAN DIST.: unspecified localities, XII. 1963.
1.1964 -2 colls.; RAWALPINDI DIST.: Ayub Natl. Park, XII.1962, 1.1964 - 2 colls.;
SIALKOT DIST.: Charwa, IX 1962 - 2 colls. ( 1 collection (29) exNesokia sp. (Rodentia,
Muridae)) KOHAT DIST.: Kohat, 310m, VIII. 1964. ( 1 collection (29) exApodemus sp.
(Rodentia, Muridae)) HAZARA DIST.: Battakundi, 2740 m, VIII. 1963. (1 collection
( 19) ex Suncus sp. (Insectivora, Soricidae) LAHORE DIST.: Near Ravi River bridge,
215m, IX. 1962.

This louse is a well documented ectoparasite of the burrowing rat, N.
indica and has been reported from Taiwan westwards across central and
southern Asia to North Africa. Polyplax asiatica has also been collected
frequently from Bandicota spp., another fossorial murid genus, and
occasionally from other commensal rodents and shrews. Except for

Vol. 101, No. 4. September & October 1990 229

collections from Nesokia and Bandicota, these records are considered to
represent accidental infestations. The type series of P. asiatica designated
by Ferris (1923) includes specimens recovered from N. indica (as N.
hardwickei) in Quetta, Balunchistan Province, Pakistan.

\2) Polyplax brachyrrhyncha Cummings, 1915.

Specimens examined: (2 collections (34cf,689) ex Acomys cahirinus (Desmarest)
(Rodentia, Muridae)) SIND PROV.: Karachi, Khadeji Fal'ls, XI. 1976.

Polyplax brachyrrhyncha was originally described from A. cahirinus
from Asyut, Egypt and is a common ectoparasite of this spiny mouse in
that country (Johnson, 1960). It has also been recorded previously from
various species of Acomys (principally^, cahirinus) from Iran (Kim and
Emerson, 1971) and from North and East Africa but the present records
are the first ones from Pakistan.

13) Polyplax calomysci Kim and Emerson, 1971.

Specimens examined: (5 collections (5c? ,69) ex Calomvscusbailwardi Thomas (Roden-
tia, Cricetidae)) BALUCHISTAN PROV.: Quetta. Ziarat, 2315 m, XXI. 1963 - 3 colls.;
Sibi, 1X1975 - 1 coll.; Kalat, 2135 m. 1X1975 - 1 coll.

This species was previously documented only by the type series from
C. bailwardi from Iran (Kim and Emerson, 1971). The rodent host is con-
fined to mountain steppe regions of Turkmenistan (USSR), Iran,
Afghanistan and Pakistan (Roberts, 1977).

14) Polyplax chinensis Ferris, 1923.

Specimens examined: (5 collections (5cf,49) ex Meriones crassus Sundevall (Rodentia,
Cricetidae)) BALUCHISTAN PROV.: 48 km SSW of Surab, IV.VII.1965.

Polyplax chinensis was originally described from the gerbil, Meriones
meridianus, from Shaanxi (as Shensi) Province, People's Republic of
China (Ferris, 1923) but has also been reported from the southeastern
USSR (Sosnina, 1982). The present collections are the first ones docu-
mented from Pakistan.

15) Polyplax hurrianicus Mishra 1981.

Specimens examined: (5 collections (ltf,349, 28 nymphs) ex Meriones hurrianae Jer-
don (Rodentia, Cricetidae)) SIND PROV.: Rani Kot, XXI, 1976, 1.1977 - 3 colls.;
Karachi, Malir Cantonment, X1976, 1.1977 - 2 colls. (1 collection (29) ex Gerbillus
nanus Blanford (Rodentia, Cricetidae)) SIND PROV.: Rani Kot, VII. 1976.

Until now, P. hurrianicus was known only from the type series from
the gerbil, M hurrianae, in Gujarat State, India (Mishra, 1981).

16) Polyplax kaiseri Johnson, 1960.

Specimens examined: (1 collection (29) ex Gerbillus nanus Blanford (Rodentia.
Cricetidae)) DERA ISMAIL KHAN DIST.: unspecified locality, 1.1964. (12 collec-
tions (18cT,319) ex Gerbillus sp. (Rodentia. Cricetidae)) BALUCHISTAN PROV.: 18
km SE of Kharan, 11.1965 - 7 colls.; LASBELA DIST.: 8 km S of Bela. III. 1965 - 5
colls.

Polyplax kaiseri is well-known from various gerbils of the genus Ger-

230 ENTOMOLOGICAL NEWS

billus in North and East Africa, although Ledger (1980) -has questioned
records of this louse in sub-Saharan Africa. Kim and Emerson (1971)
reported P. kaiseri from both G. nanus and G. cheesmani Thomas in Iran
but the records given here represent the most easterly ones to date for this
louse and the first ones for Pakistan. It is expected that P. kaiseri also
parasitizes gerbils between the North Africa and Iran/Pakistan extremes.

17) Polyplax kondana Mishra, 1981.

Specimens examined: (3 collections (2c?,59, 1 nymph) ex Millardia meltada (Gray)
(Rodentia, Muridae)) SIND PROV.: Badin, 1.1977 - 2 colls.; Thatta, VIII. 1975 - 1
coll.

This louse was previously documented only by the type series recovered
from Millardia kondana Mishra and Dhanda in Maharashtra State,
India (Mishra, 1981).

18) Polyplax paradoxa Johnson, 1960.

Specimens examined: (1 collection (29) ex Meriones persicus (Blanford) (Rodentia,
Cricetidae)) BALUCHISTAN PROV.: unspecified locality, 1.1963.

Polyplax paradoxa parasitizes various gerbils belonging to the genus
Meriones in North and East Africa (Ledger (1980) questioned some of
these records) and the Near East. Although P. paradoxa has not prev-
iously been reported from Pakistan, it has been recorded from M. persicus
and M. tristrami Thomas in Iran by Kim and Emerson ( 197 1) and Kaneko
(1972), respectively.

19) Polyplax reclinata Nitzsch, 1864 (synonyms: deltoides Fahrenholz,

1938; shimizui Kaneko, 1957).

Specimens examined: (11 collections (44c7,689, 2 nymphs) exSuncus murinus (Linn.)
(Insectivora. Soricidae)) GUJRAT DIST.: Mangowal, VI. 1965 - 1 coll.; HAZARA
DIST.: Balakot, 1X1962 - 2 colls.; LAHORE DIST.: Near Ravi River bridge, 215 m,
VI. 1965 - 2 colls.; Model Town, XI. 1962 - 1 coll.; SIALKOT DIST.: Charwa, 275 m,
X.1962 - 1 coll.; SIND PROV.: Karachi, town area, XII.1975. IIJIUV.1976 - 4 colls. (1
collection (19) ex Suncus sp. (Insectivora, Soricidae)) LAHORE DIST.: Lahore,
VH.1964.

This species is widespread in Eurasia and Africa (including Mada-
gascar) as a parasite of numerous species of shrews (Soricidae).

20) Polyplax serrate (Burmeister, 1839) (svnonyms: qffinis Fahrenholz,

\938;serrata paxi Eichler, 1952).

Specimens examined: (1 collection (29) ex Cricetulus migratorius (Pallas) (Rodentia,
Cricetidae)) GILGIT AGENCY: Kohighizar, Phandar, 3050 m, 1X1963.

This louse is a well-known cosmopolitan ectoparasite of the house
mouse, Mus musculus Linn., although it is sometimes collected from
other murid rodents (particularly Apodemus spp.). The present Pakistan
record from a migratory hamster is considered atypical.

Vol. 101, No. 4, September & October 1990 231

2\)Polyplax spinulosa (Burmeister, 1839) (synonyms: denticulatus

Nitzsch, 1864; campylopteri Zavaleta, 1945).

Specimens examined: (3 collections (6c?,319) ex Rattus rattus (Linn.) (Rodentia,
Muridae)) LAHORE DIST.: Luliani, 1.1964 - 2 colls.; Ravi road, 1X1963 - 1 coll. (1
collection (1 nymph) ex Rattus norvegicus (Berkenhout) (Rodentia, Muridae))
BALUCHISTAN PROV.: Kalat,IV.1976.(5collections(12cf,189, 2 nymphs) e\Ranus
sp. (Rodentia, Muridae)) LAHORE DIST.: 6.2 km NW of Luliani, XII. 1963, 1. 1964 - 4
colls.; RAWALPINDI DIST.: Ayub Natl. Park, XII. 1962 - 1 coll. (1 collection (Icf) ex
Apodemus sp. (Rodentia, Muridae)) HAZARA DIST.: Kaghan valley, Burawai, 3050
m, IX.1963.

Polyplax spinulosa is a cosmopolitan ectoparasite of domestic Rattus
and is probably widely distributed in Pakistan.

22) Polyplax stephensi (Christophers and Newstead, 1906.)

Specimens examined: (62 collections (229cf,4539, 63 nymphs) ex Tatera indica
(Hardwicke) (Rodentia. Cricetidae)) RAWALPINDI DIST.: Ayub Natl. Park,
XXII.1962, III.1963, 1.1964 - 15 colls.; on Murree-Rawalpindi road, 1.1964 - 1 coll.;
DERA ISMAIL KHAN DIST.: unspecified localities, XI.XII.1963. 1.1964 - 6 colls.;
LAHORE DIST.: Near Ravi River bridge, 215 m, 1X1963 - 4 colls.; Luliani. 1X1963,
1.1964 - 3 colls.; 6.2 km NW of Luliani, IX.1963 - 1 coll.; Sodhana, 17.6 km NW of
Luliani, X.1963 - 1 coll.; Lahore, Jahangir's tomb, 1X1962 - 1 coll.; Model Town.
XII.1963 - 1 coll.; Balloki, XII..1962 - 1 coll.; Bari Doab canal. 16 km W of highway.
VI.1964 - 1 coll.; Changa Manga Forest, V. 1965 - 1 coll.; unspecified localities, VII. 1965
-2 colls.; SIALKOT DIST.: Charwa, 275 m. X.1962 - 3 colls.; Marala, III. 1965 - 1 coll.:
BAHAWALPUR DIST.: Uch-Sharif. Abbraria canal. XII.1963. 1.1964 - 3 colls.; Islam
Headworks, VIII. 1963 - 1 coll.; unspecified localities. 1.1964 - 3 colls.; MULTAN
DIST.: Mian Channun. XII.1963, XII.1970 - 3 colls.; SIND PROV.: Karachi: TPX
Godowns. 11.1975, 1.1976 - 2 colls.; port area, 1.1977 - 1 coll.; GUJRAT DIST.:
Mangowal, V1.1965 -2 colls.; HAZARA DIST.: Mansehra, VII.1965 - 2 colls.; LAS-
BELA DIST.: 8 km S of Bela, III. 1965 - 1 coll.: MUZAFFARGARH DIST.: Taunsa
barrage. III. 1964 - 1 coll.; ZORALAI DIST.: Zoralai. XI. 1963 - 1 coll. (4 collections
(7c?,49, 14 nymphs) exMillardia meltada (Rodentia, Muridae)) SIND PROV.: Gharo,
VIII.1976-2colls.; MahroBula Khan, VIII.1976- 1 coll.; Rani Kot, VIII. 1976- 1 coll.(l
collection (Icf) ex Nesokia indica (Rodentia, Muridae)) LAHORE DIST.: Luliani.
1X1963. (1 collection (Id1 ,29, 6 nymphs) ex Gerbillus nanus (Rodentia. Cricetidae))
SIND PROV.: Rani Kot, VII. 1976. (1 collection (29,1 nymph) ex Canis aureus Linn.
(Carnivora, Canidae)) SIND PROV.: Karachi, Hawkes Bay, VIII. 1963. (1 collection
(29) ex Vulpes bengalensis (Shaw) (Carnivora. Canidae)) SIND PROV.: Thatta.
IV. 1976.

Clearly, P. stephensi is principally parasitic on the gerbil, T. indica.
Records from other hosts cited here are presumed to represent acciden-
tal associations although the carnivore infestations could have been
acquired as a consequence of recent predation on gerbils. There are
numerous records of P. stephensi from India, and Kim and Emerson
(1971) reported this species from Iran. Curiously, P. stephensi does not
appear to have been documented previously from Pakistan although it is
possible that some pre-1948 records given as 'India' actually refer to
localities now within Pakistan's borders.

232 ENTOMOLOGICAL NEWS

DISCUSSION

Twenty-two species of sucking lice, at least 14 of them new for the
country, are reported here from wild land mammals of Pakistan. The
large number of new records for Pakistan can be attributed to the scant
literature on the Anoplura of the region. Six additional species of suck-
ing lice have been reported from wild land mammals in Pakistan but
were not recorded during the present survey. Three of these species were
described from the palm squirrel, Funambulus pennanti: Hoplopleura
funambuli Bilquees,//. karachiensis Khanum, and Neohaematopinus qad-
rii Khan and Khanum; the first two were from Karachi in Sind Province
(Bilquees, 1976; Khanum, 1983) and the last was from Thatta District
also in Sind Province (Khan and Khanum, 1980). Two more polyplacids,
Polyplax sindensis Shafi, Samad and Rehana, from the shrew, Suncus
murinus, and P. humae Khan and Khan, from the murid rodent, Crem-
nomys blanfordi (Thomas) have been described from the Karachi area
(Shafi etal., 1984; Khan and Khan, 1985). Lastly, the linognathid louse,
Linognathus vulpis Werneck, was described from the fox, Vulpes ruppelli
(Schinz), near Karachi (Werneck, 1952); L. vulpis has also been collected
from Vulpes vulpes Linn, in Iran (Kim and Emerson, 1971). Detailed
ectoparasite collections are not available for many of Pakistan's native
mammalian species and it is estimated that about 10 additional species
of sucking lice actually parasitize this fauna.

The zoogeographical affinities of Pakistan's anopluran fauna are
worthy of brief consideration here. While most of Pakistan is situated
within the Palearctic Region, the extreme southeastern section of the
country is usually considered to be within the boundaries of the Oriental
Region (Roberts, 1977). Traub et al. ( 1 983) point out that topography and
other factors such as climate, often have a profound effect on zoogeography;
this is especially true for Pakistan's diverse terrain which they categorize
into an 'Arid Southwestern Sector', and two 'Montane Central Asian
Sectors' of the Palearctic and an 'Inter-Indian Sector' of the Oriental
Region.

Predictably, most Anoplura occurring in Pakistan have largely
Palearctic or Oriental distributions. Three species (Hoplopleura affmis, H.
merionidis, Linognathoides palaearctus) have wide, more or less exclu-
sively Palearctic distribution patterns. Nine (Hoplopleura alticola, H.
chinensis, H. funambuli, H. karachiensis, H. pavlovskyi, Linognathus vulpis,
Neohaematopinus qadrii, Polyplax calomysci, P. sindensis} have more limited
geographical and/or altitudinal Palearctic distributions. Three species
of Pakistan gerbil lice (Polyplax brachyrrhyncha, P. kaiseri, P. paradoxd)
are principally Palearctic and are widespread across the Near and Mid-
dle East and North Africa but also extend to varying degrees into the

Vol. 101, No. 4, September & October 1990

Ethiopian Region. Six species are considered here to represent the
Oriental fauna: Hoplopleura rnaniculata, H. ramgarh, Neohaematopinus
echinatus. Polypi ax humae, P. hurrianicus, P. kondana. At least two species
(Polyplax asiatica, P. stephensi} are widespread in both Palearctic and
Oriental Regions. The remaining five species of Anoplura are cos-
mopolitan or nearly so; Polyplax reclinata occurs throughout much of the
Palearctic. Oriental and Ethiopian Regions: Hoplopleura captiosa, H.
pacificia, Polyplax serrata and P. spinulosa are distributed throughout
much of the world.

Many of the louse distributions outlined above clearly mirror those
of their hosts. This is true for the Palearctic H. alticola/A. roylei, P. calo-
mysci/C. bailwardi (both of these associations are at high altitude), H.
ajfinis/Apodemus sp., L. palaearctus/M. caudata associations, the Pal-
earctic-North African P. brachyrrhyncha/Acomys sp. association, and the
Oriental-Palearctic H. maniculata/Funambulus sp., H. ramgarh/Mus sp..
N. echinatus/Funambulus sp., P. asiatica/Nesokia sp. and Bandicota sp., P.
hurrianicus/M. hurrianae, P. kondana/Millardia sp. and P. stephensi/T.
indica associations. The very widespread distributions of P. reclinata on
numerous species of shrews and of the four cosmopolitan lice mainly on
peridomestic murid rodents are likewise understandable. However, cer-
tain species of lice occupy ranges smaller in size than those of their hosts,
possibly reflecting climatic or topographic factors. For example. P.
kaiseri and P. paradoxa parasitize gerbils (Gerbillus and Meriones, respec-
tively) from North Africa to Pakistan but have not been recorded further
east despite the availability of seemingly suitable gerbil hosts. The oppo-
site trend is apparent for H. merionidis and P. chinensis both of which
parasitize Meriones spp. in the southeastern Palearctic but do not extend
to congeneric North African gerbils. To some extent, P. kaiseri and P.
paradoxa replace H. merionidis and P. chinensis as parasites of Meriones
spp. gerbils in the southwestern Palearctic. Similarly, although Rattus
turkestanicus is widely distributed in the south-central Palearctic Region
(Honacki <?/#/., 1982), its characteristic hoplopleurid louse, H.pavlovskyi.
has been reported only from the USSR and (now) Pakistan. The six
Pakistan louse species that were not collected during this survey appear
to have restricted Palearctic distributions but they are known principally
from their respective type series and further collecting may reveal their
presence elsewhere. Only L. vulpis is represented by more than the type
collection and is known from outside of Pakistan. One of these, P. humae,
is known only from Cremnomys blanfordi from the Karachi area; this
murid host is known from India and Sri Lanka (neither Roberts (1977)
nor Honacki et al. (1982) list it for Pakistan) and P. humae is here assumed
to have Oriental affinities.

It is instructive to consider the number of anopluran species (assoc-

234 ENTOMOLOGICAL NEWS

iated with wild land mammals) shared by Pakistan and its neighboring
countries/regions. Extensive literature searches have revealed that the
following numbers of species are shared as part of the 28 species reported
here for Pakistan: India (15 shared species), Iran (14), southwestern
USSR (10), People's Republic of China (10), North Africa (9), Afghanis-
tan (6). Thus, the geographically adjacent Indian (Oriental) and Iranian
(Palearctic) faunas appear to be most similar to the Pakistan fauna. It
must be pointed out, however, that not all of the faunas have received
equal study. Overall, while the Pakistan anopluran fauna has significant
Oriental and minor Ethiopian and cosmopolitan elements, this fauna
principally has Palearctic affinities.

ACKNOWLEDGMENTS

The Pakistan field collections directed by Robert Traub were made under the auspices
of Grant AI-04242 of the National Institutes of Health and of Contract DA-49-193-MD-
2277 of the Army Medical Research and Development Command (AMRDC). Field collec-
tions undertaken by Robert G. Tuck were supported by Contract DA-49-193-63-G74 of the
AMRDC with the Smithsonian Institution (H. W. Setzer, Principal Investigator). We thank
Richard G. Robbins (Department of Health and Human Services, Public Health Services,
National Institutes of Health. National Institute of Allergy and Infectious Disease, c/o
Department of Entomology, Museum Support Center, Smithsonian Institution, Washing-
ton, DC), John O. Whitaker, Jr. (Indiana State University, Terre Haute), and Nixon Wilson
(University of Northern Iowa, Cedar Falls) for reviewing earlier drafts of this paper. Yiau-
Min Huang and Aileen N, Hsu (both with the Department of Entomology, Smithsonian
Institution, Washington, DC) translated Chinese literature and supplied current informa-
tion on Chinese place names.

LITERATURE CITED

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from Funambulus pennanti (Wroughton). Pak. J. Zool. 8: 43-45.
Chin, T.-M. 1985. Studies on Chinese Anoplura VIII. The polyplacid genera Linogna-

thoides and Neohaematopinus. Entomotaxonomia, 7: 39-46.
Ferris, G.F. 1921. Contributions toward a monograph of the sucking lice. Part II. Stanford

Univ. Pubs. Univ. Sen, Biol. Sci. 2: 52-133.
Ferris, G.F. 1923. Contributions toward a monograph of the sucking lice. Part IV. Stanford

Univ. Pubs. Univ. Sen, Biol. Sci. 2: 179-270.
Honacki, J.H., K.E. Kinman and J.W. Koeppl. (eds.) 1982. Mammal species of the

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Johnson, P.T. 1960. The sucking lice (Anoplura) of Egypt I. Species infesting rodents. J.

Egypt. Publ. Hlth. Assoc. 35: 203-228.
Kaneko, K. 1963. Notes on two new sucking lice (Hoplopleuridae; Anoplura) found on

northern palm squirrel from India. Bull. Tokyo Med. Dent. Univ. 9: 129-137.
Kaneko, K. 1972. The murine lice (Anoplura) from Iran. Jap. J. Sanit. Zool. 23: 57-58.
Khan, M.AJ. and R.J. Khan. 1985. A new species of the genus Polyplax Enderlein

(Anoplura: Hoplopleuridae) parasitizing Rattus blanfordi (Thomas) (Rodentia:

Muridae). Pak. J. Zool. 17: 15-20.

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Khanum, Z. 1983. A new species of the genus Hoplopleura Enderlein (Anoplura: Hoplop-

leuridae) parasitizing Funamhulus pennanti Wroughton (Rodentia. Sciuridae). Pak. J.

Zool. 15: 45-49.
Kim, K.C. and P.H. Adler. 1982. Taxonomic relationships of Neohaematopinus to

Johnson pthirus and Linognathoides. J. Med. Entomol. 19: 615-627.
Kim, K.C. and K.C. Emerson. 1971. Sucking lice (Anoplura) from Iranian mammals. J.

Med. Entomol. 8: 7-16.
Kim, K.C. and H.W. Ludwig. 1978. The family classification of the Anoplura. Syst.

Entomol. 3: 249-284.
Ledger, J.A. 1980. The arthropod parasites of vertebrates in Africa south of the Sahara.

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Mishra, A.C. 1980. Sucking lice as a definitive index in the rodent systematics. pp. 23-31.

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Mishra, A.C. 1981. The hoplopleuridliceofthe Indian subcontinent (Anoplura: Hoplop-

leuridae). Rec. Zool. Surv. India, Misc. Publ. Occ. Pap. 21: 1-128.
Mishra, A.C. and H.R. Bhat. 1972. Hoplopleura vandeleuria sp. nov. and Hoplopleura

alticola sp. nov. (Anoplura: Hoplopleuridae) from Indian rodents. Oriental Insects 6:

521-530.
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Hoplopleura sinhgarh sp. nov. (Anoplura: Hoplopleuridae) parasitizing Mus spp.

(Rodentia: Muridae) in India. Parasitology 65: 11-21.

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Pratt, H.D. and C.J. Stojanovich. 1961. Notes on the Indian sucking lice Hoplopleura

maniculata (Neumann) and Neohaematopinus echinatus (Neumann) (Anoplura: Hop-
lopleuridae). J. Kansas Entomol. Soc. 34: 79-83.

Roberts, T.J. 1977. The mammals of Pakistan. Ernest Benn Ltd.. London, xxvi + 361 p.
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Commun. Dis. 19: 179-181.
Shafi, M.M., K. Samad and R. Rehana. 1984. Polyplax sindensis. new species (Anoplura:

Hoplopleuridae) parasitizing Suncus murinus sindensis (Insectivora: Soricidae) in

Pakistan. Pak. J. Zool. 15: 45-49.
Sosnina, E.F. 1951. A new species of Turkestan rat louse from Tadzhikistan. Doklady

Akad. Nauk SSR. 77: 365-368.
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ectoparasites on the host's habitat (on the example of Rattus turkestanicus). Wiad.

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Ceratophyllidae: key to the genera and host relationships. With notes on their evolu-
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12: 69-78.

236 ENTOMOLOGICAL NEWS

A CHECKLIST OF THE CADDISFLIES
(TRICHOPTERA) FROM WEST VIRGINIA1

Donald C. Tarter2

ABSTRACT: County distributional records and dates of adult capture are provided for 176
species, representing 15 families and 60 genera, collected in West Virginia. Hydro-
psychidae ranked first in number of species (39), followed by Limnephilidae (30), Lep-
toceridae (27), Hydroptilidae (16), Rhyacophilidae (14), and Polycentropodidae (13); the
other nine families accounted for 37 species. Of the species reported, 42 are new state
records. The largest number of species (62) was collected in Pocahontas County.

This paper represents the first checklist of caddisfiies for West
Virginia. Ross (1944) listed 13 species for the state. Over the last two
decades, several publications, including Applin and Tarter (1977), Hill
et. al. (1977, 1978), Hill and Tarter (1978), Tarter and Hill (1979, 1982),
Nugen and Tarter (1983), Tarter and Donahoe (1989), and Glover and
Tarter (1989), have contributed additional records. The collection dates
and records, based on adults unless otherwise mentioned, listed in this
paper are a compilation of literature records, West Virginia Department
of Agriculture (WVDA) and West Virginia University (WVU) specimens,
unpublished records from the National Museum of Natural History
(NMNH) and Army Corps of Engineers, Huntington District (USCE),
and individual collecting efforts by (Ralph Kirchner-(RFK) and Donald
Tarter, West Virginia Benthological Survey, Marshall University
(WVBS/MU). The purpose of this checklist is to provide a baseline for
additional work on the West Virginia caddisfly fauna.

One hundred and seventy-six species of caddisflies, representing 15
families and 60 genera, were recorded for West Virginia; 42 species are
new state records (*). I followed the classification of Wiggins (1977). The
following families in descending order contained caddisfly species:
Hydropsychidae (39), Limnephilidae (30), Leptoceridae (27), Hydrop-
tilidae (16), Rhyacophilidae (14), and Polycentropodidae (13) (Fig. 1).
Nine additional families accounted for 37 species. Species records are
listed for counties (Fig. 2). In descending order, the following counties
contained the greater number of caddisfly species: Pocahontas (62),
Randolph (54), Summers (45), Pendleton (47), Monroe (42), Kanawha
(31), Jefferson (30), Braxton (28), Wayne (26), Hardy (22), Tucker (23),
Hampshire (21 ), and Putnam (23). The remaining 36 counties contained
at least one caddisfly record (Fig. 2). Numbers in parentheses at the end
of each citation, corresponding with those in Figure 2, indicate the
county records for the species.

1 Received February 7, 1990. Accepted April 20, 1990.

^ Department of Biological Sciences, Marshall University, Huntington, WV 25755.

ENT. NEWS 101(4): 236-245, September & October, 1990

Vol. 101, No. 4, September & October 1990 237

Philopotamidae

Chimarra aterrima Hagan; Ross (1944), USCE, NMNH. August; (8,30).
C. augusia Morse; Lago and Harris (1987), no date; (8, 27, 28, 45).

* C. obscura (Walker); USCE. June; (27, 44).

C. soda Hagen; USCE, NMNH. WVBS/MU, MVU. May to August; (5, 8. 27, 44).

* Dolophilodes distinctus (Walker); MVBS/MU. July, August; (7, 8, 10. 27, 47).
Wormaldia moesta (Banks); Ross ( 1944, as Dolophilus. no county record), NMNH. May to
July; (9, 27, 29).

