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ISSN 0033 2615

PSYCHE

A JOURNAL OF ENTOMOLOGY

founded in 1874 by the Cambridge Entomological Club

Vol. 93 1986 Nos. 1-2

CONTENTS

Functional queens in the Australian greenhead ant, Rhytidoponera metallica

(Hymenoptera: Formicidae). Philip S. Ward 1

South American and Floridian disjuncts in the Sonoran genus Compsocryptus

(Hymenoptera: Ichneumonidae). Charles C. Porter 13

The orb-weaver genus Witica (Araneae: Araneidae). Herbert W. Levi 35

An eyeless subterranean beetle ( Pseudanophthalmus ) from a Kentucky coal

mine (Coleoptera:Carabidae:Trechinae). Thomas C. Barr, Jr 47

Biconus in Peru, with notice of an endemic species from the coastal desert

(Hymenoptera: Ichneumonidae). Charles C. Porter 51

A synonymic generic checklist of the Eumeninae (Hymenoptera: Vespidae).

James M. Carpenter 61

Review of the fossil Tiphiidae, with description of a new species ( Hymenoptera).

A. P. Rasnitsyn 91

An early record of tandem running in leptothoracine ants: Gottfrid Adlerz,

1896. Robin J. Stuart 103

Notes on the behavior of the dimorphic ant, Oligomyrmex overbecki

(Hymenoptera: Formicidae). Mark W. Moffett 107

Pupation in mycetophilid flies: a correction. William G. Eberhard 117

New Pselaphidae from New Hampshire (Coleoptera). Donald S.

Chandler 121

A presumptive pheromone-emitting structure in wolf spiders (Araneae:

Lycosidae). Torbjorn Kronstedt 127

A new arboricolous Thyreodon from Costa Rica (Hymenoptera: Ichneu-
monidae: Ophioninae). Charles C. Porter 133

Distinguishing the jumping spiders Eris militaris and Eris flava in North

America (Araneae: Salticidae). Wayne Maddison 141

Evidence of workers serving as queens in the genus Diacamma (Hymenoptera:

Formicidae). Mark W. Moffett 151

New species and genera of Amiseginae from Asia (Chrysididae, Hymenoptera).

Lynn Siri Kimsey 153

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1985-1986

President

Vice-President

Secretary

Treasurer

Executive Committee

Norman C. Carlin
James M. Carpenter
Kathryn Hoy
Frank M. Carpenter
Wayne Maddison
Heather Hermann

EDITORIAL BOARD OF PSYCHE

F. M. Carpenter, (Editor), Fisher Professor of Natural History,
Emeritus, Harvard University

W. L. Brown, Jr., Professor of Entomology, Cornell University and
Associate in Entomology, Museum of Comparative Zoology
B. K. HOLLDOBLER, Professor of Biology, Harvard University
H. W. Levi, Alexander Agassiz Professor of Zoology, Harvard University
M. D. BOWERS, Assistant Professor of Biology, Harvard University
ALFRED F. Newton, Jr., Curatorial Associate in Entomology, Harvard
University

E. O. WILSON, Baird Professor of Science, Harvard University

J. M. CARPENTER, Assistant Professor of Biology, Harvard University

PSYCHE is published quarterly by the Cambridge Entomological Club, the issues
appearing in March, June, September and December. Subscription price, per year,
payable in advance: $15.00, domestic and foreign. Single copies, $4.00

Checks and remittances should be addressed to Treasurer, Cambridge
Entomological Club, 16 Divinity Avenue, Cambridge, Mass. 02138.

Orders for missing numbers, notices of change of address, etc., should be sent to the
Editorial Office of Psyche, 16 Divinity Avenue, Cambridge, Mass. 02138. For
previous volumes, see notice on inside back cover.

IMPORTANT NOTICE TO CONTRIBUTORS
Manuscripts intended for publication should be addressed to Professor F. M.
Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. 02138.

Authors are required to bear part of the printing costs, at the rate of $29.00 per
printed page. The actual cost of preparing cuts for all illustrations must be borne by
contributors: the cost for full page plates from line drawings is ordinarily $10.00
each, and for full page half-tones, $12.00 each; smaller sizes in proportion. There is
ordinarily no additional charge for setting tables of less than six columns; for tables
of six or more columns the cost is $25.00 per page.

Psyche, vol. 92, no. 4, for 1985, was mailed April 27, 1986

The Lexington Press. Inc.. Lexington. Massachusetts

PSYCHE

Vol. 93

1986

Nos. 1-2

FUNCTIONAL QUEENS IN THE
AUSTRALIAN GREENHEAD ANT,
RHYTIDOPONERA METALLICA
(HYMENOPTERA: FORMICIDAE)*

By Philip S. Ward

Department of Entomology,

University of California,

Davis, CA 95616, U.S.A.

Introduction

In most species of the Indo-Australian ant genus, Rhytidoponera,
deciduously winged queens are rare or absent, their place being
taken by reproductively functional workers (Whelden, 1957, 1960;
Haskins & Welden, 1965; Ward, 1981, 1984; Pamilo et al., 1985). A
polygynous colony structure, with several mated workers in lieu of a
queen, is the normal mode of colony organization in the common
Australian greenhead ant, Rhytidoponera metallica F. Smith
(Whelden, 1960; Haskins & Whelden, 1965; Haskins & Haskins,
1983), and queenright colonies of this species have not been
reported. A few alate or dealate females are known in collections,
and Haskins & Whelden (1965) noted the sporadic production of
alate queens in laboratory colonies of R. metallica. However beha-
vioral observations by these authors suggested that the queens had
lost the ability to found colonies. In this paper I document the
occurrence of functional queens in R. metallica , describe colony
foundation and growth under laboratory conditions, and discuss the
significance of occasional queen production in this species.

* Manuscript received by the editor February 26, 1986.

Psyche

Methods

[Vol. 93

Field observations were made at several Queensland localities in
August-September, 1983, of which the following sites are discussed
below: (1) 10 km SE Kenilworth (26°40'S, 152°47'E), 340 m, dense
Eucalyptus forest; (2) Mt. Coot-tha, near Brisbane (27°29'S,
152°58'E), 160 m, mixed wet sclerophyll forest; and (3) St. Lucia,
Brisbane (27°30'S, 153°01'E), 15 m, urban parkland on the Univer-
sity of Queensland campus. Voucher specimens of Rhytidoponera
metallica from these localities have been deposited in the Australian
National Insect Collection (ANIC), CSIRO, Canberra and the
Museum of Comparative Zoology (MCZ), Harvard University.
Evidence suggests that R. “metallica” is composed of a complex of
sibling species (Crozier, 1981; cf. Brown, 1958), and the southeast-
ern Queensland populations may not be conspecific with R. metal-
lica sens. str. (type locality: Adelaide, South Australia).

Field-collected queens of Rhytidoponera from St. Lucia were
maintained in the laboratory in moist plaster-of-Paris nests. Each
nest consisted of a glass-covered chamber with the dimensions 40 X
25 X 5 mm, in a block of plaster measuring 85 X 55 X 10 mm. A
single exit, 4 mm wide, led to a foraging arena 85 X 1 10 mm in area.
After a colony size of approximately 50 workers was attained, colo-
nies were provided with larger nests. Colonies were fed small
arthropods (mostly Drosophila) on a daily basis and droplets of
honey about once a week. A small quantity of clean sand was pro-
vided to allow construction of a cocoon-spinning matrix for the first
larvae. Censuses of brood and adults were taken every 3 weeks for
the first 9 weeks of colony development, and at weekly intervals
thereafter for the first year of growth.

Results

Field observations

While conducting field work in eastern Queensland in August-
September, 1983 I frequently encountered foraging workers of Rhy-
tidoponera metallica (s.l.), and I dissected several typical, worker-
reproductive colonies, i.e. colonies with workers and (sometimes)
males, but no queens. At three locations in southeastern Queensland
I unexpectedly encounted alate queens of R. metallica :

(1) While collecting for a period of one hour in Eucalyptus forest
10 km SE Kenilworth (25 August, 1983), I located a single Rhyti-

1986]

Ward — Rhytidoponera me tallica

doponera metallica colony under a rotten log; a partial excavation
(about two-thirds of the colony) yielded 157 workers, 17 alate
queens, and numerous larvae. No males or dealate females were
seen.

(2) During several hours of field work in wet sclerophyll forest
on Mt. Coot-tha (1 September, 1983), devoted primarily to the task
of locating colonies of the very timid species, R. anceps Emery, I
noted more than a dozen, scattered, individual alates of R. metallica
resting on low vegetation (leaves, grass stalks, tree roots, etc.),
apparently in the aftermath of one or more mating flights. About
half of these alates were females (five queens were collected and
preserved).

(3) On the University of Queensland campus, St. Lucia, between
28-31 August, 1983, there was considerable flight activity of R.
metallica alates. Most of these alates were males: they were observed
in moderate numbers (30-40 males at any given time) around R.
metallica nest entrances on a campus lawn at mid-day. Most indi-
viduals were dispersing skyward, but a few males were observed
approaching nests in a low, cruising flight, 20-50 cm above the
ground. Four alate females of R. metallica were also noted: three of
these were running on campus sidewalks, the fourth was resting on a
grass stalk. The alate queens were observed between noon and 3:00
p.m., and none was associated with a specific nest. Three of the R.
metallica queens were collected; one died within 5 days, and subse-
quent dissection showed that she was uninseminated. The two
remaining queens (acc. nos. 6280 and 6281) were kept in vials with a
small quantity of earth and leaf litter. They shed their wings, exca-
vated crude cells, and began laying fertile eggs. On September 17,
1983 the queens were relocated in plaster-of-Paris nest chambers. I
also collected a single dealate queen of R. chalybaea Emery on 1
September, 1983 in a University of Queensland lecture hall (acc. no.
6297). This queen was treated in the same manner as the R. metal-
lica queens, and provided a convenient standard for colony growth
and development, since colony-founding queens are a normal occur-
rence in this species (Ward, 1983).

Development of queenright colonies: incipient stages

The preceding observations established that the early stages of
colony-founding behavior have been retained in R. metallica
queens, i.e. they can mate, disperse, undergo dealation, and exca-

Psyche

[Vol. 93

vate nests. Laboratory observations demonstrated that this can be
followed by normal haplometrotic colony development.

Both the R. metallica and R. chalybaea queens readily accepted
the plaster-of-Paris nests, and began raising worker brood. The
queens of both species foraged in their arenas for food, and accepted
both honey and fresh arthropods. Struggling Drosophila adults
(held in the foraging arena with a pair of fine forceps) were
approached with outstretched mandibles, captured, stung, and
returned to the nest.

The R. metallica queens appeared to be no less dexterous than the
R. chalybaea queen in capturing and handling prey, or in caring for
larval brood. As the larvae matured, queens of both species used
sand grains to construct cocoon-spinning matrices for the larvae.
Initially the development of brood proceeded at a similar rate in all
three colonies, with eggs, larvae, and cocoons present by the tenth
week (late November, 1983; Table 1).

Some behavioral differences were noted between the two species:
the R. metallica queens were observed foraging more frequently
during daytime hours than the R. chalybaea queen; the R. metallica
queens established their middens in the nest entrance, thus partially
closing it, whereas the R. chalybaea queen scattered most of her
refuse just outside the nest entrance; and the R. metallica queens
defecated widely (frequently in the foraging arena) whereas the R.
chalybaea queen concentrated her fecal deposits at one location (c.
25 mm2) inside the nest chamber. These minor (and perhaps idiosyn-
cratic) differences hardly diminish the overriding similarity between
the two species in early colony development.

After about twelve weeks, and just prior to the eclosion of
workers, colonies of the two species of Rhytidoponera began to
diverge in their patterns of development. The first R. metallica
workers appeared to have difficulty eclosing from their cocoons —
possibly because of inept assistance on the part of the queens — and
there was appreciable early worker mortality both as pharate adults
in cocoons and as eclosed adults. No such difficulties were evident in
the R. chalybaea colony, whose worker population increased at
considerably faster rate than that of the two R. metallica colonies
(Table 1). Moreover the R. chalybaea colony displayed regular
(although increasingly dampened) cycles of brood development,
with bouts of egg-laying followed by pulses of larval growth, cocoon
formation, and adult eclosion, whereas such cycles appeared to be

1986] Ward — Rhytidoponera me tallica 5

Table 1. Development of incipient queenright colonies of Rhytidoponera under
laboratory conditions. Under columns E, L, C, and W are given the numbers of eggs
(approximate), larvae, worker cocoons, and adult workers observed, respectively, at
each census period.

Date

chalybaea 6297
E L C

metallica 6280
E L C

metallica 628 1
E L C

12.x. 83

—

2.xi.83

—

22.xi.83

29.xi.83

—

6.xii.83

—

13.xii.83

—

20.xii.83

—

27.xii.83

—

3.L84

—

10.i.84

—

17.i.84

—

24.i.84

—

31.i.84

—

7.ii.84

—

14.ii.84

21.ii.84

—

28.ii.84

—

6.iii.84

13.iii.84

20.iii.84

disrupted in the R. metallica colonies (compare respective columns
of Table 1).

Because of the delay in successful emergence of workers, the R.
metallica queens continued to forage for about two months after the
R. chalybaea queen ceased such activity. In both species the forag-
ing activity of the queen declined gradually, over a period of several
weeks after the first successful eclosion of workers. For three weeks
after her first daughter appeared the R. chalybaea queen continued
(with decreasing frequency) to capture and sting prey ( Drosophila
adults) held at, or near, the nest entrance. During the equivalent
transition period, the R. metallica queens continued to make forays
into the foraging arena and to capture prey. The sequence of events
in colony #6280 is summarized in Table 2; similar observations were
made on colony #6281.

Psyche

[Vol. 93

Table 2. Observations on foraging activity of the queen and first eclosing workers
in R. metallica colony #6280. Day #1 (3.iii. 1 984) is the day of first successful eclosion
of a worker.

Day #

No. of adult workers
in colony

Foraging activity of queen and workers

1-2

1 (callow)

Queen foraging. Worker confined to nest.

2 (1 callow)

Queen foraging. Workers confined to nest.

Queen took prey ( Drosophila ) at nest entrance.

11-12

Queen foraging. Workers confined to nest.

Worker foraging in arena (first time), captured a
subdued Drosophila adult; queen removed prey
from worker at nest entrance, then proceeded to
forage in arena herself.

14-17

4-5

Queen and one worker in foraging arena.

22-27

6-8

Queen and several workers foraging and taking
prey, the workers more active than the queen.

Last observation of queen in foraging arena
(thereafter queen confined to nest, and all
foraging conducted by workers).

Subsequent growth and development of queenright colonies

The growth rates of the R. metallica colonies were rather slow
and uneven, relative to that of R. chalybaea (Figure 1). One year
after colony initiation, the two R. metallica colonies had worker
populations of 41 and 27 individuals, respectively, while the R.
chalybaea colony had a worker population exceeding 200. Since the
colonies were fed ad libitum, food availability is not likely to have
been a limiting factor in the slower growth of the R. metallica
colonies. In fact, all three colonies grew at a rate faster than that
inferred for incipient queenright colonies of R. chalybaea (and a
related species, R. confusa Ward) in the field (Ward, 1981).

The R. metallica colonies appeared to function similarly during
the first year of development. Then a marked divergence took place,
apparently due to queen infertility in colony #6280. In mid-October,
1984 (week 56) this colony stopped producing eggs, and the amount
of brood began declining. By mid-January, 1985 (week 66), with a
population of 50 workers (and one male of unknown parentage),
this colony contained no eggs or larvae, and only one cocoon
(worker). On January 22, the queen was observed in a sexual calling
posture (gaster raised, head and mesosoma lowered) inside the nest;

1986]

Ward — Rhytidoponera me tallica

Figure 1 . Colony size (number of adult workers) of developing, queenright colo-
nies of R. metallica and R. chalybaea, as a function of time in weeks since colony
initiation.

at the same time she was being spread-eagled by two workers who
were tugging on opposite legs. Ten minutes later the queen was
dragged and bitten on the tip of her gaster by a worker. The follow-
ing day the queen was still being molested by workers, who bit her
on the legs and gaster. On January 24, the queen was found dead
inside the nest. A few days later her disarticulated body had been
dumped in a midden pile in one corner of the foraging arena. In the
meantime there began a spate of intersibling rivalry among a group
of 15-20 workers inside the nest who repeatedly “boxed” one
another with their antennae. These rapid antennation movements
were very similar to those which occur among mated workers in
polygynous, worker-reproductive colonies of the R. impressa group
(Ward, 1983, p. 293).

One week after the death of the queen in colony #6280, workers
began “calling” for males in the characteristic sex pheromone-

Psyche

[Vol. 93

releasing posture (Holldobler & Haskins, 1977). As many as six
workers were observed calling simultaneously, both inside and out-
side the nest. Workers calling inside the nest were subject to
repeated rapid antennation of the gaster, sides of body, and head, by
other workers. When antennated in front, the calling worker would
reciprocate the gesture, while maintaining the calling posture.
Workers calling in the foraging arena outside the nest were not the
object of rapid antennation by other workers.

The sexual calling behavior of workers continued, with increasing
intermittency, for the next six months. During this time, two addi-
tional adult males were produced, but no workers. There was no
indication that sib mating occurred — males showed no apparent
interest in their calling nestmates. The colony continued to decline
in size, no additional workers were produced, and, at time of writing
(January, 1986), it consisted of 35 workers, 1 male, 2 larvae and
several eggs.

By contrast, colony #6281 remained a viable queenright colony.
The queen continued to produce fertile eggs, and was not molested
by her daughters. There was no obvious conflict among workers (i.e.
no spate of antennal boxing or other forms of aggression), and
workers did not exhibit sexual calling behavior. At time of writing,
the colony was continuing to grow and comprised the queen, about
120 workers, and abundant brood.

Discussion

These findings demonstrate that the deciduously winged females
of Rhytidoponera metallica have not lost the potential to function
as queens, despite their sporadic occurrence in nature. Under
laboratory conditions the two R. metallica colonies remained queen-
right for at least a year, and the queens and workers adopted conven-
tional roles of egg-layer and forager, respectively. On the other hand
the R. metallica colonies grew more slowly than the incipient queen-
right colony of R. chalybaea, and the colony-founding foraging phase
of the queens was correspondingly extended. Hence there remains
some uncertainty about the efficacy of colony foundation by R.
metallica queens in nature.

One of the R. metallica colonies experienced death of the queen,
apparently a case of matricide triggered by queen infertility. Since
the workers began calling for males soon after the queen’s death,

1986]

Ward — Rhytidoponera me tallica

and continued to do so for six months, it seems likely that, under
natural conditions, replacement of the queen by mated workers
would be readily accomplished. Ward (1983) alluded to the possibil-
ity that some worker-reproductive (Type B) colonies in the Rhyti-
doponera impressa group are derived from orphaned queenright
(Type A) colonies, and the present observations provide direct evi-
dence that such a transition can occur in R. metallica. Moreover
they suggest that reproductive activity on the part of the queen,
rather than her mere presence, is necessary for the suppression of
hostile takeover attempts by her daughters.

The reverse process, production of colony-founding queens by
worker-reproductive colonies, seems certain to have occurred. No
mated dealate queen was found in the queen-producing colony from
10 km SE of Kenilworth, and indeed no functional queenright colo-
nies of R. metallica have been reported in the field, even though this
species is one of the commonest Australian ants.1 Haskins &
Whelden (1965) reported the occasional production of female alates
in worker-reproductive colonies of R. metallica which had been
maintained in the laboratory for several years. These females failed
to function as queens but this could have been due to the absence of
favorable conditions for mating and dispersal.

Queen production might be viewed as an infrequent, alternate
dispersal strategy employed by worker-reproductive R. metallica
colonies in response to environmental conditions which favor long-
range dispersal over short-range movement (colony fission). The
unusually large production of queens in Queensland in August-Sep-
tember, 1983 occurred after a period of drought associated with the
1982-83 El Nino. Alate queens appeared in one of Haskins’ labor-
atory colonies after a shift in diet (C. P. Haskins, pers. comm.). The

'Among the limited number of R. metallica queens in collections, the majority of
specimens are alates; the dealate specimens which I have examined contain no infor-
mation about their reproductive status. During a five year period of collecting ants in
eastern Australia (1974-78; 1980) I encountered (and subsequently dissected) R.
metallica queens only twice. One of these was a mated dealate female wandering on
the ground by herself (colony-founding?) in open Eucalyptus woodland, 14 km E
Grenfell, New South Wales (29. X. 1975, P. S. Ward #1406); the other was a single
uninseminated (spermatheca empty, ovaries poorly developed) dealate female in a
colony with 173 workers and brood, under a stone in dry sclerophyll forest, at
Bathurst, N.S.W. (18. X. 1975, P. S. Ward #1374).

Psyche

[Vol. 93

extreme rarity of mature queenright colonies in nature could be
attributed to a frequent transition to the worker-reproductive (Type
B) colony structure, coupled with the sporadic production of queens
in the first place. That R. metallica queens still function as dispersal
units is suggested by the widespread retention of queen production.
Among material in the ANIC and MCZ, there are alate or dealate
females of R. metallica (s.l.) from Western Australia, South Austra-
lia, New South Wales, and Queensland, i.e. throughout the range of
this species (or species complex). Queens have also been collected
throughout most of the geographical distribution of R. victoriae
Andre, another species whose mature colonies are predominantly
or entirely worker-reproductive.

It is worth reiterating that queens are entirely unknown in the
majority of Rhytidoponera species (including the large, robust-
bodied forms found primarily in xeric habitats), and in such species
aerial dispersal of females is impossible. If queens are effective aerial
dispersers in R. metallica and other occasional queen-producers
(including R. clarki Donisthorpe, R. inornata Crawley, R. tasma-
niensis Emery, and R. victoriae), then this should result in differen-
tial patterns of habitat island and offshore island occupancy by the
two groups of Rhytidoponera. There are not sufficient data availa-
ble to test this prediction — and the test would be complicated by
differing habitat preferences of members of the two groups — but
records in the ANIC do show that R. metallica and related species
are found on a variety of small islands off the coasts of Western
Australia, New South Wales and Queensland.

Summary

In colonies of the Australian greenhead ant, Rhytidoponera
metallica (s.l.), female reproductive activities are almost invariably
assumed by workers. Queens (deciduously winged females) are
rarely produced, and were heretofore considered non-functional.
Field observations in southeastern Queensland in August and Sep-
tember, 1983 revealed an unusually high frequency of alate queens
in several localities. Two of three alate queens, collected while dis-
persing in the vicinity of male mating flights, proved to be insemi-
nated. In the laboratory these mated queens both established
functional queenright colonies under non-claustral, haplometrotic
conditions. The R. metallica colonies grew more slowly than an

1986]

Ward — Rhytidoponera me tallica

incipient, queenright colony of R. chalybaea (a species in which
functional queens are common), but a clear division of labor devel-
oped between the egg-laying queen and foraging workers.

One R. metallica colony suffered death of the queen in its second
year of development. This was followed by a spate of intersibling
rivalry and frequent sexual calling behavior on the part of the
workers. The other colony continued to function as a viable queen-
right colony, and showed no signs of intracolony strife or reproduc-
tive attempts by workers.

These observations show that R. metallica queens have retained
their colony-founding and reproductive potential, despite their spo-
radic occurrence in nature. This suggests that long-range dispersal
via winged queens remains an occasional viable option for worker-
reproductive colonies of R. metallica.

Acknowledgements

The University of California provided financial support for this
work. I thank Ross Crozier, Caryl Haskins and Christian Peeters
for comments on the manuscript.

References

Brown, W. L. 1958. Contributions toward a reclassification of the Formicidae II.
Tribe Ectatommini (Hymenoptera). Bull. Mus. Comp. Zool. Harvard, 118:
175-362.

Crozier, R. H. 1981. Genetic aspects of ant evolution. In Atchley, W. R. and D.
Woodruff (Eds.). Evolution and speciation. Essays in honor of M. J. D. White.
Cambridge: Cambridge Univ. Press, pp. 356-370.

Haskins, C. P. and W. M. Whelden. 1965. “Queenlessness”, worker sibship, and
colony versus population structure in the formicid genus Rhytidoponera.
Psyche, 72: 87-112.

Haskins, C. P. and E. F. Haskins. 1983. Situation and location-specific factors
in the compatibility response in Rhytidoponera metallica. Psyche, 90: 163-174.
HOlldobler, B. and C. P. Haskins. 1977. Sexual calling behavior in primitive
ants. Science, 195: 793-794.

Pamilo, P., R. H. Crozier, and J. Fraser. 1985. Inter-nest interactions, nest
autonomy, and reproductive specialization in an Australian arid-zone ant, Rhy-
tidoponera sp. 12. Psyche, 92: 217-236.

Ward, P. S. 1981. Ecology and life history of the Rhytidoponera impressa group
II. Colony origin, seasonal cycles, and reproduction. Psyche, 88: 109-126.
Ward, P. S. 1983. Genetic relatedness and colony organization in a species com-
plex of ponerine ants. I. Phenotypic and genotypic composition of colonies.
Behav. Ecol. Sociobiol., 12: 285-299.

12 Psyche [Vol. 93

Ward, P. S. 1984. A revision of the ant genus Rhytidoponera in New Caledo-
nia. Aust. J. Zool., 32: 131-175.

Whelden, R. M. 1957. Anatomy of Rhytidoponera convexa. Ann. Ent. Soc.
Amer., 50: 271-282.

Whelden, R. M. 1960. Anatomy of Rhytidoponera metallica. Ann. Ent. Soc.
Amer., 53: 793-808.

SOUTH AMERICAN AND FLORIDIAN DISJUNCTS IN
THE SONORAN GENUS COMPSOCRYPTUS
(HYMENOPTERA: ICHNEUMONIDAE).

By Charles C. Porter1

Department of Biological Sciences, Fordham University
Bronx, NY 10458

Introduction

Taxonomy

Most Compsocryptus may be recognized at a glance by their
elegantly yellow banded brown or black wings, large and anteriorly
wide areolet, short and weak notauli, axillus intermediate in posi-
tion between the anal margin of the hind wing and the submediella,
strong ventro-lateral carina on postpetiole, and long, upcurved
ovipositor.

My concept of this genus agrees, as to species included, with
Townes’ most recent definition (1969:203-4). Several of Townes’
diagnostic features, however, do not apply to the Compsocryptus I
have examined (C.fasciipennis, C.fuscofasciatus, C. melanostigma,
C. texensis, and C. xantho stigma). All Compsocryptus I have seen
possess a sharp and strong subvertical groove externo-ventrall> near
the base of the hind coxa, while Townes describes the hind coxa as
“without a groove” (1969:203). All Compsocryptus examined by me
have, in the female only, a prominent crescentic to subtriangular
baso-lateral flange at the base of the petiole, while Townes main-
tains that the petiole is “without a lateral tooth at the base”
(1969:203). Compsocryptus forms a compact genus whose species,
despite their far-flung and discontinuous distribution, seem unusu-
ally homogeneous in color and structure. I thus suspect that all
members of the genus will turn out to have a basal first gastric
projection in the female and a strong hind coxal groove.

'Research Associate, Florida State Collection of Arthropods, Florida Department of
Agriculture and Consumer Services, Division of Plant Industry, P.O. Box 1269,
Gainesville FL 32602.

Manuscript received by the editor May 28, 1985

14 Psyche [Vol. 93

Relationships

Compsocryptus displays close affinity to Trachysphyrus, Aelio-
potes, Joppidium, and Lanugo . Superficially, in color, size, habitus,
and many details of geographic distribution, Lanugo especially
parallels Compsocryptus. However, the phylogenetic connection
seems distant, since Lanugo has only a weak groove at the base of
the hind coxa, the axillus vein closer to the hind margin of the wing
than to the submediella, and the ovipositor straight and shorter than
in Compsocryptus. The South American Trachysphyrus [now re-
garded as including only the Imperialis group, as defined by
Porter (1967:275-319)] seems directly associated with Compso-
cryptus. Important characters that separate Trachysphyrus include:
dark but never pale banded wings; body color usually metallic blue,
green, or purple; female flagellum scarcely flattened below; notauli
usually (not always) extending beyond the middle of the mesoscutum;
surface of mesoscutum shining, never extensively mat; 1st gastric
tergite without a baso-lateral expansion; and 2nd gastric tergite in
many species smooth and polished (in numerous others mat).
Aeliopotes paitensis (Porter, 1986) in some ways (especially the
petiolar tooth) seems annectant between Compsocryptus and
Trachysphyrus but in other features (epomial development) is
aberrant and deserves generic status. Finally, Joppidium seems to
be a direct offshoot of Compsocryptus. Joppidium is more slender
than Compsocryptus (postpetiole at least 1.5 as long as wide), lacks
a ventro-lateral carina on the 1st gastric tergite, and has the female
flagellum more strongly flattened below toward apex. Some of its
species have a baso-lateral tooth on the petiole and yellow banded
wings, as in Compsocryptus. Joppidium also parallels its relative in
distribution, with numerous Sonoran species, several which extend
(but not disjunctly) into the southeastern United States, and with an
isolated species group in subtropical Brasil and north Argentina (no
representatives in the Peruvian Coastal Desert).

Biogeography and Ecology

Compsocryptus belongs to the Sonoran Biogeographic category
(Porter 1980:25-7). Possibly the genus evolved during the last half

1986] Porter — Sonoran genus Compsocryptus 15

of the Tertiary somewhere in southwestern North America. Cer-
tainly, its xerophilous species would have adapted well to the
increasingly drier climates of post-Oligocene times and to the
microphyll and sclerophyll Madro-Tertiary Geoflora which then
overspread the ever-rising Sierra Madre and Rocky Mountains. On
the other hand, Compsocryptus may have originated in western and
southern South America, where so many of its relatives are centered
today. Here, the Miocene climate paralleled that of the Sonoran
region. By the end of the Miocene the Argentine pampas had
become well developed, composites and other dry-adapted plants of
open habitats were radiating vigorously, and long dry seasons began
to characterize the middle latitudes as a result of the reduced rainfall
and “the ever-increasing rain-shadow effect of the rising Andes”
(Solbrig 1976:22-3). Thus arose the Chaco, an austral Sonora.

Whatever may have been its genesis, Compsocryptus today is
centered in the western United States and northern Mexico (15
species). It also has 3 remarkable disjuncts: C. fasciipennis in tropi-
cal Florida and Cuba, C. fuscofasciatus in the Peruvian Coastal
Desert, and C. melanostigma in north Argentina and nearby areas
in Paraguay and Brazil.

Even the most geographically remote species of Compsocryptus
differ only in apparently minor features of color and sculpture. This
fact may suggest that the disjunctions noted above arose in compar-
atively recent times. Both the increasing aridity of the later Tertiary
and xerothermic episodes within the Pleistocene probably allowed
semiarid communities (Thorn Scrub, Subtropical Deciduous Forest,
etc.) to range almost uninterruptedly from the southeastern United
States to Argentina. Wet periods during the Pleistocene (glacial
maxima at higher latitudes) would have favored the expansion of
forests and probably caused the fragmented distribution that is
observed among modern Compsocryptus species.

For example, “during a past period of low rainfall a prairie type
flora, such as is found today in Texas and Arizona, was able to
extend its range into the eastern United States, and remnants of this
flora reflecting dry conditions still exist on parts of the west coast of
Florida and on Big Pine Key” and other lower Florida Keys
(Spencer and Stegmaier 1973:13). Such dry periods occurred both in
the Pleistocene and in the climatically unsettled late Tertiary. They

Psyche

[Vol. 93

allowed eastward expansion by a whole complex of Sonoran Biota,
from Opuntia, Cereus, Acacia and other xerophytes, to insects like
Compsocryptus fasciipennis, several species of Joppidium, Lanugo
retentor, Derocentrus longicaudis , Eiphosoma dentator (all Ich-
neumonidae), Eumenes smithii (Eumenidae), Stictiella (Sphecidae)
and even to vertebrates, such as the reptiles Crotalus, Sistrurus, Pit-
uophis, Sceloporus, and Gopherus.

Concurrently, similar physio-climatic events could have produced
the plausibly vicariant differentiation of Compsocryptus fuscofasci-
atus and C. melanostigma in South America. As discussed under C.
melanostigma, the common ancestor of these two species may have
ranged in Chaco vegetation from Argentina to coastal Peru at a
time before the Andes were high enough at this latitude to impede
east-west exchange of lowland biota.

As intimated throughout the above discussion, Compsocryptus
prefers semiarid or arid environments, but also may be abundant in
open, degraded subtropical humid forests. Compsocryptus mela-
nostigma, for example, has been cited from the very wet Selva
Tucumano-Boliviana and Selva Misionera in Argentina. However,
the majority of these forest records are from ecotones between forest
and Chaco or from sites in the first stages of secondary succession
(logging roads, clearings, windfalls, etc.). This fact demonstrates
how precarious is the present-day equilibrium between forest and
scrub (Selva and Chaco). Almost all modern forests are surrounded
by drier environments, whose aggressive biota tends to encroach
with the slightest ecological perturbation. Compsocryptus melano-
stigma in Argentina and C. fasciipennis in south Florida are among
the many indicator species of these often anthropogenic and usually
disastrous environmental changes, from forest to scrub and finally
to desert.

Hosts

Compsocryptus are among the most common, conspicuous, and
frequently collected of New World Ichneumonidae. Nonetheless,
practically nothing is known about their host relationships. Only
Compsocryptus melanostigma has been reared. It parasitizes noc-
tuid moths of the genera Alabama and Pseudaletia. Alabama larvae
feed on cotton and pupate in rolled leaves. Pseudaletia larvae feed at
night on many kinds of grains and grasses, hiding by day under

1986]

Porter — Sonoran genus Compsocryptus

clods of earth or in other slightly subsurface shelters. Pseudaletia
pupae are made in the ground. These data explain why females of C.
melanostigma most often are collected on the ground or from low
vegetation. Other Compsocryptus species occur in similar micro-
habitats and probably parasitize comparable hosts.

Collections

Listed below in alphabetic order are the collections in which
material from this study has been or is to be deposited. Institutional
collections are designated by the name of the city where they are
located. Individual collections are referred to by the surnames of
their owners.

Cambridge. Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge, MA 02138.

college station. Department of Entomology, Texas A & M
University, College Station, TX 77843
Gainesville. Florida State Collection of Arthropods, Bureau of
Entomology, Division of Plant Industry, Florida Department
of Agriculture and Consumer Services, P. O. Box 1269, 1911
SW 34th Street, Gainesville, FL 32602.

Lawrence. Department of Entomology, Snow Entomological
Museum, The University of Kansas, Lawrence, KS 66045.
porter. Collection of Charles C. Porter, 301 North 39th Street,
McAllen, TX 78501.

townes. American Entomological Institute, c/o Dr. Virendra
Gupta, Bureau of Entomology, Division of Plant Industry,
Florida Department of Agriculture and Consumer Services,
Gainesville, FL 32602.

Genus COMPSOCRYPTUS

Cryptoideus Ashmead, 1900. Proc. U. S. Natl. Mus. 23: 42. Type: Cryptus purpuri-
pennis Cresson.

Compsocryptus Ashmead, 1900. Proc. U. S. Natl. Mus. 23: 43. Type: Cryptus calip-
terus Say.

Callicryptus Ashmead, 1900. Proc. U. S. Natl. Mus. 23: 43. Type: (Cryptus “fascia-
ms” Brulle) — fasciipennis Brulle.

Stictocryptus Cameron, 1908. Trans. Amer. Ent. Soc. 34: 243. Type: ( Cryptus “fascia-
tipennis” Brulle) = fasciipennis Brulle.

Sophocryptus Mallo, 1961. Idia 165: 17. Nomen nudum.

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[Vol. 93

Fore wing 7.2-13.0 mm long. Wings usually dark with yellow
transverse bands. Female flagellum somewhat widened and flat-
tened below on apical 0.3. Male flagellum with linear tyloids on
many intermediate segments. Mandible usually moderately broad
with lower tooth slightly shorter than upper tooth (rarely long and
slender with lower tooth much shorter than upper). Clypeus rather
large, gently convex in profile; its apical margin always edentate and
straight to weakly convex. Occipital carina sharp and narrow.
Malar space about 1.0 as long as basal width of mandible. Prono-
tum with epomia well defined but not extending much dorsad or
ventrad of scrobe. Mesoscutum with notaulus faint, traceable less
than half its length; surface mat, dully shining, or sometimes pol-
ished and with numerous, small to medium sized, crowded to well
separated punctures (punctures sparser in males). Mesopleuron
without a ridge on prepectus below. Hind coxa with a sharp and
strong subvertical groove externo-ventrally near base. Wing vena-
tion: areolet large, symmetrically to asymmetrically pentagonal,
intercubiti slightly to definitely convergent dorsad, front side of
areolet (2nd abscissa of radius) 0.9 to more than 1.0 as long as 1st
intercubitus (mesal side of areolet); discocubitus gently arched,
without a ramellus; mediella nearly straight; axillus long, diverging
from anal margin of wing, as close to submediella as to anal margin.
Propodeum: spiracle elongate; apical trans-carina varying from
strong throughout, to strong laterad but obsolete mesad, to almost
completely absent; cristae usually defined and subcrescentic to
bluntly triangular (cristae often obsolete in males). First gastric ter-
gite in female with a prominent crescentic to subtriangular baso-
lateral flange and with the postpetiole strongly expanded but in
male without a baso-lateral expansion and with the postpetiole
slender; ventral longitudinal carina usually sharp throughout,
dorso-lateral and dorsal carinae less well developed, weakest in
males. Second gastric tergite mat with very fine and dense punctures
(sparser in males) which sometimes become more widely spaced
mesad and with short, recumbant, mostly overlapping setae. Ovi-
positor: long, sheathed portion 0.5- 1.4 as long as fore wing, gently
upcurved, cylindro-compressed, its tip elongate (0.10-0.25 as high at
nodus as long from nodus to apex), nodus low and without a notch,
ventral valve on tip with sharp and inclivously oblique ridges.

1986]

Porter — Sonoran genus Compsocryptus

Fig. 1. Compsocryptus melanostigma, $. Photograph of whole insect in lateral
view. Fig. 2. Compsocryptus fuscofasciatus, $. Photograph of whole insect in lateral

20 Psyche [Voi. 93

Key to the South American Species of Compsocryptus

1. Fore wing yellow with 2 narrow brown cross bands and with
brown on apex; 2nd abscissa of radius 0.9- 1.0 as long as 1st
intercubitus; female mesopleuron usually with some strong

longitudinal wrinkling

1 . C. fuscofasciatus (Brulle)

1'. Fore wing dark brown with a broad median yellow cross band
and a large subapical yellow blotch; 2nd abscissa of radius
1.2- 1.5 as long as 1st intercubitus; female mesopleuron with-
out any longitudinal wrinkling, almost uniformly puncto-
reticulate 2. C. melanostigma (Brulle)

1 . Compsocryptus fuscofasciatus (Brulle)

(Fig. 2, 3, 4)

Cryptus fusco-fasciatus Brulle, 1846. In Lepeletier: Histoire naturelle des insectes.

Hymenopteres 4:194. Holotype Peru, Lima (lost).

Callicryptus ornatipennis Cameron, 1902. Trans. Amer. Ent. Soc. 28:372. Holotype
$: Peru, Callao (London).

Female. Color: antenna ferruginous on scape and pedicel, yel-
lowish ferruginous on 1st (sometimes also 2nd and 3rd) flagello-
mere, mostly yellow on flagellomeres 2, 3 or 4-9, brown and yellow
on flagellomeres 10-11 or 12, and black or brownish black beyond
12th or 13th flagellomere; head ferruginous with black on apex of
mandible; palpi dull ferruginous; mesosoma ferruginous with
inconspicuous dusky staining on some margins and sutures or some-
times with rather extensive black markings (as described for male);
gaster dull ferruginous with faint dusky staining on 2nd and follow-
ing tergites or occasionally with better defined black areas toward
base on 2nd and 3rd tergites; legs ferruginous, duller on tarsi, with
dusky staining on apical tarsomeres, narrowly on apex of hind tro-
chantellus and base of hind femur, and with blackish on much of
hind tibia except toward its paler (sometimes contrastingly flavo-
ferruginous) base; fore wing light yellow with three brown areas as
follows: a broad transverse band on most of apical 0.3 of median
cell, on base of discocubital cell, on apical 0.3 of submedian cell, on
basal 0.3 of 1st brachial cell, and on adjoining region of anal cell; a
second brown cross-band covering basal 0.3 of radial cell, apical 0.3

1986]

Porter — Sonoran genus Compsocryptus

of discocubital cell, areolet, apical 0.5 of 2nd discoidal cell, and
expanding below to cover all but basal 0.3 (or less) of 2nd brachial
cell; as well as with a third brown area on apical 0.5 of 3rd cubital
cell and apical 0.5 of 3rd discoidal cell and confluent below with
dark area of 2nd brachial cell; hind wing pale yellow with apical 0.3
dusky and with dusky staining prolonged more narrowly far basad
on its hind margin, as well as sometimes with an irregular transverse
dusky area at level of nervellus.

Length of fore wing: 10.5-12.5 mm. Flagellum: 1st segment
3. 2-3. 5 as long as deep at apex; apical segments averaging 0.7-0. 8 as
long as wide. Malar space: 1.0 as long as basal width of mandible.
Mesoscutum: dully shining with abundant, dense, sharp, tiny punc-
tures which emit inconspicuous, short and mostly close-packed
setae. Mesopleuron: surface with delicate to strong, trans-biased
puncto-reticulation and, at least ventrad, usually with some strong
longitudinal wrinkling. Wing venation: radial cell 3.6-4. 1 as long as
wide; areolet about as high as broad, symmetrically pentagonal,
intercubiti weakly to moderately convergent above, 2nd abscissa of
radius 0.9- 1.0 as long as 1st intercubitus. Hind femur: 5. 6-6.0 as
long as deep. Hind tibia: below and laterally on apical 0.5 with a
few, scattered enlarged setae. Propodeum: apical face discrete from
the gently arched basal face and almost vertical; basal trans-carina
traceable throughout, uniformly fine and sharp or sometimes partly
weak and irregular. First gastric tergite: dorso-lateral carinae per-
current but faint; dorsal carinae well defined, but not sharp, on apex
of petiole and basal 0.5 of postpetiole; surface of postpetiole mat,
sometimes dully shining toward apex, with fine micro-reticulation
and with tiny, sparse, shallow punctures that are best developed
apico-laterad (where their short setae partially overlap). Ovipositor:
sheathed portion 0.69-0.81 as long as fore wing; tip 0.15-0.17 as
high at nodus as long from nodus to apex.

Male. Differs from female as follows: Color: pedicel sometimes
marked with black; flagellomeres 1-5 (sometimes up to 9) ferrugi-
nous to yellowish with dusky staining, mostly above; at least flagel-
lomeres 10-13 yellow with some ferruginous staining; rest of
flagellum black; head ferruginous with yellowish on base of mandi-
ble, much of face laterally, and narrowly on lower 0.6 of frontal
orbit as well as with black on apex of mandible, broadly above and
between antennal sockets (sometimes reaching and including stem-

Psyche

[Vol. 93

Fig. 3. Map showing geographical distribution of Compsocryptus fuscofasciatus
and C. melanostigma.

1986] Porter — Sonoran genus Compsocryptus 23

maticum) and on most of postocciput; mesosoma ferruginous with
black markings usually better developed than in female and includ-
ing areas on propleuron anteriorly, pronotal collar, spot on epomia
(sometimes contiguous with black on collar), broad band on hind
margin of pronotum, prescutellar groove, much of meso and
metanotal axillary troughs, groove at base of propodeum, all of
prepectus, mesosternal sulcus, hind face of mesosternum, broad
band on hind margin of mesopleuron — prolonged dorsad along
upper mesopleural margin to subalarum, broadly on all but dorsal
margin of lower metapleuron, and irregularly on hind margin of
propodeum; gaster dull to bright ferruginous with a little black at
base of 1st tergite and with succeeding tergites sometimes only with
irregular dusky staining and sometimes with well defined black
areas toward base of tergites 2 and 3; fore and mid tibiae and tarsi
more yellowish than in female; mid femur with some blackish stain-
ing dorsad and apicad; hind tibia black with basal 0.15 contrastingly
pale yellow; hind tarsus blackish with light yellow at least near base
of 1st segment and sometimes almost throughout on both segments
1 and 2.

Length of fore wing: 8. 5-9. 6 mm. Flagellum: linear, largely per-
current tyloids present on segments 1 1 or 12 to 19 or 20; 1st segment
2. 5-2. 7 as long as deep. Malar space: 0.77-0.90 as long as basal
width of mandible. Mesoscutum: shining with abundant, moder-
ately small, sharp punctures that are separated by 1. 0-2.0 their
diameters and which emit dense, erect, moderately long setae.
Mesopleuron: more shining than in female, with medium sized,
sharp, dense, subadjacent to reticulately confluent punctures and
some longitudinally biased reticulation. Hind femur: 6. 3-7. 4 as long
as deep. Hind tibia: with enlarged setae more abundant and con-
spicuous than in female. Propodeum: rather elongately convex in
profile; apical face not discrete from basal; apical trans-carina
weaker than in female, forming low and subcrescentic cristae or
sometimes with cristae obsolete. First gastric tergite: ventro-lateral
carina obsolete on petiole but sometimes becoming sharp toward
apex of postpetiole; dorso-lateral and dorsal carinae in great part
obsolete; surface of postpetiole smooth and shining with abundant
but well separated tiny punctures that emit long and uniformly
overlapping setae.

24 Psyche [Vol. 93

Specimens Examined. 15 9 and 47 $\ PERU, Lambayeque
Province, 33 km E. Olmos, Ruta a Jaen, 23-VII-1975, C. Porter, L.
Stange; 1 km S. Lambayeque, 24-27-VII-1975, C. Porter, L. Stange;
La Libertad Province, Laredo nr. Trujillo, 7-8-VII-1974, C. Porter,
L. Stange; Simbal nr. Trujillo, 4-7-VII-1974, C. Porter, L. Stange;
Lima Province, Cupiche, 10 km E. Chosica, 25-VI-2-VII-1974, C.
Porter, L. Stange; Palle nr. Chosica, 17-VII-1974, C. Porter, L.
Stange; San Geronimo nr. Chosica, 28-VI-5-VII-1976, C. Porter,
C. Calmbacher; nr. Surco on Carretera Central at km 59, 30-VI-
1976, C. Porter, C. Calmbacher.

Relationships. This Peruvian Coastal Desert endemic differs
only in minor chromatic and structural features from the other
South and North American Compsocryptus. It seems closely related
to the Argentine C. melanostigma (Brulle) but may be distin-
guished by the following characters: (1). Fore wing yellow with two
narrow brown cross bands and with brown on apex (vs. dark brown
with a broad median yellow cross band and a large subapical yellow
blotch), 2. Second abscissa of radius 0.9- 1.0 as long as 1st intercubi-
tus (vs. 1.2- 1.5 as long), 3. Female mesopleuron usually with some
strong longitudinal wrinkling (vs. puncto-reticulate), 4. Male
mesoscutum with punctures mostly separated by 1. 0-2.0 their
diameters (vs. 2.0 or more their diameters) and 5. Male flagellum
with tyloids extending to segments 19-20 (vs. 21-23).

Field Notes. Compsocryptus fuscofasciatus has been reported
only from the northern and central Peruvian Coastal Desert
between Lima and Piura. Here, it frequents most well watered habi-
tats between sealevel and 1500 m. I have collected it along rivers and
irrigation ditches in arid country as well as in orchards and
degraded cloud forest. Like other Compsocryptus , this species most
often occurs near or on the ground in exposed, disturbed, weedy or
grassy places.

2. Compsocryptus melanostigma (Brulle)

(Fig. 1,3,5)

Cryptus melanostigma Brulle, 1846. In Lepeletier: Histoire naturelle des insectes.
Hymenopteres 4:191. Lectotype $: Brasil: “Prov. de Misiones” (Paris
Museum).

Cryptus opaco-rufus Taschenberg, 1876. Ztschr. f. die Gesam. Natuw. Halle 48:64.
Lectotype $: (Brasil): Parana (Halle).

1986] Porter — Sonoran genus Compsocryptus

Fig. 4. Compsocryptus fuscofasciatus, $. Fore wing, showing color pattern. Fig.
5. Compsocryptus melanostigma, $. Fore wing, showing color pattern. Fig. 6.
Compsocryptus fasciipennis, $. Fore wing, showing color pattern.

Cryptus lateritus Taschenberg, 1876. Ztschr. f. die Gesam. Naturw. Halle 48:65.
Lectotype S'- (Brasil): Parana (Halle).

Callicryptus pulchrifasciatus Cameron, 1909. Trans. Amer. Ent. Soc. 35:437. Lecto-
type $: Argentina: Mendoza (London).

Sophocryptus bisulcatus Mallo, 1961. Idia 165:17. Nomen nudum.

Female. Color: antenna with scape brownish ferruginous,
pedicel dusky ferruginous with apex paler, and flagellum black with

Psyche

[Vol. 93

a yellowish white annulus on segments 4 (near apex) -9; head
brownish ferruginous with black on mandibular teeth; mesosoma
brownish ferruginous; gaster dull brownish ferruginous with vague
but often widespread dusky staining; legs brownish ferruginous with
dusky on fore tibia and tarsus, mid femur dusky dorso-apicad, mid
tibia extensively blackish or dusky, mid tarsus blackish brown,
some dusky staining on hind trochantellus, much blackish brown
(especially apicad) on hind femur, and black almost throughout on
hind tibia and tarsus; wings dark brown; fore wing with a broad
transverse median yellow band that covers dorso-apical corner of
median cell, basal 0.6 of discocubital cell, basal 0.5 of 2nd discoidal
cell, all but base of 1st brachial cell, basal 0.2 of 2nd brachial cell,
and anal cell beneath 1st and 2nd brachial cells, as well as with a
rounded yellow blotch covering most of apical 0.5 of radial cell, a
little of areolet, and basal 0.3 of 3rd cubital cell; hind wing with a
very broad yellow transverse band on its postmedian 0.25, contigu-
ous with median yellow band of fore wing.

Length of fore wing: 10.0-12.3 mm. Flagellum : 1st segment
3. 4-3. 5 as long as deep at apex; apical segments 0.6-0. 8 as long as
wide. Malar space: 1.0- 1.1 as long as basal width of mandible.
Mesopleuron: with extensive comparatively fine puncto-reticulation
and without any strong longitudinal wrinkling. Wing venation:
radial cell 3. 8-4. 5 as long as wide; areolet a little broader than high,
intercubiti weakly convergent above, 2nd abscissa of radius 1.2- 1.5
as long as 1st intercubitus. Hind femur: 6. 0-7.0 as long as deep.
Hind tibia: on apical 0.5 with numerous but widely spaced enlarged
setae. First gastric tergite: dorso-lateral carinae often sharp on peti-
ole; dorsal carinae varying from obsolete to weak; surface of
postpetiole uniformly mat with finely granular micro-reticulation,
practically glabrous, even apico-laterad. Ovipositor: sheathed por-
tion 0.76-0.83 as long as fore wing; tip 0.14-0.16 as high at nodus as
long from nodus to apex.

Male. Differs from female as follows: Color: scape yellow and
ferruginous, pedicel brown with some yellowish apicad, flagellum
with a yellowish white annulus on segments 9 or 10-12 or 13; face
rather pale ferruginous with some yellowish staining or with yellow
on facial orbits and less extensively also on frontal orbits; front
becoming dark brown to black between and above antennal sockets;
postocciput partly to mostly black; mesosoma with some black on

1986]

Porter — Sonoran genus Compsocryptus

propleuron, sometimes with a pair of black spots on pronotum
dorsally behind collar, sometimes narrowly black on anterio-lateral
margin of pronotum, sometimes narrowly black on much of hind
margin of pronotum, vaguely to extensively blackish on prepectus,
sometimes tinged with black in meso and metanotal axillary
troughs, sometimes blackish behind subalarum, sometimes black
stained in mesosternal sulcus, and sometimes blackish on margins of
lower metapleuron; gaster with slight to conspicuous blackish stain-
ing, often irregularly on 2nd tergite and always rather broadly on
tergites 5-7; fore tibia and tarsus yellowish with tarsus compara-
tively dark and dusky on last segment; mid leg with blackish in part
on trochanter and trochantellus, femur pale yellow on apical 0.2 and
otherwise blackish to brownish, tibia pale yellow, and tarsus dusky
with dirty yellow on basal 0.5 of first segment; hind leg with black
and brown staining on trochanter and trochantellus, femur black or
dark brown, and tibia black with a broad, dull yellowish-white
prebasal band covering about 0.25 of segment.

Length of fore wing: 8.6-10.3 mm. Flagellum: linear, largely per-
current tyloids present on segments 11 or 12-21, 22, or 23; 1st
segment 2.6-3. 1 as long as deep. Malar space: 0.82-0.93 as long as
basal width of mandible. Mesoscutum: shining with abundant,
small, sharp punctures that generally are separated by more than 2.0
their diameters. Mesopleuron: similar to female but with coarser,
longitudinally biased wrinkling and larger intercalated punctures.
Hind femur: 5. 5-7. 2 as long as deep. First gastric tergite: postpetiole
smooth and shining with abundant but well separated tiny punc-
tures whose setae mostly overlap laterad but become somewhat
sparser toward the meson.

Specimens Examined. 31? and 2A$\ ARGENTINA, Formosa
Province, Arroyo Eh Eh Grande, 76 km N Formosa, Rta. 11,
14-VIII-1977, C. Porter, L. Stange, P. Fidalgo, Arroyo San Hilario,
15 km S. Formosa, Rta. 11, 1 1-12-VIII-1977, C. Porter, L. Stange,
P. Fidalgo, Riacho Pilaga, 27 km N. Formosa, Rta. 11, 12-VIII-
1977, C. Porter, L. Stange, P. Fidalgo; Salta Province, Dique
Itiyuro, 70 km N. Tartagal, 30-VII-1977, C. Porter, L. Stange, P.
Fidalgo, Tartagal, 1 1 - 1 8-VIII- 1 973, C. Porter, 10 km N. Vespucio,
12-VIII-1976, C. Porter, L. Stange, Rosario de la Frontera, 19-VI-
1972, C. Porter; Tucuman Province, Rio Nio, 30-XI-1964, C. Por-
ter, San Pedro de Colalao, 19-XII-1964, C. Porter, Villa Nougues,
26-27-XI-1964, 6-7-XII-1964, C. Porter; Santiago del Estero Pro-

Psyche

[Vol. 93

vince, Termas de Rio Hondo, Dique Frontal, 3-V-1972, 2-VIII-
1973, C. Porter; La Rioja Province, Villa Union, 22-IV-1972, C.
Porter; Cordoba Province, La Lejania ca. Nono, 23-25-X-1984, C.
Porter, T. O’Neill.

Relationships. As discussed under that species, Compsocryp-
tus melano stigma much resembles C. fuscofasciatus of the Peruvian
Coastal Desert. The two species may have originated from a com-
mon ancestor that once ranged across what is now subtropical
South America from north Argentina to the Pacific coast. Warm,
seasonally dry conditions, of a type preferred by most modern
Compsocryptus, apparently prevailed across this area during the
early Tertiary (Solbrig 1976:42). Subsequent Andean uplift would
have split early Compsocryptus , populations into eastern and west-
ern isolates, setting the stage for differentiation of the modern C.
melanostigma in Argentina and C. fuscofasciatus in coastal Peru.

Field Notes. This conspicuous species occurs throughout
northern Argentina below 1500 m and ranges into adjoining parts of
Brasil and Paraguay. It occupies many forest, thorn scrub, and
desert biomes, including Southeast Brasilian Wet Forest, subtropi-
cal Andean Cloud Forest, Chaco Forest, Wet Chaco, Dry Chaco,
Montane Chaco, and Subandean Desert. In wooded areas, C. mela-
nostigma prefers disturbed situations in full sun along trails or at the
forest edge. In all habitats, it flies mostly near the ground among
grasses, forbs, or low shrubs.

Compsocryptus melanostigma often is very common during fall
and winter but may be collected in most habitats at any time of the
year.

This is the only Compsocryptus for which host information has
been obtained. It has been reared from the noctuid moths Alabama
argillacea and Pseudaletia unipunctata (Townes 1966:77).

3. Compsocryptus fasciipennis (Brulle)

(Fig. 6)

Cryptus fasciipennis Brulle, 1846. In Lepeletier: Histoire naturelle des insectes.
Hymenopteres 4:191. Lectotype $ (labeled by H. K. Townes Townes): Cuba
(Paris).

This elegant species was well characterized by Townes (1962:
282-3). It differs from other Compsocryptus by its bluish black

1986]

Porter — Sonoran genus Compsocryptus

ground color; black wings with a single yellow cross band on fore
wing; coarsely punctate to (medially) reticulo-punctate mesopleu-
ron; very densely setose 2nd gastric tergite; and sheathed portion of
ovipositor averaging only 0.67 as long as fore wing.

Like the South American Compsocryptus, C. fasciipennis is iso-
lated by more than 1000 km from its nearest congeners. It occurs
only on the Keys and in the Everglades region of tropical Florida as
well as on Cuba. Other North American Compsocryptus range both
northwest and southwest from near Houston in east Texas.

Current research has added some new information on the ecology
and geographic distribution of C. fasciipennis. These data are sum-
marized below.

New Specimens Examined: 139 and 33<5: UNITED STATES,
Florida, Monroe County, Bahia Honda Key State Park ll-X-1981,
C. Porter, L. Stange; Big Pine Key, 16-18-V-1982, 25-X-1982, C.
Porter; Fleming Key, V-1979 to V-1980, Malaise Trap, H. V.
Weems, Jr.; North Key Largo, 15-V-1982, 12-X-1981, C. Porter, L.
Stange; Stock Island, 18-V-1982, C. Porter, L. Stange.

Field Notes. Like other Compsocryptus, this species usually
occurs flying close to or crawling on the ground in early secondary
successional habitats at the edge of mature forests. In October of
1981 I netted 12 males from Bidens pilosa growing on the center
strip of a parking lot on Bahia Honda Key. My Key Largo speci-
mens also were taken from stands of Bidens. On Big Pine Key, I
swept several C. fasciipennis amid herbaceous undergrowth on a
sand ridge along a trail through a Tropical Hardwood Hammock.

Townes (1960:283) cites 75 males and 44 females of C. fasciipen-
nis from south Florida (Miami and Everglades National Park to
Key West) and indicates that the yearly activity period for this
species in Florida spans “December 28 to April 12” with 1 record for
5 December. My new records show that the species begins to fly as
early as 1 1 October and continues at least until 18 May. It is scarce in
May but often becomes abundant in October (e.g., \2$ from Bahia
Honda Key on ll-X-1981).

The Malaise Trap records from Fleming Key elicit interest for
several reasons. They are the first annual survey of Compsocryptus
(and other ichneumonid) abundance done on the Florida Keys.
They also provide an idea of ichneumonid species composition and
density in a highly disturbed part of the Keys. Fleming Key is an
artificial appendage of Key West, mainly given over to a U. S. D. A.

Psyche

[Vol. 93

animal quarantine facility and with little vegetation other than
mangroves, pioneering stage herbs, and introduced ornamental
trees, such as Casuarina. Such environments select for unusually
hardy ichneumonids and species of this type should particularly
concern the biological control specialist, who is looking for para-
sites that will thrive in climatically stressed agricultural systems.

The Fleming Key Survey, run between May 1979 and May 1980,
with a gap in September and October, amassed 631 ichneumonid
specimens belonging to 37 species. Only 9 of these species accounted
for about 89% (561 specimens) of all Ichneumonid ae trapped. Dia-
degma sp. (22 specimens) was the least abundant of the “common”
group, followed by Compsocryptus fasciipennis (23), Labena gralla-
tor (36), Mallochia agenioides (41), Anomalon sp. (43), Temelucha
sp. (68), Paraditremops albipectus (103), Calliephialtes ferrugineus
(107), and Eiphosoma dentator (118, Porter 1983).

Table 1 summarizes monthly phaenology for Compsocryptus fas-
ciipennis and the eight other common ichneumonid species of the
depauperate Fleming Key Fauna, as sampled by Malaise traps.
Compsocryptus fasciipennis is active from fall to late spring with
maxima in March and October (as shown by Malaise and hand
collected specimens). This seasonal phaenology coincides approxi-
mately with that of the Argentine C. melanostigma and agrees even
more closely with the pattern shown by C. texensis in the Lower Rio
Grande Valley (present from January to May and again in
December with greatest abundance in December, as documented by
Porter, 1977:82).

Compsocryptus fasciipennis follows a cool-season phaenologic
cycle not unlike that of many other ichneumonids which inhabit
subtropical communities from Florida and Texas to Argentina.
Among the abundant Ichneumonidae at Fleming Key, 4 species
have autumn to early spring maxima and roughly parallel C. fascii-
pennis (Calliephialtes ferrugineus, Paraditremops albipectus, Teme-
lucha sp., and Diadegma sp.), 2 peak in May (Labena grallator,
Anomalon sp.), and the other 2 become most abundant during July
and August (Eiphosoma dentator, Mallochia agenioides). Nonethe-
less January to March seem the best overall months for ichneumon-
ids at this locality. All 9 species occur during this trimester and 230

:ies of the depauperate Fleming Key Fauna. For details,

1986]

Porter— Sonoran genus Compsocryptus

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Psyche

[Vol. 93

of the 561 specimens were collected then. These data agree closely
with my earlier studies on south Texas mesostenine Ichneumonidae
(Porter 1977), which reported peak diversity (20/34 species) and
maximum abundance (138/679 specimens) for December and only
slightly less impressive statistics for January (18 species and 135
specimens).

Acknowledgments

This research was done principally under my current National
Science Foundation Grant (BSR-83 13444) and in part was sup-
ported by previous NSF awards (DEB-75-22426, GB-6925). Grants
from the Committee for Research and Exploration of the National
Geographic Society permitted field research in South America dur-
ing 1973, 74, 75, 79, and ’81. Support also came from Faculty
Fellowships conferred by Fordham University for the Spring
Semester of 1980 and the Fall Term of 1984.

As a Research Associate of the Florida State Department of
Agriculture and Consumer Services, I have received generous sup-
port from the Division of Plant Industry at Gainesville, among
whose personnel special thanks befit Dr. Howard V. Weems, Jr.,
Dr. Lionel A. Stange, and Mr. Harold A. Denmark. All my collect-
ing in South Florida was facilitated by the Division of Plant
Industry.

Material of Compsocryptus melanostigma was obtained in
Argentina during repeated periods of cooperation with the Instituto
Miguel Lillo of the Universidad de Tucuman. I am particularly
indebted to Professor Rodolfo Golbach and to Dr. Abraham Wil-
link of this institution.

I also thank Mr. Thomas J. O’Neill of Fordham University for his
assistance on fieldtrips to Argentina and Peru.

Summary

Compsocryptus is a mesostenine closely related to Trachysphy-
rus. Its short notauli, long anterior side of areolet, medially situate
axillus, long and upcurved ovipositor, and (usually) dark and yellow
banded wings distinguish Compsocryptus from most other trachys-
phyroids. There are 15 species centered in the Sonoran region of

1986]

Porter — Sonoran genus Compsocryptus

western North America and Mexico plus 1 isolated species in Flor-
ida and Cuba, another in the Peruvian Coastal Desert, and a 3rd in
the Argentine Chaco. Compsocryptus fuscofasciatus from Peru
has the fore wing yellow with 2 brown bands, while in the Argentine
C. melanostigma the fore wing is dark with a broad median yellow
cross band and a large subapical yellow blotch. Townes (1962) has
fully characterized the North American species. Compsocryptus
inhabits a variety of exposed situations at altitudes below 1500 m,
including deserts, Thorn Scrub, Subtropical and Tropical Decidu-
ous forests, and disturbed sites in humid Neotropic forests. The
species fly mostly from fall to spring. Compsocryptus melanostigma
has been reared as a solitary parasite from noctuid moth pupae.

Literature Cited

Porter, C.

1967. A revision of the South American species of Trachysphyrus. Mem.
Amer. Ent. Inst. 10:1-386.

1977. Ecology, zoogeography, and taxonomy of the Lower Rio Grande Valley
Mesostenines. Psyche 84( 1 ):28-9 1 .

1980. Zoogeografia de las Ichneumonidae latino-americanas. Acta Zool. Lil-
loana 36:5-52.

1983. Eiphosoma dentator in Florida. Florida Ent. 66: 353-358.

1986. Trachysphyrus and the new genus Aeliopotes in the coastal desert of
Peru and North Chile. Psyche 92: 513-545.

Spencer, K. A. and C. E. Stegmaier, Jr.

1973. Agromyzidae of Florida. Arthropods of Florida and neighboring land
areas 7:1-205. Division of Plant Industry, Gainesville.

Solbrig, O.

1976. The origin and floristic affinities of the South American temperate desert
and semidesert regions. In D. Goodall (ed.), Evolution of desert biota, p.
7-49. University of Texas Press, Austin.

Townes, H. K.

1962. Ichneumon-flies of America north of Mexico: Subfamily Gelinae, Tribe
Mesostenini. Bull. U. S. Natl. Mus. 216(3): 1-602.

1966. A catalog and reclassification of the Neotropic Ichneumonidae. Mem.
Amer. Ent. Inst. 11:1-367.

1969. Genera of Ichneumonidae, Part 2: Gelinae. Mem. Amer. Ent. Inst.
12:1-537.

THE ORB-WEAVER GENUS WITICA
(ARANEAE: ARANEIDAE).*

By Herbert W. Levi

Museum of Comparative Zoology,

Harvard University, Cambridge, MA. 02138

Two species of neotropical orb-weavers, “Edricus” crassicauda
and Witica talis, have each been known from a single sex, the first
from females only, the second from males. The male of Edricus
spinigerus, suspected by F.P. -Cambridge (1904) to belong with the
female Epeira crassicauda, has never been collected with it,
although Cambridge’s suspicion was the reason for placing the
female E. crassicauda in the genus Edricus. While parthenogenesis
could account for absence of males in E. crassicauda, the absence of
females in Witica was more perplexing. The large females of Epeira
crassicauda have a tail with a constriction (Fig. 1), the minute males
of Witica talis (Fig. 5) have a round, subspherical abdomen bearing a
glossy plate. The two placed in different subfamilies did not appear
to be likely mates.

Surveying our collections, I found males of Witica to have been
collected in Cuba, Puerto Rico, Central and northern South Amer-
ica, roughly the same distribution as the female specimens named
“Edricus” crassicauda. Both are fairly common on Barro Colorado
Island in Gatun Lake of Panama, from which large collections are
available.

Unexpected evidence for existence of males in E. crassicauda
turned up: a male palpal part was found in the microscope slide
preparation of the seminal receptacles. When expanding the palpus
of Witica talis, I noticed that the structure first considered to be the
conductor, and which is sometimes missing from specimens, is actu-
ally an appendage of the embolus. Further, its structure is remarka-
ble, including a hand with many fine teeth, presumably functioning
as a hold-fast inside the female genital duct (Fig. 1 1). Subsequently,

♦This is the third of a series of revisions of neotropical noncribellate orb-weaving
spiders.

Manuscript received by the editor March 17, 1986.

Psyche

[Vol. 93

I examined a female epigynum in ventral view with the pigmented
integument carefully removed. The mystery suddenly resolved itself
when I found the same structure embedded in the female genital
duct (Fig. 8), proving that Witica talis, placed in the group Witicae
close to Hypognatha and Cyrtarachneae by Simon (1895) and
Roewer (1942), is in fact the male of “Edricus” crassicauda, placed
in Cycloseae by the same two authors.

In examining all available males, I noted certain differences in the
appendage of the embolus in males from Trinidad and some South
American localities (Fig. 14). This different structure was found in
females (Fig. 13) from the same areas, further proof that Witica
males belong with females of “ Edricus , "and also providing evidence
that there are two species, the females of which look quite similar
except for the contents of the genital duct.

Only one embolus tip was found on each side in each female duct,
never two. Are they there to protect a male’s sperm and prevent
further mating by the female? Or might they be spermatophores
with sperm inside the tips? Or do they just function to block the
ducts? Only one or two males with broken emboli were in collec-
tions suggesting that males do not survive mating. Males with
broken tips could not be determined to species.

The relationship and placement of the two species of Witica is
uncertain. The male palpus lacks a median apophysis and terminal
apophysis, but I expect this to be a secondary loss rather than a
primitive absence, perhaps correlated with the minute size of the
males. The female genitalia are unusual in being lightly sclerotized
and lacking a scape and other projections; the epigynum resembles
the epigynum of Pronous. The enormous difference in size of the
sexes, the total length of females being more than 4.5 times that of
the male, is found in some other orb-weaver genera, such as Gaste-
racantha and Nephila (the latter probably belonging to the family
Tetragnathidae). Also, males of Arachnura are dwarf. The females
of Arachnura have a tail, perhaps a synapomorphy. A male
Arachnura logio Yaginuma from Japan examined also has a spheri-
cal abdomen with a sclerotized dorsal plate, but has a median apo-
physis and terminal apophysis in the palpus. The anterior median
eyes of males and females of Arachnura are more projecting than
those of Witica.

1986]

Levi — Genus Witica

Figures 1-5. Witica crassicauda (Keyserling). 1. Female, legs removed.
2. Female abdomen from side. 3. Immature female. 4. Epigynum. 5. Male in
same magnification as female.

Figures 6-7. W. cayana (Taczanowski). 6. Female. 7. Epigynum.

Size indicators: 1.0 mm, except Figures 4, 7, 0.1 mm.

Witica O.P. -Cambridge

Salassia Getaz, 1893: 105. Type species by monotypy S. tricuspis Getaz. (Name
preoccupied by Salassia Folin, 1871, a mollusk.)

Witica O.P. -Cambridge, 1895: 160. Type species by monotypy Witica talis O.P.-
Cambridge, 1895. NEW SYNONYMY.

Salassina Simon, 1895: 784. Type species by original designation and monotypy S.
crassicauda Keyserling, 1865.

Physiola Simon, 1895: 875. Type species by original designation and monotypy. P.
nigrans Simon, 1895. Synonymized with Witica by Simon, 1903.

Psyche

[Vol. 93

Bion O.P. -Cambridge, 1898: 244, pi. 30. Type species by monotypy B. brevis O.P.-
Cambridge, 1898. First synonymized with Witica by Simon, 1903.

Synonymy. Simon (1903: 1003) synonymized his Physiola pub-
lished in 1895 with Witica published the same year, as an objective
synonym. I do not know the month of the publications; Simon
presumably did and Witica was published earlier. Thus since Salas-
sina was published at the same time as Physiola it must also have
been published after Witica.

F.P. -Cambridge (1904: 500) placed Epeira crassicauda described
from a female into the genus Edricus. Edricus O.P. -Cambridge,
1890, has as type species Edricus spinigerus, 1890. Edricus spinige-
rus was described from a large male similar and perhaps congeneric
with Wagneriana tauricornis F.P. -Cambridge, 1904. F.P.-
Cambridge thought that Edricus spinigerus might be the unknown
male of Epeira crassicauda. This proved to be an error.