Psychomyiidae

* Lype diversa (Banks); USCE, NMNH, WVBS/MU. July; (15, 29).

Psychomyia flavida Hagen; Ross (1944), USCE, NMNH, WVBS/MU. May to October;
(4, 8, 44, 45).

Polycentropodidae

* Cymellusfratemus (Banks); USCE, WVBS/MU. September; (40, 41, 44, 51).

* Neureclipsis crepuscularis (Walker); USCE, WVBS/MU. MVU. June to September; (40. 44.
51,55).

Nyctiophylax affinis (Banks); Tarter and Hill (1979). July: (27).

N. celta Denning; Morse (1972); no county record.

N. moestus Banks; Morse (1972) and Tarter and Hill (1979). June; (27).

/Vu'/omjrropws/unWws (Hagen); Tarter and Hill(1979). Stout and Stout (1989), WVU. May,

June; (10. 15.27).

* P. placidus (Banks); USCE. WVBS/MU. August; (30, 51).

* Polvcentropus cinereus Hagen; USCE, NMNH. WVBS/MU. May to August; (7. 8. 30.
44,'51).

P. clinei (Milne); Tarter and Hill (1979), NMNH. June, July; (9. 27).

* P. colei Ross; NMNH. WVBS/MU. June; (9).

* P. confusus Hagen; NMNH. WVBS/MU. August; (8).

P. maculatus Banks; Tarter and Hill (1979). NMNH. June to August; (8, 9, 27).
P. pentus Ross; Tarter and Hill (1979), NMNH. June; (9, 27).

Hydropsychidae

* Cheumatopsychecampvla Ross; Phillips (1982). USCE. WVBS/MU. WVD A. MVU. April
to September; (1, 2. 4, 7, 10, 17, 30, 40, 42-45, 55).

* C. ela Denning; Phillips (1982); WVBS/MU. June, July; (2. 7. 27. 44).

C. gracilis (Banks); Gordon (1974). Phillips (1982), NMNH. June, August; (7, 8).

* C. gyra Ross; Phillips (1982), NMNH. July; (15).

* C. halima Denning; Phillips (1982), WVBS/MU, MVU. June, July; (10, 27).

C. harwoodi Denning; Gordon (1974), Phillips (1982), WVDA. MVU. May to August; (6.
27, 30, 45).

* C. helma Ross; Phillips (1982), WVBS/MU, WVU. June to August; (2, 27. 30).

* C. minuscula (Banks); Phillips ( 1982), WVBS/MU, WVDA. June to August; (2, 7. 27. 29. 42.
45, 47).

C. oxa Ross; Ross (1944), Phillips (1982), WVDA. May to June; (1. 45).

238 ENTOMOLOGICAL NEWS

* C. pasella Ross; Phillips (1982). WVBS/MU. June, July; (29, 49).

*C.pertiti (Banks); Phillips (1982), WVBS/MU. April to October; (15, 17,25,30,32,41, 42, 45,
47, 50, 51, 55).

* C. sordida (Hagen); Phillips ( 1982), WVBS/MU. WVDA, WVU. June to August; (2, 5, 15,
27, 44).

* C. wrighti Ross; Phillips (1982), WVBS/MU. August; (8).

Diplectrona modesta Banks: Ross, (1944), Tarter and Hill (1979), Phillips (1982), WVDA,
WVU, RFK. May to September; (7-9, 11, 15, 17, 26-28, 38, 39, 41-43, 45, 47, 48, 51, 52,
55).

* Homoplectra monticola (Flint); Phillips (1982), WVBS/MU. May; (27).

* Hydropsvche alhedra Ross; Phillips (1982); WVBS/MU, WVDA. July, August; (8, 27,
46, 47). '

* H. betteni Ross; Phillips ( 1982), USCE, WVBS/MU, WVDA June to September; ( 1 , 1 5, 1 7,
23,29,38,39,42,45,47,51,55).

H. bronta Ross; Phillips (1982), Nugen and Tartar ( 1983) (larva); May to September; (7,15,

23, 27, 29, 30, 41, 45, 47, 48, 52).

H. brunneipennis Flint and Butler; Flint and Butler (1983). June; (27).

* H. cheilonis Ross; Phillips (1982); WVBS/MU, WVDA May to October; (1, 7. 8, 42, 45).
H. dicantha Ross; Phillips (1982), Nugen and Tarter (1983) (larva), WVDA June to Sep-
tember; (1, 2, 9-11, 15, 29, 30, 40, 49, 53, 55).

H. hageni Banks; Phillips (1982). Nugen and Tarter (1983) (larva). WVDA June to Sep-
tember; ( 1-5, 42-44, 45).

H. hoffmani Ross; Phillips (1982). Nugen and Tarter (1983) (Irva), WVDA. May to August;
(1,44').

H. leonardi Ross; Phillips ( 1982). Nugen and Tarter ( 1983) (larva). MVDA. June to August;
(1,44).

H. macleodi Flint; Nugen and Tarter (1983) (larva); (7-9. 13, 27).

H. morosa Hagen; Ross (1944), Phillips (1982), Nugen and Tarter (1983), WVDA June to
September. (1-6. 8, 15, 27, 30, 42, 44, 45, 52, 55).
H. opthalmica Flint; Flint (1965). Phillips (1982). May, August; (1, 7).
H. orris Ross; Phillips ( 1982), USCE, WVDA. June to September; (4, 9, 15, 17, 30, 39-42, 44,
45, 50).

H. phalerata Hagen; Phillips (1982), Nugen and Tarter (1983), WVDA, WVU. May to Sep-
tember; (1, 5, 7, 27, 44).

H. scalaris Hagen; Phillips (1982), Nugen and Tarter (1983), WVDA. WVU. May to Sep-
tember; (1, 5, 7. 27, 44).

H. simulans Ross; Phillips ( 1982), Nugen and Tarter ( 1983) (larva), USCE, WVDA. June to
October; (39, 41, 42, 51, 55).

H. slossonae Banks; Phillips (1982), Nugen and Tarter ( 1983) (larva). July to September; (7,
8, 15,27,43,46,47,54,55).

H. spama Ross; Ross (1944), Hill and Tarter (1978), Phillips (1982), Nugen and Tarter
(1983) (larva). May to September; (2, 3, 7, 9, 27-30, 38, 42, 45, 47, 48, 51, 55).
H. ventura Ross; Phillips( 1982), Nugen and Tarter( 1983) (larva). May to August; (7, 11,27-
30, 44, 45).

* H. venularis Banks; Phillips (1982), USCE, WVBS/MU. no date; (26, 37).

H. walkeri (Betten and Mosely); Phillips ( 1982), Nugen and Tarter ( 1983) (larva), WVDA.
June to September (4, 6, 15, 26, 27, 29, 43-45).

Macrostemum zebratum (Hagen); Ross (1944, as Macronemum), Phillips (1982) WVBS/
MU, WVDA. June to October; (1, 15, 27, 30, 42-44).

* Parapsyche apicalis (Banks); Phillips (1982), WVBS/MU. June to September; (15, 27, 44).

* Potamviaflava (Hagen); Phillips (1982), USCE, WVBS/MU, MVDA. June to September:
(1, 17,"23, 38-42, 50, 51).

Vol. 101, No. 4, September & October 1990 239

Rhyacophilidae

* Rhyacophila banksi Ross; RFK, WVBS/MU. no date; (27).

R. Carolina Banks; Applin and Tarter (1977, larva). Tarter and Hill (1979), June. July; (7, 9.
26, 27, 42, 43).

* R. carpenteri Milne; WVBS/MU (larva); (15, 27).

RJuscula (Walker); Ross (1944), Applin and Tarter (1977, larva). Hill and Tarter (1978).

June, August, September; (8, 9. 15, 25-29. 43-46).

R. glaberrima Ulmer; Applin and Tarter (1977, larva); (15).

R. invaria (Walker); Applin and Tarter (1977, larva); (15, 27).

R. ledra Ross; Applin and Tarter (1977, larva); (51).

R. loibifera Betten; Applin and Tarter (1977. larva); (39).

R. melita Ross; Applin and Tarter (1977, larva); (15).

R. minora Banks; Roback (1975), Applin and Tarter (1977, larva). May; (7-9, 15).

R. nigrita Banks; Applin and Tarter (1977, larva); (6, 8, 15, 26, 27).

R. torva Hagen; Roback (1975), Applin and Tartar (1977, larva); (9. 43).

R. vibox Milne; Applin and Tarter (1977, larva); (15, 50-52).

R. vuphipes Milne; Applin and Tarter (1977. larva); (4).

Glossosomatidae

* Agapetus minutus Sibley; WVBS/MU, MVU. July; (10).

* Glossosoma nigrior Banks; WVBS/MU, MVU. July; (10).

* Protoptila maculata (Hagen); WVBS/MU. August; (8).

P. palina Ross; Ross (1944, no county record), USCE, WVBS/MU. July-August; (4, 44).

Hydroptilidae

* Dibusa angata Ross; USCE. June-July; (29-51).

Hydroptila ajax Ross; Tarter and Donahoe (1988). September; (42).

H. armata Ross; Tarter and Donahoe (1988). August; (4).

H. delineata Morton; Tarter and Donahoe (1988). August; (4).

H. grandiosa Ross; Tarter and Donahoe (1988), USCE. June, July, September; (2, 55).

H. hamata Morton; Tarter and Donahoe (1988). USCE. September; (30, 42, 44).

H. perdita Morton; Tarter and Donahoe (1988). August; (4).

* Leucotrichia pictipes (Banks); USCE. June, July; (27, 44).
Neotrichia vibrans Ross; Tarter and Donahoe (1988). September; (51).
Ochrotrichia dardeni Harris; Tarter and Donahoe (1988). August; (4).
O. denningi Blickle and Morse; Blickle (1979) (no county record).

0. gravsoni Parker and Voshell; Tarter and Donahoe (1988), USCE; September; (42,

44).

O. tarsalis (Hagen); Tarter and Donahoe (1988). September; (42).

Oxyethira pallida (Banks); Tarter and Donahoe (1988). August; (4).

* Palaeagapetus celsus (Ross); RFK, NMNH., June, July; (27).

Stactobiella delira (Ross); Smith (1969), Tarter and Donahoe (1988). September; (6).

Phryganeidae

Agrypnia vestita (Walker); Hill et al. (1978), Tarter and Hill (1979). April to October; (1.10.
15, 17, 26-28, 41, 42, 44, 45, 47).

240 ENTOMOLOGICAL NEWS

* Banksiola crotchi Banks; WVBS/MU, MVDA. July; (23).

B. dossuaria (Say); Hill et al. (1978), Stout and Stout (1989), RFK, USNM. WVU. May to
July; (9-1 1,15,27,45,53).

Oligostomis pardalis (Walker); Stout and Stout (1989). June; (15).

Phryganeasayi Milne; Hill etal.( 1978), WVDA. July to October;(l, 15, 17.23,25,41, 42,45-
47, 51).

Ptilostomis ocellifera (Walker); Hill et al. (1978), Tarter and Hill (1979), Stout and Stout
(1989). June to September; (2, 4, 5, 7, 10, 15, 47).
P. postica (Walker); Hill et al. (1978), WVDA. August; (2).

P. semifasciata (Say); Hill et al. (1978), Tarter and Hill (1979), USNM, WVDA. May to Sep-
tember; (4. 8. 9. 15. 17,27,44).

Brachycentridae

Brachycentrus appalachia Flint; Flint (1984). May; (15).
B. lateralis (Say); Flint (1984). May; (4).
B. nigrosoma (Banks); Flint (1984). May; (4).
*B. numerosus (Says); WVBS/MU. April; (8. 15).
B. solomoni Flint; Flint (1984). April; (8).

* Micrasema scotti Ross; WVBS/MU, NMNH. May; (8).

Limnephilidae

Frenesia difficilis (Walker); Tarter and Hill (1980), Stout and Stout (1989), WVU. May, Sep-
tember, November; (11, 15).

* Goera calcarata Banks; WVBS/MU, NMNH. August; (8).

Goerita betteni Ross; Ross (1962, no county record), Tarter and Hill (1980). June, July;
(27).

Hydatophvlax argus (Harris); Hill and Tarter ( 1978), Tarter and Hill ( 1980), WVDA, WVU.
June, July; (1, 15, 27, 29, 43, 45).

Ironoquia kaskaskia (Walker); Hill and Tarter (1978), Tarter and Hill (1980), WVDA. Sep-
tember; (15, 51).

/. punctatissima (Walker); Hill and Tarter (1978), Tarter and Hill (1980). NMNH. WVDA
June, July, September, October; (6, 9, 15, 17, 23, 26. 41, 42. 45, 47, 51).
Leptophylax gracilis Banks; Hill and Tarter (1978), WVDA. June to September; (45).
Limnephilis indivisus Walker; Hill and Tarter (1978), Tarter and Hill (1980), WVDA, WVU.
June, September; (1, 15, 30, 45).

L moestus Banks; Hill and Tarter (1978), Tarter and Hill (1980), Stout and Stout (1989),
NMNH, WVU. May to August; (9, 10, 15, 27).

L. omatus Banks; Tarter and Hill (1980), WVDA. June. July; (41, 45).
L. rhombicus (Linnaeus); Tarter and Hill (1980), WVBS/MU. May; (1).
L. submonilifer Walker; Hill and Tarter ( 1978), Tarter and Hill (1980), WVDA, WVU. June,
August, October; (10, 11, 15, 41, 49).

Nemotaulis hostilis (Hagen); Hill and Tarter (1978), Tarter and Hill (1980). Stout and Stout
(1989), WVDA, WVU. August; (9, 15, 27).

Neophylax anigua Ross; Tarter and Hill (1980), WVDA. September; (15).
N. concinnus McLachlan; Hill and Tarter (1978), WVDA. MVU. September. October; (10,
15,41,47).

N. consimilis Betten; Hill and Tarter (1978), Tarter and Hill (1980), WVDA, WVU. Septem-
ber, October; (15, 30, 45, 49).

Vol. 101. No. 4, September & October 1990 241

N. stolus Ross; Hill and Tarter (1978), NMNH. October; (8).
N. wigginsi Sykora and Weaver; Tarter and Hill (1980). WVDA. September; (15).
Platycentropus radiatus (Say); Hill and Tarter ( 1 978), Tarter and Hill (1980), WVDA, WVU.
May to October; (5, 701 1. 27. 44. 45. 47).

Pseudostenophvlax sparsus (Banks); Hill and Tarter (1978), Tarter and Hill (1980). RFIC
NMNH. WVDA, WVU. May to July; (1, 6, 7, 9-1 1. 17. 27, 34, 41).
P. uniformis (Betten); Hill and Tarter ( 1978), Tarter and Hill (1980). RFK, NMNH. WVU.
May to July; (8, 9, 1 1, 27, 43. 44).

Pycnopsvche circularis (Provancher); Hill and Tarter (1978). WVDA. September; (41-
45).

P. divergens (Walker); Hill et al. (1977). Hill and Tarter (1978). Tarter and Hill (1979, 1980).
WVBS/MU. July to August; (26. 27).

P. gentilis McLachlan; Hill and Tarter (1978). Tarter and Hill (1980). WVDA. WVU, Sep-
tember, October; (4, 8, 10, 15, 18, 29. 45. 47, 49, 51).
P. guttifer (Walker); Hill and Tarter (1978), NMNH. October: (8).
P. Indiana (Ross): Hill and Tarter (1978), NMNH. October; (8).

P. lepida (Hagen); Ross (1944). Betten (1950). Hill and Tarter(1978). Tarter and Hill (1979).
WVDA. WVU. June to October; (15. 23, 26, 27, 41, 45, 47, 49, 51).
P. luculenta (Betten); Hill and Tarter (1978), Tarter and Hill (1979, 1980). WVDA. WVU.
July to October; (8, 10, 15. 26-28. 30. 45. 47).

P. scabripennis (Rambur); Ross (1944). Hill and Tarter (1978). Tarter and Hill (1979, 1980).
WVDA. WVU. June to October: (3. 8. 10, 1 1, 15. 16. 27. 29. 30. 41. 42. 45-47. 49. 51).
P. subfasciara (Say); Hill and Tarter (1978). Tarter and Hill (1980). WVDA. August to
October; (1. 15.45.49).

Lepidostomatidae

Lepidostoma griseum (Banks); Tarter and Hill (1979), WVBS/MU. August; (27).

* L. pictile (Banks); Weaver (1988), WVBS/MU. May; (8, 9).

L. sackeni (Banks); Tarter and Hill (1979), MVBS/MU. August; (27).

* L. stylifer Flint and Wiggins; WVBS/MU. NMNH. May; (8).
L. togatum (Hagen): Weaver (1988). April; (8).

L. vemale (Banks); Weaver (1988). April; (1).

* Theliopsyche melas Edwards; WVBS/MU. June; (8).

Odontoceridae

Psilotreia labida Ross; Parker and Wiggins (1987). June: (8).
P. rossi Wallace; Parker and Wiggins (1987). NMNH. July; (27).

Molannidae

Molanna blenda Sibley; Tarter and Hill (1979). WVBS/MU. August: (27).

Helicopsychidae

Helicopsyche borealis (Hagen): Ross (1944) (no date or county record), USCE, NMNH.
WVBS/MU. June. July: (4, 8. 27. 44).

242

ENTOMOLOGICAL NEWS

Leptoceridae

Ceraclea cancellata (Betten); Glover and Tarter (1989), USCE, WVDA June to August; (1,

6, 8, 21, 22, 27, 30, 39, 40-42, 44, 45, 47, 51).

C. diluta (Hagen); Glover and Tarter (1989). July; (27).

C.Jlava (Banks); Glover and Tarter (1989), USCE. June, July; (29, 40, 41).

C. maculata (Banks); Glover and Tarter ( 1989), USCE, WVDA. WVU. June to September;

(1, 11. 22, 30, 39, 40-42, 44, 45, 51).

C. neffi (Resh); Glover and Tarter (1989), WVDA, WVU. June to August; (8, 27, 30, 44).

C. ophioderus (Ross); Glover and Tarter (1989). June, August; (44).

C. slossonae (Banks); Glover and Tarter (1989), WVBS/MU. No date; (4).

C. tarsipunctata (Yorhies); Glover and Tarter (1989), USCE, WVDA. June, July; (1, 30. 42,

44, 51).

C. transversa (Hagen); Glover and Tarter ( 1989), USCE, WVDA. June to September; (27,

29. 30, 40, 42. 44, 45).

C. wetzeli (Ross); Glover and Tarter (1989). NMNH. May; (8).

Leptocerus americanus (Banks): Glover and Tarter (1989). July; (32).

Mystacides sepulchralis (Walker); Yamamoto and Wiggins (1964), Glover and Tarter

(1989). July, August; (2, 4, 30).

Nectopsyche Candida (Hagen); Glover and Tarter (1989), USCE. July; (40).

N. exquisita (Walker); Glover and Tarter (1989), USCE, WVDA, WVU. June to August; (1,

8, 30, 40, 42, 44, 45, 50).

N. pavida (Hagen); Glover and Tarter (1989), USCE. WVDA. July, August; (1, 10, 30,

40.51).

Oecetis avara (Banks); Ross (1944), Glover and Tarter (1989), USCE, NMNH. WVDA.

May to September; (1, 6-8, 29, 30. 42. 44, 45).

O. cinerascens (Hagen); Glover and Tarter (1989). WVDA July to October; (1. 15, 40,

(text continued on page 244)

Philopotamidae

Psychomyudae

Po 1 yc en t ropod i dae

Hydropsychidae

Rhyacophilidae

Glossosomatidae

Hydrop till dae

Phryganeidae

Brachycentridae

Limnephilidae

Lepi dost omati dae

Odontoceridae

Molannidae

Helicopsychidae

Leptoceridae

0 15 30

Figure 1. West Virginia caddisfly families with number of species.

Vol. 101, No. 4. September & October 1990

243

Figure 2. West Virginia map showing sampling stations of caddisflies by counties.

244 ENTOMOLOGICAL NEWS

O. inconspicua (Walker); Glover and Tarter (1989), USCE, WVDA, WVU. June to Septem-
ber; (1, 4, 6-8, 10, 11, 13, 15, 17, 23, 27, 29, 30, 31, 40-42, 44, 45, 47, 48, 50, 54).
O. noctuma Ross; Glover and Tarter ( 1989), USCE, WVBS/MU. June, July, September; (6,
8,27,40,41,42,44,51).

O. persimilis (Banks); Glover and Tarter (1989), WVDA. June to August; (27, 30, 39, 44,
50).

Setodes incerta (Walker); Glover and Tarter (1989), WVDA. June to August; (44).
Triaenodes flavescens Banks; Glover and Tarter (1989). July, August; (17, 44, 45).
T. Ignitus (Walker); Glover and Tarter (1989). April, June; (50, 51).
T. injustus (Hagen); Glover and Tarter (1989), WVDA. May to August; (15, 41, 42, 44).
T. marginatus Sibley; Glover and Tarter (1989), WVDA July, August; (44).
T. pemus Ross; Glover and Tarter ( 1989), WVBS/MU. June; (44).
T. tardus Milne; Glover and Tarter (1989), WVDA September; (41).

ACKNOWLEDGMENTS

I am grateful to the following persons and institutions for the loan of specimens and/or
records: Linda Butler (WVU), Charles Coffman (WVDA), Oliver S.Flint, Jr., (NMNH), Jan
Hacker (WVDA), Brian Haganbuch (WVDA). Ralph F. Kirchner (USCE), and the West
Virginia Benthological Survey /Marshall University. Special thanks to Oliver S. Flint, Jr..
and Robert W. Kelley for identifications, and to Lu Ann South for typing the manuscript.
Oliver S. Flint, Jr. and Ralph F. Kirchner made suggestions and comments on the
manuscript.

LITERATURE CITED

Applin, J.S., and D.C. Tarter. 1977. A taxonomic study of the larval caddisflies in the

genus Rhyacophila in West Virginia (Trichoptera: Rhyacophilidae). Ent. News 88:2 13-2 14.
Betten, C. 1950. The genus Pycnopsyche (Trichoptera). Ann. Ent. Soc. Amer. 43:508-522.
Blickle, R.L. 1979. Hydroptilidae (Trichoptera) of America North of Mexico. Sta. Bull.

509, N. H. Agr. Expt. Sta. Durham, NH. 97 pp.
Flint, O.S., Jr. 1965. New species of Trichoptera from the United States. Proc. Entomol.

Soc. Wash. 67:168-176.
Flint, O.S., Jr. and W.L. Butler. 1983. Hydropsyche brunneipennis, new species, a member

of the scalaris group, from the Potomac River near Washington, D.C. (Trichoptera: Hyd-

ropsychidae). Proc. Entomol. Soc. Wash. 85:205-211.
Flint, O.S., Jr. 1984. The genus Brachycentrus in North America, with a proposed phylo-

geny of the genera of Brachycentridae (Trichoptera). Smithsonian Contrib. Zool. No.

398. 58 pp.
Glover, J.B., and D.C. Tarter. 1990. The Leptoceridae (Trichoptera) of West Virginia.

Ent. News 101 (1): 35-38.
Gordon, A.E. 1974. A synopsis and phylogenetic outline of the Nearctic members of

Cheumatopsyche. Proc. Acad. Nat. Sci. Philadelphia 126:117-160.
Hill, P.H., D.C. Tarter, W.D. Watkins, S. Nance. 1977. A new state record in the genus

Pycnopsyche in West Virginia (Insecta: Trichoptera). Proc. W. Va. Acad. Sci. 49:25.
Hill, P.H.,, D.C. Tarter, B. Clemeans, and M.B. Roush. 1978. State records of the family

Phryganeidae in West Virginia (Insecta: Trichoptera). Proc. W. Va. Acad. Sci. 50:24.
Hill, P.H., and D.C. Tarter. 1978. A taxonomic and distributional study of the adult cad-
disflies of the family Limnephilidae of West Virginia (Insecta: Trichoptera). Ent. News

89:214-216.

Vol. 101, No. 4, September & October 1990 245

Lago, P.K., and S.C. Harris. 1987. The Chimarra (Trichoptera: Philopotamidae) of
eastern North America with descriptions of three new species. J. N.Y. Entomol. Soc.
95:225-251.

Morse, J.C. 1972. The genus Nyctiophylax in North America. J. Kansas Ent. Soc. 45:172-181.

Nugen, C., and D.C. Tarter. 1983. Larval Hydropsyche and Symphitopsyche records from
West Virginia (Trichoptera: Hydropsychidae). Ent. News 94:18-20.

Parker, C.R., and G.B. Wiggins. 1987. Revision of the caddisfly genus Psilotreta (Tricho-
ptera: Odontoceridae). Life Sci. Contr. Roy. Ont. Mus. 144:1-55.

Phillips, T. 1982. A distributional and taxonomic study of the adults of the family Hydro-
psychidae (Trichoptera) in West Virginia. Unpub. Masters thesis. Marshall University.
Huntington, West Virginia.

Roback, S.S. 1975. New Rhyacophilidae records with some water quality data. Proc. Acad.
Nat. Sci. Pa. 27:45-50.

Ross, H.H. 1944. The caddisflies, or Trichoptera. of Illinois. Bull. Illinois Nat. Hist. Sun .
23:1-326.

Ross, H.H. 1962. Three new species of Trichoptera from eastern North America. Ent. News
73:129-133.

Smith, S.D. 1969. New species of Idaho Trichoptera with distributional and taxonomic
notes on other species. J. Kansas Ent. Soc. 42:46-53.

Stout, B.M., and J.S. Stout. 1989. Northern caddisfly (Trichoptera) fauna in a remnant
boreal wetlands of West Virginia. Ent. News 100(1 ):37-40.

Tarter, D.C., and P.L. Hill. 1979. Caddisflies (Trichoptera) of the Cranberry Glades in
West Virginia. Ent. News 90:205-206.

Tarter, D.C., and P.L. Hill. 1980. Adult limnephilid caddisfly records in West Virginia
(Trichoptera: Limnephilidae). Ent. News 91:170-172.

Tarter, D.C. and S.R. Donahoe. 1989. State records for adult micro-caddisflies from West
Virginia (Trichoptera: Hydroptilidae). Proc. W. Va. Acad. Sci. 60(2, 3. 4): 138- 144.

Weaver, J.S., III. 1988. A synopsis of the North American Lepidostomatidae (Tricho-
ptera). Contrib. Amer. Ent. Inst. 24(2). 141 pp.

Wiggins, G.B. 1977. Larvae of the North American caddisfly genera. Univ. Toronto Press.
Toronto. 401 pp.

Yamamoto, T., and G.B. Wiggins. 1964. A comparative study of the North American
species in the caddisfly genus Mystacides (Trichoptera: Leptoceridae). Can. J. Zool.
42:1105-1126.

246 ENTOMOLOGICAL NEWS

COLLECTIONS OF DROSOPHILIDAE (DIPTERA)

IN KENYA, WITH DESCRIPTION OF A NEW

SPECIES OF DETTOPSOMYIA1

H. Takada2, R.C. Woodruff3, J.N. Thompson, Jr.4

ABSTRACT: Drosophilids (Diptera) were collected by sweeping in 52 localities in Kenya
from May to August, 1988. Natural population samples of Drosophila melanogaster were
collected from 16 sites to screen for variation in mutation rates and other genetic factors. A
total of 37 other species of drosophilid flies were also captured. A new species in the genus
Dettopsomyia is described, and three new records are reported.