Diagnosis. Unlike the females of most Araneidae, the abdomen
has a tail usually constricted at its base (Figs. 1, 6) and the epigynum
is flat, lightly sclerotized, with a pair of depressions, (Figs. 4, 7). The
male is separated from other Araneidae by the minute size, 1.5- 1.9
mm (Fig. 5), sclerotized abdomen (Figs. 12, 15) and lacking a
median apophysis and conductor of the palpus and having a large
embolus tip which is transferred and plugs the female’s ducts (Figs.
9,14).

Description. Female. Carapace, sternum dark brown. Legs light
with contrasting dark rings. Dorsum of abdomen black with varia-
ble white patches, venter black with a pair of small, white spots.
Eyes subequal in size. Chelicerae with three teeth on anterior, three
on posterior margin. First legs longer than fourth, second and
fourth subequal, third shortest. Abdomen with a pair of anterior
blunt spines and a tail of variable shape (Figs. 1, 6). The tail is
constricted at its base and distally has three lobes.

Male. Carapace shiny brown, sternum, legs dark brown. Dor-
sum of abdomen shiny brown, venter black. Eyes subequal in size.
Median eyes their diameter apart. Cheliceral teeth as in female, leg
proportions as in female. Endites without tooth, palpal femora
without tooth, first coxae without hook. Abdomen with round
convex dorsal shield, sometimes wider than long or longer than
wide.

1986]

Levi — Genus Witica

Genitalia. Female epigynum has openings on each side of a flat
septum in a depression (Figs. 4, 7), short connecting ducts lead into
seminal receptacles (Figs. 8, 13).

The male palpus has a radix, embolus and, between them, a
stipes. Median apophysis and conductor have been lost, probably
secondarily (Figs. 10, 11). The embolus is large and has a distal

Figures 8-12. Witica crassicauda (Keyserling). 8. Epigynum cleared showing
embolus tip. 9. Left male palpus. 10, 11. Male palpus expanded. 10. Mesal.
11. Lateral. 12. Male.

Figures 13-15. W. cayana (Taczanowski). 13. Epigynum cleared showing
embolus tip. 14. Male palpus. 15. Male.

Size indicators: 0.1 mm, except Figures 12, 15, 1.0 mm.

Psyche

[Vol. 93

curved tip which breaks off in mating and remains in the female
connecting duct. Whether it serves only as a plug or perhaps is a
spermatophore is not known.

Almost all females had one tip on each side in the epigynum, none
were seen with two. Females appear to mate only once. Very few
males with a missing tip are in collections. Apparently they do not
survive mating.

Variation. Dorsal coloration of the abdomen of females of both
species is quite variable, sometimes all white (in alcohol). The tail of
the female abdomen may be shortened or blunt or long and is at
times turned up.

Habits. The web of Witica crassicauda was found to be fairly
common in a coffee plantation at about 1000 m altitude in Puerto
Rico. It is built between trees about 1.5 meters apart, the hub 1.5
meters above the ground, the orb 30 to 35 cm horizontal diameter.
The hub is open. There is a short vertical stabilimentum and the
frame threads below the orb have whitish decorations, flattened
threads as seen under a magnifying lens. The spider hangs in the
hub, head down (Figure 17); there is no retreat. In Panama and
Costa Rica the spider is common in low elevation forests; it does not
make a stabilimentum, nor decorations on lines. The egg-sac, made
in a vial, was fluffy, yellowish white, the size of the spider and
contained about 200-250 lemon-yellow eggs.

Key to species

1 Females 2

_ Males 3

2(1) Median septum of epigynum as wide or wider than depression
on each side (Figs. 4, 8); mated females show tubes, the ends of
embolus tip on sides of septum (Figs. 4, 8); West Indies, Mex-
ico to South America (Map) crassicauda.

Median septum of epigynum narrower than depressions (Figs.
7, 13); tip of embolus never visible in depressions. Trinidad,

South America (Map) cayana.

3(1) Base of tip of palpal embolus swollen and with spur (Fig. 14);

Trinidad, South America (Map) cavana.

Base of tip of palpal embolus a curved tube (Fig. 9); West

Indies, Mexico to South America (Map) crassicauda.

1986]

Levi — Genus Witica

Figure 16. Witica crassicauda. Female on a leaf from Panama.

Witica crassicauda (Keyserling)

Figures 1-5, 8-12, 16, 17; Map

Epeira crassicauda Keyserling, 1865; 806, pi. 18, fig. 3, 4, $. Female specimen from
New Granada (BMNH) examined.

Cyclosa crassicauda: — Keyserling, 1893: 270, pi. 14, fig. 200, $.

Witica talis O.P. -Cambridge, 1895: 160, pi. 16, fig. 13, ft. Male lectotype from Teapa,|
Tabasco, Mexico (BMNH) here designated. Simon, 1903: 1003. Petrunkevitch,!
1930: 337, figs. 225, 226, Roewer, 1942: 894. new synonymy. j

Salassia tricuspis Getaz, 1893: 105. Female holotype from Uruca, Costa Rica (P.

Biolley), lost, new synonymy.

Salassina crassicauda: — Simon, 1895: 784, fig. 853,

Salassina tricuspis: — Simon, 1895: 784.

Physiola nigrans Simon, 1895: 876, figs. 938, 939, Lectotype male, two males, one I
immature and fragments of immatures paralectotypes from forest San Esteban,
Venezuela (MNHN), here designated. First synonymized with Witica by Simon,
1903.

Psyche

[Vol. 93

Bion brevis O.P. -Cambridge, 1898: 244, pi. 30, fig. 5, Male from Teapa, Tabasco,
Mexico (BMNH), not examined. First synonymized with Witica by Simon,
1903.

Edricus crassicauda: — F.P. -Cambridge, 1904: 500, pi. 47, fig. 21, $. Roewer, 1942:
762

Edricus tricuspis: — F.P. -Cambridge, 1904: 500. Roewer, 1942: 762.

synonymy. Sdldssid tricuspis is synonymized with crdssicdudd
since the description fits the latter species with which Getaz
compares it. Also only one species is known from Costa Rica. The
immature Physiold nigrdns were thought by Simon to be adult
females.

Femdle. Total length, 7.8 mm. Carapace, 3.2 mm long, 2.7 wide.
First femur, 3.5 mm; patella and tibia, 3.7; metatarsus, 2.1; tarsus,
0.9. Second patella and tibia, 3.3 mm; third, 2.0; fourth, 3.3.

Mdle. Total length, 1.4 mm. Carapace, 0.9 mm long, 0.7 wide.
First femur, 0.8 mm; patella and tibia, 0.8; metatarsus, 0.4; tarsus,
0.3. Second patella and tibia, 0.7 mm; third, 0.4; fourth, 0.6.

Didgnosis. The median septum of the epigynum is as wide or
wider than the depressions on each side (Fig. 4); the male has a
curved tube on the base of the embolus tip (Fig. 9).

Vdridtion. Dorsal color, pattern and shape of abdomen of
females are variable. Total length of female 6.5 to 12.0 mm, males,
1.4 to 1.7.

Hdbits dnd Distribution. Forests from Mexico to Venezuela
and Peru, Greater Antilles (Map).

Records. Mexico Sdn Luis Potosi: Huichihuayan, June 1941,
immat. (H. Dybas, AMNH). Guerrero: S of Acahuizotla, 17 Nov.
1946, $(E. S. Ross, CAS). Tdbdsco: Teapa, 16 July 1947, $ (C. M.
Goodnight, AMNH). Chidpds: San Quintin, Feb. 1966, 9 (G. Ball,
D. R. Whitehead, RL); Palenque ruins, 28 May 1980, 9 (J- Cod-
dington, MCZ). Guatemala Moca, 31 Aug. 1947, 9 (C. P. Vaurie,
AMNH). Honduras Atldntidd : Lancetilla, July, 1929, 9 (A. M.
Chickering, MCZ). Nicaragua Musawas, Waspuc Riv., Sept. 1955,
9 (B. Malkin, AMNH). costa rica Heredid : La Selva, 4 $ (MCZ).
Puntdrends : Corcovado Natl. Park, 2 9 (MCZ). Limon: Rio
Reventazon, imm. (AMNH). Sdn Jose : San Jose, 3 $ (AMNH).
Cdrtdgo: Turrialba, dense jungle, 9 (EPC). panama Bocds del
Toro: Rio Changuinola, 2 9 (AMNH). Chiriqui : 9 (AMNH).
Pdndmb: Canal area, very common (MIUP, MCZ, CAS, AMNH).

1986]

Levi — Genus Witica

Map. Distribution of Witica species.

Psyche

[Vol. 93

Cuba. Pinar del Rio. common (MCZ). Oriente: common (MCZ,
AMNH). Dominican republic. Sanchez; Puerto Plata, S of Santi-
ago (all MCZ). mona isl. (MCZ). Puerto rico. very common
(MCZ, AMNH, JC)

TRINIDAD. 9 (MCZ). VENEZUELA. Monagas: Caripito, Aug.
1942, 9 (AMNH). Carabobo: San Esteban, 21 Jan. 1940, 9 (CUC).
Colombia Antioquia: Mutata, Dec. 1963, 9 (MCZ); Remedios, 20
Dec. 1984; 9 (MCZ). Meta: Cano Grande, Sept. 1944, 9 (AMNH).
Valle: Rio Jamundi, 1000 m; Anchicaya; E. of Buenaventura, 3 9
(all MCZ). Cauca: Guapi, Aug. 1975, 9 (W. Eberhard, MCZ). Peru.
Cajamarca: Nanchoc, Caserio Bolivar, 30 April 1967, 9 (C.
Mazabel, AMNH).

Witica cay ana (Taczanowski), new combination
Figures 6, 7, 13-15; Map.

Epeira cayana Taczanowski, 1873: 135, pi. 5, fig. 15, Female holotype from
Cayenne, French Guiana (PAN). Specimens examined came from Uassa (Ua$a,
Amapa, Brazil) in the Taczanowski collection, PAN.

Female. Total length, 9.0 mm. Carapace, 3. 1 mm long, 2.8 wide.
First femur, 3.6 mm; patella and tibia, 4.0; metatarsus, 2.4; tarsus,
1.0. Second patella and tibia, 3.5 mm; third, 2.0; fourth, 3.6.

Male. Total length, 1.6 mm. Carapace, 1.0 mm long, 1.0 wide.
First femur, 1.1 mm; patella and tibia, 1.1; metatarsus; 0.6; tarsus,
0.4. Second patella and tibia, 0.9 mm; third, 0.5; fourth, 0.8.

Diagnosis. The median septum of the epigynum is narrower
than the depression on each side (Fig. 7). Base of embolus tip is a
lobe (Fig. 14).

Variation. The color, pattern, and shape of the female abdomen
are variable. Females vary 6.8 to 10.5 mm total length, males 1.4 to
1.6.

Habits and Distribution. Probably from forest, Trinidad and
Venezuela to Peru (Map).

Records, trinidad 16 km from Arima, 27 Feb. 1959, Arima
Rd, 29 Dec. 1945, $ (both A. M. Nadler, AMNH); Tucuche, 12
Nov. 1944, 9 (R- H. Montgomery, AMNH). Venezuela Ara-
gua: Rancho Grande, 1945, 1946, 4 9 (W. Beebe, AMNH). brazil
Roraima: Rio Irene, Aug. 1911,9 (AMNH). Colombia Magdalena:
San Pedro, 8 Feb. 1974, 9 (J. A. Kochalka, IBNA). Meta: Villavi-
cencio, 11 March 1955, 2 9 (E. I. Schlinger, E. S. Ross, CAS).

1986]

Levi — Genus Witica

Figure 17. Witica crassicauda. Left: web of female in Puerto Rico, orb 32 cm
horizontal diameter. Right: hub of another web with female. Webs dusted wilh corn
starch.

Ecuador Pichincha: Via Pto. Quito, km 113, 1984, 1985, 4 $ (L.
Aviles, MECN); 16 km SE San Domingo Tinalandia, June, 1975,
9, $ (S. and J. Peck, MCZ). Guayas. 16 km N Manglaralto, 30 Jan.
1955, 3 9 (E. I. Schlinger, E. S. Ross, CAS). Los Rios: Juan Mon-
talvo, March 1938, 3 9 (W. Clarke-Macintyre, AMNH). Bolivar:
Balzapamba, 1938, 1939, 3 9, 8 (W. Clarke-Macintyre, AMNH,
MCZ). peru San Martin: 20 km NE Moyobamba; SE Moy-
obamba; Ekin, E. of Tarapoto, 1947, 6 9 (all F. Woytkowski,
AMNH). Hudnuco: Monson Valley, Tingo Marla, 18 Dec 1954, 9
(E. I. Schlinger, E. S. Ross, CAS).

Acknowledgements

National Science Foundation grant BSR 8312772 made the
research and publication possible. L. R. Levi and W. P. Maddison
have read the paper and made suggestions. Most specimens came

Psyche

[Vol. 93

from the collections of the Museum of Comparative Zoology
(MCZ); other collections used were those of the American Museum
of Natural History (AMNH), Cornell University Collection (CUC)
N. Platnick, curator; British Museum (Natural History), P. Hillyard
(BMNH), California Academy of Sciences, W. J. Pulawski (CAS),
Exline-Peck Collection, W. Peck (EPC), Museo Ecuatoriano de
Ciencias Naturales, Quito, L. Aviles (MECN), Museum National
d’Histoire Naturelle, Paris, J. Heurtault, J. Kovoor (MNHN), J.
Carico (JC), R. Leech (RL), Polska Akademia Nauk, Warszawa,
W. Starega, A. Riedel, J. Proszynski (PAN), Inventario Biologico
Nacional, Asuncion, J. A. Kochalka (IBNA), Museo de Inverte-
brados, Universita de Panama, D. Quintero A. (MIUP). T.
Yaginuma loaned a male Arachnura.

Literature Cited

Cambridge, O. P.- 1890. Arachnida, Araneidea. 1: 57-72 in Biologia Centrali-

Americana. Zoologia, London.

1895. Arachnida, Araneidea. 1: 145-160 in Biologia Centrali-Americana.

Zoologia, London.

1898. Arachnida, Araneidea. 1: 233-288 in Biologia Centrali-Americana.

Zoologia, London.

Cambridge, F. P.- 1904. Arachnida, Araneidea. 2: 465-545 in Biologia Centrali-

Americana. Zoologia, London.

G£taz, A. 1893. Fauna Arachnologica de Costa Rica. An. Inst, fis.- geogr. nac.
Costa Rica 4: 103-106.

Keyserling, E. 1865. Beitrage zur Kenntnis der Orbitelae. Verhandl. zool.-bot.
Gesellsch. Wien, 15: 799-856.

1893. Die Spinnen Amerikas, Nurnberg. 4: 209-377.

Petrunkevitch, A. 1930. The spiders of Porto Rico. Trans. Connecticut Acad.
Sci. 30: 159-355.

Roewer, C. Fr. 1942. Katalog der Araneae, Bremen, vol. 1.

Simon, E. 1895. Histoire Naturelle des Araignees, Paris, 1: 761-1084.

1903. Histoire Naturelle des Araignees, Paris. 2: 669-1080.

Taczanowski, L. 1873. Les Araneides de la Guyane frangaise. Horae Soc.
Entom. Rossicae 9: 64-150.

AN EYELESS SUBTERRANEAN BEETLE
( PSEUD A NO PHTH A LMUS)

FROM A KENTUCKY COAL MINE
(COLEOPTERA: CARABIDAE: TRECHINAE)*

By Thomas C. Barr, Jr.

School of Biological Sciences,

University of Kentucky
Lexington, Kentucky 40506

The trechine genus Pseudanophthalmus includes approximately
240 species from caves of the Appalachian valley, Mississippian
plateaus, and Bluegrass and Central Basin regions of eastern United
States. Although the model of cave trechine speciation which I have
developed for this fauna (Barr, 1967a, 1968, 1981, 1985) requires a
two-step process of 1) local diversification in deep soil and 2) subse-
quent isolation in nearby caves, the first stage was postulated on the
basis of an abundant edaphobitic trechine fauna in Europe and
elsewhere (see Jeannel, 1926-1930, for example). In eastern United
States a single species of Pseudanophthalmus has been described
from a non-cave habitat: P. sylvaticus occurs deep in the soil under
large stones in mountain forests near Marlinton, West Virginia
(Barr, 1967b).

Existing distributions of cave Pseudanophthalmus species strongly
suggest an ancestral Pleistocene refugium in the mixed meso-
phytic forests of the Allegheny plateau (Barr, 1981, 1985). The dis-
tinctly different lineages occupying caves of the Appalachian valley
to the east of the plateau and those of the Interior Low Plateaus to
the west of the Alleghenies indicate substantial local differentiation
prior to cave colonization (Barr, 1981); the geographic clustering of
related species suggests vicariance among cave descendants of these
locally differentiated edaphobites (Barr, 1965, 1981, 1985).

An integrated phylogeny of Pseudanophthalmus has thus far
proven elusive, as though key pieces of a jigsaw puzzle were missing.
Preliminary track analysis at the species group level thus shows a

* Manuscript received by the editor December 15, 1985

Psyche

[Vol. 93

void in the Allegheny plateau, with only the gracilis (east) and inex-
pectatus (west) groups clearly related by synapomorphic characters.
The engelhardti group (s. str., see Barr, 1981) does track through the
Allegheny plateau, but only via the Tennessee River gorge west of
Chattanooga. The large, pubescent, riparian species of the tenuis
group (IN, IL, KY) are superficially and ecologically similar to the
species of the grandis group (chiefly eastern WV), but there are
insufficient synapomorphies to provide substantive support to an
hypothesis of taxonomic affinity (Barr, 1985). It is tempting to
speculate that transitional species or species groups occupied the
non-limestone terranes of the interior of the Allegheny plateau. If
these transitional forms are extinct, no sound phylogeny of Pseuda-
nophthalmus may be possible. No caves occur in the thick sequences
of clastic rocks — sandstones, conglomerates, coals, and shales — in
this region. But if ancestral edaphobitic beetles are hypothetically
invoked throughout the Pleistocene to supply the caves on either
side of the Alleghenies, why should they suddenly become extinct
after Wisconsinan glaciation? Could some of these obligate soil
inhabitants still survive in deep, forest floor soil of this region? The
discovery of P. sylvaticus suggested that this could indeed be the
case, but two decades have elapsed without further edaphobitic tre-
chines being found.

Juberthie et al. (1980) demonstrated that “troglobitic” arthropods
exist in the “milieu souterrain superficiel” of non-karst regions in
southern France. At the interface between the soil mantle and the
bedrock there are air-filled pockets — microcaverns — from which
these authors have trapped several species of millipedes and beetles
(including trechines) that are for all intents and purposes “troglo-
bites,” even in non-calcareous terranes. However, attempts to trap
such organisms in eastern United States have met with failure. Suit-
able sites for trap insertion in France or Japan are in areas of frac-
tured rock (C. Juberthie and S.-I. Ueno, pers. comm.), unlike the
majority of karst regions in eastern United States. The traps are
baited pitfall traps containing Galt’s solution or equivalent; they are
placed about 1 m below the surface, buried, and checked at intervals
of 2-4 weeks.

On October 18, 1985, J. R. MacGregor and H. D. Bryan collected
2 female Pseudanophthalmus specimens in an abandoned coal mine

1986]

Barr — Subterranean beetle

in Floyd County, eastern Kentucky. The mine portal, designated
“D-104” in MacGregor’s notes, is located at Bosco (= Hueysville),
about 22 km SSW Prestonsburg. The beetles were found in a muddy
spot on the mine floor under rocks.

These two females are identical with females of Pseudanophthal-
mus hypolithos (Barr, 1981: 83, figs. 28, 34), a species previously
known only from Old Quarry Cave, in Pine Mountain, near Ash-
camp, Pike County, Kentucky, 45 km SE of the Bosco mine. The
hypolithos group, which includes 4 species from Pine Mountain,
KY, and a single species (P. praetermissus) near the base of Cumber-
land Mountain in Scott County, VA, belongs to the engelhardti
complex, a group of 55 largely Appalachian valley species arranged
in 7 species groups (Barr, 1981). Pseudanophthalmus hypolithos,
itself, is distinguished from other species of the group by quite deep
elytral striae and convex elytral intervals, greatly reduced pubes-
cence limited chiefly to sparse and very short rows on each elytral
interval, and falciform aedeagal apex. The aedeagal character could
not be checked, but based on my experience with species of the
genus, the absence of non-genitalic differences is decisive; only
2/240 species are determined solely on male genitalic characters.

Previously I had considered Pine Mountain as a “karst island”
within the Allegheny plateau; it is a fault block about 125 km long,
extending from Elkhorn City, Kentucky, southwest to Campbell
County, Tennessee, with a band of Newman limestone (Mississip-
pi) exposed on its northwest face. To the extent that “troglobitic”
Pseudanophthalmus species are collectable within the caves of Pine
Mountain, this is still true after a fashion, but the discovery of P.
hypolithos in a coal mine indicates that “caves” are a somewhat
artificial concept in terranes where highly fractured rocks (shales,
coals, conglomerates) exist, and that “troglobitic” trechines may
occur over a wider area than is strictly delimited by karst terrane.
The Bosco mine offers another sort of entry into the deep soil com-
munity, and the discovery of P. hypolithos there is a strong impetus
to search for other edaphobitic trechines within the interior of the
Allegheny plateau.

Acknowledgements

I thank John R. MacGregor for making these specimens available
for study. This paper was supported in part by NSF DEB-8202339.

Psyche

[Vol. 93

Literature Cited

Barr, Thomas C., Jr.

1965. The Pseudanophthalmus of the Appalachian valley (Coleoptera: Cara-
bidae: Trechini). Amer. Midi. Nat. 73: 41-72.

1967a. Observations on the ecology of caves. Amer. Nat. 101: 475-492.

1967b. A new Pseudanophthalmus from an epigean environment in West Vir-
ginia (Coleoptera: Carabidae). Psyche 74: 166-174.

1968. Cave ecology and the evolution of troglobites. Evolutionary Biology 2:
45-102. Appleton-Century-Crofts, New York.

1981. Pseudanophthalmus from Appalachian caves (Coleoptera: Carabidae):
the engelhardti complex. Brimleyana, no. 5: 37-94.

1985. Pattern and process in speciation of trechine beetles in eastern North
America (Coleoptera: Carabidae: Trechinae). IN G. E. Ball, ed., Taxon-
omy, Phylogeny, and Zoogeography of Beetles and Ants. Dordrecht,
Netherlands: Dr. W. Junk Publishers.

Jeannel, Ren£

1926-1930. Monographic des Trechinae: Morphologie comparee et distribu-
tion d’un groupe de Coleopteres. L’Abeille 32: 221-550; 33: 1-592; 34:
59-122; 35: 1-808.

Juberthie, C, B. Delay, and M. Bouillon

1980. Extension du milieu souterrain en zone non-calcaire: description d’un
nouveau milieu et de son peuplement par les Coleopteres troglobies.
Mem. Biospeol. 7: 19-52.

BICONUS IN PERU, WITH NOTICE OF
AN ENDEMIC SPECIES FROM THE COASTAL DESERT
(HYMENOPTERA: ICHNEUMONIDAE).

By Charles C. Porter1

Department of Biological Sciences, Fordham University
Bronx, NY 10458

Introduction

Taxonomy and Relationships

Townes (1969:178-9) places Biconus in his Subtribe Ischnina
(Ischnus, Trachysphyrus and allied mesostenine genera), where he
considers it related to Chromocryptus 2, Cryptopteryx, and Tra-
chysphyrus. Biconus, however, shows some features unapproached
or rarely approximated by members of the preceding genera. These
characters include absence of tyloids on the male flagellum,
medially bituberculate clypeus, profoundly cleft female 4th tarso-
mere, and tendency for loss or reduction of the brachiella vein. I
remain uncertain as to the affinities of Biconus. Comparative analy-
sis of mesostenine genera in all parts of the world probably will be
necessary to clarify this problem.

Collections

Specimens of Biconus have been or are to be deposited in the
following institutional and personal collections.

Cambridge. Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge, MA 02138.

Gainesville. Florida State Collection of Arthropods, Bureau of
Entomology, Division of Plant Industry, Florida Department
of Agriculture and Consumer Services, P. O. Box 1269, 1911
SW 34th Street, Gainesville, FL 32602.

'Research Associate, Florida State Collection of Arthropods, Florida Department of
Agriculture and Consumer Services, Division of Plant Industry, P.O. Box 1269,
Gainesville FL 32602.

Manuscript received by the editor May 24, 1985

2Townes’ concept of Chromocryptus includes the species of Trachysphyrus (sensu
Porter 1967) in which the axillus vein is close to the anal margin of the hind wing.

Psyche

[Vol. 93

Lawrence. Department of Entomology, Snow Entomological
Museum, The University of Kansas, Lawrence, KS 66045.
porter. Collection of Charles C. Porter, 301 North 39th Street,
McAllen, TX 78501.

townes. American Entomological Institute, c/o Dr. Virendra
Gupta, Bureau of Entomology, Division of Plant Industry,
Florida Department of Agriculture and Consumer Services,
Gainesville, FL 32602.

Genus BICONUS

Biconus Townes, 1969. Mem. Amer. Ent. Inst. 12: 178-9.

Type: Biconus atroruber Townes.

Fore wing 4. 1-10. 2mm long. Wings hyaline with dark brown
blotches. Female flagellum long and slender, not flattened below
apicad. Male flagellum without tyloids. Mandible moderately long
with lower tooth almost as long as upper. Clypeus 1. 5-2.0 as wide as
long, moderately and asymmetrically convex or weakly and sym-
metrically convex in profile; its apical margin subtruncate to a little
convex and usually with a pair of often inconspicuous median
preapical tubercles or swellings. Malar space: 0.72-1.0 as long as
basal width of mandible. Pronotum with epomia sharp but not
prolonged much dorsad or ventrad of scrobe. Mesoscutum with
notaulus sharp but fine, reaching more than 0.6 the length of meso-
scutum; surface mat with delicate puncto-reticulation and very
dense, short setae. Mesopleuron has no ridge on prepectus below.
Hind coxa with a strong and polished subvertical groove externo-
ventrally near base. Female tarsus with 4th segment very deeply
bilobed at apex. Wing venation: areolet large, symmetrically pen-
tagonal, intercubiti gently to moderately convergent dorsad, 2nd
abscissa of radius 1.0-1. 1 as long as 1st intercubitus; discocubitus
broadly angled, ramellus well developed to vestigial; mediella defi-
nitely arched; axillus close to and paralleling anal margin of hind
wing; brachiella sometimes short or absent. Propodeum with spira-
cle round to short-oval and with its apical trans-carina represented
only by conspicuous ligulo-cuneate, ligulate, ligulo-conic or even
conical cristae. First gastric tergite without a baso-lateral expan-
sion; ventral longitudinal carina traceable but often weak or obso-
lete on postpetiole and sometimes faint also on petiole; dorsal
carinae more or less suggested toward apex of petiole and on base of

1986]

Porter — Biconus in Peru

postpetiole, sometimes absent. Second tergite mat, usually with fine
and dense micro-reticulation but lacking discrete punctures and
almost without setae, but sometimes with fine and dense short setae
that originate in very tiny, inconspicuous punctures. Ovipositor

0. 30-0.45 as long as fore wing; straight, moderately slender to rather
stout; nodus weak but with a minute notch; ventral valve on tip with
sharp, well spaced inclivously oblique ridges.

Biconus occurs at moderate elevations in the Andes of tropical
South America from Ecuador to Bolivia. Many species inhabit
montane wet forests. They are most often collected by sweeping
undergrowth in areas with a flora characterized by tree ferns, a
woody arborescent element rich in Myrtaceae and Lauraceae, and
by strikingly diverse epiphytic bromeliads and orchids.

Biconus apoecus Porter (n. sp.) is the only species that frequents
relatively arid habitats. It is found in semihumid valleys of the west
Andean foothills along the Peruvian coast from near Lima north at
least as far as Trujillo. These valleys doubtless were much wetter
only 10,000 years ago during the most recent glacial maximum and
even today support a relict cloud forest vegitation.

Key to Peruvian Species of Biconus

1. Flagellum uniformly dark; fore wing with a median and an apical

brown area; mesosoma and gaster ferruginous with black on
mesoscutum, tegula, and most of scutellum; mesopleuron
mostly with delicate and irregular wrinkling; 2nd recurrent
0.7-0. 8 as long as 1st abscissa of cubitus; male 1st flagello-
mere with many but sparse and inconspicuous linear whitish

sensilla B. apoecus n. sp.

1/ Flagellum with a white band; fore wing with a single median
brown blotch; mesosoma and gaster brownish yellow to
orange; mesopleuron with much sharp, horizontal wrinkling;
2nd recurrent 0.4-0. 5 as long as 1st abscissa of cubitus; male
1st flagellomere with prominent and rather crowded linear
white sensilla 2. B. subflavus n. sp.

1 . Biconus apoecus Porter, new species
(Fig. 2, cf. Fig. 4)

Female. Color: antenna black with some pale brown on scape;
head black; mandible black except for dull brown subapically and

Psyche

[Vol. 93

brownish white on dorsal margin; palpi light dusky brown; meso-
scutum ferruginous with black on mesoscutum, tegula, and on most
of scutellum; gaster ferruginous with weak dusky staining on last
tergite; wings hyaline with a broad transverse median band occupy-
ing upper hind corner of median cell, basal 0.5 of discocubital cell,
2nd discoidal cell basad of ramellus, 1st brachial cell, base of 2nd
brachial cell, and (more faintly) apex of anal cell as well as with a
light brownish blotch that covers apical 0.3 of radial cell plus most
of 3rd cubital cell, 3rd discoidal cell, and (more faintly) part of
apical 0.2 of 2nd brachial cell; coxae ferruginous with blackish stain-
ing apicad or sometimes more extensively; trochantelli black with
ferruginous staining, especially below; front femur black above and
dull ferruginous below; mid femur black with dull ferruginous
throughout, or at least in part, dorso-anteriorly; hind femur black
with dull ferruginous staining basad, especially above; fore tibia
dusky ferruginous; mid and hind tibiae black; tarsi black.

Length of fore wing: 7. 8-9. 3 mm. First flagellomere: 8.0 as long

as deep at apex. Clypeus: with a pair of weak median preapical
tubercles but only slightly convex on apical margin beneath tuber-
cles; lateral margin broad and reflexed. Malar space: 0.94-1.0 as
long as basal width of mandible. Mesopleuron: in large part with
delicate and irregular wrinkling. Wing venation: radial cell 3. 2-3. 6
as long as wide; 2nd abscissa of radius 1.0 as long as 1st intercubitus;
ramellus inserted at basal 0.4 of discocubitus; bulla of 1st abscissa of
cubitus 0. 2-0.3 as long as entire vein; 2nd recurrent 0.7-0.8 as long
as 1st abscissa of cubitus; brachiella reaches 0.4-0. 6 the distance to
wing margin. Propodeum: rather short and high; basal face steeply
declivous, 0.9 as long as the almost vertical apical face; cristae large,
stout, conspicuously projecting, broadly ligulate, about 0.3 as long
as apical face of propodeum; surface on basal face distad of basal
trans-carina with at least some strong and oblique wrinkles laterad
but mesad more or less extensively more finely sculptured and on
apical face strongly trans-rugose laterad but mesad often less
strongly wrinkled or mostly smooth and shining. First gastric ter-
gite: postpetiole short and weakly expanded apicad, 1.4- 1.7 as wide
apically as long from spiracle to apex; ventro-lateral carina tracea-
ble throughout, sharp apicad on petiole and on postpetiole; dorso-
lateral carina sharp on postpetiole but gradually becoming weaker
basad on petiole; dorsal carinae traceable (not sharp) toward apex

1986]

Porter — Biconus in Peru

Figs. 1-3. Biconus. Fig. 1, Biconus subflavus. Paratype. Fore wing, showing
color pattern. Fig. 2, Biconus apoecus. Paratype. Fore wing, showing color pattern.
Fig. 3, Biconus subflavus. Paratype. Hind tarsomere 4, showing very deep median
apical emargination.

of petiole and on base of postpetiole; surface of postpetiole shining
with delicate microreticulation that is strongest laterad and fades
out toward apex. Gaster : stout fusiform. Ovipositor: sheathed por-
tion 0.35-0.44 as long as fore wing; tip 0.25-0.32 as high at notch as
long from notch to apex.

male, differs from female as follows: Color: scape brownish
white below and laterally; mandible more broadly pale brown to
whitish; palpi dull white; tegula partly reddish; front femur ferrugi-
nous with dusky staining above; mid femur extensively ferruginous
with irregular dusky staining below and apico-dorsally; hind femur
ferruginous on much of basal 0.3 (especially above) and mostly
black apically.

Length of fore wing: 9.8-10.2 mm. First flagellomere: 5. 5-5. 7 as
long as deep at apex; on apical 0.6 with numerous but inconspicu-
ous and well separated whitish linear sensilla. Clypeus: median
preapical tubercles stronger than in female; apical margin gently

Psyche

[Vol. 93

bisinuate beneath tubercles; profile weakly convex with highest
point near middle. Malar space: 0.72-0.77 as long as basal width of
mandible. Wing venation: brachiella sometimes reaches less than 0.5
the distance to wing margin. Propodeum: much as in female but a
little more elongate: basal face about 1.2 as long as the steeply
sloping apical face; cristae a little broader and stouter than in
female, ligulo-cuneate, very prominent; surface apicad of basal
trans-carina duller than in female with delicate reticulation and
more or less extensive moderately strong oblique wrinkling. First
gastric tergite: postpetiole elongate, parallel-sided, 1.0- 1.2 as wide
apically as long from spiracle to apex; dorsal carinae obsolete. Gas -
ter: rather strongly depressed.

Type Material. Holotype $: PERU, Lima Province, San
Geronimo, nr. Chosica, 1 — 5— VII-1 976, C. Porter, C. Calmbacher.
Paratypes: 5$, 2 <J, same date as holotype. Holotype in Florida State
Collection of Arthropods. Paratypes in Florida State Collection of
Arthropods (1$, 1(5), Collection of Henry K. Townes (1?), Museum
of Comparative Zoology (12), University of Kansas Collection (12),
Collection of Charles C. Porter (12, 1(5).

Relationships. This species appears closely related to the
Ecuadorian Biconus atroruber (Townes 1969:178-79), with which it
agrees in being ferruginous with black markings and in having a
median and an apical dark area on the fore wing. It differs from B.
atroruber by its entirely dark (instead of white banded) flagellum; in
having the mesosomatic black markings restricted to the mesoscu-
tum, tegula, and scutellum (instead of extending also onto the
pronotum, subalarum, mesosternum, upper metapleuron, and pro-
podeum); in its mostly ferruginous (instead of mostly black) coxae;
by its strongly (instead of finely) wrinkled apical propodeal face;
and in having the female propodeal cristae ligulate (instead of sub-
conic and decurved slightly at apex) and the male cristae ligulo-
cuneate (instead of high and cone-like).

Biconus apoecus may be distinguished from the central Peruvian
B. subflavus Porter by characters given in the key, as well as by its
shorter first flagellomere, more weakly tuberculate and apically less
convex clypeus, longer female malar space, shorter and higher
propodeum, and less definitely micro-reticulate postpetiole.

Field notes. San Geronimo, Peru, the type locality, is on the
lower west Andean slopes in the valley of the Santa Eulalia River

1986]

Porter — Biconus in Peru

not far from Chosica and Lima. The valley is well watered and
enjoys a warm microclimate because of its sheltered situation at an
altitude just above the point normally reached by nightly Pacific
coastal fogs during the coolest months of the year. Natural vegeta-
tion at San Geronimo includes Acacia, Salix, Schinus, Baccharis,
Tessaria and many other Andean, Chaquenan, and Holarctic gen-
era. Much of the valley is covered by orchards of chirimoyas, citrus,
apples, pears, and bananas. Irrigation ditches that traverse the
orchards permit growth of a lush herbaceous understory from which
Biconus apoecus and other ichneumonids may be swept.

Specific name. From the Greek adjective apoecus, ’’away from
home, abroad”.

2. Biconus subflavus Porter, new species
(Fig. 1,3).

Female. Color: antenna black with some dark brown on scape
and with a white annulus (extensively brown to black stained below)
on flagellomeres 3 (near apex) or 4-9 or 10 (basally); head black
with dark brown on clypeus and lighter brown on mandibular con-
dyle; mandible blackish with much brown to pale brown, especially
subapicad and dorsad; palpi dull white; mesosoma pale brownish
yellow, a little darker and more orangish dorsally; gaster pale brown-
ish to orangish yellow; wings hyaline with a single brown blotch that
covers basal 0.4 of discocubital cell, extends a little into base of 2nd
discoidal cell, and reaches below across most of 1st brachial cell;
legs pale brownish yellow with some darker staining, especially on
apices of trochantelli and bases of femora, as well as with 4th and
5th tarsomeres largely dark brown.

Length of fore wing: 8.1-10.1 mm. First flagellomere: 9. 3-9. 7 as
long as deep at apex. Clypeus: with a pair of broad but weak median
subapical swellings, apical margin moderately convex medially
beneath swellings, lateral margin not reflexed. Malar space:
0.80-0.87 as long as basal width of mandible. Mesopleuron: largely
with fine but sharp horizontally biased wrinkling. Lower metapleu-
ron: with strong, obliquely biased wrinkling. Wing venation: radial
cell 3.4-4. 1 as long as wide; 2nd abscissa of radius 1.0-1. 1 as long as
1st intercubitus; ramellus inserted near basal 0.3 of discocubitus;

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[Vol. 93

Fig. 4. Biconus atroruber. Female in lateral view, head in anterior view, propo-
deum and first 2 gastric tergites in dorsal view, and ovipositor tip in lateral view.
(From Townes, 1969:431).

bulla of 1st abscissa of cubitus 0.1 -0.2 as long as entire vein; 2nd
recurrent 0.4-0. 5 as long as 1st abscissa of cubitus; brachiella
reaches 0.4 or less the distance to wing margin (sometimes almost
absent). Propodeum: moderately elongate; basal face gently decli-
vous, 0.70-0.85 as long as the almost vertical apical face; cristae
stout, conspicuously projecting, conico-ligulate; surface on basal
face distad of basal trans-carina mat with uniformly strong reticu-
late wrinkling and on apical face with even stronger wrinkling. First
gastric tergite: postpetiole short but rather strongly expanded
apicad, 1.3- 1.5 as wide apically as long from spiracle to apex; ven-
tral longitudinal carina sometimes obsolete on petiole; dorsal cari-
nae weakly suggested above spiracles; surface of postpetiole
strongly shining with faint microreticulation. Gaster: moderately
elongate fusiform. Ovipositor: sheathed portion 0.35-0.41 as long as

1986]

Porter — Biconus in Peru

fore wing; tip 0.23-0.26 as high at notch as long from notch to apex.

Male, differs from female as follows: Color: white flagellar
annulus reaches from apex of 8th to base of 13th segment.

Length of fore wing: 9.6 mm. First flagellomere: 6.0 as long as
deep at apex; except near base with numerous and prominent,
rather crowded, linear white sensilla. Clypeus: tubercles more dis-
tinct and apical margin more strongly convex than in female; profile
rather strongly convex with highest point a little distad of middle.
Malar space: 0.82 as long as basal width of mandible. Wing vena-
tion: radial cell 3.1 as long as wide; brachiella absent. Propodeum:
basal face long but more strongly declivous than in female, 0.85 as
long as the almost vertical apical face; cristae a little stouter and
more conical than in female; surface distad of basal trans-carina
more coarsely and regularly wrinkled than in female. First gastric
tergite: postpetiole slender and parallel-sided, 0.91 as wide apically
as long from spiracle to apex. G aster: cylindric, not depressed.

Type material. Holotype $: PERU, Cuzco Province, Machu
Picchu, 1900 m, 4-19-IX-1964, C. Porter. Paratypes: 2?, 1<$, same
data as holotype. Holotype in Florida State Collection of Arthro-
pods. Paratypes in Florida State Collection of Arthropods (19, 1<5),
and Collection of Charles C. Porter (19).

Relationships. As indicated previously, this species differs
substantially in many points of color and structure from the other
described Biconus. It may be recognized at a glance by its orangish
ground color and unifasciate fore wing.

Field notes. The type locality is in cool tropical cloud forest.
Specimens of Biconus subflavus were swept from lush undergrowth
at the forest edge along the railway tracks which parallel the Uru-
bamba River.

Specific name. From the Latin adjective subflavus, “somewhat
yellow”.

Acknowledgments

This research was done under my National Science Foundation
Grants BSR-83 13444 and DEB-75-22426. It was also subsidized in
1974, 75, and 79 by grants from the Committee for Research and
Exploration of the National Geographic Society, which made pos-
sible fieldwork in the Peruvian Coastal Desert.

Psyche

[Vol. 93

I am also indebted to the Florida State Department of Agricul-
ture and Consumer Services, from whose Division of Plant Industry
I have received generous support mediated primarily by Dr. How-
ard V. Weems, Jr, Dr. Lionel A. Stange, and Mr. Harold A.
Denmark.

Summary

Biconus is a “trachysphyroid” mesostenine found in Andean wet
forests and in the Coastal Desert of Peru. It is recognizable by its
brown blotched wings; lack of tyloids on male flagellum; mat
mesoscutum; arched mediella; anally situate axillus; sharply grooved
hind coxal base; deeply cleft female 4th tarsomere; nearly round
propodeal spiracle; unarmed petiolar base; and subligulate to coni-
cal, prominent (but never spiniform) propodeal cristae. There are 3
species: B. atroruber Townes from Ecuador (white band on flagel-
lum, body ferruginous and black, fore wing with 2 brown areas); B.
apoecus n. sp. from the Peruvian Coastal Desert (similar to B.
atroruber but without a white flagellar band); and B. subflavus from
Peruvian montane forest (mesosoma and gaster orangish, fore wing
with 1 brown blotch).

Literature Cited

Porter, C.

1967. A revision of the South American species of Trachysphyrus. Mem.
Amer. Ent. Inst. 10: 1-386.

Townes, H. K.

1969. Genera of Ichneumonidae, Part 2: Gelinae. Mem. Amer. Ent. Inst. 12:
1-537.

A SYNONYMIC GENERIC CHECKLIST OF THE
EUMENINAE (HYMENOPTERA: VESPIDAE)*

By James M. Carpenter

Museum of Comparative Zoology, Harvard University,
Cambridge, MA 02138

The present work is an extension of a similar list in Carpenter
(1983), and arose from preparatory work for a phylogenetic analysis
of nearctic potter wasp genera (Carpenter and Cumming, 1985). The
most recent available world list of genera is over 80 years old (Dalla
Torre, 1904), and fully 57% of the genus-group names currently
used in the Eumeninae have been proposed since Bluethgen (1938;
for more detail on the history of eumenine taxonomy see Carpenter
and Cumming, 1985). The following checklist includes all the cur-
rently recognized genera of Eumeninae sensu Carpenter (1981), with
their synonyms and subgenera. The arrangement is alphabetical
based upon most recent usage, and incorporates the decisions per-
taining to eumenine generic nomenclature rendered by the Interna-
tional Commission on Zoological Nomenclature (ICZN) in Opinions
747 (1965), 893 (1970) and 1363 (1985). The format is basically that
of Krombein et al. (1979). The original citations are followed by the
type species designation. Synonyms, and subgenera with their cit-
ations and synonyms, are listed after this; the nominotypical sub-
genera are not listed separately. Where two dates are listed, the first
is the true date of publication, whereas the date listed in parentheses
is that printed on the paper. A misspelling is indicated by (!), and
quotation marks are used for incorrect names. No effort has been
made to list all misspellings; only those occurring in works consi-
dered important. Nomenclatural changes derive from ongoing work
on a catalog of neotropical eumeninae (with J. van der Vecht) and a
generic reclassification of this group: Neodiscoelius Stange is a jun-
ior objective synonym of Proto discoelius Dalla Torre (new synon-
ymy); Cephalastor Soika is raised to genus (new status), and its type
species, Hyp alast oroides depressus Soika, synonymized with Ody-
nerus relativus Fox. In addition, type species are designated for

* Manuscript received by the editor July 14, 1985

Psyche

[Vol. 93

Stenolabus Schulthess (junior subjective synonym of Ischnocoelia
Perkins) and Nesodynerus Perkins. These designations conform to
standard generic concepts. Two nomina dubia. and four nomina
nuda not otherwise placed are listed separately at the end of this
paper.

It is not to be inferred that I agree with this classification, but
considering the current confusion in eumenine taxonomy, a catalog
of the available names and their status is a prerequisite for rectifying
the situation.

Abispa Mitchell, 1838, Three. Exped. Interior Eastern Australia
1:104 (as subgenus of Vespa L.). Type species Abispa australi-
ana Mitchell, 1838. Monotypic.

Abisba (!) Ashmead, 1902, Can. Ent. 34: 208 (gives as type
Vespa ephippium Fabricius, 1775, originally not included).
Monerebia Saussure, 1852, Et. Fam. Vesp. 1: 98. Type species
Odynerus splendidus Guerin, 1838. Designated by Vecht,
1960, Nova Guinea Zool. 6: 92.

Monorebia (!) Smith, 1857, Cat. Hym. Brit. Mus. 5: 42.
Monerobia (!) Bridwell, 1919, Proc. Hawaiian Entomol. Soc. 4:
120.

subg. Parabispa Vecht, 1960, Nova Guinea Zool. 10(6): 93, 94.
Type species Pterochilus eximius Smith, 1865. Original
designation.

Acanthodynerus Gusenleitner, 1969, Boll. Mus. Civ. Ven. 19: 13.
Type species Acanthodynerus giordanii Gusenleitner, 1969.
Original designation.

Acarepipona Soika, 1985 (1983), Boll. Mus. Civ. Ven. 34: 189, 192.
Type species Acarepipona insolita Soika, 1985. Original
designation.

Acarodynerus Soika, 1962 (1961), Boll. Mus. Civ. Ven. 14: 64, 146.
Type species Odynerus clypeatus Saussure, 1853. Original
designation.

Acarozumia Bequaert, 1921, Rev. Zool. Afr. 9: 249 (as subgenus of
Montezumia Saussure). Type species Nortonia amaliae Saus-
sure, 1869. Monotypic.

Afrepipona Soika, 1965, Boll. Soc. Entomol. Ital. 95: 46. Type spe-
cies Odynerus macrocephalus Gribodo, 1894. Original desig-
nation.

1986]

Carpenter — Checklist of Eumeninae

Afreumenes Bequaert, 1926, Ann. S. Afr. Mus. 23: 486 (as subgenus
of Eumenes Latreille). Type species Eumenes melanosoma
Saussure, 1852. Monotypic.

Afrodynerus Soika, 1934, Ann. Mus. Civ. Genova 57: 25, 26 (as
subgenus of Odynerus Latreille). Type species Odynerus mon-
struosus Soika, 1934. Monotypic.

Afroxanthodynerus Soika, 1979, Boll. Mus. Civ. Ven. 30: 243. Type
species Afroxanthodynerus nigeriensis Soika, 1979. Original
designation.

Alastor Lepeletier, 1841, Hist. Nat. Ins. Hym. 2: 668. Type species
Alastor atropos Lepeletier, 1841. Designated by Ashmead,
1902, Can. Ent. 34:210.

Antalastor Saussure, 1856, Et. Fam. Vesp. 3: 328 (as division of
Alastor). Type species Alastor atropos Lepeletier, 1841.
Designated by ICZN, Opinion 893, 1970: 187.

Eualastor Dalla Torre, 1904, Gen. Ins. 19: 60. New name.
Belalastor Atanassov, 1967, Izv. Zool. Inst. Sof. 23: 167 (as
subgenus of Alastor). Type species Alastor bulgaricus Ata-
nassov, 1967 (= Alastor seidenstueckeri Bluethgen, 1956).
Original designation.

subg. Parastalor Bluethgen, 1939, Veroeff. Dts. Kolon. Uebersee-
Mus. Bremen 2: 264. Type species Alastor algeriensis
Bluethgen, 1939. Monotypic.

subg. Megalastor Bluethgen, 1951, Mitt. Muench. Entomol.
Ges. 41: 169. Type species Alastor savignyi Saussure, 1852.
Original designation.

Alastoroides Saussure, 1856, Et. Fam. Vesp. 3: 327 (as subgenus of
Alastor Lepeletier). Type species Alastor clot ho Lepeletier,
1841. Designated by Ashmead, 1902, Can. Ent. 34: 210.
Paralastoroides Saussure, 1856, Et. Fam. Vesp. 3: 328 (as div-
ision of subgenus Alastoroides Saussure of genus Alastor
Lepeletier). Type species Alastor clotho Lepeletier, 1841.
Monotypic. Rejected by ICZN, Opinion 893, 1970: 188, in
favor of Alastoroides.

Alasteroides (!) Zavattari, 1912, Arch. Naturgesch. 78A(4):
255.

Alastorynerus Bluethgen, 1938 (1937), Konowia 16: 294. Type spe-
cies Odynerus ludendorffi Dusmet, 1917. Original designation.

64 Psyche [Vol. 93

Alastodynerus (!) Parker, 1966, Misc. Publ. Entomol. Soc.
Am. 5: 157.

Alfieria Soika, 1934, Bull. Soc. Entomol. Egypte 18: 436. Type
species Eumenes anomalus Zavattari, 1909. Original designa-
tion.

Alferia (!) Neave, 1939, Nomencl. Zool. 1: 111.

Allodynerus Bluethgen, 1938 (1937), Konowia 16: 280 (as subgenus
Saussure, 1853. Original designation, (as “Lionotus floricola
Sauss. 1852”).

Delphinaloides Moczar, 1937, Folia Ent. Hung. 3: 15. Invalid;
no type designated. Made available by Bohart, 1951, in
Muesebeck et al., Cat. Hym. N. Am.: 888; with type species
Odynerus delphinalis Giraud, 1866.

Allorhynchium Vecht, 1963, Zool. Verh. (Leiden) 60: 57, 58. Type
species Vespa argentata Fabricius, 1804. Original designation.
Alphamenes Vecht, 1977, Proc. K. Ned. Akad. Wet. (C) 80: 238,
242. Type species Eumenes campanulatus Fabricius, 1804.
Original designation.

Alphamenes Bertoni, 1934, Rev. Soc. Cient. Paraguary 3: 109
(as subgenus of Eumenes Latreille). Invalid; no type desig-
nated.

Ancistroceroides Saussure, 1855, Et. Fam. Vesp. 3: 221 (as division
of subgenus Ancistrocerus Wesmael of genus Odynerus La-
treille; validated by ICZN, Opinion 893, 1970: 187. Type spe-
cies Odynerus alastoroides Saussure, 1853. Designated by
ICZN, Opinion 1363, 1985: 353.

Ancistrocerus Wesmael, 1836, Bull. Acad. Sci. Bruxelles 3: 45 (as
subgenus of Odynerus Latreille). Type species Vespa parietum
L., 1758. Designated by Girard, 1879, Traite Elem. Ent. 2(2):
900.

Aucistrocerus (!) Rudow, 1876, Arch. Ver. Freunde Natur-
gesch. Mecklenb. 30: 197.

Ancystrocerus (!) Dalla Torre, 1894, Cat. Hym. 9: 50 ff.
Euancistrocerus Dalla Torre, 1904, Gen. Ins. 19: 36. New
name.

Antamenes Soika, 1958 (1957), Boll. Mus. Civ. Ven. 10: 214. Type
species Odynerus flavocinctus Smith, 1857 (= Odynerus verna-
lis Saussure, 1853). Original designation,
subg. Australochilus Soika, 1962 (1961), Boll. Mus. Civ. Ven.

1986]

Carpenter — Checklist of Eumeninae

14: 184. Invalid; no type designated. Made available by
Soika, 1974 (1973), Boll. Mus. Civ. Ven. 24: 53; with type
species Odynerus citreocinctus Saussure, 1867.

Antepipona Saussure, 1855, Et. Fam. Vesp. 3: 244 (as division of
subgenus Odynerus of genus Odynerus Latreille; validated by
ICZN, Opinion 893, 1970: 187). Type species Odynerus silaos
Saussure, 1853. Designated by ICZN, Opinion 893, 1970: 187.
Antepiponus Saussure, 1875, Smiths. Misc. Coll. 254: xxxv,
361. Emendation.

Antepipone (!) Dalla Torre, 1894, Cat. Hym. 9: 50, 96.
Mehelyella Moczar, 1937, Folia Ent. Hung. 3: 16. Invalid; no
type designated. Made available by Bohart, 1951, in Muese-
beck et al., Cat. Hym. N. Am.: 888; with type species Odyne-
rus parvulus Lepeletier, 1841.

Odontodynerus Bluethgen, 1938 (1937), Konowia 16: 280 (as
subgenus of “Euodynerus Bluethgen”). Type species Odyne-
rus orbitalis Herrich-Schaeffer, 1841. Original designation.
Dichodynerus Bluethgen, 1938. Dts. Entomol. Z.: 444. Type
species Odynerus vagabundus Dalla Torre, 1889. Original
designation (as “ Lionotus vagus Radoszkowsi (= vagabun-
dus Dalla Torre nom. nov.)”).

Metastenancistrocerus Bluethgen, 1938, Dts. Entomol. Z.: 460.
Error for Dichodynerus; cf. Bluethgen, 1939, Veroeff. Dts.
Kolon. Uebersee-Mus. Bremen 2: 246.

Anterhynchium Saussure, 1863, Mem. Soc. Phys. Hist. Nat. Geneve
17: 205 (as division of Rhynchium Spinola). Type species Ryg-
chium synagroides Saussure, 1852. Designated by Vecht, 1963,
Zool. Verh. (Leiden) 60: 73.

Anterrhynchium (!) Dalla Torre, 1904, Gen. Ins. 19: 33.
subg. Dirhynchium Vecht, 1963, Zool. Verh. (Leiden) 60: 74,
77. Type species Ancistrocerus flavopunctatus Smith, 1852.
Original designation.

Antezumia Saussure, 1875, Smiths. Misc. Coll. 254: 1 13 (as division
of Montzumia Saussure). Type species Montezumia chalybea
Saussure, 1855. Designated by Bequaert, 1921, Rev. Zool.
Afric. 9: 240.

Pinta Zavattari, 1912, Arch. Naturgesch. 78A(4): 6, 151. Type
species Montezumia chalybea Saussure, 1855. Original
designation.

Psyche

[Vol. 93

Antodynerus Saussure, 1855, Et. Fam. Vesp. 3: 242, 287 (as division
of subgenus Odynerus of genus Odynerus Latreille; validated
by ICZN, Opinion 893, 1970: 187). Type species Vespa flaves-
cens Fabricius, 1775 (“ Odynerus punctum (Fabricius)” sensu
Saussure, 1853). Designated by ICZN, Opinion 893, 1970: 187.
Kalliepipona Soika, 1952 (1951), Riv. Biol. Colon. 11: 81 (as
subgenus of Pseudepipona Saussure). Type species Rhyn-
chium radiale Saussure, 1855 (as “ Odynerus radialis ’). Orig-
inal designation.

Pseudokalliepipona Soika, 1955, Ann. Mus. R. Congo Beige
Tervuren, Zool. 36: 366 (as subgenus of Pseudepipona Saus-
sure, 1853. Type species Odynerus bellatulus Saussure, 1853.
Original designation.

Parepipona Soika, 1957, Brit. Mus. (Nat. Hist.) Exped. S. W.
Arabia 1(31): 477 (as subgenus of Pseudepipona Saussure).
Type species Rhynchium radiale Saussure, 1855 (as “Odyne-
rus radialis9). Original designation.

Anthodynerus (!) Soika, 1961, South Afr. Anim. Life 8: 445.
Araucodynerus Willink, 1968 (1967), Acta Zool. Lilloana 22: 143,
152. Type species Odynerus tuberculatus Saussure, 1853. Orig-
inal designation.

Argentozethus Stange, 1979, Acta Zool. Lilloana 35: 729. Type spe-
cies Argentozethus willinki Stange, 1979. Original designation.
Asiodynerus Kurzenko, 1977, Ins. Mongolia 5: 557. Type species
Odynerus lucifer Kostylev, 1937. Original designation.

Astalor Schulthess, 1925, Konowia 4: 59, 207 (as subgenus of Alas-
tor Lepeletier). Type species Astalor maidli Schulthess, 1925.
Monotypic.

Astator (!) Schulthess, 1925, Konowia 4: 208.

Australodynerus Soika, 1962 (1961), Boll. Mus. Civ. Ven. 14: 65,
114. Type species Odynerus pusillus Saussure, 1856. Original
designation.

Australozethus Soika, 1969, Boll. Mus. Civ. Ven. 19: 27, 29. Type
species Australozethus tasmaniensis Soika, 1969. Original
designation.

Bidentodynerus Soika, 1977 (1976), Mem. Soc. Entomol. Ital. 55:
177. Type species Odynerus bicolor Saussure, 1855. Original
designation.

Brachymenes Soika, 1961, Verh. XI Int. Kongr. Entomol. Wien:
243. Type species Eumenes wagnerianus Saussure, 1875. Origi-
nal designation.

1986]

Carpenter — Checklist of Eumeninae

Brachyodynerus Bluethgen, 1938, Dts. Entomol. Z.: 450, 459. Type
species Odynerus magnificus Morawitz, 1867. Original desig-
nation.

Brachypipona Gusenleitner, 1967, Polskie Pismo Ent. 37: 671. Type
species Pseudepipona schmidti Gusenleitner, 1967. Original
designation.

Desertodynerus Kurzenko, 1977, Zool. Zh. 56(6): 957. Type
species Desertodynerus gratus Kurzenko, 1977. Original
designation.

Calligaster Saussure, 1852, Et. Fam. Vesp. 1: 22. Type species Cal-
ligaster cyanopterus Saussure, 1852. Designated by Ashmead,
1902, Can. Ent. 34: 205.

Cephalastor Soika, 1982 (1981), Boll. Mus. Civ. Ven. 32: 33, 40 (as
subgenus of Hypalastoroides Saussure); NEW STATUS. Type
species Hypalastoroides depressus Soika, 1969 (= Odynerus
relativus Fox, 1902; NEW SYNONYMY). Original designa-
tion.

Cephalochilus Bluethgen, 1939, Mitt. Entomol. Ges. Halle 17: 13.
Type species Pterochilus grandis Lepeletier, 1841 (= Vespa
labiata Fabricius 1798). Original designation.

Cephalodynerus Parker, 1965, Ann. Entomol. Soc. Am. 58: 364.
Type species Cephalodynerus unicornis Parker, 1965. Original
designation.

Chelodynerus Perkins, 1902, Trans. Entomol. Soc. Lond.: 136.
Type species Odynerus chelifer Perkins, 1899. Monotypic.

Chlorodynerus Bluethgen, 1951, Boll. Soc. Entomol. Ital. 81: 67, 75
(as subgenus of “ Euodynerus Bluethgen”). Type species Odyne-
rus chloroticus Spinola, 1838. Original designation.

Coeleumenes Vecht, 1963, Zool. Verh. (Leiden) 60: 16, 45. Type
species Montezumia impavida Bingham, 1897. Original desig-
nation.

Ctenochilus Saussure, 1856, Et. Fam. Vesp. 3: 323 (as division of
Pterochilus (!) Klug). Type species Epipona pilipalpa Spinola,
1851. Monotypic.

Cuyodynerus Willink, 1968(1967), Acta Zool. Lilloana22: 143, 151.
Type species Odynerus cuyanus Brethes, 1903. Original desig-
nation.

Cyphodynerus Vecht, 1971, Entomol. Ber. (Amst.) 31: 127. Type
species Odynerus dimidiatus Spinola, 1838 ( non Odynerus di-
midiatus Guerin, 1834; = Odynerus canaliculatus Saussure,
1855). Original designation.

Psyche

[Vol. 93

Cyphomenes Soika, 1978, Boll. Mus. Civ. Ven. 29: 13, 210. Type
species Eumenes infernalis Saussure, 1875. Original desig-
nation.

Cyrtolabulus Vecht, 1969, Entomol. Ber. (Amst.) 29: 1. New name
for Cyrtolabus Vecht.

Cyrtolabus Vecht, 1963, Zool. Verh. (Leiden) 60: 1 1 non Cyrto-
labus Voss, 1925. Type species Cyrtolabus suavis Vecht,
1963. Original designation.

Delta Saussure, 1855, Et. Fam. Vesp. 3: 130, 143 (as division of
Eumenes Latreille). Type species Vespa maxillosa DeGeer,
1773 (= Vespa emarginata L., 1758). Designated by Be-
quaert, 1925, Bull. Brook. Entomol. Soc. 20: 137 (as <(Sphex
maxillosus’).

Phi Saussure, 1855, Et. Fam. Vesp. 3: 132 (as division of
Eumenes Latreille) non Phi Saussure, 1854. Type species
Vespa arcuata Fabricius, 1775. Designated by Bequaert,
1926, Ann. S. Afr. Mus. 23: 487.

Erinys Zirngiebl, 1953, Mitt. Pollichia (3)1: 173 non Erinys
Rye, 1876. Type species Vespa unguiculata Villers, 1789.
Monotypic.

Deuterodiscoelius Dalla Torre, 1904, Gen. Ins. 19: 18 (as division of
Discoelius Latreille). Type species Odynerus verrauxii Saus-
sure, 1852. Monotypic.

Pseudozethus Perkins, 1914, Pr. Zool. Soc. Lond.: 622. Type
species Pseudozethus australensis Perkins, 1914 (= Odyne-
rus verrauxii Saussure, 1852). Monotypic.

Diemodynerus Soika, 1962 (1961), Boll. Mus. Civ. Ven. 14: 65, 141.
Type species Odynerus diemensis Saussure, 1853. Original
designation.

Discoelius Latreille, 1809, Gen. Crust. Ins. 4: 140 (as subgenus of
Eumenes Latreille). Type species Vespa zonalis Panzer, 1801.
Monotypic.

Discaelius (!) Leach, 1815, Edinburgh Encyc. 9: 153.

Discaelias (!) Leach, 1815, Edinburgh Encyc. 9: 166.

Dicoelius (!) Haliday, 1836, Trans. Linn. Soc. Lond. 17: 325.
Discollius (!) Froggatt, 1892, Proc. Linn. Soc. N.S.W. 2(7):
226.

Tritodiscoelius Dalla Torre, 1904, Gen. Ins. 19: 18 (as division
of Discoelius ). Type species Vespa zonalis Panzer, 1801.

1986]

Carpenter — Checklist of Eumeninae

Designated by Bequaert and Ruiz, 1942 (1940), Rev. Chil.
Hist. Nat. 64: 217.

Dolichodynerus Bohart, 1939, Pan-Pac. Ent. 15: 97, 101 (as subge-
nus of Odynerus Latreille). Type species Odynerus turgiceps
Bohart, 1939. Original designation.

Ectopioglossa Perkins, 1912, Ann. Mag. Nat. Hist. (8)9: 118. Type
species Ectopioglossa australensis Perkins, 1912 {non Eumenes
australensis Meade-Waldo, 1910; = Ectopioglossa polita aus-
tralensis (Meade-Waldo)). Monotypic.

Elimus Saussure, 1852, Et. Fam. Vesp. 1: 7. Type species Elimus
australis Saussure, 1852. Monotypic.

Elisella Soika, 1974 (1972), Boll. Mus. Civ. Ven. 25: 109, 132. Type
species Ellisella linae Soika, 1974. Original designation.
Epiodynerus Soika, 1958 (1957), Boll. Mus. Civ. Ven. 10: 195 (as
subgenus of Pseudepipona Saussure). Type species Odynerus
alecto Lepeletier, 1841. Original designation.

Epsilon Saussure, 1855, Et. Fam. Vesp. 3: 229 (as division of subge-
nus Odynerus of genus Odynerus Latreille; validated by ICZN,
Opinion 893, 1970: 187). Type species Odynerus dyscherus
Saussure, 1852. Designated by ICZN, Opinion 893, 1970: 187.
Eudiscoelius Friese, 1904, Z. Hym. Dipt. 4: 16. Type species Eudis-
coelius metallicus Friese, 1904. Monotypic.

Euchalcomenes Turner, 1908, Trans. Entomol. Soc. Lond.: 90.
Type species Euchalcomenes gilberti Turner, 1908. Original
designation.

Eumenes Latreille, 1802, Hist. Nat. Crust. Ins. 3: 360. Type species
Vespa coarctata L., 1758. Designated by Latreille, 1810, Con-
sid. Gen. Crust. Arach. Ins.: 328.

Alpha Saussure, 1855, Et. Fam. Vesp. 3: 128, 137 (as division of
Eumenes) non Alpha Saussure, 1854. Type species Vespa
coarctata L., 1758. Designated by Bequaert, 1926, Ann. S.
Afr. Mus. 23: 435.

Eumenis Kriechbaumer, 1879, Entomol. Nachr. 5: 57. Emen-
dation.

Eumenidion Schulthess, 1913, Soc. Entomol. 28: 2 (as subge-
nus). Type species Vespa coarctata L., 1758. Original
designation.

Eumenidium (!) Sharp, 1915, Zool. Rec. Ins. 1913: 275.
subg. Zeteumenoides Soika, 1972, Boll. Soc. Entomol. Ital.

Psyche

[Vol. 93

104: 110 (as genus). Type species Eumenes filiformis Saus-
sure, 1855 (= Eumenes versicolor filiformis). Original
designation.

Eumenidiopsis Soika, 1939 (1938), Mem. Soc. Entomol. Ital. 17: 87
(as subgenus of Leptomenes Soika). Type species Leptomenes
subtilis Soika, 1939. Original designation.

Eumicrodynerus Gusenleitner, 1972, Nachrbl. Bayer. Ent. 21: 74 (as
subgenus of Microdynerus Thomson). Type species Lepto-
menes europaeus Soika, 1942. Original designation.
Euodynerus Dalla Torre, 1904, Gen. Ins. 19: 38 (as section of sub-
genus “Lionotus” Thomson of genus Odynerus Latreille; vali-
dated by ICZN, Opinion 893, 1970: 187. Type species Vespa
dantici Rossi, 1790. Designated by Bluethgen, 1938 (1937),
Konowia 16: 277 .

subg. Pareuodynerus Bluethgen, 1938 (1937), Konowia 16: 278
(as subgenus of “Euodynerus Bluethgen”). Type species
Vespa notata Jurine, 1807. Original designation.

Leionotus Saussure, 1851, Et. Fam. Vesp. 1: 121 (as subgenus
of Odynerus Latreille), non Leionotus Kirby and Spence,
1828. Type species Odynerus foraminatus Saussure, 1853.
Designated by Bohart, 1951, in Muesebeck et al., Cat.
Hym. N. Am.: 887.

Lionotus (!) Thomson, 1874, Opusc. Ent. 2: 85, non Lionotus
Agassiz, 1846.

Lejonotus (!) Costa, 1882, Atti. R. Acad. Sci. Fis. Mat. Napoli
9: 37.

Eustenancistrocerus Bluethgen, 1938, Dts. Entomol. Z.: 443, 460 (as
subgenus of “ Stenancistrocerus Saussure” sensu Bluethgen,
1938). Type species Odynerus blanchar dianus Saussure, 1855.
Original designation.

subg. Hemistenancistrocerus Bluethgen, 1938, Dts. Entomol.
Z.: 443, 459 (as subgenus of “Stenancistrocerus Saussure”
sensu Bluethgen, 1938). Type species Leptochilus parvulus
Saussure, 1853 ( non Odynerus parvulus Herrich-Schaeffer,
1938; = Odynerus pharao Saussure, 1863). Original desig-
nation.

subg. Parastenancistrocerus Bluethgen, 1938, Dts. Entomol.
Z.: 444, 460 (as subgenus of “Stenancistrocerus Saussure”
sensu Bluethgen, 1938). Type species Odynerus transitorius
Morawitz, 1867. Original designation.

1986] Carpenter — Checklist of Eumeninae 71

Flammodynerus Soika, 1962 (1961), Boll. Mus. Civ. Ven. 14: 65,
124. Type species Odynerus subalaris Saussure, 1855. Original
designation.

Gamma Zavattari, 1912, Arch. Naturgesch. 78A(4): 85 (as division
of Eumenes Latreille). Type species Pachymenes ventricosa
Saussure, 1852. Designated by Bequaert, 1926, Ann. S. Afr.
Mus. 23: 486.

Gastrodynerus Bohart, 1984, Pan-Pac. Ent. 60: 12. Type species
Stenodynerus vanduzeei Bohart, 1948. Original designation.

Gioiella Soika, 1985 (1983), Boll. Mus. Civ. Venezia 34: 30, 155.
Type species Odynerus katonai Schulthess, 1913. Original
designation.

Gribodia Zavattari, 1912, Arch. Naturgesch. 78A(4): 161. Type spe-
cies Monobia cavifrons Gribodo, 1891 (= Odynerus confluen-
tus Smith, 1857). Original designation.

Gymnomerus Bluethgen, 1938 (1937), Konowia 16: 286 (as subge-
nus of “Hoplomerus (Westwood) Agassiz” sensu Bluethgen,
1938). Type species Odynerus laevipes Shuckard, 1837. Origi-
nal designation.

Hemipterochilus Ferton, 1909 (1908), Ann. Soc. Entomol. Fr. 77:
572 (as subgenus of Pterocheilus Klug). Type species Odynerus
terricola Mocsary, 1883 (= Hemipterochilus bembeciformis
terricola). Monotypic.

Pseudopterochilus Kostylev, 1940, Bull. Soc. Nat. Moscou,
Biol. (N.S.) 49: 153. Invalid; no type designated. Made avail-
able by Vecht, 1972, in Vecht and Fischer, Hym. Cat. 8: 19;
with type species Odynerus bembeciformis Morawitz, 1867.

Hypalastoroides Saussure, 1856, Et. Fam. Vesp. 3: 328 (as division
of subgenus Alastoroides Saussure of genus Alastor Lepeletier;
validated by ICZN, Opinion 893, 1970: 187). Type species Alas-
tor brasiliensis Saussure, 1856. Monotypic.

Hypalastor Saussure, 1856, Et. Fam. Vesp. 3: 328 (as division
of subgenus Alastor of genus Alastor Lepeletier; validated
by ICZN, Opinion 893, 1970: 187). Type species Odynerus
angulicollis Spinola, 1851. Designated by ICZN, Opinion
893, 1970: 187. Rejected by Soika, 1960 (1958) Boll. Mus.
Civ. Ven. 11: 35, acting as first reviser, in favor of
Hypalastoroides.

Hypalasteroides (!) Zavattari, 1912, Arch. Naturgesch. 78A(4):
253.

Psyche

[Vol. 93

subg. Larastoroides Soika, 1982 (1981), Boll. Mus. Civ. Ven.
32: 33, 40. Type species Hypalastoroides costaricensis Soika,
1960. Original designation.

subg. Ortalastoroides Soika, 1982 (1981), Boll. Mus. Civ. Ven.
32: 34, 56. Type species Alastor singularis Saussure, 1852.
Original designation.

Ortastoroides (!) Soika, 1982 (1981), Boll. Mus. Civ. Ven. 32:
57.