Natural populations of Drosophila melanogaster carry transposable
DNA elements that increase mutation rates and cause other genetic
changes (Woodruff et al., 1983). Populations differ in the number of
active elements in each individual. Since the species is thought to have
originated in Africa, collections in Kenya were made by two of us (R.
Woodruff, all sites; and J. Thompson, sites 22-31; see Figure 1) to survey
transposable DNA levels (Woodruffs al., 1989). A total of 34 locations
were sampled. In doing this, 35 other species of the subfamily Droso-
philinae and 2 species of the subfamily Steganinae were identified
among 1,211 alcohol specimens (deposited at Sapporo University, Sap-
poro, Japan) and 36 live strains (maintained at Bowling Green State
University, Bowling Green, Ohio).

Information on species distributions, collection locations and
methods are given below. All collection locations are cross-referenced to
the map of Kenya (Figure 1). Information on drosophilid species is sum-
marized in Wheeler (1981, 1986), and the Afrotropical drosophilid fauna
is discussed in Tsacas et al. ( 1 98 1 ). A new species, Dettopsomyia woodruff,
is described, and new distribution records are provided for three species
of the subgenus Drosophila: Drosophila mercatorum, D.fulvimacula, and
D. virilis.

Genus Zaprionus Coquillett

1. Zaprionus tuberculatus Malloch, 1932. Stylops. 1(1): 11.
Specimens examined: 144 males and 131 females.
Locations: 1, 4-7, 1 1, 14, 16, 20, 26, 28, 29, 31, 34

Collection methods: sweeping and trapping on banana, mango, and other fruit, rotting
tomatoes and fungus.

^Received December 29, 1989. Accepted June 1, 1990.

"Faculty of General Education, Sapporo University, Sapporo, Japan;

•'Department of Biological Sciences, Bowling Green State University, Bowling Green,

Ohio;

Department of Zoology, University of Oklahoma, Norman, Oklahoma.

ENT. NEWS 101(4): 246-255, September & October, 1990

Vol. 101, No. 4, September & October 1990 247

2. Zaprionus indianus Gupta, 1970. Proc. Ind. Nat. Sci. Acad. 36: 63.
Specimens examined: 109 males and 108 females.

Locations: 1, 4, 6-8, 10, 13, 14, 16, 18-20. 26-31, 34

Collection methods: sweeping and trapping on banana, mango, and other fruit, rotting

tomatoes and fungus.

3. Zaprionus ghesquierei Collart 1937. Bull Mus. Roy. Hist. Nat. Belg. 13(9): 8
Specimens examined: four males and three females.

Distribution: Afrotropical Region

Locations: 20, 31

Collection methods: sweeping on rotting fruits

4. Zaprionus inermis Collart, 1937. Bull. Mus. Roy. Hist. Nat. Belg. 13(9): 11
Specimens examined: two males and one female.

Distribution: Central Africa

Location: 28

Collection methods: sweeping on rotting tomatoes

Genus Drosophila Fallen

Subgenus Sophophora Sturtevant

melanogaster species group Sturtevant

melanogaster species subgroup Hsu

5. Drosophila melanogaster Meigen, 1830. Syst. Beschr. Bek. Eur. Zweifl. Ins. 6: 85
Specimens examined: 113 males and 23 females

Distribution: cosmopolitan

Locations: 1, 3, 4, 6, 7, 11, 14-16, 18, 22, 23, 25. 27, 28, 31, 32. 34
Collection methods: sweeping and trapping on banana, mango, and other fruit, rotting
tomatoes and fungus.

6. Drosophila simulans Sturtevant, 1919. Psyche 26: 153
Specimens examined: 200 males and 66 females
Distribution: cosmopolitan

Locations: 1, 4. 5. 7-11, 13, 14, 16, 18-20, 24-29, 32

Collection methods: sweeping and trapping on banana, mango, and other fruit, rotting
tomatoes and fungus.

7. Drosophila yakuba Burla, 1954. Rev. Suisse Zool. 58: 161
Specimens examined: 15 males and two females
Distribution: Afrotropical Region

Locations: 4-7, 1 1, 31

Collection methods: sweeping on bananas and other rotting fruit

montium species subgroup Hsu

8. Drosophila nikananu Burla, 1954. Rev. Suisse Zool. 58: 160
Specimens examined: two males and three females
Distribution: Central Africa

Locations: 30

Collection methods: sweeping on rotting banana

9. Drosophila seguyi Smart, 1945. Proc. Roy. Ent Soc. Lond. (B), 14: 56
Specimens examined: living culture

Distribution: widespread in Africa

248 ENTOMOLOGICAL NEWS

Locations: 11, 34

Collection methods: sweeping on rotting fruit

10. Drosophila vulcana Graber, 1957. Zool. Jahrb. Abt. Syst. 85: 309
Specimens examined: ten males

Distribution: widespread in southern Africa
Locations: 1,2,4, 16,20,21

Collection methods: sweeping and trapping on banana, other rotting fruit, fig tree,
and fungus

11. Drosophila burlai Tsacas & Lachaise, 1974. Ann. Univ. Abidjan, ser. E (Ecologie)

7:200

Specimens examined: one male and living cultures
Distribution: Central Africa
Locations: 1, 23
Collection methods: sweeping fungus and rotting banana

dentissima species subgroup Bock and Wheeler

12. Drosophila anisoctena Tsacas, 1980. Ann. Soc. Ent. Fr. (N.S.) 16: 527
Specimens examined: one male and living stocks

Distribution: Kenya and Zaire

Location: 30

Collection methods: sweeping rotting banana

13. Drosophila bahunde Tsacas, 1980. Ann. Soc. Ent. Fr. (N.S.) 16: 529
Specimens examined: five males and a living culture
Distribution: Kenya and Zaire

Location: 1

Collection methods: sweeping on fungus

ananassae species subgroup Hsu

14. Drosophila ananassae Doleschall, 1858. Natuurk. Tijds. Nederl. Indie 17: 128
Specimens examined: 32 males and eight females

Distribution: Circumtropical region
Locations: 1,3,4, 10, 11, 13, 14,26,31

Collection methods: sweeping on rotting fruits and on fresh fruits, including mango,
banana, and pineapple

15. Drosophila malerkotliana Parshad & Paika, 1964. Res. Bull. (N.S.) Panjab Univ.

15: 225

Specimens examined: five males and three females
Distribution: widespread in Oriental African regions, Brazil
Locations: 4, 11, 16
Collection methods: sweeping on rotting fruit and trapping on banana

16. Drosophila fima Burla, 1954. Rev. Suisse Zool. 61: 165
Specimens examined: one male

Distribution: Central Africa

Location: 21

Collection methods: sweeping on fig tree

17. Drosophila dimitra Tsacas & Lachaise, 1981. Ann. Soc. Ent. Fr. (N.S.) 17(3): 410
Specimens examined: one male

Distribution: Kenya, Cameroun, Burundi, Zaire

Location: 21

Collection methods: sweeping under fig tree

Vol. 101, No. 4. September & October 1990 249

Subgenus Scaptodrosophila Duda

18. Drosophila latifasciaeformis Duda, 1940. Ann. Hist-nat. Mus. Nat. Hung. 33: 2
Specimens examined: 103 males and 30 females

Distribution: widespread in Africa, U.S.A., Mexico, West Indies, South America
Locations: 1, 4-7, 11, 13, 19, 20, 23, 25, 27-29, 34

Collection methods: sweeping on rotting fruit and on mango, banana, avacodo.
and tomato

19. Drosophila dibi Burla, 1954. Rev. Suisse Zool. 61: 126
Specimens examined: four males and one female
Distribution: Kenya, Ivory Coast

Locations: 5, 10

Collection methods: sweeping on banana and other fruit and trapping on banana

20. Drosophila species 1.

Specimens examined: one male and one female

Locations: 2, 8

Collection methods: sweeping on banana and other fruit and trapping on banana

Subgenus Drosophila Fallen

21. Drosophila mercatorum Patterson & Wheeler. 1942. Univ. Texas Publ. 4213: 93
Specimens examined: 12 males and three females

Distribution: Kenya (New Record), U.S.A. South America, Europe

Location: 26

Collection methods: sweeping on rotting citrus

22. Drosophila repleta Woollaston, 1858. Ann. Mag. Nat. Hist. Ser. 3, 1: 117
Specimens examined: eight males and two females

Distribution: cosmopolitan
Locations: 1. 14.22,23,31

Collection methods: sweeping fruits, including banana, mango, pineapple, and straw-
berry jam

23. Drosophila hydei Sturtevant, 1921. Carnegie Inst. Publ. 301: 101
Specimens examined: 14 males and 13 females
Distribution: cosmopolitan

Locations: 1, 19, 20, 25, 32

Collection methods: sweeping fungus and rotting fruit

24. Drosophila fulvimacula Patterson & Mainland, 1944. Univ. Texas publ. 4445: 42
Specimens examined: live cultures

Distribution: Kenya (New Record; First Old World Record), Mexico to Brazil

Locations: 7, 25

Collection methods: sweeping rotting fruit

25. Drosophila nasuta Lamb, 1914. Trans. Linn. Soc. Lond. Ser. 2 (Zool.). 16: 346
Specimens examined: one female

Distribution: Central Africa. Seychelles. Madagascar. Mauritius

Location: 1

Collection methods: sweeping on fungus

26. Drosophila virilis Sturtevant, 1916. Ann. Ent. Soc. Am. 9: 330
Specimens examined: live cultures

Distribution: Kenya (New Record), widespread in southern Nearctic, Neotropical, and

northern Oriental regions
Location: 16
Collection methods: trapping on banana

250 ENTOMOLOGICAL NEWS

27. Drosophila immigrans Sturtevant, 1921. Carnegie Inst. Publ. 301: 83
Specimens examined: five males and three females
Distribution: cosmopolitan

Locations: 1, 19, 25

Collection methods: sweeping on rotting fruit and on fungus and on fruit, including

banana and pineapple

28. Drosophila nutrita Duda, 1935. Stylops 4: 25
Specimens examined: one male

Distribution: Kenya (New Record), Abidjan, Ivory Coast, Cameroon. Zimbabwe

Location: 19

Collection methods: sweeping on rotting fruit

29. Drosophila adamsi Wheeler, 1959. Univ. Texas Publ. 5914: 183
Specimens examined: two males

Distribution: widespread in Africa

Location: 2

Collection methods: sweeping on fungus

30. Drosophila funebris (Fabricius), 1787. Mantissa Insectorum 2: 345
Specimens examined: one male

Distribution: cosmopolitan

Location: 32

Collection methods: sweeping on rotting banana

31. Drosophila pruinosa Duda, 1940. Ann. Hist-nat. Mus. Nat. Hung. 33: 41
Specimens examined: six males and two females

Distribution: Central Africa

Locations: 23, 30

Collection methods: sweeping on rotting banana

Subgenus Dorsilopa Sturtevant

32. Drosophila busckii Coquillet, 1901. Ent. News 12: 18
Specimens examined: two males and one female
Distribution: cosmopolitan

Locations: 7, 23

Collection methods: sweeping on rotting fruit

Genus Lissocephala Malloch

33. Lissocephala species 1.
Specimen examined: one female
Location: 8

Collection methods: sweeping on rotting fruit

Genus Chymomyza Czerny

34. Chymomyza species 1.
Specimens examined: two males
Location: 4

Collection methods: sweeping on rotting fruit

Genus Scaptomyza Hardy

35. Scaptomyza species 1.
Specimen examined: one male

Vol. 101, No. 4. September & October 1990 251

Location: 33

Collection methods: sweeping on a lily

Genus Dettopsomyia Lamb

36. Dettopsomyia woodruff! Takada, n.sp.
Type area: Kiboko region. Kenya
Locations: 4, 5

Collection methods: collected by R.C. Woodruff by sweeping on banana and rot-
ting fruit

Diagnosis: Eye much oblique to body axis; ocellars inserted just outside triangle made by
ocelli; anterior reclinate orbital about 1/3 as long as proclinate; cheek about 1/3 as broad as
greatest diameter of eye; costal lappet large; costal-index 1.0; R2 + 3 strongly curved to
costa apically; R4 + 5 and M parallel; acrostichal bristle absent; tibia ringed: wing not spot-
ted; acrostichal hairs in four rows; C3-fringe more than 1/2.

Description: Male and female (Figure 2). Body about 1.5mm: wings about 1.5mm in
alcohol specimens. Eye much oblique to body axis. Arista with four upper and two lower
branches plus terminal fork. Oscellar triangle dark brown. Ocellars inserted just outside
triangle. Anterior reclinate about 1/3 as long as proclinate. situated slightly before pro-
clinate. Vibrissa strong; other orals fine. Cheeks about 1/3 as broad as greatest diameter of
eye. yellowish brown. Frons flat, yellowish brown, orbits yellowish brown: a pair of dark
brownish stripes between ocelli and orbits. Palpi dark brown with two stout apical setae.

Mesonotum dark brown, with two pairs of greyish longitudinal stripes. Acrositichal
hairs in four rows; dorsocentrals in two pairs; anterior scutellars divergent; posterior
scutellars convergent, upright. Halteres yellowish brown. Sterno-index about 0.5.

Abdomen dark brown, slightly shining. Legs yellowish brown; all tibiae with two dark
brownish rings.

Wings hyaline, distal costal incision deep, lappet black and large. R2 + 3 strongly cur-
ved to costa; R4 + 5 and M parallel; C-l bristles two, long. Costal-index about 1.0; 4V-index
about 3.0; 4C-index about 2.3; 5x-index about 2.3; C3-fringe on basal 3/4.

Periphallic and phallic organs are shown (Fig. 2d. e) and the female egg-guide (Fig. 2g)
is compared to egg-guides of some other Kenyan drosophilids (Fig. 3).
Type material: Holotype male. Kiboko by sweeping in a garden of bananas and other fruit.
R.C. Woodruff, May 17, 1988. Paratypes: one male and one female from same locality, and
one female from Makindo by sweeping rotting fruit R.C. Woodruff. May 17. 1988. The
type-series of the new species is deposited in the Biological Laboratory, Sapporo Univer-
sity, Japan.

Subfamily Steganinae

Genus Leucophenga Mik

Subgenus Leucophenga Mik

37. Leucophenga apicifera (Adams), 1905. Kansas Univ. Sci. Bull. 3: 185
Specimen examined: one female

Distribution: Central to south Africa

Location: 22

Collection methods: sweeping on banana and pineapple

38. Leucophenga disjuncta Bachli, 1971. Exploration Pare. Nat. de I' Upemba. Fasc.

71:32

Specimen examined: one male
Distribution: Central Africa
Location: 21
Collection methods: sweeping on fig tree

252

ENTOMOLOGICAL NEWS

KENYA

0 40 80 120 160

Fig. 1. Collection locations: 1, Nairobi; 2, Nairobi National Park; 3, Hunter's Lodge,
highway A109 near Kiboko; 4, Makindu, on A109 west of B7 junction; 5, National Range
Research Station Headquarters, Kiboko; 6, Lokichar, on Al west of Lake Turkana; 7,
Makutano, on Al north of Kitale; 8, Kitale; 9, Eldoret; 10, Jadini Beach Hotel, on Diani
Beach south of Mombassa; 1 1, Kwale, southwest of Mombassa; 12, Shimoni, on coast just
north of Tanzanian border; 13, Ukunda, on highway A14 south of Mombassa; 14, Namanga,
on A104 on Tanzanian border; 15, Amboseli Serena Lodge, Amboseli National Park; 16,
Ngulia Lodge, West Tsavo National Park; 17, Mt. Kenya; 18, Naro Moru River Lodge, on
A2 west of Mt. Kenya; 19, Karatina, on A2 southwest of Mt. Kenya; 20, Sagana, on A2
southwest of Mt. Kenya; 21, Mayer's Farm, on B3 between Narok and Nairobi; 22, Mara
River Camp, Masai Mara National Reserve; 23, Little Governor's Camp, Masai Mara
National Reserve; 24, Narok; 25, Naivasha; 26, Isiolo; 27, Meru; 28, Kakunga, on Al bet-
ween Kitale and Kisumu; 29, Kakamega, on Al north of Kisumu; 30, Kakamega National
Forest; 31, Kisumu; 32, North Kinagop, near Naivasha on edge of Aberdare Range; 33, Mt.
Aberdares, southeast edge of Aberdare Range; 34, Island Camp, Lake Baringo.

Vol. 101, No. 4, September & October 1990

253

Fig. 2. Dettopsomyia woodruffi Takada, n.sp. (a) dorsal view of head, (b) vental or anterior
view of head, (c) lateral view of head and first leg, (d) periphallic organs, (e) ventrolateral
aspect of phallic organs, (0 spermatheca, (g) egg-guide of female.

254

ENTOMOLOGICAL NEWS

Fig. 3. Egg-guides of females of some Kenyan Drosophilidae. (a) Drosophila melanogaster
(b) D. yakuba, (c) D. nikananu, (d) D. malerkotliana, (e) D. simulans, (f) D. seguyi, (g) Zap-
rionus tuberculatus. (h) Drosophila (Scaptodrosophila) latifasciaeformis

ACKNOWLEDGMENTS

We are grateful to Cluff Hopla and Jenna Hellack for comments on the manuscript, to
Janice Frankart, Phyllis Oster, Donna Tampurages, Helena Palka, and Laura S. Hier for
their excellent technical assistance, and to Coral McCallister for the Kenya map illustra-
tion. This material is based upon work supported by a Fulbright Research Award (87-
47122) and a Bowling Green State University Faculty Research Committee Bsic Grant to
R.C.W., a University of Oklahoma Biomedical Research Support Grant and a Faculty
Enhancement travel grant to J.N.T., and a Research Grant of the Board of Sapporo Univer-
sity to H.T.

Vol. 101. No. 4. September & October 1990 255

LITERATURE CITED

Tsacas, L, D. Lachaise, and J.R. David. 1981. Composition and biogeography of the
Afrotropical drosophilid fauna. In: The Genetics and Biology of Drosophila, volume 3a.
(M. Ashburner. H.L. Carson, and J.N. Thompson, jr.. eds.), pp. 197-259. Academic
Press, London.

Wheeler, M.R. 1981. The Drosophilidae: A taxonomic overview. In: The Genetics and Biol-
ogy of Drosophila. volume 3a. (M. Ashburner, H.L. Carson, and J.N. Thompson, jr., eds.),
pp. 1-97. Academic Press. London.

Wheeler, M.R. 1986. Additions to the catalog of the world's drosophilidae. In: The Genetics
and Biology of Drosophila. volume 3e. (M. Ashburner. H.L. Carson, and J.N. Thompson,
jr., eds.), pp. 395-409. Academic Press, London.

Woodruff, R.C., B.E., Slatko, and J.N. Thompson, jr. 1983. Factors affecting mutation
rates in natural populations. In: The Genetics and Biology of Drosophila. volume 3e. (M.
Ashburner, H.L. Carson, and J.N. Thompson, jr., eds.), pp. 37-124. Academic Press.
London.

Woodruff, R.C., J.N. Thompson, jr., H.N.B. Gopalan, W.A. Ngure, E.S. Norris, and
A.A. Szekely. 1989. Genetic and molecular analyses of P DN A elements sampled from
natural populations of Drosophila melanogaster in Kenya. In: Molecular Evolution,
UCLA Symposia on Molecular and Cellular Biology, New Series, volume 122. (M.
Clegg and S. O'Brien, eds.). 1 17-125. Alan R. Liss, New York.

256 ENTOMOLOGICAL NEWS

ERRATUM OF OMISSION

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-H.P.B.

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ENTOMOLOGICAL NEW

Drinking as a pre-oviposition behavior of wild Culex

pipiens (Diptera: Culicidae) R.G. Weber, C. Tipping 257

Distribution of ponderosa pine feeding sawflies
(Hymenoptera: Diprionidae) in the U.S.
and Canada C.S. Dunbar, M.R. Wagner 266

Neotype designation for Lebia mono, fixing the name as a
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Rearing and development of Phyllocoptes fructiphilus

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Observations pertinent to the role of sexual selection in
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The leafhopper genus Bythonia (Homoptera: Cicadellidae)

H.D. Blocker, M.D. Webb 293

New records of mayflies (Ephemeroptera) from Maine

S.K. Burian, R.G. Mack 297

Separation of insect and plant material from screen-sweep

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Vol. 101. No. 5. November & December. 1990 257

DRINKING AS A PRE-OVIPOSITION BEHAVIOR
OF WILD CULEXPIPIENS (DIPTERA: CULICIDAE)1

R.G. Weber, Chris Tipping2

ABSTRACT: In the field, C. pipiens females land on open water where they stand on their
pro- and mesothoracic legs. Females do not hold onto anything prior to. or during,
oviposition, nor do they cross their hind legs for oviposition as reported previously.
Lowering of the metathoracic legs and paralleling them on the surface behind the
abdomen for oviposition occurs only after a female drinks from the ovisite. Drinking from
the water upon which they will oviposit is a regular behavior (97.3%). Most individuals
(91.9%) drink more than once, drink for more than ten seconds total (8 1.1%). and spend an
average of five seconds or more per drink (67.7%). Total time spent drinking averages 65.7
seconds. Because drinking occurs before oviposition is begun, contact chemoreceptors on
the mouthparts could be involved in the final stages of ovisite acceptance. Time spent in
drinking exceeds that necessary for mere tasting, and drinking may also serve to distend
the abdomen and assist in movement of eggs through the oviduct.

The literature of mosquito oviposition behavior begins with
Reaumur's illustrated account of egg-laying by a species of mosquito
which he did not identify (Reaumur 1738). The mosquitoes whose egg
raft construction he observed and described are believed to have belonged
to the species Culex pipiens L. (Howard, etal. 1912, Mattingly 1970). Since
Re'aumur's report, oviposition behavior of this species and its subspecies
has been examined from several perspectives^. These include behavior
at oviposition sites (ovisites) prior to landing on the water (de Meillon
and Sebastian 1965, Kennedy 1942, Mattingly 1965), behavior after landing
(Hudson 1956, Ikeshoji 1966a), egg raft construction (Beament and
Corbet 1 98 1 , Wallis 1 954), and diel distribution of oviposition activity (de
Meillon, et al. 1967, Lowe, et al. 1973,MacDonald,e*a/. 1981,Oda 1967,
Oda and Kuhlow 1979, Subra 1971, Suleman and Shirin 1981). No one
has reported the entire sequence of behaviors exhibited by female Culex
spp. from time of landing on the water to initiation of oviposition.

Physiological aspects of the female mosquito's evaluation of ovisite
chemistry also have been examined. Much of this research has dealt with
effects of various chemicals, or bacteria, on ovisite selection (e.g. Gjullin
and Johnsen 1965, Ikeshoji 1965, Kau\, etal. 1977, Murphey and Burbutis

1 Received June 12, 1990. Accepted July 7, 1990.

2Delaware Agricultural Experiment Station, Department of Entomology and Applied
Ecology, College of Agricultural Sciences, University of Delaware, Newark. DE 19717.

Culex pipiens pipiens Linnaeus, C. p. fatigans Wiedemann, C. p. molestus Forskal. C. p.
quinquefasciatus Say, and C. p. pollens Coquillett. For representative discussions of these
taxa and possible synonymies, see Barr (1960). Mattingly. et al. (1951) and Rozeboom
(1951).

ENT. NEWS 101(5): 257-265, November & December. 1990

258 ENTOMOLOGICAL NEWS

1967, Rockett 1987). Because chemicals influence ovisite selection, there
has been interest in determining which anatomical parts contain
sensors that transduce this information. Several mosquito species,
including C. pipiens, periodically appear to touch the substrate with their
mouthparts prior to ovipositing. Thus sensors associated with the mouth-
parts could be involved in ovisite selection. This behavior, referred to as
"drinking", has been reported for several species (Detinova 1936, Hudson
1956, Ikeshoji 1966b, Pappas and Pappas 1982), but it has been con-
sidered doubtful that drinking is a regular part of ovisite selection behavior
(Hudson 1956, Ikeshoji 1966b, Kalandadze and Sagatelova 1939,
Kennedy 1942). Only Culiseta inomata (Williston) is currently known
always to contact the water surface with its proboscis, and apparently
drink, prior to oviposition (Pappas and Pappas 1982).

We report here the postural changes and behaviors observed as
unmanipulated, wild Culex pipiens females which had just landed made
their final acceptance of an ovisite and began construction of their egg
raft. We further report that drinking from the water upon which they will
oviposit is a regular behavior for females of this species and is precursory
to oviposition.

MATERIALS AND METHODS

Females of several Culex species, including C. pipiens L., oviposit at
night. There is a major peak of oviposition shortly after sunset and a
smaller peak before sunrise (de Meillon, et al 1967, MacDonald, et al.
1981, Oda 1967, Subra 1971), though this may vary with latitude and
season (Oda and Kuhlow 1979) or experimental design (Suleman and
Shirin 1981). We made our observations during the post-sunset activity
period at containers of wheat straw infusion (Weber and Weber 1985),
which is very attractive to gravid, wild Culex spp. mosquitoes. We con-
ducted this research during the summer of 1988, on the University of
Delaware Research Farm in Newark, Delaware.

Our nightly procedure was to scan our containers until a mosquito was
observed hovering above the surface in the way characteristic of gravid
Culex spp. females (de Meillon and Sebastian 1965, Kennedy 1942,
Reiter 1983). We collected data only for females which we saw as they
landed, and which we then observed continuously up to placement of
about the tenth egg in their raft. A continuous spoken record of date,
starting time, and behavior from time of beginning observation to collection
of the individual was recorded on a tape recorder. When approximately
10 eggs had been placed in the raft, we collected the individual with a
mouth-operated aspirator, placed it in a labelled vial, and froze it for

Vol. 101, No. 5, November & December. 1990 259

later identification. For identification we used the key in Darsie and
Ward (1981). Data from tapes were played back later and transcribed to
printed forms. During playback and transcription, a stop watch was used
to determine times spent in the various phases of ovisite acceptance.

After landing on the water, and before beginning to oviposit, Culex
spp. females are easily disturbed by a variety of things, especially by
white (full visible spectrum) light. However, after egg raft construction is
underway (15-20 eggs laid), females can be observed, even using white
light, with much less danger of interrupting their behavior^ (Mattingly
1970). Because our interest was in behavior during the easily-disturbed
period from landing to initiation of a raft, we made all observations
using flashlights covered with red, translucent plastic (Sandholm and
Price 1962, Wallis 1954). Red light is less disturbing than white to
mosquitoes that have not yet begun to oviposit^.

We made our observations at an eye-to-mosquito distance of 1 5.3 cm-
20.3 cm(6"-8"), and a light-to-mosquito distance of 30.5 cm-35.5 cm (12"-
14"). By holding the flashlight approximately parallel to the surface and
90° to one side of the eye-mosquito axis, we could readily observe changes
in leg or proboscis attitudes, and see clearly the dimpling caused when-
ever the proboscis made contact with the water.