Hypancistrocerus Saussure, 1855, Et. Fam. Vesp. 3: 222 (as division
of subgenus Ancistrocerus Wesmael of genus Odynerus La-
treille; validated by ICZN, Opinion 893, 1970: 187). Type spe-
cies Odynerus advena Saussure, 1855. Monotypic.
Hypancistroceroides (!) Saussure, 1856, Et. Fam. Vesp. 3,
Table des Matieres: 8.

Hypancystrocerus (!) Dalla Torre, 1894, Cat. Hym. 9: 50.
Hypodynerus Saussure, 1855, Et. Fam. Vesp. 3: 225 (as division of
subgenus Odynerus of genus Odynerus Latreille; validated by
ICZN, Opinion 893, 1970: 187). Type species Odynerus hume-
ralis Haliday, 1836. Designated by Bequaert and Ruiz, 1943
(1941), Rev. Chil. Hist. Nat. 45: 69.

Hypodernus (!) Cameron, 1908, Trans. Am. Entomol. Soc.
34: 199.

Incodynerus Willink, 1968 (1967), Acta Zool. Lilloana 22: 143, 148.
Type species Odynerus romandinus Saussure, 1853. Original
designation.

Ischnocoelia Perkins, 1908, Proc. Hawaiian Entomol. Soc. 2: 28, 32.
Type species Ischnocoelia xanthochroma Perkins, 1908. Mono-
typic.

Stenolabus Schulthess, 1910, Dts. Entomol. Z.: 189. Type spe-
cies Stenolabus fulvus Schulthess, 1910. By present desig-
nation.

Ischnogasteroides Magretti, 1884 (1883), Boll. Soc. Entomol. Ital.
15: 251; 1884, Ann. Mus. Civ. Stor. Nat. Genova 21: 603. Type
species Ischnogasteroides flavus Magretti, 1884 (= Ischnogas-
teroides leptogaster flavus). Monotypic.

Jucancistrocerus Bluethgen, 1938, Dts. Entomol. Z.: 442, 460 (as
subgenus of “ Stenancistrocerus Saussure” sensu Bluethgen,
1938). Type species Odynerus jucundus Mocsary, 1883. Origi-
nal designation.

1986]

Carpenter — Checklist of Eumeninae

lucancistrocerus (!) Bluethgen, 1951, Mitt. Muench. Entomol.
Ges. 41: 174.

subg. Eremodynerus Bluethgen, 1939, Veroeff. Dts. Kolon.
Uebersee-Mus. Bremen 2: 257 (as genus). Type species
Odynerus saharensis Soika, 1934. Original designation.
Katamenes Meade-Waldo, 1910, Ann. Mag. Nat. Hist. (8)5: 46.
Type species Katamenes watsoni Meade-Waldo, 1910. Mono-
typic.

Knemodynerus Bluethgen, 1940, Entomol. Tidskr. 61:43 (as subge-
nus of “Euodynerus Bluethgen”). Type species Odynerus
excellens Perez, 1907. Original designation.

Labochilus Bluethgen, 1939, Mitt. Entomol. Ges. Halle 17: 12. Type
species Pterochilus linguarius Saunders, 1905. Monotypic.
Leptopterocheilus Soika, 1953 (1952), Bull. Soc. Sci. Nat.
Phys. Maroc. 32: 262. Type species Pterochilus linguarius
Saunders, 1905. Original designation.

Labus Saussure, 1867, Zool. Novara 2, Hym.: 3. Type species Labus
spiniger Saussure, 1867. Designated by Bingham, 1897, Fauna
Brit. India Hym. 1: 348.

Laevimenes Soika, 1978, Boll. Mus. Civ. Ven. 29: 11, 359. Type
species Eumenes laevigatus Brethes, 1906. Original designa-
tion.

Leptochiloides Bohart, 1940, Ann. Entomol. Soc. Am. 33: 165.
Type species Leptochiloides utahensis Bohart, 1940. Original
designation.

Leptochilus Saussure, 1853, Et. Fam. Vesp. 1: 233. Type species
Pterochilus mauritanicus Lepeletier, 1841. Designated by
Ashmead, 1902, Can. Ent. 34: 209 (as “ mauritianus”\ ).
Zendalia Robertson, 1928, Flowers and Insects: 12. Type spe-
cies Odynerus zendaloides Robertson, 1928 (= Leptochilus
republicanus Dalla Torre, 1889). Designated by Bohart,
1951, in Muesebeck et al. Cat. Hym. N. Am.: 897.
subg. Euleptochilus Bluethgen, 1943, in Berland, Bull. Mus.
Hist. Nat. Paris (2)15: 316. Type species Odynerus oraniensis
Lepeletier, 1841. Original designation,
subg. Lionotulus Bluethgen, 1938 (1937), Konowia 16: 276.
Type species Odynerus alpestris Saussure, 1855. Original
designation.

subg. Neoleptochilus Bluethgen, 1961, Abh. Dts. Akad. Wiss.

Psyche

[Vol. 93

Berl. (2): 66, 100. Type species “Leptochilus medanae (Gri-
bodo i.l.) Andre” (= Odynerus medanae Gribodo, 1886).
Original designation.

subg. Sarochilus Gusenleitner, 1970, Isr. J. Entomol. 5: 57.
Type species Leptochilus alterego Gusenleitner, 1970. Origi-
nal designation.

Leptodynerus Bluethgen, 1938, Dts. Entomol., Z.: 448, 457. Type
species Leptodynerus biskrensis Bluethgen, 1938. Original
designation.

Leptomenes Soika, 1939 (1938), Mem. Soc. Entomol. Ital. 17: 87.
Type species Pachymenes congensis Bequaert, 1918 (= Odyne-
rus eumenoides Smith, 1857). Original designation.

Leptomenoides Soika, 1962 (1961), Boll. Mus. Civ. Ven. 14: 64, 171.
Type species Leptomenoides placidior Soika, 1962. Original
designation.

Leptomicrodynerus Soika, 1985, Lavori Soc. Ven. Sc. Nat. 10: 37.
Type species Leptomicrodynerus tieshengi Soika, 1985. Origi-
nal designation.

Leucodynerus Bohart, 1982, J. Kans. Entomol. Soc. 55: 442. Type
species Odynerus congressus Viereck, 1908. Original designa-
tion.

Macrocalymma Perkins, 1908, Proc. Hawaiian Entomol. Soc. 2: 28,
31. Type species Macrocalymma smithianum Perkins, 1908.
Monotypic.

Maricopodynerus Viereck, 1908, Trans. Am. Entomol. Soc. 33: 397
(as subgenus of Odynerus Latreille). Type species Odynerus
maricoporum Viereck, 1908. Monotypic.

Micreumenes Ashmead, 1902, Can. Ent. 34: 208. Type species
Micreumenes currei Ashmead, 1902 (in key). Monotypic.
Smithia Saussure, 1855, Rev. Mag. Zool. 7: 371 non Smithia
Edwards and Haime, 1851. Type species Smithia natalensis
Saussure, 1855. Monotypic.

Hymenosmithia Dalla Torre, 1904, Gen. Ins. 19: 61. New name
for Smithia Saussure.

Microdynerus Thomson, 1874, Hym. Scand. 3: 58. Type species
Odynerus exilis Herrich-Schaeffer, 1839. Designated by Jones,
1937, Entomol. Mon. Mag. 73: 15.

subg. Pseudomicrodynerus Bluethgen, 1938 (1937), Konowia
16: 276. Type species Odynerus parvulus Herrich-Schaeffer,
1838. Original designation.

1986]

Carpenter — Checklist of Eumeninae

Pachymicrodynerus Bluethgen, 1938, Dts. Entomol.: 447, 455
(as subgenus of Pseudomicrodynerus). Type species Pseu-
domicro dyner us eurasius Bluethgen, 1938. Original designa-
tion.

Pseudomycrodynerus (!) Gusenleitner, 1977, Linz. Biol. Beitr.
9: 138.

Minixi Soika, 1978, Boll. Mus. Civ. Ven. 29: 14, 367. Type species
Eumenes mexicanus Saussure, 1857. Original designation.

Mitrodynerus Vecht, 1981, Proc. K. Ned. Akad. Wet. (C) 84: 444.
Type species Mitrodynerus vitripennis Vecht, 1981. Mono-
typic.

Monobia Saussure, 1852, Et. Fam. Vesp. 1: 94. Type species
Vespa quadridens L., 1763. Designated by Ashmead, 1902,
Can. Ent. 34: 210.

Triart hr a Dalla Torre, 1904, Gen. Ins. 19: 28 (as group of
Monobia ), non Triarthra Ehrenberg, 1832. Type species
Odynerus cyanipennis Guerin, 1830. Designated by Be-
quaert, 1940, Rev. Entomol. 11: 822.

Tetrathra Dalla Torre, 1904, Gen. Ins. 19: 28. Type species
Vespa quadridens L., 1763. Designated by Bequaert, 1940,
Rev. Entomol. 11: 822.

Tetrarthra (!) Bequaert, 1940, Rev. Entomol. 11: 822.

Monodynerus Gusenleitner, 1982, Entomofauna 3: 279. Type spe-
cies Monodynerus insimilis Gusenleitner, 1982. Original desig-
nation.

Montezumia Saussure, 1852, Et. fam. Vesp. 1: 87. Type species
Montezumia rufidentata Saussure, 1852 (= Odynerus azures-
cens Spinola, 1851). Designated by Ashmead, 1902, Can. Ent.
34: 207.

Alpha Saussure, 1855, Et. Fam. Vesp. 3: 160 (as division of
Montezumia ), non Alpha Saussure, 1854. Type species Mon-
tezumia rufidentata Saussure, 1952 (= Odynerus azurescens
Spinola, 1851). Designated by Bohart, 1951, in Muesebeck et
ah, Cat. Hym. N. Am.: 885.

Beta Saussure, 1855, Et. Fam. Vesp. 3: 162 (as division of
Montezumia). Type species Montezumia morosa Saussure,
1852. Designated by Bequaert, 1921, Rev. Zool. Afric. 9:
240.

Metazumia Saussure, 1875, Smiths. Misc. Coll. 254: 114 (as
division of Montezumia ). Type species Montezumia huas-

76 Psyche [Voi. 93

teca Saussure, 1857. Designated by Bequaert, 1921, Rev.
Zool. Afric. 9: 240.

Eumontezumia Dalla Torre, 1904, Gen. Ins. 19: 27. New name.
Nesodynerus Perkins, 1901, Entomol. Mon. Mag. 37: 267. Type
species Odynerus rudolphi Dalla Torre, 1889. By present
designation.

Nortozumia Vecht, 1937, Treubia 16: 263. Type species Zethus
rufofemoratus Cameron, 1903. Original designation.

Odynerus Latreille, 1802, Hist. Nat. Crust. Ins. 3: 362. Type species
Vespa spinipes L., 1758. Designated by Shuckard, 1837, Mag.
Nat. Hist. (N. S.) 1: 494.

Odynera Illiger, 1807, Magaz. Insektenk. 6: 196. Emendation.
Epipone Kirby and Spence, 1815, Introd. Entomol. 1: 340, non
“epipone” Latreille 1802, a vernacular name. Type species
Vespa spinipes L., 1758. Monotypic.

Oplopus Wesmael, 1836, Bull. Acad. Sci. Bruxelles 3: 45 (as
subgenus of Odynerus ), non Oplopus Laporte, 1832. Type
species Vespa spinipes L., 1758. Designated by Girard, 1879,
Traite Elem. Ent. 2(2): 902.

Oplomerus Westwood, 1840, Intro. Mod. Classif. Ins. 2(Syn-
opsis): 84. New name for Oplopus Wesmael; non Oplomerus
Dejean, 1833, a nomen nudum.

Hoplomerus Agassiz, 1846, Nomencl. Zool. Index Univ.: 185.

Emendation of Oplomerus Westwood.

Hoplopus Agassiz, 1846, Nomencl. Zool. Index Univ.: 186.
Emendation of Oplopus Wesmael, non Hoplopus D’Or-
bigny, 1838.

Epiponus Saussure, 1875, Smiths. Misc. Coll. 254: 360 (as sub-
genus of Odynerus Latreille). Emendation of “Epipona
Shuckard” sensu Saussure, 1855, an incorrect spelling of
Epipone Kirby and Spence.

Hoplonus (!) Dalla Torre, 1889, Ent. Almanach.: 11.
Euepipona Dalla Torre, 1904, Gen. Ins. 19: 39. New name for
Epiponus Saussure. Type species Vespa spinipes L., 1758.
Designated by Richards, 1937, Gen. Names Br. Ins. 5: 128.
subg. Allogymnomerus Bluethgen, 1951, Mitt. Muench. Ento-
mol. Ges. 41: 174 (as subgenus of Hoplomerus Westwood).
Type species Odynerus consobrinus Dufour, 1839. Original
designation-.

1986]

Carpenter — Checklist of Eumeninae

subg. Monoplomerus Bluethgen, 1941, Arch Naturgesch. (N.
F.) 10: 308 (as subgenus of Hoplomerus Westwood). Type
species Hoplomerus caroli Morawitz, 1885. Original desig-
nation.

subg. Spinicoxa Bluethgen, 1938 (1937), Konowia 16: 285 (as
subgenus of “ Hoplomerus (Westwood) Agassiz” sensu
Bluethgen, 1938). Type species Vespa reniformis Gmelin,
1790. Original designation.

Omicroides Soika, 1935, Ann. Mus. Civ. Genova 57: 129 (as subge-
nus of Eumenes Latreille). Type species Eumenes singular is
Smith, 1857. Original designation.

Omicron Saussure, 1855, Et. Fam. Vesp. 3: 133, 148 (as division of
Eumenes Latreille). Type species Zethus ? globicollis Spinola,
1841. Designated by Bequaert, 1926, Ann. S. Afr. Mus. 23: 486.
Beta Saussure, 1875, Smiths. Misc. Coll. 254: 88 (as division of
Eumenes Latreille) non Beta Saussure, 1855. Type species
Eumenes nortonianus Saussure, 1875. Designated by Be-
quaert, 1926, Ann. S. Afr. Mus. 23: 486.

Amphimenes Bertoni, 1923, Rev. Soc. Cient. Paraguay 1: 53,
non Amphimenes Bates, 1873. Type species Eumenes toto-
nacus Saussure, 1875. Monotypic.

Onychopterocheilus Bluethgen, 1955, Mitt. Muench. Ges. 44/45: 407
(as subgenus of Pterocheilus). Type species Odynerus daw
Dusmet, 1903. Original designation.

Orancistrocerus Vecht, 1963, Zool. Verh. (Leiden) 60: 58, 99. Type
species Odynerus drewseni Saussure, 1857. Original desig-
nation.

Oreumenes Bequaert, 1926, Ann. S. Afr. Mus. 23: 488 (as subgenus
of Eumenes Latreille). Type species Eumenes harmandi Perez,
1905 (= Eumenes decoratus Smith, 1852). Original desig-
nation.

Oreumenoides Soika, 1961, Verh. XI Int. Kongr. Entomol. Wien:
245. Type species Eumenes edwardsi Saussure, 1852. Original
designation.

Ovodynerus Soika, 1985 (1983), Boll. Mus. Civ. Ven. 34: 31, 130.
Type species Odynerus capicola Meade Waldo, 1915. Original
designation.

Pachodynerus Saussure, 1870, Rev. Mag. Zool. 22: 56 (as division
of subgenus Odynerus of genus Odynerus Latreille: vali-

Psyche

[Vol. 93

dated by ICZN, Opinion 893, 1970: 187). Type species Odyne-
rus calif ornicus Saussure, 1870. Designated by Bohart, 1951, in
Muesebeck et ah. Cat. Hym. N. Am.: 892.

Pachyodynerus (!) Dalla Torre, 1894, Cat. Hym. 9: 82.
Monobiella Ashmead, 1900, Trans. Entomol. Soc. Lond.: 312
(as genus). Type species Vespa atrata Fabricius, 1798.
Monotypic.

Pachyodernus (!) Cameron, 1908, Trans. Am. Entomol. Soc.
34: 199.

Pachycoelius Soika, 1969, Boll. Mus. Civ. Ven. 19: 28, 54. Type
species Pachycoelius brevicornis Soika, 1969. Original desig-
nation.

Pachymenes Saussure, 1852, Et. Fam. Vesp. 1: 73. Type species
Pachymenes sericea Saussure, 1852. Designated by Ashmead,
1902, Can. Ent. 34: 208.

Pachimenes (!) Saussure, 1855, Et. Fam. Vesp. 3: 153.
Pachyminixi Soika, 1978, Boll. Mus. Civ. Ven. 29: 14, 387. Type
species Eumenes sumichrasti Saussure, 1875. Original desig-
nation.

Parachilus Soika, 1961 (1960), Atti Soc. Ital. Sci. Nat. 99: 389, 392.
Type species Pterochilus capensis Saussure, 1854. Original
designation.

Paragymnomerus Bluethgen, 1938 (1937), Konowia 16: 286 (as sub-
genus of “ Hoplomerus (Westwood) Agassiz” sensu Bluethgen,
1938). Type species Odynerus spiricornis Spinola, 1808. Origi-
nal designation.

Paralastor Saussure, 1856, Et. Fam. Vesp. 3: 328 (as division of
subgenus Alastor of genus Alastor Lepeletier; validated by
ICZN Opinion 893, 1970: 187). Type species Alastor tubercula-
tus Saussure, 1853, Designated by ICZN, Opinion 893, 1970:
187.

Paraleptomenes Soika, 1970, Boll. Mus. Civ. Ven. 20/21: 79. Type
species Paraleptomenes nurseanus Soika, 1970. Original desig-
nation.

Paralionotulus Bluethgen, 1938 (1937), Konowia 16: 293. Type spe-
cies Leptochilus mervensis Radoszkowski, 1887. Original
designation.

Pseudolionotulus (!) Bluethgen, 1938, Dts. Entomol. Z.: 446,
454. Type species Leptochilus mervensis Radoszkowski,

1986]

Carpenter — Checklist of Eumeninae

1887. Original designation.

Paramischocyttarus Magretti, 1884 (1883) Boll. Soc. Entomol. Ital.
15: 250; 1884, Ann. Mus. Civ. Genova 21: 600. Type species
Paramischocyttarus subtilis Magretti, 1884. Monotypic.
Tanyzethus Cameron, 1910, Wiss. Ergebn. Schwed. Zool.
Exped. Kilimandjaro (8)6: 195. Type species Tanyzethus
africanus Cameron, 1910. Monotypic.

Parancistrocerus Bequaert, 1925, Trans. Am. Entomol. Soc. 51: 64
(as subgenus of Ancistrocerus Wesmael). Type species Odyne-
rus fulvipes Saussure, 1855. Original designation.

Paranortonia Bequaert, 1940, Ann. Entomol. Soc. Am. 33: 100 (as
subgenus of Pachymenes Saussure). Type species Nortonia tol-
teca Saussure, 1875. Original designation.

Paranortonia Bertoni, 1934, Rev. Soc. Cient. Paraguay 3: 109
(as genus). Invalid; no type designated.

Pararhaphidoglossa Schulthess, 1910, Dts. Entomol. Z.: 187. Type
species Pararhaphidoglossa fulva Schulthess, 1910. Original
designation.

Pararaphidoglossa (!) Zavattari, 1912, Arch. Naturgesch.
78A(4): 5; Soika, 1941, Boll. Soc. Ven. Stor. Nat. 2: 227 and
1978, Boll. Mus. Civ. Ven. 29.

Pararrhynchium Saussure, 1855, Et. Fam. Vesp. 3: 173 (as division
of Rhynchium Spinola). Type species Rhynchium ornatum
Smith, 1852. Monotypic.

Prorhynchium Saussure, 1855, Et. Fam. Vesp. 3: 174 (as di-
vision of Rhynchium Spinola). Type species Rhynchium
smithii Saussure, 1855. Monotypic.

Prorrhynchium (!) Saussure, 1856, Et. Fam. Vesp. 3: 348, Table
des Matieres: 8.

Pararhynchium (!) Saussure, 1862, Stett. Entomol. Ztg. 23:
182.

Parrhynchium (!) Dalla Torre, 1894, Cat. Hym. 9: 42.

Paravespa Radoszkowski, 1886, Hor. Soc. Entomol. Ross. 20: 46.
Type species Hoplomerus komarowi Radoszkowski, 1886 (=
Odynerus quadricolor Morawitz, 1885). Monotypic.

Theletor Kokujev, 1912, Izv. Kavkaz. Muz. 7: 4 (under descrip-
tion of Rhynchium caucasicum). Type species Rhynchium
caucasicum Kokujev, 1912. Designated by Vecht, 1972, in
Vecht and Fischer, Hym. Cat. 8: 4.

Psyche

[Vol. 93

subg. Gestrodynerus Soika, 1961 (1960), Atti Soc. Ital. Sci.
Nat. 99: 361, 369. Type species Rygchium gestroi Magretti,
1884. Original designation.

Parazumia Saussure, 1855, Et. Fam. Vesp. 3: 166 (as division of
Montezumia Saussure). Type species Odynerus carinulatus
Spinola, 1851. Designated by Bequaert, 1921, Rev. Zool. Afric.
9: 241.

Pareumenes Saussure, 1855, Et. Fam. Vesp. 3: 133 (as division of
Eumenes Latreille). Type species Eumenes quadrispinosus
Saussure, 1855. Designated by Bequaert, 1918, Bull. Am. Mus.
Nat. Hist. 39:271.

subg. Nortonia Saussure, 1869, Stett. Entomol. Z. 30: 53 (as
genus). Type species Odynerus intermedius Saussure, 1853.
Original designation.

Notonia (!) Sonan, 1938, Arb. Morph. Tax. Ent. 5: 70.
Parifodynerus Soika, 1962 (1961), Boll. Mus. Civ. Ven. 14: 64, 167.
Type species Parifodynerus parificus Soika, 1962. Original
designation.

Parodontodynerus Bluethgen, 1938 (1937), Konowia 16: 280 (as
subgenus of “Euodynerus” Bluethgen). Type species Eumenes
ephippium Klug, 1817. Original designation.
Paradontodynerus (!) Guichard, 1978, Entomol. Gazette 31:
45.

Parodynerus Saussure, 1855, Et. Fam. Vesp. 3: 245 (as division of
subgenus Odynerus of genus Odynerus Latreille; validated by
ICZN, Opinion 893, 1970: 187). Type species Vespa bicincta
Fabricius, 1781. Designated by Soika, 1958 (1957), Boll. Mus.
Civ. Ven. 10: 214.

Pirhosigma Soika, 1978, Boll. Mus. Civ. Ven. 29: 11, 229. Type
species Eumenes simulans Saussure, 1875. Original designation.
Plagiolabra Schulthess, 1903 (March), Verh. Zool. Bot. Ges. Wien
53: 361, 365. Type species Plagiolabra nigra Schulthess, 1903.
Monotypic.

Leontiniella Brethes, 1903 (Sept.), An. Mus. Nac. Buenos Aires
(3)2: 265. Type species Leontiniella argentina Brethes, 1903.
Monotypic.

Postepipona Soika, 1974 (1972), Boll. Mus. Civ. Ven. 25: 77. Type
species Postepipona socotrae Soika, 1974. Original designa-
tion.

1986] Carpenter — Checklist of Eumeninae 81

Proepipona Soika, 1977, Steenstrupia 4: 125, 126. Type species
Vespa lateralis Fabricius, 1781. Original designation.
Protodiscoelius Dalla Torre, 1904, Gen. Ins. 19: 18 (as division of
Discoelius Latreille). Type species “Epipona chilensis Spinola,
1851 = Discoelius merula Haliday, 1836”. Designated by
Bequaert and Ruiz, 1942 (1940), Rev. Chil. Hist. Nat. 64: 217.
Neodiscoelius Stange, 1979, Acta Zool. Lilloana 35: 729; NEW
SYNONYMY. Type species Discoelius merula Haliday,
1836. Original designation.

Pseudabispa Vecht, 1960, Nova Guinea Zool. 10(6): 91, 102. Type
species Odynerus abispoides Perkins, 1912. Original desig-
nation.

Pseudacaromenes Soika, 1978, Boll. Mus. Civ. Ven. 29: 15 (in key).
Type species Eumenes alfkeni Ducke, 1904. Original desig-
nation.

Pseudoacaromenes (!) 1981, Zool. Record 115 (1978) Ins.: 262,
List of new generic and subgeneric names: 15, 39.
Pseudalastor Soika, 1962 (1961), Boll. Mus. Civ. Ven. 14: 65, 131.
Type species Odynerus concolor Saussure, 1853. Original
designation.

Pseudepipona Saussure, 1856, Et. Fam. Vesp. 3: 309 (as division of
subgenus “Epipona” of genus Odynerus Latreille; validated by
ICZN, Opinion 893, 1970: 187). Type species Odynerus herri-
chii Saussure, 1856. Monotypic.

Metepipona Bluethgen, 1951, Mitt. Muench. Entomol. Ges. 41:
193 (as subgenus of Pseudepipona). Type species Odynerus
peculiaris Morawitz, 1895. Original designation.
Trichepipona Bluethgen, 1951, Mitt. Muench. Entomol. Ges.
41: 171, 193 (as subgenus of Pseudepipona ). Type species
Odynerus lativentris Saussure, 1855. Original designation.
Leptepipona Bluethgen, 1951, Mitt. Muench. Entomol. Ges. 41:
171, 194 (as subgenus of Pseudepipona). Type species Vespa
tripunctata Fabricius, 1787. Original designation.
Pseudopipona (!) Opinion 893, ICZN, 1970, Bull. Zool.
Nomencl. 26: 187.

Pseudepipone (!) Bytinski-Salz and Gusenleitner, 1971, Isr. J.
Ent. 6: 298.

subg. Deuterepipona Bluethgen, 1951, Mitt. Muench. Entomol.
Ges. 41: 171, 194 (as genus). Type species Odynerus ionius
Saussure, 1855. Original designation.

82 Psyche [Vol. 93

Pseudochilus Saussure, 1856, Et. Fam. Vesp. 3: 321 (as division of
“ Pterochilus” Klug). Type species Pterochilus glabripalpis
Saussure, 1852. Monotypic.

Pseudodontodynerus Bluethgen, 1939, Veroeff, Dts. Kolon. Ueber-
see-Mus. Bremen 2: 249. Type species Odynerus pretiosus
Dusmet, 1928. Monotypic.

Pseudodynerus Saussure, 1855, Et. Fam. Vesp. 3: 220 (as division of
subgenus Ancistrocerus Wesmael of genus Odynerus Latreille;
validated by ICZN, Opinion 893, 1970: 187). Type species
Odynerus luctuosus Saussure, 1855. Monotypic.
Pseudoleptochilus Bluethgen, 1938 (1937), Konowia 16: 294. Type
species Odynerus frenchi Dusmet, 1917. Original designation
(as “Lionotus frenchi Dusmet”).

Pseudonortonia Soika, 1936, Ann. Mus. Civ. Genova 59: 268. Type
species Odynerus difformis Saussure, 1853. Original desig-
nation.

Subancistroceroides Bluethgen, 1938, Dts. Entomol. Z.: 441,
460 (as subgenus of “ Sub ancistrocerus Sauss ” sensu Blueth-
gen, 1938). Type species Odynerus aegyptiacus Saussure,
1863. Original designation.

Pseudopterocheilus Perkins, 1901, Entomol. Mon. Mag. 37: 266.
Type species Odynerus pterocheiloides Perkins, 1899. Original
designation.

Pseudopterochilus (!) Dalla Torre, 1904, Gen. Ins. 19: 39.
Pseudosymmorphus Bluethgen, 1938 (1937), Konowia 16: 293. Type
species Odynerus hindenburgi Dusmet, 1917. Original desig-
nation.

Pseudozumia Saussure, 1875, Smiths. Misc. Coll. 254: 128 (as di-
vision of Montezumia Saussure). Type species Montezumia
indica Saussure, 1855. Monotypic.

Pseudzumia (!) Dalla Torre, 1894, Cat. Hym. 9: 38.

Pseumenes Soika, 1935, Ann. Mus. Civ. Genova 57: 145 (as subge-
nus of Pareumenes Saussure). Type species Eumenes eximius
Smith, 1861. Original designation.

Psiliglossa Saunders, 1872, Trans. R. Entomol. Soc. Lond.: 42. New
name for Stenoglossa Saussure.

Stenoglossa Saussure, 1852, Et.Fam. Vesp. 1: 4, non Steno-
glossa Chaudoir, 1848. Type species Raphiglossa odyne-
roides Saunders, 1850. Monotypic.

Psiloglossa Dalla Torre, 1894, Cat. Hym. 9: 8. Emendation.

1986]

Carpenter — Checklist of Eumeninae

Pterocheilus Klug, 1805, Beitr. Natuurk. 1: 143. Type species Vespa
phalerata Panzer, 1797. Designated by Blanchard, 1840, in
Laporte, Hist. Nat. Ins. 3: 389.

Pterochilus (!) Illiger, 1807, Mag. Insektenk. 6: 196.

Pterochile (!) Blanchard, 1840, in Laporte, Hist. Nat. Ins. 3:
389.

Pterochylus (!) Saussure, 1853, Et.Fam. Vesp. 1: 239.
Odontopterochilus Kostylev, 1940, Bull. Soc. Nat. Moscou,
Biol. (N. S.) 49: 148 (as subgenus of Pterocheilus). Invalid;
no type designated. Made available by Vecht, 1971, Entomol.
Ber. (Amst.) 31: 127; with type species Pterocheilus heptneri
Kostylev, 1940.

Nannopterochilus Bluethgen, 1961, Ab. Dts. Akad. Wiss. Berl.
1961: 62, 86, 231. Type species Vespa phalerata Panzer, 1797.
Original designation.

subg. Megapterocheilus Bohart, 1940, Ann. Entomol. Soc.
Am. 33: 169, 173. Type species Pterochilus mirandus Cres-
son, 1879. Original designation,
subg. Onchopterocheilus Bohart, 1940, Ann. Entomol. Soc.
Am. 33: 169, 191. Type species Pterochilus comptus Cresson,
1879. Original designation.

subg. Micropterocheilus Bohart, 1940, Ann. Entomol. Soc.
Am. 33: 168, 201. Type species Pterocheilus desertorum
Bohart, 1940. Original designation.

Pteromenes Soika, 1961 (1960), Atti Soc. Ital. Sci. Nat. 99: 389, 407.
Type species Pterochilus paradisiacus Soika, 1941. Original
designation.

Raphiglossa Saunders, 1850, Trans. R. Entomol. Soc. Lond. (2)1:
71. Type species Raphiglossa eumenoides Saunders, 1850.
Designated by Ashmead, 1902, Can. Ent. 34: 206.
Raphidoglossa Dalla Torre, 1894, Cat. Hym. 9: 7. Emendation.
Raphiglossoides Soika, 1936, Boll. Soc. Entomol. Ital. 68: 77. Type
species Raphiglossoides aethiopicus Soika, 1936. Original
designation.

Rhynchalastor Meade-Waldo, 1910, Ann. Mag. Nat. Hist. (8)6(31):
110. Type species Rhynchalastor fuscipennis Meade-Waldo,
1910. Monotypic.

Rhynchium Spinola, 1806, emendation of Rygchium Spinola, 1806;
validated by ICZN, Opinion 747, 1965: 186. Type species Ryg-

Psyche

[Vol. 93

chium (!) europaeum Spinola, 1806 (= Vespa oculata Fabri-
cius, 1781). Monotypic.

Rygchium Spinola, 1806, Ins. Ligur. 1: 84 incorrect original
spelling for Rhynchium.

Rhynchium Billberg, 1820, Enum. Ins.: 109. Emendation of
Ry chium (!) Spinola.

Rynchium Sturm, 1829, Verz. Ins. Nurnberg: 12. Emendation.
Rhygchium Saussure, 1853, Et. Fam. Vesp. 1: xxxi, 276.
Emendation.

Rhynchuium (!) Saussure, 1863, Mem. Soc. Phy. Hist. Nat.
Geneve 17: 242.

Eurrhynchium Dalla Torre, 1904, Gen. Ins. 19: 33. New name.
Rygohium (!) Willink, 1982, Bol. Ac. Nac. Sci. 55: 195.
Smeringodynerus Snelling, 1975, Proc. Entomol. Soc. Wash. 77: 56.
Type species Odynerus morelios Saussure, 1857. Original
designation.

Sphaeromenes Soika, 1978, Boll. Mus. Civ. Ven. 29: 12, 225. Type
species Sphaeromenes discrepatus Soika, 1978. Original desig-
nation.

Spinilabochilus Kurzenko, 1981, Hym. Far East: 81, 97. Type spe-
cies Spinilabochilus turcmenicus Kurzenko, 1981. Original
designation.

Stellepipona Soika, 1974 (1973), Boll. Mis. Civ. Ven. 24: 106. Type
species Odynerus stellenboschensis Cameron, 1905. Original
designation.

Stenancistrocerus Saussure, 1863, Mem. Soc. Phys. Hist. Nat.
Geneve 17: 216 (as division of subgenus Ancistrocerus Wes-
mael of genus Odynerus Latreille; validated by ICZN, Opinion
893, 1970: 187). Type species Odynerus atropos Lepeletier,
1841. Designated by Bequaert, 1925, Trans. Am. Entomol.
Soc. 51: 63.

Stenancystrocerus (!) Dalla Torre, 1894, Cat. Hym. 9: 55-95.
Atr op ancistrocerus Bluethgen, 1938, Dts. Entomol. Z.: 442,
444, 461. Type species Odynerus hispanicus Dusmet, 1903.
Original designation.

subg. Paratropancistrocerus Bluethgen, 1938, Dts. Entomol.
Z.: 442. 461 (as subgenus of Atropancistrocerus). Type spe-
cies Odynerus transcaspicus Kostylev, 1935. Original desig-
nation.

1986]

Carpenter — Checklist of Eumeninae

Stenodyneriellus Soika, 1962 (1961), Boll. Mus. Civ. Ven. 14: 65, 71.
Type species Stenodyneriellus turneriellus Soika, 1962. Origi-
nal designation.

Stenodyneroides Soika, 1940, Ann. Mus. Civ. Genova 60: 471 (as
subgenus of Odynerus Latreille). Type species Odynerus corvus
Meade-Waldo, 1915. Original designation.

Stenodynerus Saussure, 1863, Mem. Soc. Phys. Hist. Nat. Geneve
17: 228 (as division of subgenus Odynerus of genus Odynerus
Latreille; validated by ICZN, Opinion 893, 1970: 187). Type
species Odynerus chinensis Saussure, 1863. Designated by
Bohart, 1939, Pan-Pac. Ent. 15: 100.

Stemodynerus (!) Rohwer, 1913, Proc. U.S. Nat. Mus. 44: 445.
Nannodynerus Bluethgen, 1938 (1937), Konowia 16: 281 (as
subgenus of “ Euodynerus Bluethgen”). Type species Liono-
tus teutonicus Bluethgen, 1937. Original designation.
Parhypodynerus Soika, 1974 (1973), Boll. Mus. Civ. Ven. 24:
110. Type species Odynerus pavidus Kohl, 1905. Original
designation.

Stenonartonia Soika, 1974 (1973), Boll. Mus. Civ. Ven. 24: 25. New
name for Paranortonia Soika.

Paranortonia Soika, 1941, Boll. Soc. Ven. Stor. Nat. 2: 25 non
Paranortonia Bequaert, 1940. Type species Nortonia poly-
bioides Schulthess, 1904. Original designation.

Stenosigma Soika, 1978, Boll. Mus. Civ. Ven. 29: 14, 407. Type
species Eumenes allegrus Zavattari, 1912. Original designation.

Stroudia Gribodo, 1892 (1891), Boll. Soc. Entomol. Ital. 23: 262.
Type species Stroudia armata Gribodo, 1892. Monotypic.

Subancistrocerus Saussure, 1855, Et. Fam. Vesp. 3: 206 (as division
of subgenus Ancistrocerus Wesmael of genus Odynerus La-
treille; validated by ICZN, Opinion 893, 1970: 187). Type spe-
cies Odynerus sichelii Saussure, 1854. Designated by Bequaert,
1925, Trans. Am. Entomol. Soc. 51: 61.

Ep ancistrocerus Saussure, 1856, Et. Fam. Vesp. 3: 352. New
name for Subancistrocerus Saussure. Type species Odynerus
sichelii Saussure, 1854. Designated by Bequaert, 1925,
Trans. Am. Entomol. Soc. 51: 61.

Symmorphoides Soika, 1977 (1976), Boll. Mus. Civ. Ven. 28: 171,
172. Type species Symmorphoides maroccanus Soika, 1976.
Original designation.

Psyche

[Vol. 93

Symmorphus Wesmael, 1836, Bull. Acad. Sci. Bruxelles 3: 45 (as
subgenus of Odynerus Latreille). Type species Odynerus ele-
gans Wesmael, 1833. Designated by Richards, 1935, Trans. R.
Entomol. Soc. Lond. 83: 162.

Protodynerus Saussure, 1855, Et. Fam. Vesp. 3: 184, 186. New
name for Symmorphus.

Synomorphus (!) Rohwer, 1917, Proc. U.S. Nat. Mus. 53: 234.
“Odynerus Latreille” sensu Bluethgen, 1938 (1937), Konowia
16: 274, 291. Type species Vespa muraria L., 1758. Desig-
nated by Bluethgen, 1938 (1937), Konowia 16: 274, 291.
Koptodynerus Bluethgen, 1943, Stett. Entomol. Z. 104: 152 (as
subgenus of “Odynerus Latreille” sensu Bluethgen). Type
species Symmorphus declivus Harttig, 1932. Monotypic.
subg. Parasymmorphus Cumming and Vecht, 1986, Entomol.
Ber. (Amst.) 46: 23. Type species Odynerus momunganensis
Schulthess, 1934. Original designation.

Synagris Latreille, 1802, Hist. Nat. Crust. Ins. 3: 360. Type species
Vespa cornuta L., 1758 (as Vespa cornuta F.). Monotypic.
Eusynagris Dalla Torre, 1904, Gen. Ins. 19: 30. New name for
Synagris.

Catilostenus Meunier, 1888, Nat. Sicil. 7: 150. Type species
Catilostenus nigroviolaceus Meunier, 1888. Monotypic. Iden-
tity doubtful.

subg. Paragris Saussure, 1855, Et. Fam. Vesp. 3: 156 (as div-
ision of Synagris). Type species Synagris humberti Saussure,
1855. Designated by Ashmead, 1902, Can. Ent. 34: 210 (as P.
hubertii !).

Hypagris Saussure, 1855, Et. Fam. Vesp. 3: 157 (as division
of Synagris). Type species Synagris abdominalis Saussure,
1855 (= Synagris analis Saussure, 1852). Designated by
Ashmead, 1902, Can. Ent. 34: 210.

Antagris Saussure, 1863, Mem. Soc. Phys. Hist. Nat. Geneve
17: 181 (as division of Synagris). Type species Synagris
aequatorialis Saussure, 1852 (= Synagris spiniventris
Illiger, 1802). Designated by Ashmead, 1902, Can. Ent. 34:
210.

subg. Pseudagris Saussure, 1863, Mem. Soc. Phys. Hist. Nat.
Geneve 17: 203 (as division of Synagris). Type species Syn-
agris carinata Saussure, 1863. Monotypic.

1986]

Carpenter — Checklist of Eumeninae

subg. Rhynchagris Maidl, 1914, Anz. Ak. Wiss. Wien 51: 91.
Type species Synagris vicaria Stadelmann, 1898. Monotypic.
Syneuodynerus Bluethgen, 1951, Boll. Soc. Entomol. Ital. 81: 67, 75
(as subgenus of “Euodynerus Bluethgen”). Type species Odyne-
rus egregius Herrich-Schaeffer, 1839. Original designation.
Syneodynerus (!) Kurzenko, 1981, Hym. Far East: 101.
Tachyancistrocerus Soika, 1952, Boll. Soc. Venez. Stor. Nat. 6: 37.
New name for Subancistrocerus Bluethgen.

“ Subancistrocerus (Saussure) nov. gen.” Bluethgen 1938, Dts.
Entomol. Z: 441, 460 non Subancistrocerus Saussure, 1855.
Type species Odynerus rhodensis Saussure, 1855. Original
designation.

Tachymenes Soika, 1983 (1982), Boll. Mus. Civ. Ven. 33: 118. Type
species Odynerus vulneratus Saussure, 1855. Original desig-
nation.

Tricarinodynerus Soika, 1952 (1951), Riv. Biol. Colon. 11: 73, 79.
Type species Odynerus guerinii Saussure, 1852. Original
designation.

Carinodynerus Soika, 1957 Brit. Mus. (Nat. Hist.) Exped. S.
W. Arabia 1(31): 478 (as subgenus of Pseudepipona Saus-
sure). Type species Odynerus guerinii Saussure, 1852. Origi-
nal designation.

Tricomenes Soika, 1978, Boll. Mus. Civ. Ven. 29: 10, 254. Type
species Eumenes pilosa Fox, 1899. Original designation.
Tropidodynerus Bluethgen, 1939, Veroeff, Dts. Kolon. Uebersee-
Mus. Bremen 2(3): 259, 260. Type species Polistes interrupta
Brulle, 1832. Original designation.

Xanthodynerus Bluethgen, 1954, Dts. Entomol. Z. (N. F.) 1: 255 (as
subgenus of “Euodynerus Bluethgen”). Type species Odynerus
octavus Soika, 1943. Original designation.

Xenorhynchium Vecht, 1963, Zool. Verh. (Leiden) 60: 111. Type
species Vespa nitidula Fabricius, 1798. Original designation.
Zeta Saussure, 1855, Et. Fam. Vesp. 3: 132, 146 (as division of
Eumenes Latreille). Type species Sphex abdominalis Drury,
1770. Designated by Bequaert, 1926, Ann. S. Afr. Mus. 23: 487.
Zeteumenes Bertoni, 1921, Rev. Soc. Cient. Paraguay 1: 117.
Type species Vespa canaliculata Olivier, 1791 (= Sphex argil-
lacea L., 1758). Designated by Vecht, 1977, Proc. K. Ned.
Akad. Weten. (C)80: 242.

Psyche

[Vol. 93

Zetamenes (!) Bertoni, 1926, Rev. Soc. Cient. Paraguay 2: 75.
Beteumenes Bertoni, 1934, Rev. Soc. Cient. Paraguay 3: 109 (as
subgenus of Zeteumenes). Invalid; no type designated.
Zetheumenidion Bequaert, 1926, Ann. S. Afr. Mus. 23: 487 (as sub-
genus of Eumenes Latreille). Type species Eumenes femoratus
Schulthess, 1910. Original designation.

Zethus Fabricius, 1804, Syst. Piez.: xii, 282. Type species Vespa
coeruleopennis Fabricius, 1798. Designated by Latreille, 1810,
Con. Gen. Crust. Arach. Ins.: 328, 438.

Didymogastra Perty, 1833, Delect. Anim. Artie. Brasil: 144.

Type species Didymogastra fusca Perty, 1833. Monotypic.
Lethus (!) Say, 1837, Boston J. Nat. Hist. 1: 387.

Heros Saussure, 1855, Et. Fam. Vesp. 3: 115 (as division of
Zethus ), non Heros Haeckel, 1840. Type species Zethus gigas
Spinola, 1841 (= Vespa coeruleopennis Fabricius, 1798).
Monotypic.

Wettsteinia Dalla Torre, 1904, Gen. Ins. 19: 13. Type species
Labus sichelianus Saussure, 1875. Designated by Bohart and
Stange, 1965, Univ. Calif. Publ. Ent. 40: 25.

Euzethus Dalla Torre, 1904, Gen. Ins. 19: 14. New name.
Laboides Zavattari, 1912, Arch. Naturgesch. 78A(4): 65. Type
species Labus sichelianus Saussure, 1875. Designated by
Bohart and Stange, 1965, Univ. Calif. Publ. Ent. 40: 25.
subg. Zethusculus Saussure, 1855, Et. Fam. Vesp. 3: 118. Type
species Zethus jurinei Saussure, 1852. Designated by Ash-
mead, 1902, Can. Ent. 34: 205.

subg. Zethoides Fox, 1899, Proc. Acad. Nat. Sci. Philad.: 436.
Type species Zethoides smithii Fox, 1899, ( non Zethus smi-
thii Saussure, 1855; = Zethus chapadensis Bohart and
Stange, 1965). Monotypic.

Baeoprymna Cameron, 1912, Timehri 2: 225. Type species
Baeoprymna rufoornata Cameron, 1912 (= Zethus minia-
tus Saussure, 1858). Monotypic.

Protozethus Bertoni, 1926, Rev. Soc. Cient. Paraguay 2: 75.
Type species Zethus olmecus Saussure, 1875. Original
designation.

1986]

Carpenter — Checklist of Eumeninae

subg. Madecazethus Soika, 1979, Boll. Mus. Civ. Ven. 30: 20,
53. Type species Labus madecassus Schulthess, 1907. Origi-
nal designation.

Nomina dubia in Eumeninae

Eumenestiferus Meunier, 1888, Nat. Sicil. 7: 300. Type species
Eumenestiferus brasiliensis Meunier, 1888. Monotypic. Un-
identified.

Micragris Saussure, 1855, Et. Fam. Vesp. 3: 158 (as division of
Synagris Latreille). Type species Synagris spinolae Saussure,
1855. Monotypic. Unidentified.

Nomina nuda in Eumeninae

Allepipona Bluethgen, 1951, Mitt. Munch. Entomol. Ges. 41: 194.
Antalastoroides Saussure, 1856, Et. Fam. Vesp. 3: 328 (hypothetical
group).

Austrodynerus Soika, 1958 (1957), Boll. Mus. Civ. Ven. 10: 119.
Lissodynerus Soika, 1974 (1973), Boll. Mus. Civ. Ven. 24: 119.

Acknowledgements

I thank J. van der Vecht for critically reviewing the manuscript.
An earlier version was submitted to Cornell University as part of a
doctoral dissertation (Carpenter, 1983). G. C. Eickwort, D. M.
Bates, J. L. Cisne and Q. D. Wheeler, members of the author’s
special committee, read and commented upon this version. Arnold
S. Menke, U.S. National Museum, provided valuable comments on
a later version.

Summary

A synonymic checklist of the genus-group names in the Eumeni-
nae is provided. Presently, 177 genera with 34 additional subgenera
are considered valid. Neodiscoelius Stange, 1979, is newly synonym-
ized with Protodiscoelius, Dalla Torre, 1904; Hy palast oroides
depressus Soika, 1969, is synonymized with Odynerus relativus Fox,
1902; the subgenus Cephalastor Soika, 1982, is raised to genus; and
type-species are designated for Nesodynerus Perkins, 1901, and
Stenolabus Schulthess, 1910.

Psyche

[Vol. 93

Literature Cited

Bluethgen, P.

1938 (1937). Systematisches Verzeichnis der Faltenwespen Mitteleuropas,
Skandinaviens und Englands. Konowia 16: 270-295.

Carpenter, J. M.

1981 (1982). The phylogenetic relationships and natural classification of the
Vespoidea (Hymenoptera). Syst. Ent. 7: 1 1-38.

1983. Phylogenetic studies in Vespoidea (Hymenoptera). PhD Thesis, Cornell
University.

Carpenter, J. M. and J. M. Cumming.

1985. A character analysis of the North American potter wasps (Hymenoptera:
Vespidae; Eumeninae). J. Nat. Hist. 19: 877-916.

Dalla Torre, K. W.

1904. Vespidae, Gen. Ins. 19: 1-108.

International Commission on Zoological Nomenclature.

1965. Opinion 747. Rygchium Spinola, 1806 (Insecta, Hymenoptera): Valida-
tion of emendation of Rhynchium. Bull. Zool. Nomencl. 22: 186-187.

1970. Opinion 893. Eumenidae names of Saussure (Hymenoptera): Grant of
availability to certain names proposed for secondary divisions of genera.
Bull. Zool. Nomencl. 26: 187-191.

1985. Opinion 1363. Ancistroceroides Saussure, 1855 (Insecta, Hymenoptera):
Type species designated. Bull. Zool. Nomencl. 42: 353-354.

Krombein, K. V., P. D. Hurd, D. R. Smith and B. D. Burks.

1979. Catalog of Hymenoptera in America North of Mexico. Smiths. Inst.
Press, Washington, D.C.

REVIEW OF THE FOSSIL TIPHIIDAE,

WITH DESCRIPTION OF A
NEW SPECIES (HYMENOPTERA)*

By A. P. Rasnitsyn

Paleontological Institute,

Academy of Sciences of the USSR,

Profusoyuznaja, 113, 117868,

Moscow USSR

Through the courtesy of Professor Frank M. Carpenter (Harvard
University, Cambridge, Mass.) and Dr. Paul E. S. Whalley (British
Museum, Natural History, London, U.K.) I have been able to study
the type specimens (good photographs of the specimen in one case)
of all described extinct species ever attributed to the Tiphiidae. Five
of them have been described as members of the subfamily Antho-
boscinae by Cockerell: in 1906 (Lithotiphia scudderi, Geotiphia fox-
iana), 1910 ( G . sternbergi, G. halictina) and 1927 ( G . pachysoma)\
while Hoplisidea kohliana was described originally as a member of
the Sphecidae (Cockerell, 1906) and later transferred to the Antho-
boscinae by Evans (1966).

From my study of these specimens I have found that the latter
species most probably belongs to the Sceliphronini (Sphecidae) and I
will treat it elsewhere. The five other species are discussed below and
one new species is described. All the species described by Cockerell
are from the Lower Oligocene of Florissant, Colorado; the new one
is from the ?Upper Oligocene of the Sikhote-Alin Mts., Maritime
Province of the USSR. Only the holotypes are known for all these
species and each specimen is a female, suggesting a female biased
tiphiid population during the Oligocene.

Only two other fossil specimens of Tiphiidae have been men-
tioned in the literature; both were found in Baltic amber collected by
A. Menge and both were identified by Brische (1886) as “Tiphia (?)”.
Unfortunately, Menge ’s collection is apparently lost (Heie, 1967,
P- 1 19).

* Manuscript received by the editor August 3, 1985

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The species treated here (figures 1-7) can be assigned to the
Tiphiidae on the basis of the strongly fossorial nature of the legs
(mid and hind tibiae thick and spiny), combined with the pleisio-
morphic wing venation; the latter differs distinctly from that of the
Scoliidae, which do have similar fossorial adaptations. In one case
(Fig. 2) this indirect evidence is confirmed by the structure of the
mesosternum, which shows the pair of lamellae that characteristi-
cally partly cover the midcoxae.

The fossil species show a habitus and female wing venation typi-
cal for the Anthoboscinae. Nevertheless, they do not belong to that
subfamily, mainly because their antennal sockets are overlain with
tubercles, clearly seen in one case (Fig. 6) and less clear in another
(Fig. 7). There are additional features distinguishing the fossils from
Anthoboscinae, viz., flagellum straight or variously bent (Figs. 1, 2,
4, 6) instead of tightly curled (as in all female Anthoboscinae stu-
died), femora lacking genual plates (Figs. 1-3, 7) or propodeum
with longitudinal lines (Figs. 4, 5).

All Tiphiidae with the antennal sockets partly covered by frontal
tubercles or ridges belong to the Myzininae and Methochinae. The
latter subfamily is not involved here, since its members have thin
tibiae bearing only weak spines. [I follow V. Gorbatovsky (personal
communication) in treating Pterombrus Smith as a member of the
subfamily Methochinae]. Therefore, the Myzininae is the only sub-
family with the characters of the fossils and in particular with those
of Geotiphia. [Lithotiphia is poorly known but I consider it similar
enough to the former genus to classify them together and not to
reject Lithotiphia as a tiphiid incertae sedis ]. Within the Myzininae
the fossils take an isolated position because of the very primitive,
male-like wing venation of the females.

Both of these extinct genera can be identified by the following
diagnoses. Lithotiphia (Fig. 1): forewing with cu-a cross-vein ante-
furcal; head capsule with a short oral cavity, distant from occipital
carina; hind tibiae very strongly swollen. Geotiphia (Figs. 2-7): fore
wing with cu-a interstitial or postfurcal; oral cavity longer, with
hypostomae reaching occipital carina; hind tibiae less swollen. The
latter genus possibly deserves to be divided into two genera, since
sternbergi and pachysoma, in contrast to other species, show mid
and/or hind femora with the genual plates, and the propodeum with
longitudinal lines. The propodeal structure is unknown in any other

Figure 1 . Lithotiphia scudderi Cockerell, holotype, no. 2022, Museum of Com-
parative Zoology, Harvard University. Wing cells are lettered. Scale line in all fig-
ures, 3 mm.

species and I hesitate to create another new genus on a sole charac-
ter. The following is a descriptive account of the species in these two
genera. The details shown in the figures are generally not described
below.

Lithotiphia scudderi Cockerell
Figure 1

Lithotiphia scudderi Cockerell, 1906, p. 51

Body length, 12.3 mm; fore wing length, about 5 mm (Length is
measured here from base to apex of cell 3r). Gastral terga with light
spots. Integumental sculpture not discernible because of covering
by Canada balsam. Holotype: M.C.Z. no. 2022.

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[Vol. 93

Figure 2. Geotiphia foxiana Cockerell, holotype, no. 2021, Museum of Compar-
ative Zoology, Harvard University.

Geotiphia foxiana Cockerell
Figure 2

Geotiphia foxiana Cockerell, 1906, p. 52

Body length, as preserved, 11 mm (probably originally 12 mm.);
fore wing length, 6.2 mm. Integumental sculpture not discernible.
Ground color moderately dark, the flagellum, tibiae, tarsi, veins,
and pterostigma less dark; metasomal sterna with light spots sub-
laterally, 2nd sternum having the spots large and contiguous. Color
pattern of terga unknown. Wing membrane not infumate. Holo-
type: M.C.Z. no. 2021.

Geotiphia halictina Cockerell
Figure 3

Geotiphia halictina Cockerell, 1910, p. 279

Body length, 18 mm; fore wing length, 3.5 mm. Venation similar to
that of foxiana, but differing in smaller size and the position of cell

1986]

Rasnitsyn — Fossil Tiphiidae

Figure 3. Geotiphia halictina Cockerell, drawing based on photograph of holo-
type, no. 18619, Museum of the University of Colorado.

3r remote from wing margin apically. Integumental sculpture and
color pattern unknown. (Description based on photograph of
holotype).

Geotiphia sternbergi Cockerell
Figure 4

Geotiphia sternbergi Cockerell, 1910, p. 277

Body length, 8 mm; fore wing length, 12 mm. Head with posterior
surface punctate dorsally and laterally, finely punctatorugose
medially. Thorax with distinct, moderately large punctures dor-
sally; lateral adscutellar depression, metanotum and propodeum
finely reticulate. Gastral terga with sculpture fine and sparse, not
clear in detail. Ground color dark (not known for fore and mid

Psyche

[Vol. 93

Figure 4. Geotiphia sternbergi Cockerell, holotype, no. 18868, American
Museum of Natural History, New York. Wing veins are lettered.

legs), anterior metasomal segments with small light spots laterally.
Fore wing apex infumate. Differs from the above species by its large
size, modified antennal segments, and in having the hind femur with
genual plate; fore wing with cell 2rm very long, and possibly in
having the propodeum with longitudinal lines. Holotype: A.M.N.H.,
no. 18868.

Geotiphia pachysoma Cockerell
Figures 5 and 6

Geotiphia pachysoma Cockerell, 1927, p. 432.

Body length, 9.2 mm; fore wing length, 6.0 mm. Head punctato-
rugose dorsomedially in part, thorax smooth, with distinct but weak

1986]

Rasnitsyn — Fossil Tiphiidae

Figure 5. Geotiphia pachysoma Cockerell, holotype, no. In. 26929, British
Museum (N.H.), London. Dorsal view.

punctures dorsally; lateral parts of metanotum striate longitudi-
nally. Body with ground color dark, without obvious light spots;
wing membrane infumate in apical two-fifths. Similar to sternbergi
in having genual plates and dissected propodeum, differing in small
size and in having cell 2 rm shorter; genual plates longer. Holotype:
B.M. (N.H.), no. In 26929.

Geotiphia orientalis, new species
Figure 7

Fore wing length about 6 mm. Pterostigma rather long, with 2r-rs
arising halfway before apex; cell 3r rounded at costal margin; RS
between RS+M and 2r-rs almost straight; cells lr, 2rm and 3rm all
relatively short; 2rm and 3rm of subequal length; lm-cu just before
the middle of 2rm; 2m-cu at the middle of 3rm, which has the

Psyche

[Vol. 93

Figure 6. Same as Fig. 5, ventral view.

posterior side very short and the distal side (3r-m) strongly arched;
crossvein cu-a at the fork of M+Cu; posterior genual plates absent
on mid and hind femora. Surface sculpturing indistinct. Body struc-
ture as preserved lacks taxonomically important features, the details
in part difficult to interpret. Ground color moderately dark; tibiae,
tarsi, venation, pterostigma, and metasomal segments 2 and 3 less
dark and without light spots (subsequent segments not preserved).
Wing membrane not infumate.

Holotype (only specimen known): no. 3429/ 100, Paleontological
Institute, Moscow, USSR; collected at Bolshya SvetloTodnaya
River, Pozharsky District, Maritime Province, USSR: ?Upper
Oligocene.

1986]

Rasnitsyn — Fossil Tiphiidae

Figure 7. Geotiphia orientalis, holotype, no. 3429/100. Paleontol. Inst. Acad.
Sciences, USSR, Moscow.

Comparison. As preserved this species is very similar to foxi-
ana, differing in having a longer pterostigma, the posterior side of
cell 3rm shorter, and the metasomal segments without light spots
[The latter difference may be meaningless because the color pattern
is known only for the metasonal sterna in foxiana and possibly only
for terga in orientalis].

The above data show considerable taxonomic and anagenetic
evolution of the subfamily Myzininae since the early Oligocene, an
interval of about 35 million years. Both fossil genera have been
replaced with a wide array of living genera, and even the most
primitive modern genus, Myzinum Latreille, is probably further
away from its Oligocene predecessors than these predecessors are
from their anthoboscine ancestor. A paleontological history is not

100

Psyche

[Vol. 93

known for any living myzinine genera, probably because of their
preference for environments unfavorable to fossilization (xeric bio-
topes or, in the case of Hylomesa, tropical forests), but all of them
can be easily derived from Geotiphia morphologically (but not from
Lithotiphia, because of the apomorphic position of the cu-a cross-
vein). Geotiphia can be characterized in short as an anthoboscine
with supraantennal tubercles, a position not consistent with the
current phylogenetic scheme showing synapomorphies for all
Tiphiidae other than Anthoboscinae and additional synapomor-
phies for all Tiphiidae except Anthoboscinae and Thynninae
(Brothers, 1975). An alternative scheme with Myzininae independ-
ent of other subfamilies (excluding Anthoboscinae and probably
Metochinae) seems to me more realistic.

The paleontological records indicate the minimal age of the
Myzininae as Early Oligocene. The records seem too scanty, how-
ever, to help in identifying the geographic area where the subfamily
arose.

Summary

Types of the previously described fossil Tiphiidae are studied.
Two genera and six species are recognized, each species known only
from the holotype: Lithotiphia Cockerell, with only one species,
scudderi Cockerell; and Geotiphia Cockerell, with foxiana Cocke-
rell (type-species), halictina Cockerell, orientalis, n.sp., sternbergi
Cockerell, and pachysoma Cockerell. The fossils are found to
represent the most primitive members of the subfamily Myzininae,
indicating that the subfamily originated from the Anthoboscinae
independently of the Thynninae, Tiphiinae, and Brachycistidinae.
Hoplisidea kohliana Cockerell is now determined as belonging to
the Sceliphronini of the family Sphecidae and will be treated else-
where. All species mentioned are from the Lower Oligocene of Flor-
issant, Colorado, except the new one, G. orientalis, which is from
the ?Upper Oligocene of Sikhote-Alin Mts., Maritime Province of
USSR.

1986]

Rasnitsyn — Fossil Tiphiidae

101

Literature Cited

Brischke, D.

1886. Die Hymenopteren des Bernsteins. Schrift. naturf. Gesellsch. Danzig,
n.f., 6: 278-279.

Brothers, D. J.

1975. Phylogeny and classification of the Aculeate Hymenoptera, with special
reference to Mutillidae. Univ. Kansas Sci. Bull., 50(1 1): 483-648.
Cockerell, T. D. A.

1906. Fossil Hymenoptera from Florissant, Colorado. Bull. Mus. Comp.
Zoology, Harvard Univ., 50: 33-58.

1910. Fossil insects and a crustacean from Florissant, Colorado. Bull. Amer.
Mus. Nat. Hist., 28: 275-288.

1927. Hymenoptera and a caddis larva from the Miocene of Colorado. Ann.
Mag. Nat. Hist., (9)20: 429-435.

Evans, H. E.

1966. The comparative ethology and evolution of the sand wasps. Harvard
Univ. Press, Cambridge, Mass., p. 1-526.

Heie, O.

1967. Studies on fossil aphids (Homoptera, Aphidodea). Spol. Zool. Mus.
Hauniensis, 26: 1-274.

AN EARLY RECORD OF TANDEM RUNNING IN
LEPTOTHORACINE ANTS: GOTTFRID ADLERZ, 1896

Robin J. Stuart1

Department of Zoology, Erindale College, University of Toronto,
Mississauga, Ontario, Canada L5L 1C6

Tandem running in ants is a recruitment technique in which one
ant leads a single follower to a particular target or target area. It has
been observed in various subfamilies, including the Myrmicinae,
Ponerinae and Formicinae, and appears to function in recruiting
nestmates to food discoveries, new nest sites, and into battle.
Detailed experimental analyses have revealed that tandem running
in some species is mediated by chemical and tactile cues, and various
authors have suggested that this recruitment strategy may have been
the evolutionary precursor of more sophisticated forms of group
and mass recruitment (see Wilson 1971, Holldobler 1978, Stuart
and Alloway 1983).

The term “tandem running” was first used by Wilson (1959) to
describe the behaviour of Cardiocondyla venustula and C. emeryi
workers as they recruited nestmates to new food sources. However,
Wilson (1959, 1971) attributed the first observation of tandem
running to Hingston (1929), and his description of foraging in
Camponotus sericeus. Nonetheless, Gottfrid Adlerz appears to
have observed this behaviour even earlier. Adlerz (1896), writing in
Swedish, described part of a nest emigration which he observed in
nature and which involved a mixed colony of the obligatory slave
maker Harpagoxenus ( =Tomognathus ) sublaevis and its Lepto-
thorax slaves. In translation, Adlerz described the event as follows
(see p. 9 of the original text):

“On one occasion, I observed a Tomognathus-Leptothorax
community being moved. The move had already started when I
arrived. The distance moved was only from one side of the stump

■Present Address: Museum of Comparative Zoology Laboratories, Harvard
University, Cambridge MA 02138 U.S.A.

Manuscript received by the editor October 10, 1985.

103

104

Psyche

[Vol. 93

to the other and the move was obviously caused by a nearby
community of stack ants ( Formica rufa) which disturbed the ants
at their previous location. During a period of 20 minutes, 8
Tomognathus workers were seen being carried in the usual
manner by the Leptothorax workers. In addition, one Tomo-
gnathus worker was seen walking at the heels of a Leptothorax
worker toward the new nest. The former held its head and anten-
nae on the abdomen of the Leptothorax worker and seemed to
get very agitated if it lost its guide during an unexpected turn and
did not find it immediately. As is usual during this kind of guid-
ance, the following ant carefully duplicated every little turn made
by the guide.”

The last few lines of this passage are a fairly accurate description of a
tandem run; and the last line indicates that Adlerz was quite familiar
with this recruitment technique.

Recent studies of the nest emigration behaviour of various Har-
pagoxenus species by Stuart and Alloway (1985) tend to confirm
Adlerz’s observations. Slaves in these mixed colonies are generally
responsible for the bulk of the moving effort during nest emigra-
tions: they transport brood and their adult nestmates, and lead
tandem runs between the two nests. Slave-maker workers some-
times follow in slave-led tandem runs, and H. americanus and H.
canadensis followers are relatively common. However, Stuart and
Alloway did not observe any H. sublaevis followers in their study.
Nonetheless, H. sublaevis followers have been observed in slave-led
tandem runs to food (Buschinger and Winter 1977), and they prob-
ably occur occasionally during nest emigrations as well.

Various species of nonparasitic leptothoracine ants use tandem
runs for recruiting nestmates to food (Moglich et al. 1974), during
nest emigrations (Moglich 1978), and for recruitment into battle
(Stuart and Alloway 1983); and certain leptothoracine slave makers,
including H. sublaevis and H. canadensis, lead tandem runs during
their slave raids (Buschinger et al. 1980, Stuart and Alloway 1983).
Other slave makers in this group, including H. americanus, le&d
processions during their raids (Wesson 1939, Alloway 1979,
Buschinger et al. 1980) and these processions constitute one of the
more advanced recruitment techniques thought to be evolutionarily
derived from tandem running (Wilson 1971, Stuart and Alloway

1986]

Stuart — Tandem running

105

1983). H. canadensis appears to be an unusual obligatory slave
maker, in that it will also lead tandem runs to food and during nest
emigrations; behaviours which may be indicative of the relatively
primitive nature of this species (Stuart and Alloway 1985).

Thus, Adlerz may have been the first to report tandem running in
ants; and the tandem run he described apparently involved a Lepto-
thorax slave leading a Harpagoxenus sublaevis slave maker during a
nest emigration.

Acknowledgments

The author thanks Vivian Sterne for translating the Adlerz arti-
cle. Financial support was provided by an Ontario Graduate Schol-
arship to the author and a Natural Sciences and Engineering
Research Council of Canada grant to T. M. Alloway.

References

Alloway, T. M. 1979. Raiding behaviour of two species of slave-making ants,
Harpagoxenus americanus (Emery) and Leptothorax duloticus Wesson (Hyme-
noptera: Formicidae). Anim. Behav. 27: 202-210.

Adlerz, G. 1896. Myrmecologiska studier III. Tomognathus sublaevis Mayr.
Bihang Till K. Svenska Vet.-Akad. Handlingar 21 (IV-4): 1-76. (Translation)

Buschinger, A., and U. Winter. 1977. Rekrutierung von Nestgenossen mittles
Tandemlaufen bei Sklavenraubzugen der dulotischen Ameise Harpagoxenus
sublaevis { Nyl.). Ins. Soc. 24: 183-190.

Buschinger, A., W. Ehrhardt and U. Winter. 1980. The organization of slave
raids in dulotic ants — a comparative study (Hymenoptera; Formicidae). Z.
Tierpsychol. 53: 245-264.

Hingston, R. W. G. 1929. Instinct and intelligence. Macmillan Company, New
York, xv + 296 pp.

Holldobler, B. 1978. Ethological aspects of chemical communication in ants.
Adv. Study Behav. 8: 75-115.

MOglich, M. 1978. Social organization of nest emigration in Leptothorax
(Hymenoptera; Formicidae). Ins. Soc. 25: 205-225.

MOglich, M., U. Maschwitz, and B. HOlldobler. 1974. Tandem calling: A
new kind of signal in ant communication. Science 186: 1046-1047.

Stuart, R. J., and T. M. Alloway. 1983. The slave-making ant, Harpagoxenus
canadensis M. R. Smith, and its host-species, Leptothorax muscorum (Ny-
lander): Slave raiding and territoriality. Behaviour 85: 58-90.

Stuart, R. J., and T. M. Alloway. 1985. Behavioural evolution and domestic
degeneration in obligatory slave-making ants (Hymenoptera: Formicidae: Lep-
tothoracini). Anim. Behav. 33: 1080-1088.

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[Vol. 93

Wesson, L. G. 1939. Contributions to the natural history of Harpagoxenus ame-
ricanus Emery (Hymenoptera: Formicidae). Trans. Amer. Entomol. Soc. 65:
97-122.

Wilson, E. O. 1959. Communication by tandem running in the ant genus Cardi-
ocondyla. Psyche 66: 29-34.

Wilson, E. O. 1971. The insect societies. Harvard Univ. Press, Cambridge, Mass,
x + 548 pp.

NOTES ON THE BEHAVIOR OF THE DIMORPHIC ANT
OLIGOMYRMEX OVERBECKI*
(HYMENOPTERA: FORMICIDAE)

By Mark W. Moffett

Museum of Comparative Zoology,

Harvard University,

Cambridge, Massachusetts 02138

Species of the myrmicine genus Oligomyrmex are common in
tropical Asia, although the ants are easily overlooked because of
their small size and inconspicuous activities. The genus is of special
interest because of the well developed worker dimorphism shown by
all species. Some natural history information is available on Ere-
bomyrma (Eidmann, 1936; Wilson, 1962, 1986), the American sister
group to Oligomyrmex which has only recently been resurrected
from synonomy with that genus (Wilson, 1986). However, the natu-
ral history of Old World Oligomyrmex ants has never been
investigated.

I have made preliminary behavioral observations on a colony of
Oligomyrmex overbecki Viehmeyer collected in Singapore (fig. 1).
This species is clearly one of the world’s smallest ants, with minor
workers having head widths of 0.29-0.32 mm, while the “miniature”
majors have head widths of 0.42-0.45 mm.

Materials and Methods

The study colony was collected on the grounds of the Botanic
Gardens of Singapore, under bark still firmly attached to the trunk
of a large Eugenia grandis tree (Myrtaceae), within 50 cm of ground
level. The colony was placed in a plastic box 20 X 10 X 7 cm deep,
with a moistened paper-mache bottom gouged towards one end
with several small, shallow chambers, which were then covered with
a sheet of glass. The ants moved into the artificial nest chambers,
where they could readily be observed through the glass.

* Manuscript received by the editor January 26, 1986

107

108

Psyche

[Vol. 93

Fig. 1 . Portion of Oligomyrmex overbecki study colony, showing the queen (center), minor and major workers, and brood.
Scale bar = 1.0 mm.

1986]

Moffett — Oligomyrme x overbecki

109

A behavioral repertoire of the workers was compiled during 14
hours within a four day period beginning five weeks after the colony
was collected. Estimates of total repertory size were made by fitting
the observed behavioral frequencies to a lognormal Poisson distri-
bution as described by Fagen and Goldman (1977), using a compu-
ter program supplied by R. M. Fagen. Additional behavioral data
was gathered during roughly 25 hours of observations before the
repertoire study.

While collecting the repertoire data, light-colored (callow)
minors, which were uniformly golden-yellow to light brownish yel-
low, were distinguished from more darkly pigmented minors (vary-
ing from yellowish brown to brown, with antennae, legs and gaster
lighter). In addition, the non-callows were subdivided into “re-
pletes,” which had their gasters moderately expanded with yellowish
fluid, and non-repletes, which had small, contracted gasters. (By this
criterion, all major workers and all callow minor workers were
judged to be “replete.”)

Voucher specimens from the study colony have been deposited in
the Museum of Comparative Zoology (Harvard University).

Results

Nesting habits: The workers, queen and brood were tightly

massed together between two small adjacent pieces of superficial
bark. No food was seen within the nest. The nest area was originally
estimated to contain about 400 workers, but upon return to the
United States for study, 31 majors and about 180 minors remained.
The original proportion of major workers probably approached ten
percent.