Early in the study it became apparent that proboscis contact with the
water was a characteristic behavior. To more closely examine this behavior
we caged eight wild, gravid females (obtained at ovisites) individually
over oversite water colored with a flourescing, red dye". These indi-
viduals were observed at frequent intervals and were collected after they
had begun rafts.

RESULTS AND DISCUSSION

We observed 37 ovipositing, wild C pipiens L. females. All were collected
and identified. Both C. pipiens L. and C. quinquefasciatus Say separate out
at the same location in the key we used for identification (Darsie and
Ward 1981). We consider the individuals whose behavior we report here
to belong to C. pipiens rather than C. quinquefasciatus because C. pipiens is
the more northerly species, occurring north of 39° N (Barr 1957). This

Beament and Corbet (1981) were able to transport individual ovipositing C. pipiens

females, picked up in clear plastic dishes of water, from an outdoor container into their

laboratory for examination under a microscope.
-'Adult insects of a variety of species cannot perceive wavelengths above ca. 600-650 nm

("red") (Burkhardt 1964) and larvae of C. pipiens are not attracted to red light (Weber

1989).
^'Intracid Rhodamine WT Liquid", (product #4517100). Manufactued by Crompton &

Knowles Corporation, P.O. 341, Reading, PA 19603.

260 ENTOMOLOGICAL NEWS

study was conducted at 39° 41' N (U.S.G.P.O. 1984). Further, all 90 male
Culex sp. from Newark, Delaware, examined by Barr (1957) during his
study of C. pipiens and C. quinquefasciatus distribution were found to be
C. pipiens.

All individuals we observed landed on open water, where they stood
and eventually oviposited. One individual walked ca 5 mm from where it
landed before beginning to oviposit; the others oviposited where they
landed. In contrast with Reaumur's (1738) report, no females held onto
anything during oviposition, although particles of straw and other veg-
etable detritus were usually present on the water.

Females landed and stood on their pro- and mesothoracic legs. The
metathoracic legs were held up, with their tarsi curving upward above
the abdomen, as has been reported for Culex molestus by Hudson (1956).
During this time the abdomen was parallel to the surface. Usually the
proboscis was directed approximately 45 degrees downward, but in 15
instances (40.5%) was more nearly horizontal. Some time after landing
(which we were unable to measure accurately), the proboscis was lowered
to put its tip in contact with the surface. When contact was made, the
meniscus was plainly visible, which allowed us to readily determine
duration of contact, or "drinking". After drinking one or more times of
variable duration, females lowered their metathoracic legs to the surface
and brought them close together behind the tip of the abdomen.
Metatarsi were brought approximately parallel; we saw no instances of
crossed hind tarsi as described and illustrated by Reaumur ( 1 738). Soon
after lowering the legs and bringing the tarsi back, the tip of the abdomen
was lowered nearly to the surface, between the tarsi. Shortly after this, the
first egg appeared and was placed, on its end, on the water between the
hind tarsi. Data we present about drinking, below, was collected prior to
appearance of the first egg.

All but one of the 37 C. pipiens females we observed (97.3%) drank from
the ovisite before they began to oviposit. Females drank at least once
before beginning to lay eggs, and one drank 1 1 times (mean = 4.4 times).
The one individual which did not drink landed on the surface and began
to lay almost immediately. Because this happened at 108 minutes post-
sunset, we believe that this individual may have gone through the pre-
oviposition drinking phase earlier, had been disturbed, and merely
resumed ovipositing when it relanded. Seventeen other individuals
landed at post-sunset times of 1 1 6 to 1 63 minutes, but these drank from 2
to 10 times before beginning to oviposit.

Of the 37 mosquitoes, 9 1 .9% drank more than once (Fig. 1 ), 8 1 . 1% spent
more than ten seconds drinking (Fig. 2), and 67.6% spent an average of
five seconds or more per drink (Fig. 3). Total time individuals spent

Vol. 101. No. 5. November & December, 1990

261

0 34567891011

Figure 1. Number of times gravid, wild Culex pipiena L. females drank before they began to
oviposit.

5 10 15 20 25 30 35 40 45 >45
SECONDS PER DRINK

Figure 2. Total seconds spent in drinking by gravid, wild Culex pipiens L. females before
they began to oviposit.

262

ENTOMOLOGICAL NEWS

drinking ranged from 3 to 343 seconds (ave. = 65.7 seconds), but only
21.6% of our sample spent more than 80 seconds drinking (Fig. 2).
Average time/drink ranged from 1.3 to 76.0 seconds (Fig. 3). Only 13.5%
of the sample had an average time/drink greater than 30 seconds.
Drinking from the ovisite prior to oviposition thus appears to be a
normal, possibly necessary, component of C. pipiens preoviposition
behavior, at least for the population in our study area.

>
Q

1 20 40 60 80 100 120 140 160

10 30 50 70 90 110 130 150 >160

SECONDS SPENT DRINKING

Figure 3. Average time per drink, in seconds of gravid, wild Culex pipiens L. females.

All eight C. pipiens females caged over red water and collected after
they began to oviposit (but before raft completion), had red liquid in
their abdomens which fluoresced under UV illumination. The quantity
of red liquid in each individual varied, but each contained enough that
fluorescence was plainly visible through the intact ventral abdomen.
Dissections showed the liquid to be within the gut tract, but we did not
attempt to determine whether it was in the ventral diverticulum or the
ventriculus (Snodgrass 1959).

We have not yet examined the role of mouthpart chemoreception in
ovisite acceptance. Because drinking occurs before oviposition is begun,
the possibiliy exists that labral, labellar, or cibarial contact chemo-
receptors (Mclver 1982) could be involved in the final stages of ovisite
acceptance. Hudson (1956) doubted that drinking is a necessary pre-
oviposition behavior of Culex molestus. However, Ikeshoji ( 1966a) found
"when the proboscis was extirpated and taking up the water was prevented

Vol. 101. No. 5, November & December. 1990 263

there was hardly any oviposition" by Culexfatigans. Even if mouthpart
chemosensors are involved in final ovisite acceptance, the drinking we
observed could have additional importance to the gravid mosquito.
Most individuals drank more than once, drank for appreciable time, and
females on dyed water imbibed considerable amounts of ovisite water,
which suggests that if drinking is used to test ovisite suitability, this is not
its only function. Following their study ofCuliseta inornata oviposition
behavior, Pappas and Pappas (1982) suggested that "drinking before egg
laying may build up pressure in the abdomen and help release eggs."
Considering the constancy and duration of drinking we observed, this
also may be true for C. pipiens.

ACKNOWLEDGMENTS

We thank Robert Lake for assistance in identifications. Dana Marshall and David
Carter gave valuable aid in spotting incoming females and in data collection. This research
was supported in part by Hatch Funds. This is Miscellaneous Paper no. 1316 of the
Delaware Agricultural Experiment Station, Contribution no. 613 of the Department of
Entomology and Applied Ecology, University of Delaware, Newark. DE.

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America. Am. J. Trop. Med. Hyg. 6:153-165.
Barr, A.R. 1960. A review of recent findings on the systematic status of Culex pipiens. Calif.

Vector Views 7: 17-21.
Beament, J. and S.A. Corbet. 1981. Surface properties of Culex pipiens pipiens eggs and the

behaviour of the female during egg-raft assembly. Physiol. Entomol. 6:135-148.
Burkhardt, D. 1964. Color discrimination in insects. In.J.E. Beament, J.E.Treherne and V.

Wigglesworth (eds.): Advances in insect physiology. Vol. 2., Academic Press. London-
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feeding and time of oviposition. Bull. Wld. Hlth. Org. 36:39-46.
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maculipennis messeae Fall. [In Russian). Med. Parasit. 3:525-543. (Not seen: abstr. in

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in mosquito breeding waters of a factor which stimulates oviposition. Japan. J. Exp.

Med. 36:67-72.

264 ENTOMOLOGICAL NEWS

Ikeshoji, T. 1966b. Studies on mosquito attractants and stimulants. Part I. Chemical

factors determining the choice of oviposition site by Culex pipiens fatigans andpallens.

Japan. J. Exp. Med. 36:49-59.
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field waters. Wld. Hlth. Org. Vector Control 130:1-9.
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maculipennis maculipennis |In Russian). Med. parasit. 8:50-55. (Not seen; abstr. in Rev.

Appl. Entomol. (B) 28:184).
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fatigans breeding waters in areas around Delhi. J. Comm. Dis. 9:8-21.
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Entomol. Res. 32:279-301.
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Culex pipiens quinquefasciatus released into a natural population. Mosq. News 33:221-

227.
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oviposition by Culex pipiens and Culex restuans in southern Ontario. Proc. Entomol.

Soc. Ont. 112:39-40.
Mattingly, P.P. 1965. Intercurrent resting, a neglected aspect of mosquito behavior. Proc.

1st Internal. Congr. Parasit., Rome 2:910-91 1.
Mattingly, P.P. 1970. Mosquito Eggs VI. Genera Eretmapodites and Culex. Mosq. Syst.

Newsletter 2: 17-21.
Mattingly, P.P., L.E. Rozeboom, H.L.K.L. Knight, F.H. Drummond, S.R.

Christophers and P.G. Shute. 1951. The Culex pipiens complex. Trans. R. Entomol.

Soc. Lond 102:331-342.
Mclver, S.B. 1982. Sensilla of mosquitoes (Diptera: Culicidae). J. Med. Entomol. 19:489-

535.
Murphey, FJ. and P.P. Burbutis. 1967. Straw infusion attractiveness to gravid female

Culex salinarius. J. Econ. Entomol. 60:156-161.
Oda,T. 1967. Hourly and seasonal distribution of the number of eggs rafts of Culex pipiens

pollens deposited in earthen jars. Trop. Med. 9:39-44.
Oda, T. and F. Khulow. 1979. Tages- und jahreszeitliche Veranderungen der

Eiablageaktivitat von Culex pipiens pipiens L. Tropenmed. Parasit. 30:443-445.
Pappas, C.D. and L.G. Pappas. 1982. Observations on the egg raft formation behavior of

Culiseta inornata. Ann. Entomol. Soc. Am. 75:393-394.
Reaumur, R.A. d. 1738. Memoirs pour servir a 1'histoire des insects, Vol IV, Mem XIII.

L'Imprimerie Royale, Paris.
Reiter, P. 1983. A portable, battery-powered trap for collecting gravid Culex mosquitoes.

Mosq. News. 43:496-498.
Rockett, C.L. 1987. Bacteria as ovipositional attractants for Culex pipiens (Diptera:

Culicidae). Great Lakes Entomol. 20:151-155.
Rozeboom, L. E. 1951. The Culex pipiens complex in North America. Trans. R. Entomol.

Soc. Lond. 102:343-353.
Sandholm, H.A. and R.D. Price. 1962. Field observations on the nectar feeding habits of

some Minnesota mosquitoes. Mosq. news 22:346-349.
Snodgrass, R.E. 1959. The anatomical life of the mosquito. Smithsonian Misc. Coll. 139,

No. 8.
Subra, R. 1971. Etudes ecologiques sur Culex pipiens fatigans Weidemann, 1828(Deiptera.

culicidae) dans une zone urbaine de savane soudaniene ouest-africaine. Rythme de

ponte et facteurs conditionnant 1'oviposition. Cah. O.R.S.T.O.M.. Ser. Entomol. Med.

Parasitol. 9:317-322.

Vol. 101, No. 5, November & December, 1990 265

Suleman, M. and M. Shirin. 1981. Laboratory studies on oviposition behavior of Culex

quinquefasciatus Say (Diptera: Culicidae): Choice of oviposition medium and

oviposition cycle. Bull. Entomol. Res. 71:361-369.
U.S.G.P.0. 1984. The national gazetteer of the United States of America: Delaware, 1983.

Geological Survey Professional Paper 1200-DE. Washington, DC: U.S. Government

Printing Office, 1984. 101 pp.

Wallis, R.C 1954. A study of oviposition activity of mosquitoes. Am. J. Hyg. 60:135-168.
Weber, R.G. 1989. Response of larval Culexpipiens (Diptera: Culicidae) to light produced

by light emitting diodes. Entomol. News 100:104-110.
Weber, R.M. and R.G. Weber. 1985. The egg raft seam as an indicator of species in Culex

pipiens and Culex restuans. Mosq. Syst. 17:373-370.

BOOK REVIEW

THE BIOLOGY OF SCORPIONS. Gary A. Polis, ed. 1990. Stanford
University Press. $85.00

This long-awaited book greets the reader with an eerie photograph on its dust jacket of a
scorpion brightly fluorescing under ultraviolet light. Fluorescence of the cuticle is but one
of many fascinating and sometimes enigmatic featues of this group of animals. Scorpions
are all viviparous; their young are nourished to a greater or lesser extent in the uterus of the
mother, and are born alive. They then climb aboard their parent's back for a further period
of protection, and in a few species even older young may share the parental burrow. In one
species this has developed into a rudimentary society, a dozen or more scorpions of both
sexes sharing a common home deep within a termite mound. Scorpions may be found in
habitats ranging from the wrack of the shore to elevations of 16,000 ft. or more, but are most
diverse in deserts. Accordingly, they are capable of better water conservation than any
other arthropod that has been examined. Sensory hairs on the legs can detect Rayleigh and
compressional waves in sand, so that scorpions can locate even burrowing prey with great
accuracy.

Deservedly called living fossils, scorpions assumed their present body form at least 450
million years ago, and probably even earlier. Predominantly a marine group through
much of their history, the single surviving clade of scorpions is now entirely terrestrial;
textbook statements to the contrary, scorpions were latecomers (Late Devonian?) to land.
Scorpion fossils are rare but often include unprecedented detail because an unusual layer
in scorpion cuticle, the hyaline exocuticle, survives fossilization and persists, evidently
little changed, for tens of millions of years.

Only about 20 species of the more than 1 500 known ones are dangerous to man, but those
few take a toll: they cause more human deaths each year than any other non-parasite.
Given all these reasons to be interested in scorpions, it is remarkable that a book like this
one has not appeared sooner.

Gary Polis edited the book, wrote the introduction, and also wrote or contributed to five
of the 1 1 chapters. Other contributors include Cloudsley-Thompson (mythology). Hadley
(environmental physiology). Hjelle (anatomy), McCormick (prey, predators, parasites).
Root (neurobiology). Simard and Watt (venoms), Sissom (systematics and phylogeny ). and
Warburg (biorhythms). The clarity of the writing is uniformly good and organization of
each chapter is tight, overcoming some of the major problems often experienced with
multiauthor books. Illustrations, with a few exceptions, are clearly and cleanly executed.

(Continued on page 272)

266 ENTOMOLOGICAL NEWS

DISTRIBUTION OF PONDEROSA PINE
(PINUS PONDEROSA) FEEDING SAWFLIES

(HYMENOPTERA: DIPRIONIDAE)
IN THE UNITED STATES AND CANADA1

Clarence S. Dunbar, Michael R. Wagner^

ABSTRACT: Six species of diprionid sawflies are known to feed on ponderosa pine, Pinus
ponderosa, in western United States. Collection records were compiled from 29 insect
collections throughout the United States and Canada. Based on these records, distri-
butions of the species on ponderosa pine are reported.

Six species of diprionid sawflies (Diprionidae), five in the genus
Neodiprion, feed on ponderosa pine, Pinus ponderosa Dougl. ex Laws., the
most widely distributed pine in North America (Powells 1965).
Diprionid sawflies are major economic pests of natural and plantation
forests in the eastern United States (Wilson 1977.) As forest management
intensifies in the West, sawflies are increasingly important economic
pests (Dahlsten 1961, 1966). This research was undertaken to determine
the distribution of diprionids feeding on ponderosa pine. Records have
never been compiled for these insects. Keys for identification of North
American genera are provided by Ross (1955) and Smith (1974).

METHODS

Twenty-nine federal and state research stations, museums and
universities provided 97 collection records for sawflies found feeding on
ponderosa pine. These localities were plotted on a map with the dis-
tribution of ponderosa pine (Fig. 1). The distribution of ponderosa pine
was taken from Powells, 1965. The distribution of each species of sawfly
was then determined by considering both the collection records and the
distribution of its host (Fig. 2)

RESULTS

Locality data for each species are as follows:

'Received March 16, 1990. Accepted July 10, 1990.

2School of Forestry, Box 4098, Northern Arizona University, Flagstaff, AZ 8601 1

ENT. NEWS 101(5): 266-272, November & December, 1990

Vol. 101. No. 5. November & December. 1990 267

Zadiprion townsendi (Cockerell)

ARIZONA: Flagstaff, 28 Jun 1916; Santa Rita Mts.. Jul 1924; North Rim. Grand Canyon. 7
Dec 1964; Mt. Lemmon. Sta. Catalina Mts.. 3 Aug 1967. COLORADO: Jefferson Co..
Evergreen. 26 Aug 1937; Teller Co.. Woodland Park. 2 Jun 1938; Boulder Co.. Boulder. 2
Mar 1963; Chambers Lake, Roosevelt N.F., 9200 ft.. 2 Aug 1968; Larimer Co.. Rist Canyon.
Mar 1979; El Paso Co., Black Forest. NEBRASKA: Dawes Co., Crawford 12-28 Jul 1910;
Dawes Co., Pine Ridge, Jul 1910; Sioux Co., War Bonnet Canyon; Sioux Co.. Hat Creek;
Lancaster Co., Lincoln 1 May 1914. NEW MEXICO: Bernalillo Co., Rio Arriba Co.,
Vallecitos, 30 June 1924; Socorro Co., Magdalena, 13 Jan 1961; Tijeras. 19 Dec 1969;
Bernalillo Co., Sandia Park. 19 Dec 1969; San Jaun Co., 8.7 mi. N. Navajo. 30 Jun 1972.
SOUTH DAKOTA: Custer Co.. Custer State Park, nr Custer, 1911. UTAH: San Juan Co..
Blanding. Chippean Ridge 2 Nov 1972.

Neodiprion autumnalis (Smith)

ARIZONA: Yavapai Co.. Camp Verde, 1 8 May 198 1 ; Coconino Co., Flagstaff, 26 May 198 1 ;
Apache Co.. Springerville. 13 Aug 1981. CALIFORNIA: Butte Co.. Jarbae Pass. 12 May
1949; Shasta Co., Mt. Shasta. 14 Aug 1957; Humbolt Co., Orleans, 8 May 1961 ; Trinity Co.,
Ruth, Oct 1976. COLORADO: Teller Co.. Woodland Park. 5 Oct 1914; El Paso Co.. Husted
12 Oct 1914. IDAHO: Kootenai Co., Coeurd'Alene, 18Sep 1922;IdahoCo.,Grangeville. 19
Jun 1971. MONTANA: Sanders Co.,Camas 28 Aug 1913; LakeCo..BitterrootN.F..Ravalli.
1 1 Oct 1959; Phillips Co.. Landusky. 24 Sept 1959. NEBRASKA: Cherry Co.. Valentine, 23
Jul 1971; Dawes Co., Nebraska N.F., Chadron, 20 Jun 1973. NEW MEXICO: Cibola Co.,
Grants, Jun 1957. OREGON: Allison R.S.. reared 1943; Klamath Co., Diamond Lake, 25
Sep 1963. SOUTH DAKOTA: Custer Co., Pringle, Sep 1935; Harding Co., Camp Crook, 1
Sep 1971; Perkins Co., Lemmon, 6 Jun 1972; Todd Co., Olsenville, 23 Jun 1972; Todd Co.,
Rosebud, 14 Jun 1973. WASHINGTON: Spokane Co., Spokane, 26 Sept 1960.

Neodiprion fulviceps (Cresson)

ARIZONA: Coconio Co., Flagstaff nr 1-40, 21 Jul 1982. NEVADA.

Neodiprion gillettei (Rohwer)

ARIZONA: Yavapai Co.. Prescott, 1 1 Jun 1928; Yavapai Co., Oak Creek Canyon. 7 Oct
1951; Chiricahua Mts.. 21 Aug 1961; Sitgreaves N.F.. Chevelon Rd.. 7 Jul 1969; Oak Creek,
vicinity of Sedona, 1 Oct 1970; Kaibab N.F., 22 Sep 1974; Apache Co., Springerville 27 May
1987. COLORADO: Grand Co., Granby, 9 Nov 1961; Larimer Co.. Rist Canyon 21 Jul
1986.

Neodiprion mundus (Rohwer)

UNITED STATES. IDAHO: Valley Co.. Cascade, 9 Jul 196 1 ; Idaho Co.. White Bird, 2 1 Oct
1961. OREGON: Corvallis. 13 May 1951; Benton Co.. Corvallis 3 Jun 1944.
CANADA. Kelowna. B.C.

Neodiprion ventral is (Ross)

ARIZONA: Coconino Co., Flagstaff, 30 Jun 1982. COLORADO: Phillips Co.. Holyoke;
Jefferson Co., Plainview, Oct 1935. NEBRASKA: Lancaster Co.. Lincoln. Oct 1977.

268

ENTOMOLOGICAL NEWS

L AUTUMNALIS * N. VENTRALIS
N. GILLETTE! X N. MUNDUS

+ N. FULVICEPS

200

Figure 1. Distribution of ponderosa pine feeding sawflies from collection records. The
distribution of ponderosa pine is indicated by stippled areas.

Vol. 101, No. 5, November & December, 1990

269

N. AUTUMNALIS — •

N. GILLETTEI
•• Z. TOWNSENDI

N. VENTRALIS
N. MUNDUS

N. FULVICEPS

Figure 2. Hypothesised distribution of ponderosa pine feeding sawflies extrapolated from
collection records and host distribution.

270 ENTOMOLOGICAL NEWS

NORTH DAKOTA: Slope Co., Burning Coal Vein Area, 1 8 Jul 1 973; Slope Co., 6 Jun 1 974.
WYOMING: Platte Co., Glendo, 25 Oct 1961.

TAXONOMIC NOTES

The Neodiprion fulviceps complex was defined by Ross (1955). The
complex was then separated into two species, N. autumnalis and N.
fulviceps (Smith and Wagner 1986). N. autumnalis overwinters as eggs,
whereas N. fulviceps overwinters as cocoons (Wagner et. al. 1986).

Many of the specimens previously identified as Neodiprion fulviceps
complex, are actually TV. autumnalis (Smith and Wagner 1986). Sawflies
that were incorrectly identified as Neodiprion edwardsii and Neodiprion
demoides by B.D. Burks, have been identified as N. autumnalis (Smith,
personal communication). The following records are from specimens
identified as N. fulviceps, but need to be reexamined for correct identity.
Many may be N. autumnalis especially those collected late in the season,
if the dates refer to adult emergence or collection records.

Neodiprion fulviceps (complex)

UNITED STATES. ARIZONA: Santa Rita Mts., 26 Sept 1925. CALIFORNIA: Siskiyou
Co., Dorris, 20 Jun 1912; Mendocino Co., Ft. Bragg, 8 May 1936; Mendocino Co., Pygmy
Forest, 4 mi. East Mendocino City, 2 1 May 1938; Plumas Co., West of Milford, 6000 ft.. 8 Jul
1942; Mendocino Co., Mendocino, 1 Oct 1957. COLORADO: Jefferson Co., Plainview , 1
Jul 1937; Larimer C., Ft. Collins, Jul 1964 as larvae, adults Sep 1964. MONTANA: Rosebud
Co.. Forsyth,22;Jul 1922; Rosebud Co., Lee, 24 Jul 1922 (as larvae), adults emerged Sep 13-
22 1922; Phillips Co., Little Rockies, Zortman, Sep 1960; Ravalli Co., Stevensville, 15 Jun
1973. NEBRASKA: Thomas Co., Halsey, 8 Oct 1936. NEVADA: Clark Co., Charleston
Mts., Kyle Canyon, 7500 ft. OREGON: Benton Co., Peoria, 21 Apr 1940. UTAH: Garfield
Co., Panguitch, Utah State University Farm. 17 Jul 1970; Millard Co., Kanosh, East Side of
Clear Creek. 5 Aug 1970; Cache Co.. Cove Fort, Clear Creek. 15 Oct 1970.
CANADA Cascade, B.C.; Falkland, B.C.; Fountain Creek, B.C.; Kelowna. B.C.; Okanagan
Mission, B.C.; Osoyoss, B.C.; Pritchard, B.C.; Rock Creek, B.C.; Winfield. B.C.

DISCUSSION

Neodiprion autumnalis is the most widely distributed species reaching
from the Southwest, north through the east side of the Rockies, across
Montana to the Northwest and south to northern California. Zadiprion
townsendi is found in the Southwest and as far north as the southwest
corner of South Dakota. N. gillettei is located in the Southwest and as far
north as northern Colorado. N. ventralis has been collected in northern
Arizona, northern Colorado, southeastern Wyoming, and the southwest
corner of North Dakota. N. fulviceps occurs in the Southwest and N.
mundus in the Northwest (Fig. 2).

Vol. 101, No. 5, November & December, 1990 271

Five of the six species of sawflies that feed on ponderosa pine occur
near Flagstaff, Arizona. We have observed some interesting features of
the food resource allocation for four of these sympatric species. N. gilletti
prefers small trees less than three feet tall or branches of older trees that
touch the ground. N. autumnalis and N. ventralis generally occur on
medium pole sized trees, while N.fulviceps typically feeds on older more
mature trees. This type of stratification along with differing life cycles
allows these sympatric sawflies to feed on the same species of pine
without competing.

VOUCHER SPECIMENS

Depositories for voucher specimens are as follows: National Museum of Natural
History, Washington, DC; Arizona State University, Tempe; Biosystematics Research
Centre, Ottawa, Ontario; Bureau of Plant Industry, Lincoln, NE; California Academy of
Sciences, San Francisco; California Department of Food and Agriculture, Sacramento;
Colorado State University, Ft. Collins; Forest Service, Albuquerque, NM; Forest Service.
Berkeley, CA; Forest Service, Ft. Collins, CO; Forest Service, Missoula, MT; Forest Service,
Ogden, UT; Forest Service, Portland, OR; Forest Service, Washington, DC; Forestry and
Range Sciences Laboratory, LaGrande, OR; Forestry Sciences Laboratory, Lincoln, NE;
Montana State University , Bozeman, MT; Natural History Survey, Champaign.IL;
Oregon Department of Agriculture. Salem; Oregon State University. Corvallis; Pacific
Forestry Centre, Victoria, BC; University of Arizona;, Tucson; University of California.
Berkeley; University of Nebraska, Lincoln; University of Wyoming. Larmie.

ACKNOWLEDGMENTS

We are grateful to the following people who contributed collection records: John
Anhold, Donald L. Dahlsten, Clarence J. DeMars.Jr.JeraldE. Dewey, James A. DiGiulio.
Mary Ellen Dix, Henri Goulet, James Hatfield, Mark Harrell, Frank F. Hasbrouck, Leland
M. Humble, M. Ivie, Stephen V. Johnson, B.C. Kondratieff, Robert J. Lavigne, Garland N.
Mason. Douglas L. Parker, Wojciech J. Pulawski, John M. Schmid, Edward L. Smith.
David R. Smith, Torolf R. Torgersen, Marius S. Wasbauer, Floyd Werner, and Richard L.
Westcott. We would also like to thank, C.D. Johnson and Jon Nealson for reviewing the
manuscript.