Repertoire: The complete behavioral repertoire of the worker
castes and subcastes is presented in Table 1. During the period in
which the worker data was collected, 27 behavioral acts were
observed for the queen, including 19 instances of nipping at im-
matures (described below), five self-grooming events and three
instances of licking large larvae. The total repertoire size is
estimated to be between 32-36 for the minor caste (data from all
subcastes combined), and between 6-11 for the majors (95%
confidence intervals).

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Table 1. Repertoiries of Oligomyrmex overbecki worker castes, including subdi-
visions of the minor caste (see text). Numbers represent the proportion that each
behavior represented of the total number of acts observed for each type of worker.

Replete

Minor

Non-replete

Minor

Callow

Minor

Major

Self-grooming

0.2443

0.2632

0.2237

0.6074

Allogroom minor

0.1401

0.1219

0.0461

Allogroom major

0.0104

0.0042

0.0066

Allogroom queen

0.0048

0.0014

Lick eggs

0.0248

0.0028

0.0921

Lick small larva

0.0200

0.0125

0.0066

0.0123

Lick large larva

0.2284

0.1759

0.1645

0.1411

Lick pupa

0.0587

0.0111

0.1974

0.0061

Carry eggs

0.0483

0.0069

0.1908

Carry small larva

0.0041

0.0028

0.0197

Carry large larva

0.0352

0.0457

Carry pupa

0.0028

0.0014

Carry minor worker

0.0035

Pull on queen

0.0014

Nip at immature
Assist in:

0.0076

0.0263

0.0066

0.2209

larval ecdysis

0.0021

0.0014

ecdysis to pupa

0.0104

0.0042

adult eclosion

0.0035

0.0042

meconium removal

0.0035

Manipulate meconium

0.0193

0.0208

0.0197

Remove liquid waste

0.0062

0.0042

Handle nest material

0.0200

0.0706

0.0066

Forage

0.0179

0.1371

Retrieve solid food

0.0007

0.0014

Eat solid food

0.0248

0.0180

Feed on immatures

0.0304

0.0291

Feed larva solid food
Regurgitate to:

0.0041

larva

0.0048

0.0028

0.0197

minor worker

0.0110

0.0248

0.0123

major worker

0.0048

0.0042

queen

Carry or eat

0.0014

dead nestmate

0.0007

0.0014

No. acts observed

1449

722

152

163

1986] Moffett— Oligomyrmex overbecki 111

The most conspicuous difference between the minor worker sub-
castes was that darkly pigmented non-repletes formed the bulk of
the foragers. The repertoire data indicate several other differences in
the frequency of behaviors (differences judged significant when p <
0.05 with chi-square test). Callow workers carried and licked eggs
with greater frequency than did darker colored minors, but carried
large immatures less frequently than did the latter. In comparison to
darkly pigmented minors, callows rarely fed on solid foods and
rarely allogroomed other workers. They also regurgitated to larvae
more often than did the darker subcastes, yet apparently seldom
regurgitated to other adult ants (difference in frequencies was not
significant in the latter case).

Darkly-pigmented replete minors were intermediate between cal-
low and non-replete minors in the frequencies of performance of
many of those behaviors that varied most markedly between the
minor subcastes. This suggests the possibility that these minors
could be intermediate in age between callow minors (which were
consistently replete) and non-replete minors.

Majors rarely foraged. During my observations only four majors
were seen outside the nest of the captive colony, and one major was
observed on a foraging route near the nest entrance in the field.
Major workers apparently only fed by regurgitation.

The O. overbecki queen did not attract a large retinue of workers,
but commonly one or two minors climbed onto her alitrunk or
gaster. In addition, twice I observed replete minors briefly pulling
on an antenna or mandible of the queen. Only rarely would a major
climb onto the queen, and the density of majors was not noticeably
greater near the queen than elsewhere.

Occasionally a major, minor, or the queen briefly appeared to try
to grip or bite immatures, most commonly large larvae (“nip at
brood” in Table 1). The function of this behavior is unclear, for
although consumption of brood by minor workers was common,
this biting behavior was most frequently performed by majors and
apparently never damaged the immatures.

Larvae fed directly on fragments of insect corpses and from food
regurgitated to them by minors.

Foraging Pattern and Diet: During my field observations

columns of minor workers extended at least 30 cm from the nest on

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Psyche

[Vol. 93

the bark of the tree. In captivity, foragers often followed trunk
routes at least 3-5 cm long before departing from them to forage
singly.

Foraging minor workers fed at crushed fruit flies, fragments of
freshly killed cockroaches, honey water baits, and Bhatkar diet
(Bhatkar and Whitcomb, 1970). The ants avoided wounded fruit
flies, and did not recruite minor and major workers to wounded
prey as has been observed for Erebomyrma nevermanni (Wilson,
1986).

Soon after most large baits were presented, ants began arriving at
the bait using a well-defined route, suggesting an odor trail had been
laid down. However, recruitment behavior was difficult to docu-
ment because of the tiny size of the ants and their weak response to
food, even following periods of food deprivation.

Typically food was torn into small pieces and carried into the nest
by solitary individuals. Whole dead fruit flies near the nest entrances
were sometimes dragged into the nest by groups of 2-5 workers.
However, this group transport behavior was poorly coordinated, as
workers often pulled in conflicting directions.

Repletes: The O. overbecki majors were mildly replete (“semi-

replete”), with their gasters never expanding to a size much greater
than that of their heads. Moreover, the majors were no more replete
than replete minor workers (judging by the volume of the gaster
relative to that of the trunk).

Emigrations: Two shifts in nest location were documented in

the laboratory. These followed periods of mild stress in which a 60
watt bulb was positioned 25 cm above the glass-covered nest
chamber, while an unoccupied shaded chamber was provided 4-5
cm away. Within ten minutes the ants became more active, with
darkly pigmented minors and a few majors leaving the nest
chambers to explore the nest environs. Gradually more and more
workers moved back and forth between the nest chambers and the
shaded chamber, until it was clear that a set route had been estab-
lished. Traffic along the emigration route was relatively steady
throughout the period of brood transfer, with the number of ants
passing an arbitrary point on the route exceeding 20 per minute.

The first immature was carried out of the nest 50 minutes into the
second emigration; the sequence of brood transfer is documented in
Figure 2. There was no group transport of immatures and no adult

larvae

paujeo jaqiuriN

Fig. 2. Transfer of brood during an Oligomyrmex overbecki emigration. Times are given as the number of minutes si
strong light was first shined on the ants.

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[Vol. 93

transport (although adult transport of minor workers was observed
at other times; see Table 1). Eggs were completely transferred early in
both emigrations. The last immatures to be transferred were larvae,
not because workers selected pupae over larvae, but because the
clumped larvae were difficult to pull apart for transport.

Only minor workers carried brood. Callow minors aided in pull-
ing larvae and pupae free of piles of brood, but were clumsy at
carrying larger immatures, which were quickly turned over to
darker workers. Callows did, however, occasionally carry small lar-
vae and eggs, taking egg clusters at a higher frequency than did
other minors (p <0.01, Fisher’s exact probability test). Both replete
and non-replete darkly pigmented minors transferred brood, and
there were no significant differences between the frequency with
which these subcastes carried different brood stages (for each brood
stage p > 0.05).

The queen emigrated soon after brood transfer began in the first
emigration, and ten minutes before the start of brood transfer dur-
ing the second emigration. She moved rapidly within a small en-
tourage of minors, but no workers rode on her during her journey.

Alarm and Defense: In three trials in which a small Solenopsis

geminata worker with excised gaster was dropped into the brood
area, most workers and the queen fled to adjacent nest chambers,
with some minor workers carrying brood. Usually several major
workers and a few minors stayed close to the intruder, mandibles
open and facing the Solenopsis. Sometimes the ants attempted to
bite the intruder. As described for Erebomyrma nevermanni (Wil-
son, 1986), the proportion of major workers near the intruder was
clearly higher than in the colony as a whole. The ants responded
similarly to freshly crushed minor heads presented on applicator
sticks, suggesting the head as a source of alarm pheromones. Majors
were particularly attracted to crushed minor heads, approaching
them with their antennae directed ahead and mandibles open. There
was virtually no response to crushed thoraxes and gasters.

Discussion

The major workers of Oligomyrmex overbecki apparently func-
tion primarily in colony defense and as repletes. The replete condi-
tion is very poorly developed (the ants are “semi-replete” in the
sense of Wilson, 1986). Major workers also participated to a limited

1986]

Moffett — Oligomyrmex overbecki

115

extent in brood care. It is possible that the repertoire of majors is
normally more restricted, but that high minor worker mortality in
the captive colony and the resulting altered caste ratios led to an
expansion of the major worker repertoire. The relationship between
worker caste ratios and major repertoires for dimorphic ants is only
beginning to be explored (see Wilson 1984, 1986).

Observations on a Oligomyrmex cf. solidaris colony collected in a
rotten log from Bako National Park in Sarawak indicates that the
majors of this species also are semi-replete and are crucial to colony
defense. O. cf. sodalis majors were quick to attack Pheidologeton
silenus and Pheidole megacephala workers dropped into the nest
areas, and were much more efficient than minor workers in inflict-
ing damage on the enemy. The importance of rapid and effective
response to workers of these ant species was dramatized when the
artificial nest container housing the O. cf. sodalis colony was raided
by Pheidole megacephala ants. Within a four hour period the Phei-
dole had completely destroyed the Oligomyrmex colony of several
hundred individuals and emigrated into their nest container.

Minor workers of O. overbecki show a pattern of temporal poly-
ethism common for ants (Wilson, 1971), caring for immatures (par-
ticularly smaller immatures) as callows and shifting towards
foraging activities as they age. Probably only younger workers are
semi-repletes, with the ants losing their replete condition at about
the time they begin to forage.

Oligomyrmex overbecki (as well as O. cf. sodalis, pers. obser.)
forms trunk trail foraging routes, as do a variety of other pheido-
logetine ants: Erebomyrma nevermanni (Wilson, 1986); Pheidolo-
geton diversus (Moffett, 1984) and all other Pheidologeton species
(pers. obser.); and Lophomyrmex bedoti (Moffett, 1986).

Acknowledgements

I thank E. O. Wilson and D. H. Murphy for encouragement and
advice. The research was supported by grants from the National
Geographic Society and Harvard University.

Literature Cited

Bhatkar, A. and W. H. Whitcomb.

1970. Artificial diet for rearing various species of ants. Fla. Entomol. 53:
229-232.

116

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[Vol. 93

Eidmann, H.

1936. Okologisch-faunistische Studien an sudbrasilianischen Ameisen. Arbeit,
phys. angew. Ent. Berlin-Dehlem 3: 26-48, 81-113.

Fagen, R. M. and R. Goldman.

1977. Behavioral catalogue analysis methods. Anim. Behav. 25: 261-274.

Moffett, M. W.

1984. Swarm raiding in a myrmicine ant. Naturwissenschaften 71: 588-589.

1986. Observations on Lophomyrmex ants from Kalimantan, Java, and
Malaysia. Malayan Nat. Journ., in press.

Wilson, E. O.

1962. The Trinidad cave ant Erebomyrma (= Spelaeomyrmex) urichi (Wheeler),
with a comment on cavernicolous ants in general. Psyche 69: 63-72.

1971. The Insect Societies. Belknap Press of Harvard University Press, Cam-
bridge, 548 pp.

1984. The relation between caste ratios and division of labor in the ant genus
Pheidole (Hymenoptera: Formicidae). Behav. Ecol. Sociobiol. 16:
89-98.

1986. Caste and division of labor in Erebomyrma, a genus of dimorphic ants
(Hymenoptera: Formicidae: Myrmicinae). Insectes Sociaux, in press.

PUPATION IN MYCETOPHILID FLIES:
A CORRECTION

By William G. Eberhard

Smithsonian Tropical Research Institute and
Escuela de Biologia, Universidad de Costa Rica
Ciudad Universitaria, Costa Rica

In a previous paper (Eberhard 1970) I made several claims re-
garding two species of the mycetophilid fly genus Leptomorphus: 1)
the larval cuticle is not shed prior to pupation; 2) the last two and
one half segments of the larva are discarded at pupation; and 3) the
larval head capsule is engulfed by the pupa during pupation (Eber-
hard 1970). Recent, more detailed observations of Leptomorphus
sp. have shown that points 1 and 3 are probably wrong, and this
note is an attempt to present a more accurate account of pupation.

Observations were made during Sept. 1984 near San Jose, Costa
Rica on larvae living on the undersurface of a fungus-covered
board, where they inhabited silken sheets with slime trails similar to
those of L. bifasciatus and L. subcaeruleus (Eberhard 1970). One
observation of the process of pupation was made under a dissecting
microscope. This larva hung on an approximately horizontal pupal
line fastened at either end to a glass slide, and was observed from
above (i.e. from the larva’s ventral surface); occasionally I tilted the
slide so as to check the larva in lateral view. Species identification in
the genus Leptomorphus is not presently possible (R. Gagne, pers.
comm.); voucher specimens of adults reared from the larvae
observed are deposited in the U.S. National Museum.

Results

The overall sequence of events was the same as that described for
L. bifasciatus and L. subcaeruleus (Eberhard 1970) except that lar-
vae were on lines for somewhat less than 24 hours before pupating.
Although the head capsule was nearly engulfed by the swollen ante-
rior portion of the larva’s body when pupation began, it did not
disappear. Instead, as the anterior end of the animal’s body assumed

Manuscript received by the editor October 2, 1985.

117

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the new (pupal) shape, the head capsule moved smoothly posteriorly
along the center line of the animal’s ventral surface. The capsule
paused briefly when it reached the “collar” or the anterior end of the
band of silk that fastened the larva to the pupal line, then moved on
smoothly, passing beneath the mat of silk threads holding the larva
to the pupal line. As the head capsule neared the posterior end of the
body, the cuticle there began to wrinkle during each contraction of
the animal’s body, also as noted previously (Eberhard 1970). When
the posterior end of the pupa broke free from the remains of the
larva, the head capsule was left as part of the mass of larval material
that remained attached to the line. Careful dissections of some of
these masses in water revealed the presence of not only the head
capsule but also a long tubular sheath of very thin, transparent
cuticle that bore the rows of dark denticles found near segmental
boundaries on the ventral surfaces of larvae (Eberhard 1970). Thus
the entire larval cuticle was shed during pupation, and the head
capsule was not engulfed.

With respect to point 2 (posterior segments of larval body dis-
carded during pupation), the new evidence does not clearly contra-
dict previous descriptions. Several minutes prior to the migration of
the head capsule to the posterior end of the larva, the last two and
one half segments of the larva’s body had darkened to a caramel
brown color, and the material inside was amorphous and inert
when viewed through the larval cuticle. In contrast, there were clear
internal movements of well defined structures just anterior to this
area, and it appeared that the posterior tip of the pupa had already
formed and was being repeatedly pushed posteriorly against the
inert brown material. When the larval cuticle was finally discarded
(above), these posterior two and one half segments did not wrinkle
or contract as did the rest of the larval cuticle, but retained their
form, and the rows of denticles marking the segmental boundaries
on their ventral surface remained clearly visible and as far apart as
they had been in the intact larva.

Discussion

Probably the pupation process in the Leptomorphus species of
previous reports was the same as that described here. The larval
head capsule is small and partially transparent, and difficult to see
without magnification. The observations of larval head capsules on

1986]

Eberhard — Pupation in my cetophilid flies

119

the ventral surfaces of pupal abdominal segments (Eberhard 1970)
probably represent cases in which the larval skin was only partially
shed, and broke near the tip of the pupal abdomen.

It has been argued that silk attachments to larval cuticle should be
shed along with the larval cuticle (Eberhard 1970, Malloch 1917).
Although this seems reasonable, it is clearly not the case in Lepto-
morphus sp. How the larval skin is shed so smoothly without dis-
turbing, as far as can be seen, the silk lines that form the only
attachment of the animal hanging on its pupation line remains a
mystery.

Acknowledgements

I am grateful to R. Gagne for kindly identifying specimens, and
the Vicerrectoria de Investigation of the Universidad de Costa
Rica for financial support.

References

Eberhard, W. G.

1970. The natural history of the fungus gnats Leptomorphus bifasciatus (Say)
and L. subcaeruleus (Coquillett) (Diptera: Mycetophilidae). Psyche 77:
361-383.

Malloch, J. R.

1917. A preliminary classification of Diptera, exclusive of Pupiparia, based
upon larval and pupal characters, with keys to imagines in certain fami-
lies, Part I. Bull. 111. State Lab. Nat. Hist. 12(3): 161-407, pis. 28-57.

NEW PSELAPHIDAE FROM NEW HAMPSHIRE
(COLEOPTERA)1

By Donald S. Chandler

Department of Entomology,

University of New Hampshire,

Durham, NH 03824

Two species of undescribed Pselaphidae were discovered during a
comparison of the fauna of an uncut and a 40 year-old forest. The
species are described here to provide names for a forthcoming paper
comparing the pselaphid fauna of these two sites. Holotypes were
cleared, disarticulated, and mounted on slides in Canada Balsam.
Both are placed in the Field Museum of Natural History, Chicago.
All measurements of specimens are in millimeters.

I would like to thank certain individuals for the loan of speci-
mens, greatly extending the known ranges of these two new species.
The abbreviations used to indicate specimen deposition follows the
individual’s affiliation: Rickard Baranowski, Lund University,
Sweden (RBC); J. Milton Campbell, Biosystematics Research Insti-
tute, Ottawa, Canada (CNCI); Michael A. Ivie, Montana State
University, Bozeman (DZEC); and Alfred F. Newton, Jr., Field
Museum of Natural History, Chicago (FMNH). Specimens other-
wise lacking an indication of deposition are in the collections of the
author and the University of New Hampshire. I would like to thank
J. F. Burger and R. Marcel Reeves, University of New Hampshire,
for reviewing the manuscript.

Euplectus silvicolus n. sp.

(Figs. 1-3)

Length 1.36-1.44. Head glabrous, punctures indistinct, vertex
with arms of distinct U-shaped impression originating from nude
vertexal foveae; mandibles with five teeth on inner margin, third

'Scientific Contribution Number 1413 from the New Hampshire Agricultural Exper-
iment Station.

Manuscript received by the editor March 11, 1986.

121

122

Psyche

[Vol. 93

Figs. 1 -3. Euplectus silvicolus n. sp.. male. 1 . ventral view last leg. 2. ventral view
sternite VI. 3a. dorsal view aedeagus; 3b. left lateral view aedeagus.

Fig. 4. Actizona borealis n. sp., male aedeagus. a. dorsal view; b. left lateral view.

tooth largest. Elytra with four basal foveae. Tergites I-III with basal
carinae, depressions between basal carinae only conspicuously
setate on tergites I-II, I-III equal in length, IV half again as long as
III.

Males with small spur at apices of all tibiae, metatrochanters at
base with large medially directed spur; sternites IV-V simple, convex
medially, VI obscurely depressed at middle, short aciculate setae in
depression forming arc, division of sternite VII arcuate to left.

Females lacking spurs of tibiae and metatrochanters; sternites
evenly convex.

Specimens examined, 27. HOLOTYPE male, New Hampshire,
Carroll Co., The Bowl, 2.5 mi NW Wonalancet, VIII-6-1985, D. S.

1986]

Chandler — New Pselaphidae

123

Chandler, sift conifer logs. PARATYPES : 1 male, 1 female, same
data except IX-1-1984; 1 male, same data except VIII-21-1985; 6
males, 10 females, same data except VI-8/ 14-1984 (2), VI-15/20-
1984 (1), VI-28/ VII-4-1984 (1), VIII-2/ 10-1984 (2), VIII-1 1/ 16-1984
(1), V-23/ VI-4-1985 (1), VII-2/ 10-1985 (6), VII-24/ 30-1985 (1),
VIII-22 / 28-1985 (1), flight intercept trap; 1 male, 1 mi N Wona-
lancet, East Fork Spring Brook, 1900', VII-23-1985, D. S. Chandler,
sift hemlock logs; 1 male, 1 female, same data except VII-2/ 10-1985,
VII-31/ VIII-6-1985, flight intercept trap. Coos Co.: Norton Pool, 2
mi E East Inlet Dam, IX-7-1984, D. S. Chandler, sift rotten
spruce/ fir logs. Canada: Nova Scotia: 1 male. Cape Breton
Highlands National Park, MacKenzie Mountain, PG648868,
VII-4-1983, R. Vockeroth, pan traps (CNCI); 1 female, same data
except Lone Shieling, PG729861, VI-25- 1983, Y. Bousquet, pans
(CNCI).

Biology: This uncommon species was only found in rotten conifer
logs in an extensive litter survey at The Bowl. Most specimens were
collected by flight intercept traps.

Discussion: This species is quite distinct among the Nearctic spe-
cies of Euplectus by the presence of basal carinae on tergite III, spur
of the male metatrochanters, simple sternites IV-VI, and smooth
vertexal area. Since two species of Euplectus have been introduced
to North America from Europe, the major faunal works of Jeannel
(1950) for France and Besuchet (1974) for Central Europe were
checked to be certain this species had not been previously described.
In Wagner’s (1975) recent revision of the Nearctic species of Euplec-
tus, this species would be placed in the calif ornicus- group. Silvicolus
may be separated at couplet 5 of Wagner’s key by the lack of any
papilliform setae in the depression of sternite VI. This species differs
from the generic diagnosis of Grigarick and Schuster (1980) in pos-
sessing basal carinae on tergite III, which are lacking in all other
Nearctic species and also in the twenty Palearctic species in my
collection.

Actizona borealis n. sp.

(Fig. 4)

Length 1.20-1.32. Head with pubescent vertexal foveae, penulti-
mate antennomeres symmetrical, antennal club with parallel mar-
gins, twice as long as wide. Elytra with three basal foveae.

124

Psyche

[Vol. 93

Promesocoxal foveae present, metasternal foveae separated by over
two foveal diameters. Tergite lengths subequal, I-II with faint short
basal carinae.

Males with protrochanters angulate on posterior margin, pro-
tibiae with small preapical spur; mesotrochanters posteriorly angu-
late, apical spur on inner margin of mesotibiae; sternites II-III with
small setate tubercle near postero-lateral margins, VI simple, VII
oval and setae over surface.

Females lacking spurs of tibiae and trochanters, lacking tubercles
of sternites II-III.

Specimens examined, 8. HOLOTYPE male, New Hampshire,
Coos Co., Jefferson Notch, 910 m, VII-14/ 3 1-1982, A. Newton &
M. Thayer, window trap. PARA TYPES: New Hampshire: Carroll
Co.: 2 males, 2 mi NW Wonalancet, VI-8/ 14-1984, VI-15/20-1984,
D. S. Chandler, window trap; 1 male, The Bowl, 2.5 mi NW
Wonalancet, VIII-1 6-1 984, sift rotten wood; 1 female, same data
except XI-23-1984, R. M. Reeves, sift birch stump; 1 male, same
data except, VII-23- 1985, D. S. Chandler, sift rotten beech logs.
Canada: British Columbia: 1 male, Princeton, South Wash Creek,
VII-22-1983, Lindgren funnel trap (DZEC); 1 male, Monashee
Mountain near Cherryville, 1400-1600 m, VIII-12-1982, R. Bara-
nowski (RBC), sifting litter and moss in spruce forest.

Biology: Collected in rotten beech and birch logs in uncut forests
in New Hampshire.

Discussion: This species is very similar to Actizona chuskae
Chandler from Arizona (Chandler 1985) in appearance and male
characters. The genitalic form of borealis is identical in the British
Columbia and New Hampshire specimens, and differs from that of
chuskae in the form of the apex and internal spines of the aedeagus.
These genitalic differences and the pubescent vertexal foveae of
borealis readily separate the two species.

Summary

Two undescribed species of Pselaphidae, Euplectus silvicolus n.
sp. and Actizona borealis n. sp., were discovered during a faunal
comparison of the forest floor Coleoptera of cut and uncut forests m
New Hampshire.

1986]

Chandler — New Pselaphidae

125

Literature Cited

Besuchet, C.

1974. 24. Familie: Pselaphidae, pp. 305-362. In: Die Kaefer Mitteleuropas.
Band 5, Staphylinidae II (Hypocyphtinae and Aleocharinae), Pselaphi-
dae, 381 pp. Eds. H. Freude, K. W. Harde, and G. A. Lohse. Goecke &
Evers, Krefeld.

Chandler, D. S.

1985. The Euplectini of Arizona (Coleoptera: Pselaphidae). Entomography 3:
107-126.

Grigarick, A. A. and R. O. Schuster.

1980. Discrimination of genera of Euplectini of North and Central America
(Coleoptera: Pselaphidae). Univ. Calif. Pubis Ent. 87: vi + 56 pp., 79
plates.

Jeannel, R.

1950. Coleopteres Pselaphides. Faune Fr. 53: iii T 421 pp.

Wagner, J. A.

1975. Review of the genera Euplectus, Pycnoplectus, Leptoplectus, and Aco-
lonia (Coleoptera: Pselaphidae) including Nearctic species north of Mex-
ico. Entomologica am. 49: 125-207.

A PRESUMPTIVE PHEROMONE-EMITTING STRUCTURE
IN WOLF SPIDERS
(ARANEAE, LYCOSIDAE)*

By Torbjorn Kronestedt

Department of Entomology,

Swedish Museum of Natural History,

S-104 05 Stockholm, Sweden

The occurrence of pheromones in lycosid spiders has long been
indicated on behavioural grounds. (For a review on chemical com-
munication in spiders, see Tietjen and Rovner 1982.) There are
bioassay evidences for (1) contact sex pheromones deposited on the
substrate by females (Bristowe and Locket, 1926; Rovner, 1968;
Hegdekar and Dondale, 1969; Richter et al., 1971; Dijkstra, 1976;
Robert and Krafft, 1981), (2) contact sex pheromones associated
with draglines laid by females (Kaston, 1936; Engelhardt, 1964;
Richter et al., 1971; Dondale and Hegdekar, 1973; Tietjen, 1977,
1979b; Tietjen and Rovner, 1980; Robert and Krafft, 1981), (3)
contact sex pheromones associated with female integument (Kas-
ton, 1936), and (4) airborne sex pheromones given off by females
(Tietjen, 1979a). Candidates for contact pheromone perception are
chemosensitive hairs occurring on legs and palps. The number of
these hairs is considerably increased in adult males in comparison to
immatures and adult females (Tietjen and Rovner, 1980, 1982), and
in certain lycosid genera this increase is rather drastic (Kronestedt,
1979a). No site of production and release of pheromones in wolf
spiders has so far been found (Tietjen and Rovner, 1982). The pres-
ent note focuses on a type of structure which is presumably involved
in the release of pheromones in this spider family.

Studies on courtship behaviour in various lycosid species have
been undertaken for supplementing morphological data in taxo-
nomic contexts as well as for finding connections between adult
male secondary sex characters and species-specific behavioural ele-
ments. Among the species studied, the adult male of Alopecosa
cuneata (Clerck) has a unique character in its first tibiae being tumid

* Manuscript received by the editor January 8, 1986.

127

128

Psyche

[Vol. 93

(Fig. 1), the significance of which was unknown until the complete
courtship sequence was observed (Kronestedt, 1979b, and ms. in
prep.). Unlike what is common in lycosids, the female in this species
plays a ritualistic active part in the premating display. The male is
unable to mount the female before the following sequence has been
passed through. The female will grasp one of the male’s first tibiae
with her chelicerae and pull him towards her, all the time holding
her grip around his first tibia. This phase will last for approx. 10 s.
After being released, the male will immediately mount the female.

On each side of the swollen first tibiae there is an oblique depres-
sion which may aid the female in maintaining her grip. Moreover,
these tibiae are black and strongly sclerotized (except for the depres-
sions). Their unique shape is evidently essential in the premating
display of A. cuneata, and thus also a strong isolating mechanism
when connected to behaviour. What releases the grasping behaviour
of the female? No definite answer can be given until further exten-
sive experiments have been made. However, in trying to find
whether there is any chemical cue involved, the male tibia was exam-
ined using SEM.

The cuticle of the first tibia in male A. cuneata is equipped with
numerous pores (Fig. 2a), a condition hitherto unknown among
lycosids. These pores could well be the emitting site for some type of
aphrodisiac. In the closely related species A. pulverulenta (Clerck),
with normal first tibiae in the adult male, less abundant and more
scattered leg pores were observed (Fig. 2b). The latter condition was
also found in adult females of the mentioned Alopecosa species and
in both sexes of other lycosid species as well (Fig. 2c, d). Therefore,
it is assumed that the situation in male A. cuneata is a special
adaptation of a commonly occurring contact pheromone releasing
system in lycosid spiders. If these presumptive pheromones are, at
least in part, volatile, they are also candidates in olfactory commu-
nication, for which other receptors may operate (e. g. the tarsal
organ: Dumpert, 1978).

Most investigators have focused on the means by which males
find and recognize females. However, it is of utmost importance for
the female to identify the proper male, as males are often less dis-
criminant. Chemical recognition of males by females in lycosids is
little studied but probably of significance (Tietjen and Rovner,

Fig. 2. (Right). Presumably pheromone-releasing pores dorsally on adult male first tibia, a) Alopecosa cuneata, b) A.
pulverulenta, c) Pardosa fulvipes, d) Trochosa spinipalpis. Scale: 2 pm (a), 1 pm (b-d).

130 Psyche [Vol. 93

1982), as already assumed for nocturnal species of the genus Tro-
chosa (Engelhardt, 1964).

Semiochemicals play an indispensible role in spider communica-
tion. Locating sites of pheromone production and perception is
essential for understanding behaviours and morphological adapta-
tions of sexual significance.

Specimens of Alopecosa cuneata (Clerck), A. pulverulenta
(Clerck), Pardosa fulvipes (Collett), and Trochosa spinipalpis
F.O.P. -Cambridge were all collected in pitfall traps with formalin in
the vicinity of Stockholm, Sweden. The material was stored in
ethanol, and parts used for SEM were dehydrated in an ethanol
series, kept in xylene for one or two days, cleaned in ultrasonic
cleaner, air-dried, mounted on SEM stubs, and sputter-coated with
Pd-Au. Examination was carried out with a JEOL JSM-35 at 15kV.

Summary

The cuticle of lycosid spider legs is shown to be equipped with
pores presumably involved in the release of sex pheromones. The
pores occur in both sexes. The male of Alopecosa cuneata has an
increased number of pores on its first tibiae, and the premating
behaviour in this species speaks in favour of the male producing
some aphrodisiac from the leg pores.

Literature Cited

Bristowe, W. S. and Locket, G. H. 1926. The courtship of British lycosid spi-
ders, and its probable significance. Proc. zool. Soc. Lond. 1926: 317-347.
Dijkstra, H. 1976. Searching behaviour and tactochemical orientation in males
of the wolfspider Pardosa amentata (Cl.) (Araneae, Lycosidae). Proc. K. ned.
Akad. Wet. (C) 79: 235-244.

Dondale, C. D. and Hegdekar, B. M. 1973. The contact sex pheromone of
Pardosa lapidicina Emerton (Araneida: Lycosidae). Can. J. Zool. 51: 400-401.
Dumpert, K. 1978. Spider odor receptor: Electrophysiological proof. Experien-
tia 34: 754-755.

Engelhardt, W. 1964. Die mitteleuropaischen Arten der Gattung Trochosa C.
L. Koch, 1848 (Araneae, Lycosidae). Morphologie, Chemotaxonomie, Biologie,
Autokologie. Z. Morph. Okol. Tiere 54: 219-392.

Hegdekar, B. M. and Dondale, C. D. 1969. A contact sex pheromone and some
response parameters in lycosid spiders. Can. J. Zool. 47: 1-4.

Kaston, B. J. 1936. The senses involved in the courtship of some vagabond spi-
ders. Entomologica am. (N. S.) 16: 97-167.

1986]

Kronstedt — Wolf spiders

131

Kronestedt, T. 1979a. Study on chemosensitive hairs in wolf spiders (Araneae,
Lycosidae) by scanning electron microscopy. Zool. Scr. 8: 279-285.

Kronestedt, T. 1979b. Etologiska karaktarer vid taxonomiska studier av
vargspindlar. Ent. Tidskr. 100: 194-199.

Richter, C. J. J., Stolting, H. C. J. and Vlijm, L. 1971. Silk production in
adult females of the wolf spider Pardosa amentata (Lycosidae, Araneae). J.
Zool., Lond. 165: 285-290.

Robert, T. and Krafft, B. 1981. Contribution a l’etude des mechanismes de
la communication tacto-chimique intervenant dans le rapprochement des sexes
chez Pardosa hortensis Thorell (Araneae, Lycosidae). Atti Soc. tosc. Sci. nat.
Memorie (B) 88 (Suppl.): 143-153.

Rqvner, J. S. 1968. An analysis of display in the lycosid spider Lycosa rabida
Walckenaer. Anim. Behav. 16: 358-369.

Tietjen, W. J. 1977. Dragline-following by male lycosid spiders. Psyche 84:
165-178.

Tietjen, W. J. 1979a. Tests for olfactory communication in four species of wolf
spiders (Araneae, Lycosidae). J. Arachnol. 6: 197-206.

Tietjen, W. J. 1979b. Is the sex pheromone of Lycosa rabida (Araneae: Lycosi-
dae) deposited on a substratum? J. Arachnol. 6: 207-212.

Tietjen, W. J. and Rovner, J. S. 1980. Trail-following behaviour in two species
of wolf spiders: Sensory and etho-ecological concomitants. Anim. Behav. 28:
735-741.

Tietjen, W. J. and Rovner, J. S. 1982. Chemical communication in lycosids and
other spiders. In Spider communication. Mechanisms and ecological signifi-
cance (P. N. Witt and J. S. Rovner, eds.): 249-279. Princeton Univ. Press,
Princeton, N.J.

A NEW ARBORICOLOUS THYREODON
FROM COSTA RICA

(HYMENOPTERA ICHNEUMONIDAE: OPHIONINAE).

By Charles C. Porter1

Department of Biological Sciences, Fordham University
Bronx, NY 10458

Through courtesy of Daniel H. Janzen of the Department of
Biology at the University of Pennsylvania, I have received for study
a new Thyreodon of the Atricolor group (Porter 1984), reared by
him in Costa Rican Tropical Deciduous Forest at Santa Rosa Na-
tional Park. I herewith describe this ecologically aberrant Thyreodon.

1. Thyreodon santarosae Porter, new species
(Figs. 1, 2)

Female. Color : antenna varying from almost all black to exten-
sively dusky, brown, or dull yellowish brown; head and body shin-
ing black to brownish black (more lustrous on gaster) and with
variably developed, diffuse, dull to (occasionally) light brown stain-
ing that usually is best developed on mandible and on gastric ter-
gites 2 and 3 in part; legs sometimes entirely black or often with
variable brownish suffusion on coxae and trochanters, trochantelli
and femora shining medium brown with some dusky staining, and
tibiae and tarsi dull pale brown with dusky only on last tarsomere;
wings varying from almost entirely blackish to subdued golden yel-
low with blackish on apical 0.3 of fore wing, sometimes also near
base of fore wing, as well as on apical 0.3 of hind wing and conspic-
uously (but often not extensively) in anellan cell of hind wing.

Length of fore wing: 15.6-19.0 mm. Flagellum: with 57-60 seg-

ments; 1st segment 2.0-2. 3 as long as deep at apex. Mandible: with
numerous, medium sized to large, basally denser, but mostly well
discrete punctures. Malar space: 0.54-0.63 as long as basal width of
mandible. Temple: 0.70-0.88 as long as eye in dorsal view; rounded

'Research Associate, Florida State Collection of Arthropods, Florida Department of
Agriculture and Consumer Services, Division of Plant Industry, P.O. Box 1269,
Gainesville FL 32602.