LITERATURE CITED

Dahlsten, D.L. 1961. Life history of a pine sawfly, Neodiprion sp., at Willits, California

(Hymenoptera: Diprionidae). Can. Entomol. 98: 1055-1083.
Dahlsten, D.L. 1966. Some biological attributes of sawflies in the Neodiprion fulviceps

complex in a brushfield pine plantation. (Hymenoptera: Diprionidae). Can. Entomol.

98: 1055-1083.
Powells, H.A. 1965. Silvics of Forest Trees of the United States. U.S. For. Serv.. Agric.

Handbook No. 271.762pp.
Ross, H.H. 1955. The taxonomy and evolution of the sawfly genus Neodiprion. For. Sci. 1:

196-209.

272 ENTOMOLOGICAL NEWS

Smith, D.R. 1974. Conifer Sawflies, Diprionidae: Key to North American genera,

checklist of world species, and new species from Mexico (Hymenoptera). Proc.

Entomol. Soc. Wash. 76 (4); 409-418.
Smith, D.R. and M.R. Wagner. 1986. Recognition of two species in the pine feeding

"Neodiprionfulviceps complex" (Hymenoptera: Diprionidae) of western United States.

Proc. Entomol. Soc. Wash. 88 (2): 215-220.
Wagner, M.R., D.G. McCullough and J.M. DiMatteo. 1986. Life history ofNeodiprion

fulviceps (Cresson). A ponderosa pine feeding sawfly (Hymenoptera: Diprionidae).

Proc. Entomol. Soc. Wash. 88 (2): 221-226.
Wilson, L.F. 1977. A guide to insect injury of conifers in the Lake States. U.S. For. Serv.,

Agric. Handbook 501. 218 pp.

(Continued from page 265)

but the photographs survive being printed on matt-textured, cream-colored paper only
because the originals were undoubtedly very sharp.

The nearly 500 pages of text can probably be said to sum up all that is currently known of
scorpion biology, but may not be entirely up to date; the references and wording in some
chapters suggest strongly that they were completed several years ago, perhaps as early as
1983. On the other hand, the bibliography contains numbers of references from 1987, and a
handful from 1988.

As a systematist, I was particularly interested in the chapter on systematics and phy-
logeny, and found it a gem. It includes keys to all the known genera of scorpions, and
diagnoses and other notes for each family. The section on fossil history relies almost
entirely (and understandably) on the posthumous monograph of Kjellsevig-Waering,
which now is seen as containing some serious errors and misinterpretations (for example,
the "gills" of the Devonian Tiphoscorpio are in reality parts of an extinct myriapod, and the
"carapace" of the same animal has no features of a scorpion). The life history and ecology
chapters likewise are excellent, and the one on venoms morbidly fascinating.

The subject of fluorescence arises again in a chapter on field and laboratory methods,
where it is remarked that 500 to 1000 scorpions can easily be captured in a single night using
UV light. Sisson, Polis, and Watt warn, however, that while scorpions fluoresce, rattle-
snakes do not. The scorpionologist must exercise caution!

In summary, this is truly a landmark book: the first real synthesis of the biology of a
group of important and exciting organisms. It now becomes the standard reference on
scorpions, and will remain so for many years.

William A. Shear

Department of Biology,

Hampden-Sydney College,

Hampden-Sydney, VA 23943

Vol. 101, No. 5, November & December, 1990 273

NEOTYPE DESIGNATION FOR LEBIA MORIO

DUFTSCHMID, FIXING THE NAME AS A

JUNIOR SYNONYM OF DROMIUS AGILIS (F.)

(COLEOPTERA: CARABIDAE)1

James K. Liebherr^

ABSTRACT: A neotype is designated for Lebia mono Duftschmid, nomen dubium. which
fixes it as a junior subjective synonym of Dromius agilis (F.) (NEW SYNONYMY), and
removes it from synonymy with Sericoda bogemannii (Gyllenhal).

In 1812 Caspar Duftschmid published volume 2 of his Fauna Austriae,
in which he described Lebia morio (Table 1). In 1868, Gemminger and
Harold synonymized Lebia morio under "Platynus BogemannT
(Gyllenhal, 1813), which had been described as ""Harpalus Bogemannii".
Csiki (1931) retained morio Duftschmid as a junior synonym of the
younger bogemannii Gyllenhal, but placed a "?" before the entry,
presumably indicating his uncertainty regarding the prior action of
Gemminger and Harold. I have recently completed a revision of the
genus Sericoda Kirby (Liebherr, in press), which contains bogemannii as
a member species. In the present note I designate a neotype for L. morio,
fixing it as a junior subjective synonym of Dromius agilis (F.) (NEW
SYNONYMY). This action removes morio from the synonymy of the
younger bogemannii, preserving the stability of that name.

That Gemminger and Harold did not see any specimens of L. morio
prior to their 1868 publication is supported by Gusenleitner's (1984)
explanation of the fate of the Duftschmid collection, which I summarize
below. Duftschmid's collection was purchased by Josef Knbrlein, and in
1861, the Knbrlein collection was purchased by the Oberbsterreichisches
Landesmuseum, Linz. Subsequent examination of the collection by
Landesmuseum curators established that Knorlein had removed the
original labels from the Duftschmid material, and uniformly substituted
his own labels. Some time after 1933, the specimens in the Duftschmid-
Knbrlein collection were incorporated into the main collection of the
museum, removing any positional information of the Knbrlein labeled
specimens. These actions eliminate the possibility of positively identifying
which specimens Duftschmid used as the basis for his Fauna Austriae,
effectively rendering Duftschmid's species nomina dubia. Specimens
held at the Ob. Landesmuseum derived from the Duftschmid-Knbrlein
collection are identifiable by the label — "Alte Sammlung."

Deceived April 16, 1990. Accepted May 22, 1990.

2Dept. of Entomology, Comstock Hall, Cornell Univ., Ithaca. New York 14853-0999.

ENT. NEWS 101(5): 273-275, November & December, 1990

274 ENTOMOLOGICAL NEWS

Duftschmid placed morio in his "FUnfte Familie" of his interpretation
of Lebia (Table 1 ). Duftschmid's generic concept was considerably broader
than that of currently recognized genera. Duftschmid's "Familie", which
in his usage was a rank lower than genus, roughly corresponds to a
present-day genus. Only his "Familie" 4 is heterogeneous phylogeneti-
cally (Table 1). These species also exhibit the smallest body size of any
species Duftschmid included in Lebia. All species in "Familie" 5 are
currently placed in the genus Dromius, with Duftschmid's "agilis F." and
""velox F. (sic = Olivier) currently considered conspecific (Csiki, 1932).

Duftschmid's concept of velox differs from that of agilis chiefly by the
former being black, and agilis being rufous. Duftschmid's morio may be
diagnosed from velox only by having the thorax subtly transversely
rugulose, versus transversely rugulose. It seems likely that Duftschmid's
morio was described from a dark specimen of what would currently be
considered the variably colored Dromius agilis.

The male neotype selected and designated here to represent Lebia
morio Duftschmid bears the labels: Austria (typescript label) / alte
Samml. (typescript label) / agilis F., Aust. (handwritten label) / agilis
Fab., det. M. Priesner / NEOTYPE, Lebia morio Duftschmid,
J.K. Liebherr 1990 (red label). It is deposited in the Oo. Landesmuseum,
Linz. This designation establishes Lebia morio Duftschmid, 1812 as a
junior subjective synonym of Dromius agilis (Fabricius, 1787) (NEW
SYNONYMY), and removes L. morio from synonymy with Sericoda
bogemannii (Gyllenhal), thus ensuring the nomenclatural stability of
this well-known Holarctic species.

Table 1. Listing of species, with "Familie" rank designations, included by Duftschmid
(1812) in Lebia. The doubtful status of Lebia morio Duftschmid (Csiki, 1931) is clarified by
the neotype designation herein. Other current senior synonyms, combinations, and authors
are based on Csiki (1928, 1932), supplemented by Freude et al. (1976) and Cameron
(1988).

"Familie" species epithet and author current status

1 fasciolata F. Polystichus connexus (Geoffrey)

2 axillarisF. Cymindis axillaris (F.)

2 humeralis F. Cymindis humeralis (Geoffroy)

2 homagrica Duftschmid Cymindis axillaris (F.)

2 lunaris Dahl Cymindis angularis Gyllenhal

2 miliaris F. Cymindis variolosa F.

3 crux-minor L. Lebia cruxminor L.

3 cyanocephala L. Lebia cyanocephala L.

3 chlorocephala Hoffman Lebia chlorocephala (Hoffman)

3 haemorrhoidalis F. Lebia marginata (Geoffroy)

3 turcica F. Lebia scapularis (Geoffroy)

Vol. 101, No. 5, November & December. 1990

275

'Familie" species epithet and author

4 quadrillum Creutzer

4 quadriguttata Paykull

4 truncatella L.

4 punctatella Megerle

4 glabrata Megerle

4 obscuro-guttata Andersch

4 plagiata Megerle

5 quadrimaculala L.
5 agilis F.

5 ve/ox F.

5 mono Duftschmid

5 quadrinotata Panzer

5 fascitata F.

6 atricapilla L.

6 elongatula Zenker

6 punctato-striata Duftschmid

current status

Lionychus quadrillum (Duftschmid)
Bembidion quadrimaculatum (L.)
Syntomus truncatellus (L.)
Syntomus foveatus (Geoffroy)
Microlestes minutulus (Goeze)
Syntomus obscuroguttatus (Duftschmid)
Microlestes plagiatus (Duftschmid)

Dromius quadrimaculatus (L.)
Dromius agilis (F.)

Dromius agilis (F.)
9

Dromius quadrinotatus (Panzer)
Dromius sigma (Rossi)

Demetrius atricapillus (L.)
Demetrias atricapillus (L.)
Dromius linearis (Olivier)

'Authorship listed is that attributed by Duftschmid (1812) even though incorrect in som>
cases.

ACKNOWLEDGMENTS

I thank Mag. Fritz Gusenleitner, Abtlg. Botanik und Evertebrates, Oo.Landesmuseum.
Linz, for gracious access to information and specimens. I thank J.D. Oswald and E.R.
Hoebeke for advice about nomenclarural matters, and for critical reviews of the manuscript.

LITERATURE CITED

Cameron, H.D. 1988. Geoffroy in Fourcroy. 1785. Sphecos 17: 6-7.

Csiki, E. 1928. Ofoleopterorum Catalogus (part 97). Carabidae 2. Mormolycinae.

Harpalinae 1: 1-226. W. Junk. Berlin.
Csiki, E. 1931. Coleopterorum Catalogus (part 115), Carabidae 2, Mormolycinae.

Harpalinae 5: 739-1022. W. Junk. Berlin.
Csiki, E. 1932. Coleopterorum Catalogus (part 124). Carabidae 3. Harpalinae 7: 1279-1 598.

W. Junk, Berlin.
Duftschmid, K. 1812. Fauna Austriae, oder beschreibung der osterreichischen Insekten

fur angehende Freunde der Entomologie 2: 1-311. K.K. priv. akad. Junst- Musik- und

Buchhandlung, Linz and Leipzig.

Fabricius, J.C. 1787. Mantissa Insectorum 1: 1-348. Hafniae.
Freude, H., K.W. Harde, and G.A. Lohse. 1976. Die Kafer Mitteleuropas, 2 (Adephaga

1): 1-302. Goecke und Evers, Krefeld.
Gemminger, M., and B. de harold. 1868. Catalogus Coleopterorum 1 (Cicindelidae

Carabidae): 1-424. E. Deyrolle fils. Paris.
Gusenleitner, F. 1984. Das Raisel um den Verblieb der Caspar Erasmus Duftschmid-

Kollektion. Koleopt. Rundschau 57: 93-95.
Gyllenhal, L. 1813. Insecta Suecica, Classis 1. Coleoptera sive Eleuterata 1(3): 1-730. FJ.

Leverentz, Scaris.
Liebherr, J.K.. in press. Phylogeny and Revision of the Anchomenus Clade: the genera

Tetraleucus. Anchomenus. Sericoda, and Elliptoleus (Coleoptera: Carabidae: Platynini).

Bull. Amer. Mus. Nat. Hist.

276 ENTOMOLOGICAL NEWS

REARING AND DEVELOPMENT OF

PHYLLOCOPTES FRUCTIPHILUS

(ACARIiERIOPHYIDAE)1'2

Abdullah Kassar, James W. Amrine Jr.

ABSTRACT: A mass rearing program for Phyllocoptesfructiphilus, the eriophyid vector of
rose rosette disease, was developed. The mites inhabit shoot tips and leaf petiole bases of
several varieties of roses. A rearing arena, allowing observation of individual mites, was
designed. Developmental periods for life stages, and wintering form and sites are
reported.

Rose rosette disease (RRD) is a disease of many wild and domestic
roses, especially Rosa multiflora Thunb. It was orginally described and
reported from western states and Canada (Thomas and Scott 1953) and
more recently from midwestern states (Allington etal. 1968, Crowe 1982,
Gergerich and Kim 1983, Hindal and Amrine 1987, 1989). It also is
known from Ohio in 1988 and West Virginia in 1989 (Amrine,
unpublished).

Rose rosette disease is transmitted by the eriophyid mite, Phyllocoptes
fructiphilusKeikr(A\\'mgtonetal. 1968, Amrine el al 1988, Gergerich and
Kim 1983). Little is known about the mite's biology, but its life history is
thought to be similar to that of other eriophyids (Allington et al. 1968). It
lives and breeds in protected areas between leaf petiole bases and lateral
buds, within small developing leaflets and particularly on the tips of
rapidly growing shoots. Many eriophyids that overwinter as adults are
known to develop deuterogynous females. However, none have been
reported for this species. Conventional methods of rearing individual
eriophyid mites (Rice and Strong 1962, Tashiro 1967, Slykhuis 1969)
proved unsatisfactory foTP.fructiphilus, since the mite could not feed and
breed satisfactorily. Also, rearing large colonies in the greenhouse was
not possible because spider mites generally invade and must be controlled.

This paper describes a reliable method for rearing large colonies of P.
fructiphilus and a chamber for observation of individual mites. Life
history data are also reported.

^Received March 10, 1990. Accepted June 13, 1990

-Published with the approval of the Director of the West Virginia Agricultural and Forestry

Experiment Station as Scientific Article #2217. This research was supported in part with

funds appropriated under the Hatch Act.
^Division of Plant and Soil Sciences, P.O. Box 6108, West Virginia University, Morgantown

WV 26506-6108.

ENT. NEWS 101(5): 276-282. November & December, 1990

Vol. 101, No. 5, November & December, 1990 277

MATERIALS AND METHODS

Large colonies of mites were maintained year-round on potted R.
multiflora plants in illuminated rearing stands. Each rearing stand was
constructed of wood 5x10 cm (2"x4"'s) with dimensions of 1.2x0.8x2.3 m
to accommodate two shelves, one at the bottom and one at the center
(Figure 1). Lighting for each shelf consisted of five pairs of 1.2m fluor-
escent tubes (40 W each), three pairs at the top (l.lm above each shelf)
and one on each side, and six incandescent lights ( 100 W each) in two
rows of three each at the top between the fluorescent tubes. Each shelf
had a separate switch and timer for each type of light.

Large R. multiflora were dug, pruned to the crown, potted in sterilized
soil medium, placed on the stands, and maintained at 16L:8D under
fluorescent lights. Incandescent lights were used for 4 hrs in mid-
photoperiod to stimulate mid-day higher temperatures (33°C) and light
intensity. Temperature ranged from 27°C to 33°C and relative humidity
averaged 45%. The plants were artificially infested with noninfected P.
fmctiphilus obtained from R. multiflora in Morgantown, WV, and with
RRD-infected mites from R. multiflora in Madison, IN. Noninfected and
RRD-infected colonies were kept in separate rooms and mites were
regularly identified to insure species purity. Special precautions were
taken to assure exclusion of spider mites: trimming plants to two or three
canes 1 5-20 cm long; spraying these basal canes with spraymount to trap
any attached mites; and restricting entry, especially if person(s) had
visited the greenhouse.

To rear and observe individual mites, a 12x12 mm plastic coverslip
(0.17-0.25 mm thick) was cut into four, 6x6 mm sections, and a 3 mm
diameter hole was punched into the center of each section. The 6x6 mm
section was cemented to a young host plant leaflet using Elmers Glue- All
(Figure 2). An egg or female P.fructiphilus was placed in the arena, and a
circular, 12 mm diameter glass coverslip (0.13-0.17 mm thick), treated
with Spritz anti-fog spray, placed on top. Each leaf with arenas and
damp facial tissue around the leaf base was then placed on damp filter
paper in a petri dish and maintained at 16L:8D under fluorescent lights.
Test leaves were maintained at room temperature (23±1°C).

To determine the duration of each life stage, younger leaves from
healthy multiflora rose were prepared with 5-10 arenas, depending on
leaf size (Figure 3). For each trial of three replicates, 20 adult mites (live
mites can not be sexed, but males made up only 10-30% of population
(Amrine, unpublished)) were placed singly in separate arenas. Adults
were removed after egg depositon. Eggs were checked at 6-hr intervals,
and emerging larvae were transferred to separate arenas (one per arena)
on fresh leaves. The number of stages and time required for each

278

ENTOMOLOGICAL NEWS

Figure 1. An illuminated rearing stand.

Vol. 101, No. 5. November & December. 1990

279

A
B
C

KXXXXXXXXXXXXXXXXXXXXXXXX^^

Figure 2. Cross section of a rearing arena (12X). A, coverslip treated with anti-fog;
B, plastic coverslip with 3 mm diameter hole in center; C, host plant leaflet.

Figure 3. The arrangement of individual arenas on a host leaf. A, arena: B.
damp tissue.

280 ENTOMOLOGICAL NEWS

developmental period were recorded.

To determine wintering stages and sites, plant material was collected
in December 1988 at Madison IN from plants known to have been
infested with P. fructiphilus and returned to the lab and kept in 5°C
refrigerator to be examined thoroughly for over-wintering mites.

RESULTS AND DISCUSSION:

Mite Rearing. The rearing stands produced good growth of multi-
flora rose plants and supported large colonies of P. fructiphilus which
lasted for about 3-4 months under these conditions. The rearing arena
was successful in caging all stages of mites, for observation. However,
most adult P. fructiphilus moved about and climbed the arena wall and
onto the coverslip. Adults apparently fed little and survival was low; this
is reflected in the scarcity of adult data in Table 1 . The low acceptance of
arena habitus by adult mites probably reflects their preference for rapidly
growing shoot tips.

Life Cycle. PI fructiphilus has a typical eriophyid mite life cycle which
consists of egg, protonymph, deuto nymph and adult. Development times
for each stage are presented in Table 1 . Newly laid eggs were transparent
and became milky white as they aged. Single eggs were deposited ran-
domly within the feeding arena. The egg stage averaged 4.3 days.

Newly emerged protonymphs were transparent and also became white
with age. Protonymphs were active and fed for about 2.4 days before
transforming into the stationary, swollen and shiny "phararte" form.
Deutonymphs resemble adults in size and shape. As they mature, the
color changes from white to yellowish white. They actively move and
feed for about 2.5 days before the mities become quiescent, swollen and
shiny, pharate forms.

The yellowish white adults are active and start feeding almost im-
mediately after emerging. Females start laying eggs within 12-24 hrs.
after emergence. In our study, females laid an average of 1 egg/day for the
first few days and then no eggs for the remainder of the adult stage, a
period which varied from 10 to 50% fo their life span producing an
average of 0.64 eggs/day (Table 2).

Hibernation Sites. Examination of whole branches in December
revealed an orange form of the mite which had been noticed in field
populations during the 1987 and 1988 seasons. This form is thought to
represent either a facultative change in color and shape as a response to
adverse conditions, or it may be a deutogyne. Aside from the orange
color and more trapezoidal shape, there were no obvious anatomical
differences from typical P. fructiphilus to support a deutogyne desig-

Vol. 101, No. 5, November & December, 1990 281

Table 1. Life stage intervals for Phyllocoptes fructiphilus

Mean Duration
Stage N (days) Range

egg 37 4.31 ±0.1 6 1.29-6.04

Protonymph 31 3.29±0.13 2.17-4.33

Deutonymph 32 3.29±0.18 1.83-4.63

adult 7 14.14±0.96 6.00-26.00

Table 2. Egg production of Phyllocoptes fructiphilus K.

Mite No.

Sex

Days
Observed

Total
Eggs Laid

Eggs/Day

0.83

0.00

0.25

0.86

0.63

0.79

0.50

Mean

14.14

7.17

0.64

Variance

6.64

2.79

0.24

Standard Error

0.97

0.63

0.19

nation (Amrine, unpublished). P. fructiphilus wintered in groups or singly
in various sheltered places but mostly under bud scales of small lateral
buds and occasionally under the loose bark of the previous year's growth.

SUMMARY

A method of rearing P. fructiphilus using artificial lighting and
transplanted multiflora rose plants proved successful in reducing spider
mite infestations and eliminating the need to control spider mites.

A special arena was designed utilizing plastic slide coverslips glued to
rose leaflets which allowed study of the life stages from egg to the adult.
The average duration of life stages was: egg 4.31 days, first nymph 3.29
days, second nymph 3.29 days and adult 14.14 days.

282 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

We wish to thank the West Virginia Department of Agriculture, Plant Protection
programs for providing financial support for this research. We also thank Terry Stasny for
her contributions to the research, and L. Butler, J. Weaver and H. Hogmire for comments
on the manuscript

LITERATURE CITED

Allington, W.B., R. Staples and G. Viehmeyer, 1968. Transmission of rose rosette virus

by the eriophyid mite Phvllocoptesfructiphilus. J. Econ. Entomol. 61:1 132-1 140.
Amrine, J.W., D.F. Hindal, T.A. Stasny, R.L. Williams and C.C. Coffman. 1988.

Transmission of the rose rosette disease agent to Rosa multiflora by Phyllocoptes

fructiphilus (Acari:Eriophyidae). Ent. News 99:239-252.
Crowe, F.J. 1982. A recent outbreak of Witches' Broom of rose in eastern Kansas and

western Missouri. Phytopath. 72:976-977.
Gergerich, R.C. and K.S. Kim. 1983. A description of the causal agent of rose rosette

disease. Arkansas Farm Res. 32: 7.
Hindal, D.F. and J.W. Amrine. 1987. New findings of rose rosette disease. Phytopath.

77:987 (abstract).
Hindal, D.F., J.W. Amrine, R.L. Williams and T.A. Stasny. 1988. Rose rosette

disease on multiflora rose (Rosa multiflora) in Indiana and Kentucky. Weed

Technology 2: 442-444.
Rice, R.E. and F.E. Strong. 1962. Bionomics of the tomato russet mite, Vasates lycopersici

(Massee). Ann. Entomol. Soc. Amer. 55:431-435.

Slykhuis, J.T. 1969. Methods for experimenting with mite transmission of plant viruses,
pp. 347-368. In: K. Maramorosch & H. Koprowski. Methods in Virology, Vol.1,

Academic Press. 640 pp.
Tashiro, H. 1967. Self-watering acrylic cages for confining insects and mites on detached

leaves. J. Econ. Entomol. 60:354-356.
Thomas, E.A. and C.E. Scott. 1953. Rosette of rose. Phytopathology 43: 218-219.

Vol. 101, No. 5, November & December. 1990 283

OBSERVATIONS PERTINENT TO THE ROLE OF
SEXUAL SELECTION IN THE STONEFLY

PTERONARCELLA BAD I A
(PLECOPTERA: PTERONARCYIDAE)1

David D. Zeigler^

ABSTRACT: Virgin males of Pteronarcella badia delivered semen volumes of just over one
percent of their body weight. Mated females oviposited the great majority of their eggs
shortly after mating and thereafter remained unreceptive to male drumming calls and
tactile contacts. Implications of these findings are discussed in relation to sexual selection
theory.

Much work and even more theorizing has appeared over the past two
decades in the area of sexual selection. Perhaps not surprising, much of
this work has involved insects (Thornhill and Alcock 1983, Alexander
and Borgia 1979, Kaneshiro 1983, Carson 1978,West-Eberhard 1984, and
many others). Some workers imply that sexual selection is, in varying
forms and to varying degrees, essentially ubiquitous in sexually repro-
ducing animals (West-Eberhard 1984, Thornhill and Gwynne 1986).

My work with the reproductive calling behavior of stoneflies
(drumming) has led to the question of sexual selection's possible role in
shaping adult behavior(s) in this interesting but inadequately studied
group. This paper will deal with two aspects of reproduction which bear
on the nature and degree of sexual selection pressures to be expected in
stoneflies. The first question is that of semen volume transferred by
males during mating. The second is the relative degree of polygyny/
ployandry typical of stoneflies.

MATERIALS

Mature Pteronarcella badia (Hagen) nymphs were collected from the
Conejos River, Conejos Co., CO in late June 1988 just before peak
emergence of the adults. Nymphs were transported to the lab (Southwest
Texas State University, San Marcos, TX) for rearing in chilled styrofoam
ice chests. Adults were separated at emergence to insure virgin condition.
Small triangular enclosures of sheet balsa wood (7.5 mm/side and 1.5
mm deep) with clear plastic lids were used to contain adults during
observations. All virgin contacts, including matings. were between
adults two to three days old. Pre-and post-mating weights of males were
made using an American Scientific Products S/P 120 scale. Values re-
ported below are means plus or minus sample standard deviations.

^Received March 1. 1990. Accepted June 1. 1990.

-Biology Department. Pembroke State University. Pembroke. NC 28372

1-NT. NEWS 101(5): 283-287. November & December. 1990

284 ENTOMOLOGICAL NEWS

RESULTS AND DISCUSSION

Semen Weight — Six intersexual pairings of virgin P. badia adults were
made in order to estimate the amount of semen transferred in a first
mating situation. Males were weighed just before and after mating. Prior
to mating, males weighted .043 1 ± .0042 gr. while postmating weight was
.0426 ± .0041 gr. This amounts to only a 1.12 ± .022 percent loss in body
weight. In numerous other insect groups, the male transfers large semen
volumes or spermatophores (from 20 to 40 percent of the male's pre-
mating weight) which are apparently used by the females in part as a
nutritional resource (reviewed by Thornhill and Alcock 1983, Gwynne
1983, Thornhill and Gwynne 1986). This nourishment may be essential
for final development and formation of the female's eggs, and, when in
the form of a spermatophore, may actually be eaten by the female
(Gwynne 1983, Thornhill and Gwynne 1986). Movement of most sper-
matophore/semen nutrients into the female's eggs can require from less
than 24 to over 70 hours (Gwynne 1983). Generally, females are the more
choosy of the two sexes due to their larger relative investment in the
offspring. In species where males deliver substantial nutrients with their
semen, females might be expected to mate preferentially with males
capable of donating large spermatophore/semen volumes. However,
this expectation will be counterbalanced by an increase in male selec-
tivity for "optimal" females due to the male's increased material
investment in the offspring (Gwynne 1983, Thornhill and Gwynne 1986).
Such "role reversal", typified by choosy males, seems unlikely in P. badia
judging from the small weight of semen transferred, and from the short
time period between mating and egg laying (see below).

Males of P. badia show no obvious signs of choosiness in mate
selection. They typically attempt mounting virgin females, non-virgin
females, and even other males. Also, in two out of eight separate inter-
specific pairings of P. badia males with Isogenoides zionensis Hanson
females (a co-emerging species at the Conejos River), the males mounted
the females and attempted copulation. Each of these two males remained
mounted for over five minutes before dismounting. These observations
suggest a lack of identification contact pheromones in P. badia as well as
a lack of selectivity and discrimination by the P. badia males.

The six intraspecific matings were not timed, but all lasted approx-
imately 30-45 minutes. After mating, females were gently transferred to
styrofoam cups with screen lids. Each cup contained three cm of river
water for oviposition. All six females laid eggs shortly after mating (395 ±
54 eggs). This egg data compared favorably with that from 16 other P.
badia females (354 ± 74 eggs) which were mated in the course of other
experiments. Seven of the latter were observed to lay second clutches of

Vol. 101. No. 5. November & December. 1990 285

eggs (55.1 ± 23.4 eggs) 2-5 days after the initial batches were laid. Two of
these seven females were remated by virgin males prior to their second
laying, but the other five had no secondary male contacts. These data
suggest that females require only one mating for the fertilization of most,
if not all, of their eggs, most of which are laid in the first oviposition event.
The six females mated in the semen weight experiments lived for 1 1.4 ±
0.9 days.

Mated Female Response to Male Calls — Five P. badia males were
seen to resume calling within minutes after their first mating, and two
similar observations were made after second matings. Males of
Taeniopteryx nivalis (Fitch) have also been observed to resume calling just
after mating (Stewart and Zeigler 1984). These observations indicate that
male stoneflies are polygynous, though to what extent is still an
unanswered question. Mated female stoneflies have not been observed
to answer male calls (Rupprecht 1967, Stewart and Zeigler 1984), but this
observation has often been based on a female's exposure to male calls
only minutes or hours after mating. To test the hypothesis that females
might again become responsive to male calls after egglaying and/or
some refractory period, mated females (which laid eggs within 6 hours
after mating) were exposed to male calls as follows. Five females were
exposed to male calls (her chamber acoustically coupled to a chamber
containing a signaling male) 24 hours after mating, six females were
exposed to calls 48 hours after mating, and seven females were exposed
to calls 96 hours after mating. In none of these 1 8 exposures was a mated
female observed to answer male calls. Similar acoustically coupled
chambers, with the male and female in two separate but coupled chambers,
have been used in previous studies to record the drumming signals of
virgin stonefly pairs and have a "proven" track record of providing
efficient signal transfer between the sexes (Stewart and Zeigler 1984.
Zeigler and Stewart 1985).

Observations of tactile contacts between other mated females and
virgin males also indicate an un willing or at least "disinterested attitude"
on the part of the females. One mated female (56 hrs post-mating) ran
from a virgin male on initial contact and would probably have escaped
from him in nature. As both were contained in a common chamber, the
male eventually mounted her and mated. This female continued to move
about the chamber while the male was engaged (virgin females typically
remain motionless during the mating process). Another female, minutes
after mating but before egglaying, successfully rejected a second male's
mounting attempts by raising her abdomen at about a 90 degree angle
from her body axis and wagging it laterally during mounting attempts.

286 ENTOMOLOGICAL NEWS

Similar rejection responses have been noted in other stoneflies
(Rupprecht 1967, Zeigler and Stewart 1977, Stewart and Zeigler 1984).

In terms of sexual selection theory, a female who puts all her eggs in
one basket (fertilizes all her eggs with sperm from one male) should
profit by being choosy as to which male fertilizes her eggs (Thornhill and
Gwynne 1986). This is especially so for insects such as stoneflies where
the male contributes nothing to the female or offspring except sperm
(Borgia 1 98 1 ). But, if adults are short-lived and females typically encounter
males infrequently, mating with the first male encountered may be the
optimal strategy due to the costs of locating two or more males and
making a comparison (Wittenberger 1983). Predation, bad weather, or
other environmental hazards could also make waiting around for a
second more "attractive" or fit male a suboptimal strategy (Borgia 1979.
Wittenberger 1983). Alternately or concurrently, if males are typically of
equal or near-equal fitness and meet some minimum threshold specifi-
cations (a proposition I am now addressing in an unfinished manuscript),
the female would on average, be best served by mating with the first male
encountered (Wittenberger 1983). Based on years of drumming studies, I
can state qualitatively that as a group virgin stonefly females are not
obviously coy or choosy in selecting a mate. Females typically answer the
drumming call of, and mate with, the first male encountered (these
statements derive largely from laboratory observations).

In nature, females may feed and so fuel the continued development of
a second large egg batch, but several hours of observations along the
Conejos River during peak emergence have not yielded evidence of adult
feeding (Ed Dewalt, personal communication). The data on feeding in
adult stoneflies as reviewed by Hitchcock (1974) and Hynes (1976) in-
dicate that adults of the family Pteronarcyidae, which includes
Pteronarcella, do not feed but do drink water (and may take in honeydew
or nectar). Adults of both sexes were provided with water and were seen
drinking during the course of observations. In some other families,
adults do feed, and in some genera the females apparently require food
in order to fuel initial egg development and maturation, which can take
many days (Hynes 1976), but this is obviously not the pattern in P. badia.
Two of the 16 P. badia females mentioned above were mated within two
hours of emergence and went on to lay a typical clutch of eggs within five
hours of mating.

In conclusion, it appears likely that females of P. badia rely on single
matings to fertilize most, if not all, of their eggs. The possibilry exists that
second matings may occur shortly after the initial mating but prior to egg
laying, possibly with sperm displacement by the second male, but the
non-responsiveness of mated females to male calls and observations of

Vol. 101, No. 5. November & December. 1990 287

male rejections by recently mated females argue against this being a
typical event.

ACKNOWLEDGMENTS

I wish to thank Ed Dewalt who assisted in this project with nymph collection, egg counts,
field observations, and "stimulating" conversation.

LITERATURE CITED

Alexander, R.D. and G. Borgia. 1979. On the origin and basis of the male-female

phenomenon, pp. 417-440. In M.S. Blum and N.A. Blum (eds.). Sexual Selection and

Reproductive Competition in Insects. Academic Press. New York.
Borgia, G. 1979. Sexual selection and the evolution of mating systems, pp. 19-80. in M.S.

Blum and N.A. Blum (eds.). Sexual Selection and Reproductive Competition in Insects.

Academic Press. New York.

Borgia, G. 1981. Mate selection in the fly Scatophaga stercoraria: female choice in a male-
controlled system. Anim. Behav. 29: 71-80.
Carson, H.L. 1978. Speciation and sexual selection in Hawaiian Drosophilia. pp. 93-107. In

P.P. Brussard (ed.). Ecological Genetics: The Interface. Springer- Verlag. New York.
Gwynne, D.T. 1983. Male nutritional investment and the evolution of sexual differences

in theTettigoniidae and other Orthoptera. pp. 337-366. In D.T. Gwynne and G.K. Morris

(eds.). Orthopteran Mating Systems: Sexual Competiton in a Diverse Group of Insects.

Westview Press. Boulder. Colorado.
Hitchcock, S.W. 1974. Guide to the Insects of Connecticut: Part VII. The Plecoptera or

Stoneflies of Connecticut. Bulletin Number 107. State Geological and Natural History

Survey of Connecticut.

Hynes, H.B.N. 1976. Biology of Plecoptera. Ann Rev. Entomol. 21: 135-153.
Kaneshiro, K.Y. 1983. Sexual selection and direction of evolution in the biosystematics of

Hawaiian Drosophilidae. Ann. Rev. Entomol. 28: 161-178.

Rupprecht, R. 1967. Das Trommeln der Plecopteren. Z. Vergl. Physiol. 59: 38-71.
Stewart, K.W. and D.D. Zeigler. 1984. Drumming behavior of twelve North American

stonefly (Plecoptera) species: first descriptions in Peltoperlidae. Taeniopterygidae. and

Chloroperlidae. Aquatic Insects. 6: 49-61.
Thornhill, R. and D.T. Gwynne. 1986. The evolution of sexual differences in insects. Am.

Sci. 74: 382-389.
Thornhill, R. and J. Alcock. 1983. The Evolution of Insect Mating Systems. Harvard

University Press. Cambridge. Massachusetts.

West Eberhard, MJ. 1984. Sexual selection, competitive communication and species-
specific signals in insects, pp. 283-324. In T. Lewis (ed.). Insect Communication.

Academic Press, New York.
Wittenberger, James F. 1983.Tacticsofmatechoice. pp. 435-447. In P. Bateson(ed.). Mate

Choice. Cambridge University Press. Cambridge.
Zeigler, D.D. and K.W. Stewart. 1977. Drumming behavior of eleven Nearctic stonellv

(Plecoptera) species. Ann. Entomol. Soc. Am. 70: 495-505.
Zeigler, D.D. and K.W. Stewart. 1985. Drumming behavior of five stoneflv (Plecoptera)

species from central and western North America. Ann. Entomol. Soc. Am. 78: 717-

722.

288 ENTOMOLOGICAL NEWS

DEVELOPMENT OF THE BLACK WILLOW SCALE,

CHIONASPIS SALICISNIGRAE
(HOMOPTERA: DIASPIDIDAE), IN TENNESSEE1

Paris L. Lambdin^

ABSTRACT: Weekly collections of the black willow scale, Chionaspis salicisnigrae, on Salix
nigra were made from 7 January 1984 to 3 1 December 1989 to determine the life history of
the species. This species overwintered as eggs and had three generations per year in
Tennessee. Each female deposited a mean of 152 (18-265) eggs beneath a waxy, tentlike
covering. Adult males emerged in May, August and September. Behavioral aspects of each
instar are discussed.

The black willow scale, Chionaspis salicisnigrae (Walsh), was originally
described in 1868 from specimens collected on black willow, Salix nigra
Marsh. Species of Chionaspis are predominatly mono- or oligophagous
insects (Takagi 1 969). The black willow scale has been collected from the
bark and leaves of several ornamental trees in the United States that
include: Amelanchier canadensis (L.), (Rosaceae); Cornus pubescens Nutt.
and C. asperifolia Michx., (Cornaceae); Fraxinus americana L. (Oleaceae);
Populus canadensis Moench., P. candicans Ait., P. deltoides Marsh., P.
grandidentata Michx., P. tremuloides Michx., Salix interior Rowlee, S. nigra
Marsh., and Salix sp. (Salicaceae), (Kosztarab 1963, Dekle 1976). Willow
trees, planted to enhance the aesthetic beauty of residential areas, are
often infested with this pest. Damage to the host is caused by sap extraction
which results in loss of vigor, dieback, stunting and eventual death of the
affected plant.

The black willow scale was reported to infest native willows in Indiana
and occasionally became an economic problem (Diez and Morrison
1916). Langford (1926) concluded that this species had two generations
in Colorado with eggs hatching in late April and July, respectively. Also,
Houser ( 1 9 1 8) and Kosztarab ( 1 963) reported that the overwintering eggs
hatched in mid-May in Ohio and that C salicisnigrae may have two
generations per year. Because little information is known concerning the
biology of the black willow scale, my objective was to determine the life
history of this species in Tennessee.

^Received June 12. 1990. Accepted August 1. 1990.

"Department of Entomology and Plant Pathology, University of Tennessee, Knoxville.TN
37901

ENT. NEWS 101(5): 288-292, November & December, 1990

Vol. 101, No. 5, November & December, 1990 289

MATERIALS AND METHODS

Chionaspis salicisnigrae on S. nigra, was studied under both field and
laboratory conditions from 1984-89 at The University of Tennessee,
Knoxville. Greenhouse populations were maintained on black willow
cuttings ( 6 to 26 cm long) grown in an equal mixture of clay, sand and
peat in plastic pots (10 cm in diam.). Field populations of C. salicisnigrae
on 6 infested black willow trees and 1 2 potted saplings, located at the U.T.
Plant Science Farm, were sampled weekly. Collections of scale insects
from infested branches (2 to 4 cm long) were taken to the lab to be
processed, stained, and mounted on slides to construct the life history of
the species. In the greenhouse, fecundity was determined by counting the
number of eggs deposited under the test of 100 females, and by placing
the females in Hoyer's solution to count the remaining number of eggs
visible through the derm. Number of generations per year and develop-
mental rates were monitored by transferring a minimum of 100 (100-145)
newly emerged crawlers to each of 1 2 uninfested host plants upon eclosion.
The tests were replicated 4 successive years from 7 Jan. 1984 to 31 Dec.
1989.

RESULTS AND DISCUSSION

The black willow scale is a multivoltine species that overwintered as
eggs under the parental tests. Females undergo three developmental
stages, the mobile crawler stage followed by the sessile second and adult
stages. Males have additional prepupal and pupal stages before devel-
opment into the mobile adult stage. Waxy tests of the immature stages
were enlarged by moving their abdomens side to side while secreting wax
through the pygidial macroducts as described by Stoetzel (1976).

Populations in the greenhouse developed one to two weeks earlier
than those in the field. Due to the mild winter and spring in 1989.
overwintering eggs began to hatch about two weeks earlier than for
previous years. Females deposited a mean of 152 (18-265) eggs over a 10-
14 day period. Absolute fecundity was not obtained. At eclosion, the
chorion split anteromedially and the crawler freed itself with leg move-
ments. Eclosion of eggs within the population was staggered and may
account for the overlapping of stages within a generation. The smooth
chorion of each egg was covered with minute, waxy filaments that prevent
the eggs in a cluster from adhering to one another. Eggs were 0.25 mm
long, 0. 10 mm wide, and reddish-purple. Overwintering eggs were present
from mid-November to 1st week in May (Fig. 1). Eggs were deposited in
June, August and in mid-November.

290 ENTOMOLOGICAL NEWS

The mobile crawlers began emerging the 3rd week in April, 2nd week
in June, and the 4th week in August. Color of the newly emerged crawler
was similar to that of the egg. Several of the female crawlers settled under
the old parental tests which eventually resulted in a heavy encrustation
on the branches. Heaviest infestations on willow trees occurred on the
trunk and branches in the lower 1/3 to 1/4 region of the tree or sapling. As
competition for space increased on older branches, the distance crawlers
settled from each other diminished resulting in complete encrustation of
these branches. Those females that settled on new growth were often
found near the twig nodes. Most males migrated to the leaves where they
settled on the top surface near the veins. Those male crawlers remaining
on the twigs and branches frequently settled adjacent to a cluster of
females. Once settled on a suitable site, the crawlers began exuding fine,
waxy filaments from the dermal microducts and macroducts that gave
the specimens a fuzzy appearance. Second instars were found the 1st
week in May, 2nd week in July, and 1st week in September. The snow-
white tests of the second stage females were subcircular and slightly
convex, while those of the males were distinguished by being more
elongated with parallel sides and rounded posteriorly. Prepupal males
were present from 2nd-to-4th week in May, the 2nd week in July to 1st
week in August, and the 2nd-to-4th week in September. Prepupal males
developed into pupal males after ca. one week. The immature male tests
possessed a flexible posterior exit flap that allowed the adult male to
emerge by backing out of the test.

The ephemeral adult males began emerging the 4th week in May, 1st
week in August and the 2nd week in September. Upon emergence, males
immediately began to seek out and fertilize adult females. Flight activity
in males was rare. However, when they came in contact with the edge of a
leaf or twig, they would often take to flight. Two adult males have been
found among the field population that had no wings. Males were observed
to walk over the surface of an infested branch constantly tapping the
female tests with their antennae. When a suitable female was encount-
ered, the male would raise the posterior of the abdomen bringing
forward the genitalia between the legs and positioned it beneath the
female test at the anal groove in search of the vulvar orifice. After mating,
males often groomed themselves by rubbing the genitalia between the
prothoracic legs before beginning a search for other females. Males
observed died within 24-36 hours after emergence.

The adult females were present the 3rd week in May, 4th week in July
and 3rd week in September. The lady beetle, Rhyzobius lophanthae
(Blaisdell) (Coleoptera: Coccinellidae), was commonly found feeding
on eggs deposited beneath the test of the female. Upon discovery of a

Vol. 101. No. 5, November & December, 1990

291

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292 ENTOMOLOGICAL NEWS

gravid female, the lady beetle chewed an irregular hole in the dorso-
posterior section of the scale test to feed on the eggs.

The black willow scale was found to have three generations per year in
Tennessee which differs somewhat from the two generations reported by
Hollinger (1923), Langford (1926) and Kosztarab (1963) for studies on
more northern and western populations. The report of 33 (1 1 - 54) eggs
per female by Langford (1926) appears quite low in comparison to some
152 (18-265) eggs per female found in this study. Additonal studies are
needed to determine the influence of temperature on development and
morphology of the various stages of the black willow scale.

ACKNOWLEDGMENTS

I wish to thank Dave Paulsen and Feng Pingzhang, Department of Entomology and
Plant Pathology, University of Tennessee, Knoxville.TN, for their most valuable assistance
in collecting and processing specimens for this study, and J. B. Chapin, Department of
Entomology, Louisiana State University, Baton Rouge, LA, for her help in the identi-
fication of the ladybug found feeding on the black willow scale.

LITERATURE CITED

Dekle, W.W. 1976. Florida armored scale insects. In Arthropods of Florida and

neighboring land areas. Fla. Dept. Agric. Dir. PI. Ind. 3:345p.
Dietz, H.F. and H. Morrison, 1916. The Coccidae or scale insects of Indiana. Office of St.

Entomologist. 195-321.

Hollinger A.H. 1923. Scale insects of Missouri. Univ. of Mo. Agric. Exp. Sta. Bull. 58:71p.
Houser, J.D. 1918. The willow scale. Ohio Exp. Sta. Bull.: 332p.
Kosztarab, M. 1963. The armored scale insects of Ohio (Homoptera: Coccoidea:

Diaspididae). Bull, of the Ohio Biol. Sur. 2:120p.
Langford, G.S. 1926. The life history of the willow scale (Chionaspis salicis-nigrae Walsh).

In Colo. Sta. Entomol. Cir. 51:50-58.
Stoetzel, M.B. 1976. Scale-cover formation in the Diaspididae (Homoptera: Coccoidea).

Proc. Entomol. Soc. Wash. 78:323-332.

Takagi, S. 1969. A new species of Chionaspis from Florida. Kontyu. 37:269-271.
Walsh, B.D. 1868. First report on noxious and beneficial insects of Illinois. Rpt. Acting

State Entomologist. 40p.

Vol. 101, No. 5, November & December, 1990 293

THE LEAFHOPPER GENUS BYTHONIA
(HOMOPTERA: CICADELLIDAE)1> 2

H. Derrick Blocker3, M.D. Webb4

ABSTRACT: Available specimens of the genus Bythonia are examined, and the subfamily
status of Bythoniinae is reviewed. A new species of Bythonia from Brazil is described, and
the genus is assigned to the subfamily lassinae.

The genus Bythonia Oman (1936) was described from a female
specimen from Bolivia, Nionia rugosa Osborn. Linnavuori (1959)
described the male of rugosa from a specimen from Peru and one new
species, kalypso, from a single male specimen from Brazil. He described
the subfamily Bythoniinae based on these three specimens. A third
species is described here from a single male specimen collected in Brazil
and deposited in the British Museum (Natural History).

Bythonia is here assigned to the subfamily lassinae based on a review
of this subfamily by Blocker (1979a), which agrees with the subfamily
description of Kramer ( 1 963). Some of the characters used by Linnavuori
(1959) to establish a new subfamily are present in some of the more
primitive genera of lassinae, e.g., absence of a coronal suture, swollen
clypeus (e.g., Gargaropsis), postfrontal suture present (e.g., Gargaropsis.
Baldriga), and ocelli in anterior margin of head (several genera).
Bythonia is probably most closely related to Pachyopsis and Scaropsia (see
Blocker 1979b). A worldwide reclassification of the higher categories of
lassinae is needed.

Bythonia Oman

Bythonia Oman, 1936:358. Type species. Nionia rugosa Osborn, 1923:32 by monotypy.

Vertex short, slightly shorter medially than next to eye; face short and
broad, ocellocular area with a distinct ledge above antennal pit; post-
frontal suture extending past antenna, curving mesad of ocellus; ocellus
remote from eye, approximately 3X its diameter; forewing with 3 ante-
apical cells; hindwing with second anteapical cell narrow. The original
generic description (Oman 1936) and the subsequent description of
Linnavuori (1959) should be consulted.

^Received May 7, 1990. Accepted June 4, 1990

Contribution 90-508-J from the Kansas Agricultural Experiment Station.
Department of Entomology, Kansas State University, Manhattan, KS 66506.
Department of Entomology. British Museum (Natural History). Cromwell Road.
London SW7 5BD.

ENT. NEWS 101(5): 293-296. November & December. 1990

294 ENTOMOLOGICAL NEWS

Bythonia rugosa (Osborn)
(Fig. 1)

Nionia rugosa Osborn, 1923:32.

Bythonia rugosa (Osborn), Oman 1936:358.

Length of male 7.0 mm, female 8.0 mm; head width of male 2.7 mm,
female 3.0 mm; pronotal width of male 2.6 mm, female 2.9 mm; pronotal
length of male 1.3 mm, female 1.4 mm; vertex length approx 0.2 mm,
slightly longer next to eye in both sexes.

Female abdominal sternum VII (fig. 1) elongate, with a spine on
anterior third of each lateral margin and caliper-like area at each
caudolateral margin, caudal margin dentate. Description and illustra-
tion of male in Linnauuori (1959).

Holotype, female, Sta. Cruz de la Sierra, Bol. (J. Steinbach), Ace. 4549,
in Carnegie Museum. The type and a male specimen from Peru (locality
and collector unknown), in The American Museum of Natural History,
have been studied.

Bythonia kalypso Linnavuori

Bythonia kalypso Linnavuori, 1959:15.

Length of male 9.0 mm. This specimen was described and illustrated
by Linnavuori (1959) from a single specimen from Brazil. The holotype
has not been studied. It is reportedly deposited in the Hungarian Natural
History Museum but could not be located in that collection (Dr. Tamas
Vasarhelyi, personal communication); neither is it present in the
Linnavuori collection at the American Museum of Natural History (Dr.
M.D. Schwartz, personal correspondence).

Bythonia consensa, new species

(Figs. 2-7)

Length of male 7.0 mm; head and pronotal width 2.5 mm; vertex width 0.2 mm next to
eye, slightly shorter medially; pronotal length 1.2 mm; female unknown.

Color dark red with vertex fuscous, face with median fuscous band; pronotum and
scutellum with a fuscous pattern, legs with fuscous markings; forewings uniformly dark
red.

Ocellus 2Vi X its diameter from eye, on anterior margin of vertex; face tumid; preapical
tarsomere of hind leg reduced, without apical setal row. Abdominal sternum VIII (fig. 7)
with conspicuous process on lateral margins.

Pygofer (fig. 2) elongate, with a heavily sclerotized diagonal internal ridge located at
midlength; a bifurcate process at midlength on ventral margin which extends caudad;
apical half heavily setose, especially along ventral margin. Plates (fig. 3) linear, elongate,
with numerous microsetae. Style (fig. 4) elongate, apex caliper-like, with a retrorse spine

Vol. 101, No. 5, November & December. 1990

295

v f r , T ? ?

, f t ' T r T

Figs. 1 -7. Bythonia rugosa: 1 , female sternum VII. ventral view. Bythonia concensa: 2, pygofer.
lateral view; 3, plate, ventral view; 4. style, broad aspect; 5. connective, dorsal view; 6.
aedeagus, lateral view; 7. sternum VIII. ventral view.

296 ENTOMOLOGICAL NEWS

subapically, numerous microsetae on dorsolateral surface; connective (fig. 5) elongate,
thickened anteriorly in lateral view, curving ventrad; aedeagus (fig. 6) with conspicuous
apodeme, with a medial spine at midlength, bifurcate at midlength, arms broadly U-
shaped; shaft elongate, widened at apical 1A, with a pair of lateral spines; apex recurved,
with numerous microspines on caudal surface.

Holotype, male, Tijuco Preto, Esp. Santo [Brazil] deposited in the British Museum
(Natural History). Described from this single specimen.

B. concensa is related to rugosa but can be distinguished by the shape of the apex of the
aedeagal shaft and the shape of the style apex.

ACKNOWLEDGMENTS

The type of Bythonia was lent by the Carnegie Museum courtesy of Chen W. Young;
the male specimen of B. rugosa was lent by the American Museum of Natural History
courtesy of M.D. Schwartz. C.A. Viraktamath, during a visit, examined the types of concensa
and rugosa and offered advice on subfamily placement.

LITERATURE CITED

Blocker, H.D. 1979a. The lassinae (Homoptera: Cicadellidae) of the Western Hemis-
phere. J. Kansas Entomol. Soc. 52:1-70.

Blocker, H.D. 1979b. A proposed phylogeny of New World lassinae (Homoptera:
Cicadellidae). Ann. Entomol. Soc. America 72:857-862.

Kramer, J.P. 1963. A key to the New World genera of lassinae with reviews ofScaroidana
and Pachyopsis (Homoptera: Cicadellidae). Bull. Brooklyn Entomol. Soc. 58:37-50.

Linnavuori, R. 1959. Revision of the Neotropical Deltocephalinae and some related
subfamilies (Homoptera). Ann. Zool. Soc. 'Vanamo' 20:1-370,

Oman, P.W. 1936. A generic revision of American Bythoscopinae and South American
lassinae. Univ. Kansas Sci. Bull. 24:343-420.

Osborn, H. 1923. Neotropical Homoptera of the Carnegie Museum, parts 1 and 2. Ann.
Cam. Mus. 15:8-79.

Vol. 101, No. 5, November & December, 1990 297

NEW RECORDS OF MAYFLIES
(EPHEMEROPTERA) FROM MAINE

Steven K. Burian2, Ronald G. Mack3

ABSTRACT: Five species of mayflies (Ephemeroptera) are reported from Maine for the
first time: Ameletus lineatus, Acerpenna macdunnoughi, Acentrella ampla. Centroptilum
helium, and Leucrocuta aphrodite. The occurrence of the primarily southern species
Acentrella amplua in central and coastal Maine represents the northernmost record for the
species.

Maine is a large geographically diverse area for which the mayfly
fauna is poorly understood. A current review of the historical records for
Maine indicate that only about 100 species were reported (Burian 1990).
This number represents only half the species expected to occur in the
region based on continental distribution records (Edmunds et al. 1976).

To improve our knowledge of the diversity and distribution of mayflies
in Maine an extensive survey was conducted from 1 985-87. In addition to
inland sampling (Burian 1990), the offshore islands of Mount Desert
and Isle au Haut were intensively sampled (Mack 1988). These sampling
programs produced several thousand new specimens for study. The
purpose of this paper is to report the occurrence of five species (Ameletus
lineatus Tr aver, Acerpenna macdunnoughi (\&Q), Acentrella ampla (Traver),
Centroptilus bellum (McDunnough), and Leucrocuta aphrodite
(McDonnough)) for the first time from Maine.

Abbreviations for life stages, locations, and collectors used in species
records are: nymph (N), adult (A), males (M), females (F), at ((a ), Mount
Desert Island (MDI), Acadia National Park (ANP), S.K. Burian (SKB),
K.E. Gibbs (KEG), and R.G. Mack (RGM). Distribution records are
listed separately for nymphs and adults. Records are arranged by county,
and habitat information for sites listed is provided by Mack (1988) for
Mount Desert Island and Burian (1990) for inland areas. Specimens
listed in this paper were deposited in the Aquatic Insect Collection of the
Department of Entomology, University of Maine, Orono, Maine, U.S.A.

^Received February 15. 1990. Accepted June 18, 1990.

"Dept. of Biology, Southern Connecticut State University, 501 Crescent Str.. New Haven,
CT 06515.

-'Maine Dept. of Marine Resources, Mount Desert Island Biological Laboratory. Salisbury
Cove. ME 04672.

ENT. NEWS 101(5): 297-300, November & December, 1WO

298 ENTOMOLOGICAL NEWS

Siphlonuridae
Ameletus lineatus Traver

(N) FRANKLIN: 3rd small brook crossing golf course road. Sugarloaf Ski Area 23-V-85
SKB; HANCOCK: Duck Brk. ft Rt. 233 MDI-ANP 2-V-87 ROM; Great Brk. 1 km south of
outlet to Great Long Pond MDI-ANP 19-V-87 RGM; Stanley Brk. along Park Loop Rd.
MDI-ANP 19-V-87 RGM.

(A) HANCOCK: Stanley Brk. along Park Loop Rd. MDI-ANP 19-V-87 (F) & 23-V-87 (F)
RGM.

In addition to the records noted here for the Moosehead Plateau and
costal Maine, early instar nymphs that are either Ameletus ludens or A.
lineatus were collected from Washington County. Mature nymphs or
adults are necessary to clarify these additional records.

Baetidae
Acerpenna macdunnoughi (Ide)

(N) FRANKLIN: Carrabassett R. east of rest area (« Rt. 16 22-V-86 SKB; Carrabasett
R. ft' confluence with West Branch Carrabassett R.. Kingfield 20-VI-86 SKB;
HANCOCK: Breakneck Brk. ANP boundary. Hulls Cove, MDI 31-111-86 SKB; Duck Brk.
outlet of Eagle Lake ft Rt. 223 MDI-ANP 8-V-86 SKB; Duck Brk. ft Rt. 3 MDI-ANP 2-V-87
RGM; Lurvey Brk. ft Seal Cove Rd. MDI-ANP 13-VII-87 RGM; Great Brk. 100 m south of
the mouth of brook MDI-ANP 19-V-87 RGM; Brook (unidentified) entering Aunt Betty
Pond on east side of carriage path MDI-ANP 25-V-87 RGM; Stanley Brk. along Park Loop
Rd. MDI-ANP 7 & 18-V-87 & 6-VI-87 RGM; PENOBSCOT: Sunkhaze Str. ft Studmill Rd.
bridge, Costigan 9-VI-86, 3-VII-86, & 5-VIII-86 SKB; Baker Brk. upstream & downstream of
culvert on County Rd., Milford 27-VI-86 SKB; Birch Str. upstream & downstream of bridge
on County Rd., Milford 19-V-86, 4 & 5-VIII-86 SKB; PISCATAQUIS: Lazy Tom Str. ft
spillway bridge on Spencer Bay Rd. 24- VI- 86, 28-V-86, 25-VI-86, & 29-VII-86 SKB; Lily Bay
Brk. above confluence with North and South Brooks 26-VIII-86 SKB; Tussel Brk. above
inlet to Moosehead Lake 29-V-86; WALDO: Small stream ft intersection of Back Rd. &
North Palermo Rd., Palermo 14-IV-86 SKB; WASHINGTON: Flood Brk. downstream of
culvert ft Rt. 64-VI-86 & 12-VIII-86 SKB; Small brook ft rest area on East Musquash Lake,
upstream of culvert 5-VI-86 SKB; Tomah Str. upstream & downstream of bridge ft Rt. 6 13-
VIII-86 SKB.

(A) FRANKLIN: Small tributary (unnamed) to Carrabassett R. at base of Sugarloaf Mtn.
27-VI-86(M) SKB; PENOBSCOT: Sunkhaze Str. ft Studmill Rd. bridge, Costigan 1 l-V-86
(F) SKB; PISCATAQUIS: Lazy Tom Str. ft spillway bridge on Spencer Bay Rd. 26-VIII-86
(F) SKB; Lagoon Brk. upstream & downstream of culverts on road to Big Lyford Pond 29-
V-86 (M,F) SKB; Lily Bay Brk. confluence with Moosehead Lake 26-VIII-86 SKB.

This species is widely distributed across the state and is one of the most
frequently encountered baetids in cool rocky streams.

Vol. 101, No. 5. November & December. 1990 299

Acentrella ampla (Traver)

(N) HANCOCK: Duck Brk. (a bridge off of New Eagle Lake Rd. MD1-ANP 1 l-V-86 KEG
&9-V-87 ROM; Stanley Brk. MDI-ANP 6-IX-87 ROM: PENOBSCOT: Souadabscook Str.
(o Rt. 9 29-IV-86 KEG: WASHINGTON: Small brook Co rest area on East Musquash Lake
do Rt. 6 5-VI-86 SKB.

(A) HANCOCK: Duck Brk. (g bridge off of New Eagle Lake Rd. MDI-ANP 19-V-87 (M.F)
RGM.

Primarily a southeastern species, Acentrella ampla is restricted to the
coastal and central southern interior parts of Maine. The species was
most abundant at Duck Brk. on Mount Desert Island where it occurred
in dense Fontinalis mats in swift rock-bottom streams.

Centroptilum bellum McDunnough

(N) FRANKLIN: Reed Brk. above confluence with Carrabassett R. (« Rt. 16 west of
Kingfield 2 l-VIII-86; HANCOCK: Breakneck Brk. Co ANP boundary Hulls Cove. MDI 10-
Vl-86 SKB: Breakneck Brk. (a Rt. 3 MDI-ANP 13-V1-87 RGM: PENOBSCOT: Sunkhaze
Str. Co Studmill Rd. bridge, Costigan 9-VI-86. 3-V1I-86. & 5-VIII-86 SKB; Birch Str.
upstream & downstream of bridge on County Rd., Milford 9-VI-86 & 19-VI-86 SKB: Baker
Brk. upstream & downstream of culvert on County Rd.. Milford 4-VIII-86 SKB:
SOMERSET: Carrabassett R. 2.4 km west of North Anson along Rt. 16 21-VI-86 & 13-VII-
86 SKB: WASHINGTON: Tomash Str. upstream & downstream of bridge Co Rt. 6 l-VIII-86
SKB.

(A) FRANKLIN: Reed Brk. above confluence with Carrabassett R. (o Rt. 16 12-VIII-86
(M.F) SKB; WASHINGTON: Tomah Str. (o bridge on Rt. 6 l-VII-86 SKB.

This species is broadly distributed across the state and frequently occurs
in cool rocky streams and rivers.

Heptageniidae
Leucrocuta aphrodite (McDunnough)

(N) LINCOLN: Sheepscot R. (mainstem) Rt. 126 North Whitefield 15-VIII-86 SKB.

(A) HANCOCK: Duck Brk. Co Rt. 3 MDI-ANP ll-VII-87 (M) RGM; PENOBSCOT:
Sunkhaze Str. Co Studmill Rd. bridge 5-V1II-86 (M) SKB: Birch Str. (o confluence with
Stillwater R. Co Rt. 116 bridge 12- VI-86(M,F) SKB; Lights (» Mobil Gas Station (o Stillwater
Ave. exit of 1-95 7-VII-86(M. F) SKB; Lights Co Maineway Store. Old Town 16-V1I-86 (M.F)
SKB; Lights behind Nutting Hall, University of Maine campus. Orono 1 1 & 30-VI-86 & 4-
V1II-86 (M.F) SKB.

Broadly distributed across eastern Canada (Edmunds et al. 1976).
Leucrocuta aphrodite is restricted in Maine to the central and southern
interior areas. The species probably dispersed into Maine from the south
and west (Burian 1990), and it is expected that additional collections
made in the southwestern parts of the state will uncover other populations.

300 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

Support for this project was provided by the Maine Department of Inland Fisheries and
Wildlife Nongame and Endangered Species program; the Maine Department of
Environmental Protection; K.E. Gibbs through the Maine Agricultural Experiment
Station; the Department of Entomology of the University of Maine; and the United States
Department of the Interior National Parks Service. Much appreciated technical help was
provided by William L. Peters, Jan Peters, R.W. Flowers of the Department of Entomology,
Florida A&M University and David Funk of the Stroud Water Research Center. Special
thanks are extended to all past collectors who contributed specimens to the mayfly
collection and this study.

LITERATURE CITED

Burian, S.K. 1990. The Mayflies (Ephemeroptera) of Maine: A Faunal Survey, Part 1 and
Taxonomy of the Leptophlebiidae.Potamanthidae, Polymitarcyidae, and Ephemeridae,
Part 2. Ph.D. Dissertation, University of Maine, Orono. ME, U.S.A. 409 pp.

Edmunds, G.F., Jr., S.L. Jensen, and L. Berner 1976. The Mayflies of North and Central
America. University of Minnesota Press, Minneapolis, MN. 330 pp.

Mack, R.G. 1988. Faunal Survey and Observed Biology of the Mayflies of Acadia
National Park. M.S. Thesis, University of Maine, Orono, ME, U.S.A. 102 pp.

Vol. 101. No. 5, November & December. 1990 301

SEPARATION OF INSECT AND PLANT MATERIAL
FROM SCREEN-SWEEP SAMPLES1

Gregory Zolnerowich, John M. Heraty, James B. Woolley^

ABSTRACT: An alcohol technique which separates plant debris from insect material in
screen-sweep samples is described. Data show this method can be up to 100% efficient at
separating Chalcidoidea (Insecta: Hymenoptera) from bulk plant material when 95%
ethanol is used. This technique opens up new habitats for collecting, decreases time needed
to sort a sample, and reduces storage space required to house samples.

Screen-sweeping, a technique pioneered by Lubomir Masner,
(personal communication), is an effective method for collecting large
numbers of small insects (e.g., see Noyes 1989), and the use of a screen-
sweep net is the first step in reducing plant debris in a sample. Triangular
net heads are used when collecting to maximize the surface area of the
net in contact with the ground. A V* inch galvanized hardware cloth
screen fitted over the net opening effectively prevents many leaves,
stems, stones, and twigs from entering the net bag (L. Masner, pers.
comm., Noyes 1982, 1988). Polyethylene matting used in place of hard-
ware cloth results in fewer damaged specimens (L. Masner, pers. comm.)
but is less durable under rugged collecting conditions. In the field, the
net bag is periodically emptied into gallon size heavy duty plastic freezer
bags. The sample in the bag is sprayed with a saturated salt solution
(NaCl in H^O) which drowns and preserves small insects. We are con-
cerned primarily with collecting the superfamily Chalcidoidea
(Hymenoptera), parasitic wasps which are usually 1-5 mm long. Of
special interest are Encyrtidae, many of which are found in dense grasses
at ground level. However, even with a screen, sweeping such habitats
quickly results in a ball of plant material inside the net which can
damage delicate specimens. In addition, processing and subsequent
laboratory sorting of such samples can be time-consuming. We describe
here an alcohol flotation technique which further separates plant debris
from insect material in a screen-sweep sample. This technique opens up
new habitats for collection, decreases time needed to sort a sample, and
reduces storage space required to house samples.

APPARATUS

®
The individual parts needed are easily constructed of Rubbermaid

'Received June 4. 1990. Accepted July 13. 1990.

"Department of Entomology. Texas A&M University. College Station. Texas 77843

ENT. NEWS 101(5): 301-306. November & December. 1990

302

ENTOMOLOGICAL NEWS

Servirf Saver^plastic containers. The main units employed by this
method are a fine mesh strainer, a separation chamber, and a transport/
pouring container (Fig. 1).

Fine mesh strainer (Fig. 1 , A). The strainer is comprised of two tall 1 .4
liter containers with their bottoms removed. The internal dimensions of
each container are 12 x 12 x 13 cm. A suitable fine mesh material is
stretched across the bottom of one container, which is then pushed
tightly into the second container to form a taut screen. A small section of
panty hose makes an excellent straining material, as it is strong, has an
intricate weave which prevents even tiny insects from passing through,
and insects or plant material do not cling to it.

Separation chamber (Fig. 1, B). The separation chamber is formed
by a 2.4 and 4.5 liter square container. The bottom is removed from the
smaller container and its sides are shortened so that it fits inside the
larger container. The internal dimensions of the smaller container are 21
x 21 x 7.5 cm. A screen bottom made of 1A inch mesh galvanized hardware
cloth is attached near the bottom of the smaller container. Silicone
rubber aquarium sealer can be used to affix the hardware cloth screen 1
cm above the bottom rim. The internal dimensions of the 4.5 liter con-
tainer are 24.5 x 24.5 x 1 1 cm.

Figure 1. Apparatus used in separation procedure. From left to right are the fine mesh
strainer (A), separation chamber (B), and transport/pouring container (C).

Vol. 101, No. 5, November & December, 1990 303

Transport/pouring container (Fig. 1, C). A tall 5 liter container
measuring 20 x 20 x 22 cm is used for transportation in the field, storage,
and as a receptacle when pouring alcohol.

Other equipment. Washing bottles are needed for rinsing specimens
from the fine mesh strainer. Different sizes of funnels are used to facil-
itate the transfer of insects from the strainer into storage containers and
for filtering dirty alcohol. An alcohol hydrometer and graduated cylinder
are needed to monitor the concentration of alcohol used in the process.

PROCEDURE

Samples should be processed the day of collection to insure maximum
efficiency of separation. The contents of a single freezer bag are trans-
ferred into the fine mesh strainer. This is easily done by cutting a bottom
corner of the bag and pouring the sample into the strainer. If the bag is
very full, the sample should be divided in half for processing. Once in the
strainer, a gentle stream of water should be played over the sample for 3-4
minutes to flush out the brine. The sample is allowed to drain for a few
minutes.

The sample is then transferred to the separation unit and enough 95%
ethanol added to nearly fill the nested containers. Insects sink through
the bottom screen of the inner container into the larger 4.5 liter con-
tainer. Plant material such as flowers, leaves, seeds, and stems floats to
the surface, or sinks and is restrained by the hardware cloth screen. To
insure that the maximum number of insects sinks through the mesh, the
sample is agitated for 4-5 minutes by gently shaking the inner container
and stirring the debris. This breaks up any plant material packed together
on the surface or obstructing the mesh bottom. The inner unit is then
lifted out along with the bulk of the plant material, which can be
discarded.

The alcohol and insects left in the 4.5 liter container are poured
through the fine mesh strainer into the 5 liter container. The strainer
traps the insects in the alcohol, and from there the sample is spooned or
flushed with alcohol into a suitable container and stored in ethanol for
later sorting in the laboratory. The alcohol in the storage containers
should be changed after 24 hours. Insects stored in alcohol keep best if
housed in a freezer (Masner and Goulet 1981).

Alcohol employed in the separation process which remains clean can
be used again. To prevent the deposition of dirt onto specimens, dirty
alcohol can be filtered and then reused for the next sample. A vacuum
filtering flask is most effective for filtering in the laboratory. In the field,
large cone-shaped paper coffee filters inserted into a wide funnel are

304 ENTOMOLOGICAL NEWS

effective at removing dirt from alcohol. Paper coffee filters are thinner
and work faster than standard laboratory filter paper.

FIELD TRIALS

Tests were conducted to determine the effectiveness of this technique.
Screen-sweep samples were collected and subjected to the separation
process. Floating plant portions were retained and examined to ascertain
the number of insects that would potentially be lost with this technique.
To save time, we only sorted to certain categories of Hymenoptera (Tables
1 and 2). Of those categories, "Other Chalcidoidea" includes families
such as Eulophidae, Eurytomidae, Pteromalidae, and Torymidae. "Other
Microhymenoptera" are small non-chalcidoid wasps such as Cynipoidea,
Proctotrupoidea, and Scelionoidea. The first three tests used the same
70% ethanol (Table 1).

Specimens which sank would have been retained while those which
floated with the plant material would have been discarded. In categories
with large sample sizes the percentage of chalcidoid specimens which
floated and would have been discarded ranged from 0.7-5.6 percent
(Table 1).

In additional tests using the same alcohol, the loss rate reached 10-18%
for some categories of Hymenoptera. We hypothesized that the alcohol
used in the separation process was becoming diluted with water and
allowing more insects to float. In addition to any moisture inherent in the
samples, they were being subjected to the brine and a water rinse, all of
which could introduce water into the alcohol. To test this hypothesis, two
of us swept a local grassy meadow for one hour each. This collecting site
is characterized by having a rich chalcidoid fauna in very dense, mature
grasses. The total weight of the samples collected was 1.14 kg. The
samples were combined and divided into six equal portions based on
weight and subjected to the separation process using 50, 70, and 95%
ethanol with two repetitions, each using fresh alcohol. The percentage of
the total number of insects from each category that would have been
discarded is shown in Table 2.

The test using 50% ethanol gave poor results, with losses often
exceeding 20% of the total collected. The test with 70% ethanol reflected
our earlier findings, with chalcidoid losses ranging from about 1-7% for
categories with large sample sizes. The test with 95% ethanol produced
the best results, and insect loss was from 0 - 2.8% for all but one category
of Hymenoptera (Table 2).

Although alcohol used in the separation process will become diluted,
using an alcohol hydrometer to monitor concentration will prevent the

Vol. 101, No. 5, November & December. 1990 305

undue loss of valuable specimens. Based on 2 1 freezer bag samples, each
bag processed decreases alcohol concentration by approximately 4 per-
cent. We discard our alcohol when the concentration reaches 70-75%, the
level where our data show we may begin to lose 5% of the specimens.

Table 1. Results using the same 70%ethanol for consecutive tests in the separation process.
Data are expressed as % (N) where % is the percentage of total insects (N) from each
category which would have been discarded with the plant material. Specimens collected
from Texas, Jim Wells Co.. La Copita Research Station. 20.V.1987.

Test 1 Test 2 Test 3

Aphelinidae &
Trichogrammatidae 0.0(89) 1.8(108) 0.7(409)

Encyrtidae 5.6(125) 4.6(108) 3.7(463)

Eucharitidae 0.0(1) 25.0(4) 18.7(16)

Eupelmidae 0.0(3) 0.0(1) 0.0(7)

Mymaridae 1.3(223) 4.5(111) 1.0(380)

Other Chalcidoidea 5.6(531) 2.3(683) 5.5(2023)

Other Microhymenoptera 14.7(68) 14.0(71) 14.0(410)

Ichneumonoidea 3.3(30) 4.0(94) 8.6(29)

Aculeates 75.0(4) 12.5(8) 24.4(41)

Table 2. Percentage of the total number of insects from each category which would have
been discarded in two repetitions using fresh 50, 70, and 95% ethanol in the separation
process. Data are expressed as % (N) where % is the percentage of total insects (N ) from each
category. Specimens collected from Texas. Brazos Co., Lick Creek Park. 4.VIII.1987.

50% Ethanol 70% Ethanol 95% Ethanol

Rep. 1 Rep. 2 Rep. 1 Rep. 2 Rep. 1 Rep. 2
Aphelinidae &

Trichogrammatidae 26.7(45) 9.3(172) 3.1(159) 2.2(92) 0.0(106) 2.8(143)

Encyrtidae 22.4(85) 6.5(138) 4.6(132) 1.3(76) 0.0(128) 2.6(117)

Eupelmidae 0.0(6) 0.0(6) 7.7(13) 25.0(4) 0.0(6) 0.0(12)

Mymaridae 23.4(77) 5.4(167) 6.6(211) 5.4(110) 0.7(136) 1.4(146)

Other Chalcidoidea 21.5(195) 6.3(319) 2.5(318) 3.2(190) 0.4(260) 0.0(244)

Other Microhymenoptera 41.0(22) 11.9(42) 2.9(34) 22.0(18) 0.0(28) 13.3(15)

Ichneumonoidea 25.0(4) 15.4(13) 18.8(16) 9.1(11) 0.0(14) 0.0(2)

Aculeates 0.0(1) - — 0.0(4) 0.0(5)

306 ENTOMOLOGICAL NEWS

SUMMARY

There are many advantages of this plant separation technique. The
required parts are inexpensive, easy to assemble, and readily trans-
portable in the field. We have employed this technique in campgrounds,
motels, and the laboratory. The effective removal of extraneous plant
material from screen-sweep samples dramatically decreases the time
and space required to sort and house them. Although this technique has
been shown to be effective for small Hymenoptera, it is hoped that other
collectors will utilize it. Using this method, we commonly collect many
Collembola, Microcoryphia, Thysanoptera, Hemiptera, Homoptera,
Coleoptera, Diptera, and Arachnida. The use of this technique may
encourage workers to sample habitats which in the past were
unpalatable.

ACKNOWLEDGMENTS

We thank Lubomir Masner of the Biosystematics Research Centre. Ottawa, for
continued suggestions and inspiration regarding all aspects of collecting. He. Robert
Wharton of TAMU, and two anonymous reviewers kindly critiqued this paper. This paper
is Technical Article No. 25661 from the Texas Agricultural Experiment Station.

LITERATURE CITED

Masner, L. and H. Goulet. 1981. A new model of flight-interception trap for some

hymenopterous insects. Entomological News 92(5): 199-202.
Noyes, J.S. 1982. Collecting and preserving chalcid wasps (Hymenoptera: Chalcidoidea).

Journal of Natural History 16: 315-334.
Noyes, J.S. 1988. Encyrtidae (Insecta: Hymenoptera). Fauna of New Zealand 13. 188

pages.
Noyes, J.S. 1989. A study of five methods of sampling Hymenoptera (Insecta) in a tropical

rainforest, with special reference to the Parasitica. Journal of Natural History 23: 285-

298.

STATEMENT OF OWNERSHIP. MANAGEMENT & CIRCULATION

1. Title of publication: ENTOMOLOGICAL NEWS

2. Date of filing: October 5. 1990

3. Frequency of issue: Bimonthly (every other month) except July and August

4. Location of known office of publication: 232 Oak Shade Rd.. Tabernacle Twp.,
Vincentown PO, New Jersey 08088

5. Location of the headquarters or general business offices of the publishers: 1900 Race St
Philadelphia, PA 19103

6. Name and address of publisher, editor and managing editor:

Publisher: American Entomological Society, 1900 Race St. Philadelphia.
PA, 19103. Editor: Howard P. Boyd, 232 Oak Shade Rd., Tabernacle Twp.,
Vincentown PO, New Jersey, 08088

(Continued on page 315)

Vol. 101, No. 5, November & December. 1990 307

A SURVEY OF THE TIGER BEETLES
(COLEOPTERA: CICINDELIDAE) OF COLORADO1

Michael G. Kippenhan2' 3

ABSTRACT: Thirty-three species of tiger beetles representing two genera are recorded
from Colorado. This richness of species is a combination of the great geomorphological
diversity of Colorado, and the close proximity of Colorado to the area of highest North
American tiger beetle diversity, the southwest. Taxonomic clarifications are given for
several literature records.

The general distributions of North American tiger beetles are rela-
tively well known (Boyd et al. 1982). However, no checklist is presently
available for the Colorado species. Wickham (1902) in his catalogue of
Colorado beetles provides a list of species. However, the taxonomy is
out-of-date and unreliable. The richness of the cicindelid fauna of
Colorado is especially interesting because of two factors: ( 1 ) the diverse
geomorphology of Colorado, with elevations ranging from 1020 m at the
Kansas border to numerous peaks exceeding 4000 m in the Rocky
Mountains, and (2) the close proximity of Colorado to the center of the
highest North American tiger beetle diversity, the southwest (Willis
1972). Colorado is readily divided into three broad physiographic regions:
plains, mountains and the plateaus (Fig. 1 ). The eastern two-fifths of the
state is the western extension of the Great Plains. The southern Rocky
Mountains rise abruptly, with the eastern most range, the Front Range
displaying some of the most striking high-altitude scenery in the world
(Chronic and Chronic 1972). The western quarter of the state is charac-
terized by flat-topped plateaus overlooking steep gorges cut by rivers.

Many species of tiger beetles have distinctive distribution patterns
associated with these land forms in Colorado. For example. Cicindela
denverensis Casey, C. p. pukhra Say. C. s. scutellaris Say and C. o. obsoleta
Say are restricted to plains. Cicindela f. formosa Say and C. scutellaris Say,
both common widespread eastern and central North American species
reach their western limit of distribution in Colorado along the Front
Range, except for isolated populations of C/ gibsoni Brown and C. .v.
yampae Rumpp in Moffat County (Fig. 1). Cicindela duodecimguttata
Dejean also reaches its western limit of distribution in Colorado, whereas
C. o. guttifera LeConte reaches its eastern most range limit in the Front
Range. North-south range limits of several species occur in Colorado.
Cicindela 1. limbata Say reaches its most southern limit and C. o. obsoleta

'Received March 2. 1990. Accepted May 14, 1990.

-Colorado State University, Department of Entomology, Fort Collins. Colorado 80523
Present Address: 15185 Deby [Drive. Colorado Springs. Colorado X0921

ENT. NEWS 101(5): 307-315. November & December. 1990

308 ENTOMOLOGICAL NEWS

Say, C. o. santaclarae Bates, C. marutha Dow, C. n. nigrocoerulea LeConte,
and C. n. bowditchi Leng reach their northern limit in Colorado. Cicindela
theatina Rotger, orginally described from the Great Sand Dunes, Alamosa
County, is only known from a small area of southern Colorado.

The following preliminary checklist follows the classification of Boyd
et al. (1982). Generally, Willis' (1968) (adaptation of Rivalier, 1954)
taxonomic treatment of the species of Cicindela was followed. The
subfamily Cicindelinae is represented in Colorado by 2 genera and 33
species. The number of species reported herein is less than New Mexico
(38) but higher than other surrounding states of Kansas (30), Wyoming
(28) and Utah (24) (Boyd et al. (1982)). Boyd et al. (1982) listed
Megacephala virginica (L.) from Colorado, but this record is considered
here doubtful (R.L. Huber, personal communication), and is not included.
This species was listed for Oklahoma near the Colorado border by Drew
and Van Cleave (1961). Three additional subspecies records (C. n.
nigrocoerulea LeConte, C. o. santaclarae Bates and C. p. cinctipennis
LeConte), not originally listed by Boyd et al. (1982) from Colorado are
included and clarified for the first time. The following species have been
collected in surrounding states and may eventually be collected in
Colorado: C. cursitans LeConte (Kansas and Nebraska), C. celeripes
LeConte (Kansas, Nebraska, and Oklahoma), C. sedecimpunctata Klug
(Arizona and New Mexico), C. sperata LeConte (Arizona, New Mexico,
Oklahoma, and Utah), C. tenuisignata LeConte (Arizona, New Mexico,
Oklahoma, and Utah), and C. willistoni LeConte (Kansas, New Mexico,
Oklahoma, Utah, and Wyoming). County records (numbers) following
each species refer to Fig. 1.

Tiger Beetles of Colorado

Subfamily Cicindelinae Fisher

Tribe Megacephalini W. Horn

Subtribe Omina W. Horn

Genus Amblychelia Say

A. cylindriformis(Say)

This species is known only from the plains physiographic region of
Colorado. Counties: 6, 15, 32, 38, 40, 59, 60, 61, 62, 63
A. picolominii Reiche

No specimens of this species were examined during this study. Vaurie
(1955) listed a single specimen from Montezuma County. County 50

Vol. 101, No. 5, November & December. 1990 309

Tribe Cicindelini Sloane
Subtribe Cicindelina W. Horn

Genus Cicindela L.
Subgenus Cicindela(s. s.)

C. longilabris laurentii Schaupp

The geographic variation of this species was documented by Spanton
(1988). Counties: 1, 3, 5, 18, 21, 26, 28, 42, 50
C. nebraskana Casey

Spanton (1988) did not cite specimens from Colorado, but recent
collecting has established its presence well into Colorado. Counties:
4,32
C. repanda repanda Dejean

This fluvial species can be usually collected wherever suitable habitats
occur throughout the state. Counties: 1,4,5,7, 11, 14, 16,20,24,25,30,32,
38,48,50,58,59,61,62
C. duodecimguttata Dejean

This fluvial species reaches its western range limit along the Front
Range. Counties: 5, 11,41
C. oregona LeConte

It is expected that this species inhabits all of the western half of the
state. The subspecies C. o. oregona LeConte, C. o.guttifera LeConte, and C.
o. navajoensis Van Dyke were listed for Colorado by Boyd et al. (1982),
however, C. o. oregona does not occur in Colorado (Freitag 1965).
C. oregona guttifera LeConte

The subspecies ranges from the Front Range to the Western Slope.
Counties: 1,5, 16,17,20,22,23,24,27,29,30,31,32,33,41,43,46,4748,51.

52, 53, 54, 55, 56, 59

C. o. guttifera LeConte X C. o. navajoensis Van Dyke

Specimens of this hybrid form (sensu Freitag 1965) have been collected
in Mesa County, especially in Colorado National Monument. No speci-
mens clearly assignable to C. o. navajoensis were examined from Colorado.
Interestingly, Freitag (1965) also did not list any material of this form
from Colorado, despite indicating that Colorado is included in the
geographical range of this subspecies (his Fig. 18). County 24
C. hirticollis shelfordi Graves

This recently described subspecies occurs throughout Colorado except
the extreme western edge (Graves etal. 1988). Counties: 5, 6. 9. 14, 15,38,
61
C. hirticollis corpuscula Rumpp

Twelve specimens from Mesa County, Colorado River, south of Loma

310 ENTOMOLOGICAL NEWS

in the University of Colorado collection are assignable to this sub-
species. Rumpp (1961) did not examine any Colorado material. County
24
C. limbata limbata Say

This species reaches it southernmost limit in northeastern corner of
Colorado. Counties: 9, 10, 11
C. theatina Rotger

The type locality of this species is the Great Sand Dunes National
Monument, Alamosa County, and is restricted to these sand dunes and
adjacent areas. Counties: 43, 55, 57
C. formosa formosa Say

This species reaches its western limit in Colorado along the Front
Range, and can be expected to occupy most suitable habitats of the
plains physiographic region of Colorado. Counties: 5, 6, 7, 10, 1 1, 14, 17,
33,36,37,38,41,55,61,62
C. formosa gibsoni Brown

This is the only population of C. formosa west of the Front Range and
only known from Moffat Co. Wallis (1961) listed the only other popu-
lations of C.f. gibsoni Brown from Saskatchewan, Canada. The Colorado
population has been recognized as another distinct subspecies by
Gaumer (1977) in an unpublished thesis. County 1
C. purpurea group

Various workers disagree on the exact taxonomic status of
C. p. audubonni LeConte, C. p. cimarrona LeConte, C. s. splendida Hentz,
C. denverensis Casey, C. /. limbalis and other forms in the purpurea group.
Until a comprehensive revision treating all these taxa becomes avail-
able, Boyd et al (1982) will be followed.
C. purpurea audubonii LeConte

The complete range of this common subspecies in Colorado is
unknown. Counties: 5, 6, 9, 14, 16, 17, 20, 32, 33, 56
C. purpurea cimarrona LeConte

Counties: 16,41,48,52,59
C. splendida splendida Hentz

The plains of Colorado is the western limit of this species, and is
apparently uncommon in Colorado. Counties: 6, 7
C. denverensis Casey

The type locality of this species is Denver (Denver County) (Casey
1897). Counties: 5, 6, 7, 13, 39, 60
C. limbalis limbalis Klug

Counties: 3, 5, 17,21,32,61
C. decemnotata Say

This species may be restricted to the western portion of Colorado.
Counties: 1, 51

Vol. 101. No. 5. November & December. 1990 31

C. pulchra pulchra Say

This grassland species inhabits most or all of the plains physiographic
region of Colorado, and is more common in southeastern Colorado.
Counties: 5, 6, 14, 32, 40, 41, 52, 58, 60, 61, 62
C. fulgida fulgida Say

Counties: 5, 6, 8, 9, 14, 21, 32, 39, 41, 56, 58, 60, 61, 62
C. scutellaris scutellaris Say

This grassland species reaches its western limit of its range in Colorado.
Counties: 6, 7, 8, 9, 10, 12, 32, 40, 41, 58, 61, 62
C. scutellaris yampae Rumpp

The type locality of this apparently endemic subspecies is Moffat
County (Rumpp 1986). County 1
C. lengi W. Horn

The three subspecies (C. I. lengi W. Horn, C. I. versuta Casey, and C. I.
jordai Rotger) were listed for Colorado by Eoydetal. ( 1982). However, C. /.
jordai does not occur in Colorado (R.L. Huber. personal communi-
cation). The exact status of C. /. lengi and C /. versuta in Colorado is
unclear. Some populations appear to have both forms represented.
Therefore, localities will not be separated except for C. I. versuta.
C. lengi W. Horn

Counties: 4, 5, 6, 7, 14, 32, 33, 34, 58, 59 ,61
C. lengi versuta Casey

The following are literature records only (Willis and Stamatov 1971;
Lawton 1972), and no specimens were examined. Counties: 1, 55
C. tranquebarica Herbst

Colorado has two subspecies, the eastern C. t. tranquebarica Herbst
and western C. t. kirbyi auct. These subspecies were not separated in this
study because of apparent hybrids and incomplete distribution records
throughout the range of these forms in Colorado. Counties: 1,5,6, 7, 9. 14,
17,22,24,32,36,48,55,58,62

Subgenus Cicindelidia Rivalier

C. nigrocoerulea nigrocoerulea LeConte

This subspecies was previously reported from adjacent states of New
Mexico and Kansas (Boyd etal. 1982), but not Colorado. However, Leng
(1902) and W. Horn (1930) listed Colorado as part of this species dis-
tribution. Willis and Stamatov ( 197 1 ) reported this species from Boulder
County, which is apparently the most northern record of this species.
Counties: 17,32,40,58,60,62
C. nigrocoerulea bowditchi Leng

The type locality of this subspecies is near Durango (La Plata County)
(Leng 1902). Relatively unmarked individuals occur in populations of

312 ENTOMOLOGICAL NEWS

this subspecies form. Counties: 50, 51, 52
C. obsoleta obsoleta Say

Counties: 6, 14, 32, 39, 40, 58, 59, 61, 63
C. obsoleta santaclarae Bates

Boyd et al. (1982) did not list this form from Colorado, however, Leng
(1920) (as C. o. anita Dow) and W. Horn (1930) recorded specimens from
Colorado. Specimens of this subspecies were only examined from counties
bordering New Mexico. Counties: 51, 52
C. punctulata punctulata Olivier

Probably the most common and widespread Colorado tiger beetle.
Numerous large series of individuals were examined from light trap
collections, especially from the plains. Counties: 1 , 5, 6, 8, 1 1, 1 2, 17, 23, 24,
29, 32, 36, 37, 38, 39, 40,41, 42, 47, 49, 50, 52, 55, 58, 59, 61, 62, 63
C. haemorrhagica haemorrhagica LeConte

Boyd et al. (1982) listed this species for Colorado, but no specimens
were examined during this study.

Subgenus Habroscelimorpha Dokhtourow

C. circumpicta johnsoni Fitch
Counties: 60, 61,62

Subgenus Eunota Rivalier

C. togata globicollis Casey

Colorado is the western most range extension for the form. Counties:
29, 60, 61

Subgenus Cylindera Westwood

C. pusilla pusilla Say

Boyd et al. (1982) listed this species as C. terricola Say. A revision of the
C. pusilla group is needed to clarify the proper usage of these names. No
specimens of C. p. pusilla were examined during this study.
C. pusilla cinctipennis LeConte

Boyd etal. (1982) did not include this subspecies from Colorado. Leng
(1902) listed Colorado as part of the range of this form, but in 1920 did not
include it. Counties: 1, 5, 23, 24, 32, 50, 52, 54

Subgenus Ellipsoptera Dokhtourow

C. nevadica knausi Leng
Counties: 29, 38, 58, 60, 61,62

Vol. 101, No. 5, November & December, 199()

313

'at
u

.c
c.

•5

314 ENTOMOLOGICAL NEWS

C. cuprascens LeConte

Counties: 5, 1 1, 15,38, 39, 40, 59, 60, 61, 62
C. macra macra LeConte

Willis (1967) provided records for this species. Apparently Colorado is
the western limit of its range. Counties: 5, 15
C. marutha Dow

Counties: 24, 39, 61,62
C. lepida Dejean

Counties: 6, 60, 61, 62

ACKNOWLEDGMENTS

I would like to thank B.C. Kondratieff, Colorado State University for providing facilities
and specimens during this study. Michael Weissmann, University of Colorado; Richard S.
Peigler, Denver Museum of Natural History also made specimens available for study.
Howard P. Boyd, Tabernacle, New Jersey; and Ronald L. Huber, Prairie Village, Kansas for
helping identify material. B.C. Kondratieff and R.L. Huber provided helpful comments on
earlier drafts of the manuscript. The following persons contributed additional material for
study: Timothy A. Ebert. Paul A. Opler, Judith L. Welch, and especially Howard E. Evans.
Two reviewers provided helpful suggestions which improved the manuscript.

LITERATURE CITED

Boyd, H.P. and Associates. 1982. Checklist of the Cicindelidae. The tiger beetles. Plexus

Publ. Co., Marlton, New Jersey. 31 pp.

Casey, T.L. 1987. Coleopterological notices, VII. Ann. New York Acad. Sci. 9: 285-684.
Chronic, J. and H. Chronic. 1972. Prairie, peak and plateau. A guide to the geology of

Colorado. Colorado Geol. Surv. Bull. 32. 126 pp.
Drew, W.A. and H.W. Van Cleave. 1961. The tiger beetles of Oklahoma (Cicindelidae).

Proc. Oklahoma Acad. Sci. 41: 101-122.
Freitag, R. 1965. A revision of the North American species of the Cicindelu maritime! group

with a study of hybridization between Cicindela duodecimguttata and oregona. Quest.

Entomol. 1: 87-170.
Gaumer, G.C. 1977. The variation and taxonomy of Cicindela formosa Say (Coleoptera:

Cicindelidae). Ph.D dissertation, Texas A & M Univ. 253 pp.
Graves, R.C., M.E. Krejci, and A. C. F. Graves. 1988. Geographic variation in the North

American tiger beetle, Cicindela hirticollis Say, with a description of five new subspecies

(Coleoptera: Cicindelidae). Can. Entomol. 120: 647-678.
Horn, W. 1930. Notes on the races ofOmus californicus and a list of the Cicindelidae of

America north of Mexico (Coleoptera). Trans. Amer. Entomol. Soc. 56: 73-86.
Lawton, J.K. 1972. Collecting notes on the tiger beetles in the southwestern and

southcentral United States. Cicindela 4(2): 35-48.
Leng, C.W. 1902. Revision of the Cicindelidae of Boreal America. Trans. Amer. Entomol.

Soc. 28: 93-186.
Leng. C.W. 1920. Catalogue of the Coleoptera of America North of Mexico, compiled by

John D. Sherman Jr., Mount Vernon, New York. 470 pp.
Rivalier, E. 1954. Demembrement du genre Cicindela Linn. II. Faune americaine. Rev. Fr.

Entomol. 21:249-268.

Vol. 101, No. 5, November & December, 1990

315

Rumpp, N.L, 1961. Three new tiger beetles of the genus Cicindela from southwestern

United States (Cicindelidae-Coleoptera). Bull. S. Calif. Acad. Sci. 60: 165-187.
Rumpp, N.L. 1 986. Two new tiger beetles of the genus Cicindela from western United States

(Cicindelidae: Coleoptera). Bull. S. Calif. Acad. Sci. 85(3): 139-151.
Spanton, T.G. 1988. The Cicindela sylvatica group: geographic variation and classification

of the Nearctic taxa, and reconstructed phylogeny and geographical history of the

species (Coleoptera: Cicindelidae). QuaesL Entomol. 24: 51-161.
Vaurie, P. 1955. A review of the North American genus Amblycheila (Coleoptera:

Cicindelidae). Amer. Mus. Nov. 1724: 1-16.

Wallis, J.B. 1961. The Cicindelidae of Canada. Univ. Toronto Press. 74 pp.
Wickham, H.F. 1902. A Catalogue of the Coleoptera of Colorado. Univ. Iowa Bull. Nat.

Lab. 5:217-310.
Willis, H.L, 1967. Bionomics and zoogeography of tiger beetles of saline habitats in the

central United States (Coleoptera: Cicindelidae). Univ. Kansas Sci. Bull. 47: 145-313.
Willis, H.L. 1968. Artificial key to the species of Cicindela of North America North of

Mexico (Coleoptera: Cicindelidae). J. Kansas Entomol. Soc. 41: 303-317.
Willis, H.L. 1972. Species density of North America Cicindela. Cicindela 4(2): 29-34.
Willis, H.L. and J. Stamatov. 1 97 1 . Collecting Cicindelidae in the Northwest Cicindela 3

(3): 41-51.

(Continued from page 306)

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316

ENTOMOLOGICAL NEWS

INDEX: Volume 101

Acalyptrate Diptera reared from 117

higher fungi in northeastern Ohio

Acanthochalcis, redescrip. of males, 75
biol. notes. & iden. of American

Acari 276

A.E.S. meeting reports 12. 122. 153. 160

Allen, R.T.

Eurypauropus spinosus from Arkan- 95
sas & key to No. Amer. sp. of
Eurypauropus

Amrine, J.W., Jr. 276

Anisotoma basilis breeding in spo- 129
rocarps of slime mold, Lycogala
flavofuscum

Announcements 80, 87, 128, 140, 256

Anoplura 225

Anthopotamus, new genus for No. 200
Amer. sp. previously known as
Potamanthus

Ascalaphidae 158

Ascalobyas, first U.S. record & 158

range extension from northeastern
Mexico to Texas

Bae, Y.J. 200

Behningiidae 219

Berzark, L.G. 69

Biogeographic affinities of Eph- 193

emeroptera of Black Hills, S. Dakota

Blocker, H.D.
New lassinae from Mexico & Peru 207

Blocker, H.D., M.D. Webb
Leafhopper genus Bvthonia 293

Book reviews 1 36, 1 57, 1 69, 265

Books rec'd. & briefly noted 1 16, 256

Bowles, D.E., K. Stephan, M.L. Mathis
New method for collect, adult 222

phryganeid caddisflies

Brown, J.W.

Taxonomicdistrib.&phylogenetic 109
significance of male foreleg hair-
pencil in Tortricidae

Brown, J.W.

Additions to Papilionoidea of 167

Revillagigedo Is., Mexico

Brushwein, J.R. 23

Bunyard, B., B.A. Foote
Acalyptrate Diptera reared from 117
higher fungi in northeastern Ohio

Biol. notes on Drosophila gutti- 161

fera. a consumer of mushrooms

Buprestidae 213

Burian, S.K., R.G. Mack

New records of mayflies from 297

Maine

Bythonia, leafhopper genus 293

Caddisflies from West Virginia, 236

a checklist

Carabidae 211,273

Cassidinae, marking in field for 216

population dynamics studies

Chalcididae 75

Chionaspis salicisnigrae, black 288

willow scale, develop, of in Tenn.

Christiansen, B. 48

Chrysomelidae 216

Cicadellidae 20, 93, 207, 293

Cicindelidae 307

Coccinellidae 164

Coleoptera 98, 129, 133, 137, 147, 164,
211,213,216,273,307

Coreidae 203

Crickets, relation between hearing 29
& flying in

Culex pipiens, drinking as a pre- 257

oviposition behavior of

Culicidae 257

Curtara, n.sp. from Bermuda 20

Darling, D.C., R.C. Plowright

HPLABEL: a program & micro- 143
font for generation of date/locality
labels using a laser printer

Dettopsomyia, n.sp. from Kenya 246

Deyrup, M., D. Manley
Seasonal flight activity of male 99

velvet ants in south Florida

Diaspididae 288

Dietz, A. 123

Vol. 101. No. 5, November & December. 1990

317

Dilaridae
Dindal, D.L.
Diprionidae
Diptera

155

170

266

69. 117. 161.246.257

Dolania americana, sand-burrowing 219
mayfly, new distrib. record for

Drecktrah, H.G.

Larval & pupal descrip's. of 1

Marilia fusca

Dromius agilis. senior synonym of 273

Lebia morio
Drosophila guttifera, a consumer 161

of mushrooms, biol. notes on

Drosophilidae 161

Dufourea versatilis, coll. of, from 67

Idaho

Dunbar. C.S., M.R. Wagner

Distrib. of ponderosa pine saw- 266
flies in U.S. & Canada

Durden, LA. 39

Durden. L.A., B.V. Peterson, N. Wilson,
B. Christiansen

Some ectoparasites of bats from 48
Seram Is., Indonesia

Durden, L.A.. R. Traub. K.C. Emerson
Sucking lice from Pakistan 225

mammals, with notes on zoogeography

Dyscinetus morator feeding on roots 98
of azaleas

Ectoparasites of bats from Halma- 39
hera Is., Indonesia

Ectoparasites of bats from Seram 48

Is., Indonesia

Eischen. F.A.. A. Dietz

Improved culture techniques for 123
mass rearing Galleria mellonella

Elmidae 147

Emerson, K.C. 225

Ephemeroptera 193,200.219.297

Eriophyidae 276

Eurypauropodidae 95

Eurypauropus spinosus from Arkansas 95
& key to No. Amer. sp. of Eurypauropus

Fee, F.D.

Flint, O.S., Jr., P.A. Harp

Lepidostoma (Nosopus) ozarkense. XI
n.sp. from Arkansas

Foote. B.A. 117.161

Freytag, P.H.

New sp. ofCurtara from Bermuda 20

New sp. Zonana from Venezuela 93

Galleria mellonella. improved cul- 123
ture techniques for mass rearing

Garcia. M.A.. L.M. Paleari

Marking Cassidinae larvae in field 216
for population dynamics studies

Glover. J.B., D.C. Tarter

Leptoceridae of West Virginia

Gryllidae 29

Hairpencil, male foreleg, in Tor- 109

tricinae

Halictidae 67

Halstead, J.A.

Redescrip. of males, biol. notes, 75
& iden. of Amer. Acanthochalcis

Harp, P.A.

Harris, S.C., P.K. Lago
Annotated checklist of Rhyacophil- 57
oidea & Integripalpia of Alabama

Hemiptera 203

Heraty, J.M. 301

Hermann, S.J.

New record & range extension for 141
Rhvacophila wallowa from Rocky Mt.
Nat. Pk., Colorado

Hoebeke, E.R. 129

Antennal anomaly in Oxypoda 133
opaca from New York

Hoffmann. K.M.

Distrib. notes on No. & Cent. 155

Amer. Dilaridae

Hoffmann, K.M., J.R. Brushwein
Spider taxa assoc'd. with immature 23
stages of Mantispa interrupta

Homoptera 20. 93. 207. 288. 293

HPLABEL: a program &microfont for 143
generation of date/locality labels
using a laser printer

318

ENTOMOLOGICAL NEWS

Hydroptilidae
Hymenoptera

88
9,13,67,75,99.154,266

lassinae. new from Mexico & Peru 207

Immigrant synanthropic flower flies 69
(two) new to No. Amer.

Integripalpia & Rhyacophiloidea, an- 57
notated checklist of, in Alabama

Invertebrate populations in nests 170
of a screech owl & an Amer. kestrel
in central New York

Jacobs, W.L.

New distrib. record for sand- 219

burrowing mayfly, Dolania americana

Kassar, A, J.W. Amrine, Jr.
Rearing & develop, of Phyllocoptes 276
fructiphilus

Keirans, J.E. 39

Kippenhan, M.G.
Survey of tiger beetles of Colo-
rado

307

Lago, P.K.

Lambdin, P.L.

Develop, of black willow scale, 288
Chionaspis salicisnigrae in Tenn.

Lariviere, M.-C. 211

Larochelle, A, M.-C. Lariviere
Notiophilus palustris, a European 211
carabid beetle new to No. Amer.

Larval & pupal descrip's. of Marilia 1
fusca

Lebia morio, neotype designation for, 273
fixing the name as a junior synonym
of Dromius agilis

Leiodidae 129

Lepidoptera 109, 123, 167

Lepidostoma (Nosopus) ozarkense, 8 1

n.sp. from Arkansas

Lepidostomatidae 81

Leptoceridae 35

Leptoceridae of West Virginia 35

Leptoglossus zonatus, cytology of 203

Liebherr. J.K.

Neotype designation for Lebia 273

morio, fixing the name as a junior
synonym of Dromius agilis

Mack, R.G. 297

Mailing dates 320

Manley, D. 99

Manley, G.V.
A new Mastogenius from Jamaica 213

Mantispa intemipta, spider taxa 23

assoc'd. with immature stages of

Mantispidae 23

Marilia fusca, larval & pupal des- 1

crip's, of

Mastogenius, a n.sp. from Jamaica 213

Mathis. M.L. 222

Mayflies, new records from Maine 297

McCafferty, W.P.

Biogeographic affinities of 193

Ephemeroptera of Black Hills. So.
Dakota

McCafferty, W.P., Y.J. Bae

Anthopotamus, new genus for No. 200
Amer. sp. previously known as
Potamanthus

McDearman, W. 13

Menke, AS.

Status of Pison doggonum 154

Microcylloepusformicoideus, a new 147
riffle beetle from Death Valley Nat.
Mon.. Calif.

Mutillidae 99

Neotrichia, n.sp. from Colorado 88

Neuroptera 23, 155, 158

Notiophilus palustris, a European 211

carabid beetle new to No. Amer.

1
29

Odontoceridae
Orthoptera

Otte, D.

Relation between hearing & flying 29
in crickets

Oxypodaopaca, antenna} anomaly in, 133
from New York

Vol. 101. No. 5, November & December, 1990

319

Packauskas, RJ
Cytology of Leptoglossus zonatus 203

Paleari, L.M 216

Papilionoidea 167

Pauropoda 95

Peterson. B.V. 48

Peterson, B.V, L.A. Durden. J.E.
Keirans, P.M. Taylor
Some ectoparasites of bats from 39
Halmahera Is., Indonesia

Philips, JJL, D.L. Dindal
Invertebrate populations in nests 1 70
of a screech owl & an Amer. kestrel
in cent New York

Phryganeid caddisflies, new method 222
for collect adults

Phryganeidae 222

Phyllocoptesfructiphilus, rearing 276

& develop, of
Pison doggonum, status of 154

Plecoptera 283

Plowright R-C. 143

Potamanthidae 200

Propylea quatourdecimpunctata: 164

add'l. U.S. records of an adventive
lady beetle

Psephenidae 137

Psephenops, n.sp. & new record of 137
genus from Costa Rica

Pteronarcella badia. obs. on role 283

of sexual selection in

Pteronarcyiidae 283

Publisher's statement 306

Pyralidae 123

Riffle beetle, Microcylloepus formi- 147
coideus. new from Death Valley NaL
Mon., Cali£

Ruiter, D.E.

N-Sp. ofNeotrichia from Colorado, 88
with add's. & corrections to distrib.
& records of Colorado Trichoptera

Scarabaeidae 98

Separation of insect & plant mater- 301
ial from screen-sweep samples

Shepard, W.D.

Microcylloepusformkoideus, a new 147
riffle beetle from Death Valley NaL
Mon., Calif.

Smith, D.R.
A new Xyela from western U.S. 9

Smith. RTX W. McDearman

A new Rhadinoceraea feeding on 1 3
Zigadenus in southeastern U.S.

Society meeting reports 12, 122, 153, 160

Spangler. PJ.

New sp. & new record of water- 137
penny genus Psephenops from Costa Rica

Sphecidae 154

Spider taxa assoc'd. with immature 23
stages of Mantis pa interrupta

Staines, C.L.. Jr.

Dvscinetus morator feeding on 98

roots of azaleas

Staphylinidae 133

Stephan. K. 222

Synanthropic flower flies, two 69

immigrant, new to No. Amer.

Syrphidae 69

Relation between hearing & flying 29
in crickets

Rhadinoceraea. n.sp. feeding on 13

Zigadenus in southeastern U.S.

Rhyacophila wallowa, new record & 141
range extension for. from Rocky Mt.
Nat. Pk.. Colorado

Rhyacophilidae 141

Rhyacophiloidea & Integripalpia, 57

checklist of, in Alabama

Takada, H., R.C. Woodruff. J.N.
Thompson

Coll. of Drosophilidae in Kenya. 246
with descrip. of a n.sp. of Dettop-
somvia

Tarter. D.C.

Checklist of caddisflies from 236

West Virginia

Taylor, P.M. 39

Tenthrcdinidae 13

320

ENTOMOLOGICAL NEWS

Thompson, F.C., F.D. Fee, L.G. Ber-
zark

Two immigrant synanthropic flower 69
flies new to No. Amer.

Thompson, J.N. 246

Tiger beetles of Colorado, a survey 307

Tipping, C. 257

Tortricidae 109

Traub, R. 225

Trichoptera 1, 35. 57, 81, 88, 141,

222, 236

Velvet ants, seasonal flight acti- 99

viry of males in south Florida

Vogtsberger, R.C.

First U.S. record of Ascalobyas, 158

a range extension from northeastern
Mexico to Texas

Wagner, M.R.

Walsh, S.L.

Coll. of Dufourea versatilis
from Idaho

Webb, M.D.

266

293

Weber, R.G., C. Tipping

Drinking as a pre-oviposition 257

behavior of wild Culex pipiens

Wheeler, AG., Jr.

Propylea quatourdecimpunctata: 164

add'l. U.S. records of an adventive
lady beetle

Wheeler, Q.D., E.R. Hoebeke

New host record for Anisotoma 129

basilis breeding in sporocarps of
slime mold, Lycogala flavofuscum

Wilson, N. 48

Woodruff, R.C. 246

Woolley, J.B. 301

Xyela, new from western U.S. 9

Xyelidae 9

Ziegler, D.D.

Obs. on role of sexual selection 283
in stonefly Pteronarcella badia

Zolnerowich, G., J.M. Heraty, J.B.
Woolley

Separation of insect & plant 301

material from screen-sweep samples

Zonana, n.sp. from Venezuela 93

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