Manuscript received by the editor September 20, 1985

133

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[Vol. 93

Figs. 1 and 2. Thyreodon santarosae, 9- Paratypes. Dorsal views of entire
insects. Fig. 1. Morph with dark wings and dark legs. Fig. 2. Morph with largely
yellowish wings and partly pale legs. Note that males resemble females in dorsal view.

1986] Porter — Thyreodon from Costa Rica 135

off and not receding or slightly expanded behind eyes. Occipital
carina: bent mesad well above base of mandible, not approaching
hypostomal carina. Clypeus: with abundant, commingled small to
large punctures, most of which are separated by conspicuous
smooth interspaces. Lateral ocellus: 0.77-0.90 as long as OOL.
Mesoscutum: notauli not crested near base, broad and shallow,
often (not always) becoming much weaker apicad, traceable 0.8
length of mesoscutum, scarcely convergent rearward; surface shin-
ing with numerous, small to medium sized, sharp punctures that are
mostly separated by at least their diameters and sometimes in gen-
eral by more than their diameters. Scutellum: high, convex, and
shining with mostly subadjacent to adjacent small to medium sized,
sharp punctures and with lateral carinae developed only at its base.
Mesopleuron: sternaulus faint but usually percurrent; surface shin-
ing and with abundant, medium sized to small, sharp punctures that
are mostly subadjacent to a little reticulately adjacent on lower 0.5
but which average slightly sparser on upper 0.5; speculum smooth
and polished. Lower metapleuron: dully shining with small to
medium sized, sharp, subadjacent punctures and some coarse periph-
eral wrinkling. Propodeum: swollen, contours rounded, the dorsal,
lateral, and apical faces not sharply discrete; basal face shining with
long and dense appressed grayish setae, its median field gently
swollen and with the punctures very dense and tiny, its lateral field
more shining with larger and more widely spaced punctures that
expose much polished integument basad; lateral face with long and
dense appressed setae and sometimes with variably developed mod-
erately strong and mostly longitudinal wrinkles, as well as always
with abundant medium sized, mostly subadjacent to adjacent (or
sometimes extensively adjacent) punctures; apical face in compari-
son to rest of propodeum at least in large part contrastingly smooth
and brilliantly polished with tiny punctures that emit long but little
overlapping setae and often with many long and oblique, well
separated and rather fine wrinkles.

Male. Color: shows same range of variation as noted for
female and, in addition, is marked with dull to bright yellow as
follows: usually on basal 20-25 flagellomeres below (becoming
duller distad); with a small to large ventral blotch on scape; on as
little as 0.5 to as much as almost all of face (and sometimes also on

136

Psyche

[Vol. 93

much of interantennal crest), except for brown on antennal sockets,
brown also on a large to very large, quadrangular to (more often)
dorsally narrowed median facial blotch (which is occasionally
reduced to a small pale brown tinge and which sometimes, when
conspicuous, surrounds a yellow area along clypeo-frontal suture),
and also brown on a large to small or even obsolete area in and
(frequently) above and below anterior tentorial pit, which may be
confluent dorsally with the median brown facial area; sometimes
also with yellow in malar space and broadly bordering hind orbit to
as much as upper 0.2 of eye; and yellow also on most of clypeus
except for its pale brown apical margin (clypeus rarely in large part
brown with yellow only laterad); on most of basal 0.7 of mandible;
on maxillary palpomeres 1-3; sometimes on an anterio-ventral fore
coxal blotch; occasionally on a small dorso-lateral mid coxal blotch;
on a broad anterio-dorsal stripe on fore and sometimes mid tro-
chanters (yellow on mid trochanter often dull and weakly de-
veloped); sometimes also anterio-dorsally on fore and mid trochan-
telli; on a broad anterio-dorsal front femoral stripe; and rarely also
on part of mid femur anterio-dorsally.

Length of fore wing: 14.6-18.5 mm. Malar space: 0.63-0.71 as
long as basal width of mandible. Hind tarsus: segments 1-4 beneath
with setae longer and denser than in female, pale gray, obliquely
outstanding, closely packed, 0.4 as long as depth of tarsomeres.
Clasper: in lateral view with dorsal margin on apical 0.46 broadly
concave; dorso-apical angle semi-acute (not spiniform) and slightly
upcurved; apical margin reclivously oblique; apico-ventral angle
blunt. Other characters as described for female.

Type Material. Holotype $\ COSTA RICA, Guanacaste Pro-
vince, Santa Rosa National Park, D. H. Janzen, 1984 (Washing-
ton). Paratypes: 135 and 9$\ same data as Holotype: 2$ (Wash-
ington), 1? and 1(5 (Cambridge), 19 (College Station), 1$ and 1<5
(Gainesville), 1$ and \$ (Lawrence), 1$ and 1(5 (London), 1? and
1(5 (Los Angeles), 19 and 1(5 (New York), 1? and \$ (Ottawa); 1?
and 15 (Philadelphia); 1$ and 1(5 (Townes); 1? (Porter).

Variation. Thyreodon santarosae shows unusually marked
intrapopulation variability in wing and leg color. This variation
correlates appreciably but imperfectly with sex. Of the 139 exam-
ined, 10 have the wings predominantly yellow and in 9 of these

1986]

Porter — Thyreodon from Costa Rica

137

specimens all the tibiae and tarsi are pale brown (legs wholly black
in the 10th yellow-winged $), whereas both wings and legs are black
in the 3 remaining $. Among the 10 $, 1 has yellow wings but dark
legs, 1 black wings but pale legs, and 8 both black legs and wings.

Relationships. Thyreodon santarosae belongs to the Atricolor
group of Thyreodon (Porter 1984). This assemblage includes robust
species with inflated temples, often weakly impressed notauli, and
without a transverse or longitudinal crest at the anterior end of the
notauli. It has several undescribed Sonoran, Middle American,
Caribbean and South American species plus the Nearctic T.
atricolor (Olivier), the Sonoran T. fernaldi Hooker, and T. orna-
tipennis Cresson from the Mexican wet tropics.

Thyreodon santarosae differs most trenchantly from its relatives
in the extensively smooth and polished apical propodeal face (hind
face of propodeum coarsely reticulo-rugose in T. atricolor and T.
fernaldi, finely and densely puncto-reticulate in T. ornatipennis).
Other diagnostic features are its laterally almost ecarinate scutel-
lum; smooth speculum; and relatively sparse (mostly subadjacent or
more distant) mandibular, clypeal, mesoscutal, and mesopleural
punctures.

Field Observations and Hosts. Santa Rosa National Park,
the type locality, is in Tropical Deciduous Forest at 250-350 m on
the Pacific Coast of Guanacaste Province, Costa Rica. Daniel H.
Janzen reared the entire type series from “larvae of Saturniidae in
the Subfamily Ceratocampinae. . .collected at 3-20 m above the
ground” (personal communication). The parasites emerged during
April to December 1984. No individuals of T. santarosae were
obtained by hand nets or Malaise Traps.

This species appears to be unique among Thyreodon for its
apparent restriction to intermediate and higher strata of a Tropical
Forest community and because it attacks ceratocampine caterpil-
lars. Most other Thyreodon fly close to the ground or around
understory shrubs at no more than 2 m altitude, and the only pre-
vious rearing data for this genus involve sphingid Lepidoptera that
pupate in the ground (Porter 1984).

Specific Name. For Costa Rica’s Santa Rosa National Park,
where Dan Janzen has found enthusiastic support for his ecological
studies.

138

Psyche

Collections

[Vol. 93

Listed below are the collections in which type material of T.
santarosae is to be deposited. Institutional collections are coded by
the names of the cities where they are housed, individual collections
according to the surnames of their owners.

Cambridge. Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge, MA 02138.

college station. Department of Entomology, Texas A&M Uni-
versity, College Station, TX 77843.

Gainesville. Florida State Collection of Arthropods, Division of
Plant Industry, Florida Department of Agriculture and Con-
sumer Services, Gainesville FL 32602.

Lawrence. Department of Entomology, Snow Entomological
Museum, The University of Kansas, Lawrence, KS 66045.
London. Department of Entomology, British Museum (Natural
History), Cromwell Road, London, SW7 5BD, England.
los angeles. Natural History Museum, Los Angeles County
Museum of Natural History, Exposition Park, 900 Exposition
Boulevard, Los Angeles, CA 90007.
new york. Department of Entomology, American Museum of
Natural History, Central Park West at 79th Street, New York,
NY 10024.

Ottawa. Canadian National Collection, Biosystematics Research
Institute, Agriculture Canada, Ottawa, K1 A 06C, Canada.
Philadelphia. Department of Biology, University of Pennsylva-
nia, Philadelphia, PA 19104.

townes. American Entomological Institute, c/o Dr. Virendra
Gupta, Division of Plant Industry, Florida Department of
Agriculture and Consumer Services, Gainesville FL 32602.
porter. Collection of Charles C. Porter, 301 North 39th Street,
McAllen, TX 78501.

Washington. Department of Entomology, U. S. National Museum,
NHB 168, Washington, DC 20560.

Acknowledgments

This paper was supported by Servicio de Parques Nacionales de
Costa Rica and by Daniel H. Janzen’s National Science Foundation
Grant BSR 8403531.

1986] Porter — Thyreodon from Costa Rica 139

Summary

Thyreodon santarosae n. sp. differs from its relatives in the Atri-
color species group by having the hind propodeal face broadly pol-
ished. It was obtained only by rearing from ceratocampine saturniids
(Lepidoptera) in Tropical Deciduous Forest at Santa Rosa National
Park in northeast lowland Costa Rica. Host larvae were collected at
3-20 m in the forest overstory. Other known Thyreodon are active
near ground level and those few that have been reared parasitize
sphingid Lepidoptera.

Literature Cited

Porter, C. 1984. Laticinctus group Thyreodon in the northern Neotropics.
Wasmann Journal of Biology 42: 40-71..

DISTINGUISHING THE JUMPING SPIDERS
ERIS MILITARIS AND ERIS FLA VA
IN NORTH AMERICA (ARANEAE: SALTICIDAE)*

By Wayne Maddison

Museum of Comparative Zoology,

Harvard University,

Cambridge, Massachusetts 02138

The jumping spiders now identified as Eris marginata are among
the most frequently encountered in North America, for they are
common on trees, shrubs and herbs throughout much of the
continent. However, two species have been confused under this
name. One is an abundant transamerican species whose proper
name is Eris militaris; the other is Eris flava, widely distributed in
eastern North America though common only in the southeast. In
this paper I describe how they may be distinguished. The abbrevia-
tion MCZ refers to the Museum of Comparative Zoology; ZMB to
the Zoologisches Museum, Humboldt-Universitat zu Berlin.

Eris militaris (Hentz), new combination
Figures 2-7, 14

Attus militaris Hentz 1845: 201, pi. xvii, fig. 109, 11<3- Type material lost or de-
stroyed (see Remarks, below), from North Carolina and Alabama. Neotype here
designated, 1$ in MCZ from North Carolina with label “NC: jackson co.,
Coyle Farm, 1.5 mi SW of Webster, 7 Sept. 1975; F. Coyle.”

Plexippus albovittatus C. L. Koch 1846: 118, fig. 11789- Syntypes in ZMB 19 with
labels “P. albovittatus 1739” and “1739”, and 19 with label “P. albovittatus
ZMB 1739”, examined. Type locality Pennsylvania (Koch, 1846). new synonymy.
Eris aurigera C. L. Koch 1846: 189, fig. 1237 S- Syntypes in ZMB 1$ with carapace
and abdomen in alcohol with labels “Eris aurigera C. L. Koch*, 1774” and
“Typus” and remaining body parts mounted on cover slip in small box with label
“(Eris aurigera Koch*) Dendryphantes marginatus Walck., ZMB 1774a, D. mil-
itaris Hentz, XI, Syntypus” and \$ mounted on cover slip with label “Eris
aurigera*, C. L. Koch, $ Rf.?, 1774b, Syntyp., Paraphidippus”, both examined.
Euophrys humilis C. L. Koch 1846: 217, fig. 12629- Holotype 19 in ZMB with labels
“Holotypus”, “1804”, “ZU 1804”, “Euophrys humilis”, “Pennsylvanien, Zim-
mermann leg.”, “Zool. Mus. Berlin”, examined.

Manuscript received by the editor January 15, 1986.

141

142

Psyche

[Vol. 93

Icius albovittatus Keyserling 1884: 502, fig. 10$. Syntypes 1$ 1 immature in MCZ
with labels “15 Icius albovittatus Keys., $ Massachusetts”, “15”, examined.
(Junior homonym of Icius albovittatus Keyserling, 1883.)

Icius moestus Banks 1892: 77, pi. V, fig. 33 3- Holotype in MCZ 1$ with labels “Icius
moestus Bks”, “Dendryphantes moestus Bks type”, “Ithaca, N.Y.”, “Nathan
Banks Coll.” examined.

Dendryphantes marginatus: — Simon 1901: 624 (not Attus marginatus Walckenaer;
see Remarks below).

Dendryphantes louisianus Chamberlin 1924: 34, fig. 51$. Holotype in MCZ l$with
label “ Dendryphantes louisianus Ch. $ Type, La.: Kenner, R. V. Chamberlin
Coll.” examined.

Phidippus molinor Chamberlin 1925: 133, fig. 49$. Holotype in MCZ 1$ with label
“ Dendryphantes molinor Chamb., $ holotype, Utah: Mill Creek Canyon, R. V.
Chamberlin Coll. 1071”, examined.

Paraphidippus marginatus: — Chickering 1944: 180 (in part), figs. 78-82.
Paraphidippus marginatus: — Kaston, 1948: 479.

Eris marginata: — Kaston 1973: 118 (in part), figs. 51-54.

Remarks on synonymy: It is unfortunate that most workers since
about 1930 have accepted without question Simon’s (1901: 624)
synonymy of Attus militaris Hentz 1845 with Attus marginatus
Walckenaer 1837, for the synonymy is incorrect: Walckenaer’s orig-
inal description (p. 466) and Abbot’s figure (number 444) clearly
refer to Hentzia palmarum (Hentz). Walckenaer refers to an elon-
gate abdomen, a fawn-brown first pair of legs, yellow posterior legs,
and chelicerae elongate and held in front, whereas Eris militaris has
an abdomen of typical width, posterior legs strongly marked with
dark brown, and chelicerae robust and divergent. Abbot’s drawing
(see Figure 1), on which Walckenaer based his description of A.
marginatus, unambiguously portrays a male Hentzia palmarum,
given that his specimen was from Georgia. Because the name margi-
natus is inappropriate for the transamerican Eris species, another
name must be used. The type material for the next oldest name,
Attus militaris, is apparently lost or destroyed. Burgess (1875, vii)
said that only 60 specimens glued on cards remained of Hentz’s
collections, the remainder having been destroyed. The surviving
specimens were in the collection of the Boston Society of Natural
History, which has subsequently become the Boston Museum of
Science. The Museum of Science no longer has these specimens nor
any record of them (D. Salvatore, pers. comm.), nor does the MCZ,
which received many of the Society’s collections. I presume Hentz’s
types to have been lost or destroyed. Without the type material the
interpretation of Attus militaris is not entirely clear, for Hentz’s

1986]

Maddison — Jumping spiders

143

Fig. 1. Abbot’s figure 444 on which Walckenaer (1837) based his description of
Attus marginatus. Abbot’s legend reads “444. Aranea. Taken 4th April, two upon a
Myrtle on the side of a Pond in the Oak Woods of Burke County. Rare.” From a
color slide taken by Allen Brady of Abbot’s (1792) original in the British Museum
(Natural History).

1845 description might refer to either the transamerican or the east-
ern species. Still, his failure to describe a white marginal band in the
male, and his illustration showing a dark femur on the male palp
(better seen in his original color drawing) both suggest that he had
the transamerican species. Therefore, I have designated a male of
this species as neotype for Attus militaris. This is advantageous for
nomenclatural stability, for Hentz’s name was the only name com-
monly used before 1930 for the abundant transamerican species. In
contrast, I have been unable to find any use of Koch’s names albo-
vittatus, aurigera, and humilis since 1864, except in synonymies and
catalogues.

Male carapace margin and clypeus brown (Fig. 7), without white
scales, or if the clypeus has white scales, then they only rarely extend
along margin past palps. Longitudinal white bands extending back

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Psyche

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Figs. 2-7. Eris militaris 2. Left palp, ventral view (Port Elgin, Ontario).
3. Epigynum, ventral view (Dwight, Ontario). 4. Cleared epigynum, dorsal view
(Dwight, Ontario); arrow shows flowerlike gland opening. 5. Left palp, oblique
view from the ventral-retrolateral-distal (Pine Lake, Michigan), and 6. Same (For-
syth, Georgia); arrow shows lack of wrinkles on retrolateral half of embolar base.
7. Male carapace and chelicerae (Walloon Lake, Michigan), oblique view. Scale bars
0.1 mm for 2-6; 1 mm for 7.

from anterior lateral eyes usually broad. Palp femur and patella as
dark as the more distal segments. Embolus shorter and stouter, and
more broadly joined to the embolar base (Figs. 5, 6) than in flava.
Wrinkles on the ventral surface of the embolar base usually straight,
and absent from the retrolateral half (Figs. 2, 5, 6; see arrow in Fig.
6).

Female carapace generally with a continuous covering of white
scales above margin beneath anterior lateral eyes. Epigynal open-
ings usually smaller and more laterally facing (Fig. 3) than in flava.

1986]

Maddison — Jumping spiders

145

Figs. 8-13. Eris flava 8. Left palp, ventral view (Florida City, Florida).
9. Epigynum, ventral view (Point Pelee, Ontario). 10. Cleared epigynum, dorsal
view (Point Pelee, Ontario); arrow shows flowerlike gland opening. 11. Left palp,
oblique view from the ventral-retrolateral-distal (Pine Lake, Michigan), and
12. Same (Florida City, Florida); arrow shows curled wrinkles on retrolateral half of
embolar base. 13. Male carapace and chelicerae (S. of St. Joseph, Michigan),
oblique view. Scale bars 0.1 mm for 8-12; 1 mm for 13.

Each duct proceeds medially to a flower-like structure (apparently
gland openings; see arrow in Fig. 4), then posteriorly.

Habitat varied; common on trees and shrubs. Distribution shown
in Fig. 14.

Eris flava (Peckham and Peckham)

Figures 8-13, 15

Dendryphantes flavus Peckham & Peckham 1888: 39, pi. I, fig. 27$, pi. Ill, figs. 27,
27a$. Syntypes in MCZ 3$, 1 immature $ with label “Dendryphantes flavus

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Pkm., 1888. New York. Type. 9”, examined. Type vial also contains one imma-
ture Phidippus.

Dendryphantes armatus Banks 1909: 167, fig. 5$. Syntypes in MCZ 7$ with labels
“S. de las Vegas, Cuba 10-17. 07”, Dendryphantes armatus Bks.”, “type”, “D.
militaris H, armatus B, Cuba”, “Nathan Banks Coll.” examined, new

SYNONYMY.

Paraphidippus militaris: — Bryant 1940: 502.

Paraphidippus marginatus: — Chickering 1944: 180 (in part).

Eris marginata: — Kaston 1973: 118 (in part).

Eris flava: — Kaston 1973: 120, figs. 66-67.

Remarks on synonymy: Eris flava was thought to be an uncom-
monly collected species known only from females (Kaston, 1973).

1986]

Maddison — Jumping spiders

147

While less common than militaris, many males are available in col-
lections (including the Peckham and Banks collections), identified
as militaris or marginata. Chickering’s Michigan collections are
mixed E. militaris and E. flava. Though most of Kaston’s identifica-
tions were correct, at least some Floridian males he identified as
E. marginata prior to his 1973 paper are E. flava.

Male carapace with marginal band of white scales extending
across clypeus (Fig. 13) and usually back well past the palps. Longi-
tudinal white bands extending back from ALE usually narrower
than in militaris. Palp femur and often patella distinctly paler than
more distal segments. Embolus longer and thinner than in militaris,
arising more abruptly and more directly behind the embolar base
(Figs. 11, 12). Wrinkles on embolar base, especially the more retro-
lateral ones, are distally curled retrolaterally (Figs. 8, 11, 12; see
arrow in Fig. 12).

Fig. 15. Distribution of Eris flava. The specimen from the North Platte River at
Bridgeport, Nebraska was collected by me, and the identification checked carefully.

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[Vol. 93

Female carapace with patch barren of scales just above the mar-
ginal white band at a point below anterior lateral eyes. Epigynal
openings wider and face more anteriorly (Fig. 9) than in militaris.
Each duct first proceeds posteriorly and then laterally to the flower-
like structure (Fig. 10, arrow), then posteriorly. The epigynal ducts
are the best distinguishing feature.

Habitat information is sparse, but the species appears to prefer
marshes and fields. Found in cedar swamp (Mass.), sweeping grass
and herbs near river (Nebr.), on vegetation in marshy area (Ont.),
meadow (111.), on Nelumbo lutea and in fields (Fla.). Distribution
shown in Fig. 15.

Acknowledgements

All but a few of the specimens examined are in the MCZ. The
types of Koch names were kindly loaned by M. Moritz and S.
Fischer of the ZMB. For the loan of the remaining specimens, I
thank E. Schlinger and D. Wagner (Essig Museum, University of
California, Berkeley) and W. J. Gertsch (American Museum of
Natural History). H. W. Levi and D. R. Maddison gave useful
comments on the manuscript.

References

Abbot, J.

1792. Drawings of the Insects of Georgia, in America, vol. 14, Spiders. Unpub-
lished manuscript in the British Museum (Natural History) (Copy seen).

Banks, N.

1 892. The spider fauna of the Upper Cayuga Lake Basin. Proc. Acad. Nat. Sci.
Philadelphia, 1892: 11-81, 5 pi.

1909. Arachnida of Cuba. Estacion central agronomica de Cuba, Second
Report, Part II, pp. 150-174.

Bryant, E. B.

1942. Cuban spiders in the Museum of Comparative Zoology. Bull. Mus.
Comp. Zool., 86: 259-532+22 pi.

Burgess, E.

1875. Preface, pages v-xiii, in A collection of the arachnological writings of
Nicholas Marcellus Hentz, M. D., Boston Soc. Nat. Hist.

Chamberlin, R. V.

1924. Descriptions of new American and Chinese spiders, with notes on other
Chinese species. Proc. U.S. Nat. Mus., 63(13): 1-38.

1925. New North American spiders. Proc. California Acad. Sci., (4) 14(7):
105-142.

1986]

Maddison — Jumping spiders

149

Chickering, A. M.

1944. The Salticidae (jumping spiders) of Michigan. Papers Michigan Acad.
Sci., Arts & Letters, 29: 139-222.

Hentz, N. M.

1832. On North American spiders. American J. Sci., 21: 99-122.

1845. Descriptions and figures of the Araneides of the United States. Boston J.
Nat. Hist., 5: 189-202.

Kaston, B. J.

1948. Spiders of Connecticut. Connecticut State Geol. and Nat. Hist. Survey,
70: 1-874.

1973. Four new species of Metaphidippus, with notes on related jumping spi-
ders (Araneae: Salticidae) from the eastern and central United States.
Trans. Amer. Micros. Soc., 92: 106-122.

Koch, C. L.

1846. Die Arachniden. Dreizehnter Band. Nurnberg, pp. 1-234.

Peckham, G. W. and E. G. Peckham

1888. Attidae of North America. Trans. Wisconsin Acad. Sci., Arts & Letters,
7: 3-104.

1909. Revision of the Attidae of North America. Trans. Wisconsin Acad. Sci.,
Arts & Letters 16: 355-646.

Simon, E.

1901. Histoire Naturelle des Araignees. Deuxieme edition. Tome 2, fasc. 3,
pp. 381-668. Paris.

Walckenaer, C. A.

1837. Histoire naturelle des Insectes. Apteres. Tome 1. Paris, pp. 1-682.

EVIDENCE OF WORKERS SERVING AS QUEENS
IN THE GENUS DIACAMMA
(Hymenoptera: Formicidae)

By Mark W. Moffett

Museum of Comparative Zoology, Harvard University,
Cambridge, Massachusetts 02138

There is no morphologically distinguishable queen caste known in
the ponerine genus Diacamma. Wheeler and Chapman (1922)
observed a typical Diacamma worker copulating with a normal
male, and it has been assumed that some workers are functioning as
reproductives. I report an experiment that supports this view.

Ants in the D. rugosum complex at Sullia in Karnataka State,
southern India, live in polydomous colonies; foragers move freely
between nests within a colony, which are separated by one to several
meters. Each nest is a blind-ended tunnel 10-25 cm deep containing
brood and between about 50-120 workers. When individual nests
within a colony were collected and kept in captivity, some workers
foraged frequently, while the remainder never left the artificial nest
tubes.

In a preliminary experiment conducted during February and
March, 1982, the ants taken from one nest were sorted into foraging
and non-foraging behavioral types and then further divided into
groups of 5-6, with eight groups of foragers (total 45 ants) and four
groups of non-foragers (total 21 ants); every group was provided a
separate test tube “nest” with stoppered water source and no brood.
The foraging ants continued to come and go from their nest tubes,
and in none of these groups were any eggs produced over a period of
a month. Non-foraging ants continued to stay within their nest
tubes and eventually had to be provided food within the tubes. In all
four non-foraging groups the test tubes soon held brood, and the
five immatures that survived to the pupal stage (three from one tube
and two from another) were workers.

This indicates that part of the worker population is fertilized and
is serving as queens, as is the case with the African ponerine
Ophthalmopone berthoudi (Peeters and Crewe, 1984, 1985), which
also lacks winged gynes.

I am grateful to R. Gadagkar and M. Gadgil for aid during my
stay in India.

151

152

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Literature Cited

Peeters, C. and R. Crewe. 1984. Insemination controls the reproductive division
of labor in a ponerine ant. Naturwissenschaften 71: 50-51.

1985. Worker reproduction in the ponerine ant Ophthalmopone berthoudi :

an alternative form of eusocial reproduction. Behav. Ecol. Sociobiol. 18: 29-37.
Wheeler, W.M. and J. W. Chapman. 1922. The mating of Diacamma. Psyche 29:
203-211.

NEW SPECIES AND GENERA OF AMISEGINAE
FROM ASIA (CHRYSIDIDAE, HYMENOPTERA)*

By Lynn Siri Kimsey

Department of Entomology,

University of California, Davis 95616

In a large shipment of miscellaneous, non-American chrysidids
sent to me by Henry Townes of the American Entomological Insti-
tute, Gainesville, Florida, all of the Amiseginae turned out to be
new species. The majority of these were collected by E. and M.
Becker in the Pasoh Forest Reserve in Malaysia. The new species of
Cladobethylus and Isegama represent range extensions for both
genera. Cladobethylus was previously known from Sri Lanka and
Mindanao, Philippines. Isegama has been previously described only
from Sri Lanka.

Holotypes have been deposited in the American Entomological
Institute, Gainesville, Florida.

A variety of structures, dimensions and abbreviations, used
below, need explanation. The malar space is the distance between
the base of the mandible and the ocular margin. On the mesopleu-
ron there are 2 possible carinae and/or sulci. The scrobal sulcus
extends transversely across the mesopleuron from the scrobal pit.
The oblique mesopleural carina originates below the pronotal lobe,
and extends ventrally. Subantennal distance is the length between a
line drawn across the lower edge of the antennal sockets and the
clypeal apex. Abbreviations used below are: F = flagellum, MOD =
midocellus diameter, PD = puncture diameter and T = gastral
tergum.

Atoposega simulans Kimsey, new species
(Figs. 1, 6)

Holotype female. Body length 5 mm. Face (fig. 1); scapal basin
with numerous coarse cross-ridges, bordered along ocular margin
by large punctures less than 0.6 PD apart; malar space 3 MOD;

* Manuscript received by the editor April 4, 1986.

153

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head width 1.3 times length; midocellus 2.5 MOD from ocular mar-
gin; subantennal distance 1 MOD; ocelli arranged in broad triangle;
hindocellus 1.2 diameters from ocular margin; F-I length 3X
breadth; F-II length 0.6X breadth; pronotum 0.5X as long as com-
bined median lengths of scutum, scutellum and metanotum; meso-
pleuron with large dense punctures, without scrobal sulcus; scutal
punctures coarse and contiguous, somewhat arranged in rows;
metanotum 0.8X scutellar length; forewing (fig. 6) densely setose
with dark bands across wing at medial vein and at apex of RS (fig.
6), entire wing brown-stained; hindfemur ventral surface coarsely
punctate; T-I and II polished and impunctate medially with lateral
zone of fine scratches and punctures; T-III-IV with apical band of
tiny punctures. Head black; scape light brown; pedicel and F-I whit-
ish, except apex of F-I blackish; F-II-XI blackish; thorax red,
except dorsal and posterior face of propodeum black; legs and
coxae red, except foretarsomeres, hindtibial apex and venter of
hindfemur dark brown; abdomen shiny black, with faint green tints
on T-I-II laterally.

Holotype female: MALAYSIA: Pasoh Forest Res., Negri S., 17
April 1980, P. and M. Becker (Gainesville). Paratypes: 10 females,
same data as type, except various dates from 8 July 1978 to 3
November 1979.

Discussion. This species appears to be structurally intermediate
between the other species of Atoposega: lineata Krombein and rieki
(Krombein). A. simulans has the long pronotum, patterned wings
and larger size of lineata, and T-I-II laterally “scratched” with
metallic tints and the forefemur rough and coarsely punctate, as in
rieki.

Bupon Kimsey, new genus

Diagnosis. Malar space with vertical sulcus; vertex with coarse
close punctation; brow with strongly projecting transverse ridge
(figs. 3, 4); eyes encircled by irregular carina; occipital carina well-
developed; scapal basin coarsely cross-ridged; male flagellum short
and cylindrical; pronotum about half as long as combined lengths of
scutum, scutellum and metanotum, with oblong pit posteromedially
and on lateral lobe; mesopleuron without scrobal sulcus or oblique

1986]

Kimsey — New Amiseginae from Asia

155

Figs. 1, 3, 7-9. Front view of face. Fig. 2.
Fig. 4. Lateral view of head. Figs. 5, 6. Forewing.

Dorsal view of thorax.

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mesopleural carina; forewing (fig. 5) with long slender stigma + Rl,
RS extended by evenly curved streak, medial vein arises before cu-a;
metanotum 0.9X as long as scutellum, medial enclosure punctate,
differently sculptured from lateral area; propodeum with short dor-
sal surface, abruptly declivous posteriorly, lateral angles short and
blunt; hindcoxa with dorsobasal carina; terga sharp-edged laterally,
covered with dense small punctures; tarsal claw with large perpen-
dicular submedial tooth.

Type: Bupon pasohanus Kimsey.

Etymology. Bu - great, pons - bridge (Latin, masculine).

Discussion. The most unusual diagnostic feature of Bupon is
the strongly projecting transverse frontal carina. The only other
amisegine genus with any indication of such a carina is Perissosega
where it is faint by comparison. In other respects Bupon more
closely resembles Cladobethylus, based on the lack of most of the
derived characteristics found in the Amiseginae. Two derived char-
acteristics that are found in Bupon and will immediately distinguish
this group are the transverse facial carina and the short, relatively
broad male flagellum.

Bupon pasohanus Kimsey new species
(Figs. 2-5)

Holotype male. Body length 4.5 mm. Face (figs. 3, 4); scapal basin
with coarse cross-ridges, deeply sunken below transverse shelf-like
ridge, punctures 0.2-0. 5 PD apart; eye encircled by carina; clypeal
apex broadly rounded; subantennal distance 1 MOD; malar space
2.1 MOD, with vertical sulcus; ocelli arranged in broad triangle;
hindocelli 0.8 diameter from ocular margin; midocellus 1.8 MOD
from ocular margin; occipital carina complete; pronotum 0.5X
combined lengths of scutum, scutellum and metanotum; with
oblong pit posteromedially and on lateral lobe; thorax (fig. 2), with
dorsal punctures coarse and contiguous; mesopleuron sculptured
like pronotum, without scrobal sulcus; propodeal posterior face
finely and densely rugose, lateral angle short and blunt; terga sharp-
edged laterally, with coarse small punctures 0.5 PD apart. Body
black; legs including coxae yellow, except hindtarsomeres and
apices of hindfemur and tibia blackish; antenna dark brown, except
scape paler beneath.

1986]

Kimsey — New Amiseginae from Asia

157

Holotype male: MALAYSIA: Pasoh Forest Res., Negri S., E.
and M. Becker, 27 July 1979, secondary forest (Gainesville). Para-
types: 30 males, collected from June 1978 to April 1980.

Cladobethylus aquilus Kimsey, new species
(Fig. 7)

Holotype male. Body length 3 mm. Face (fig. 7); scapal basin
primarily smooth with short strip of cross-ridges on either side of
broad medial stripe; clypeus long and rounded apically, subantennal
distance 1.1 MOD; malar space 3.5 MOD long; head about as long
as wide; midocellus 2 MOD from ocular margin; ocelli arranged in
nearly equilateral triangle; hindocelli separated from ocular margin
by 0.8 diameters; pronotum about 1.2X as long as scutum; meso-
pleuron with long parallel-sided scrobal sulcus, punctures slightly
larger than on pronotum; metapleuron smooth below hindwing
base; propodeum with posteromedial stripe smooth but somewhat
irregular, bordered by carina laterally; terga with basal zone of tiny
punctures 0.5-2 PD apart. Body black, except pronotum and scu-
tum with faint blue tint; legs including coxae yellow; antennae dark
brown; mandibles yellowish brown.

Female unknown.

Holotype male: PAPUA NEW GUINEA: Bulolo, 900 m, 13
February- 13 March 1979, J. Sedlacek (Gainesville). Paratypes: 4
males, same data as type, 1 male Baiyer River, 6-25 February 1979,
1 100 m.

Discussion. The face of aquilus resembles that of C. ceylonicus
Krombein based on the strongly converging lower sides of the face
and the greatly reduced cross-ridging in the scapal basin. However,
aquilus can be distinguished from this and other Cladobethylus
species by the long clypeus, smooth metapleuron, dorsum with faint
blue tints, dark brown antenna, and pronotum longer than the
scutum.

Cladobethylus gilbus Kimsey, new species
(Fig. 8)

Holotype male. Body length 4 mm. Face (fig. 8); scapal basin with
numerous fine cross-ridges, bordered by large punctures less than 0.6
PD apart; head venter with 2 ovoid foveae along midline of genal

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bridge; malar space 3.2 MOD long; head 1.2X as wide as long;
midocellus 2.5 MOD from ocular margin; ocelli arranged in a nearly
equilateral triangle; hindocelli separated from ocular margin by 1
diameter; subantennal distance 0.9 MOD; clypeal apex truncate;
pronotum with fine short posteromedial line; mesopleuron without
scrobal sulcus, punctation same as pronotum; metapleuron with
zone of cross-ridging below hindwing base; propodeum with broad,
polished, impunctate, vaguely margined, posteromedial stripe; terga
polished and impunctate. Body black, except pronotum and scutum
with faint blue tints; antennae, legs including coxae yellow, and
mandibles yellow with red tips.

Female unknown.

Holotype male: MALAYSIA: Pasoh Forest Res., Negri S., 5
November 1978. P. and M. Becker (Gainesville). Paratypes: 106
males and 50 females, same data as type, except dates from May
1978 to May 1980.

Discussion. C. gilbus males have two ovoid foveae underneath
the head, one on either side of the genal bridge. I have seen no other
Cladobethylus males with this modification. Otherwise, gilbus can
be distinguished by the densely cross-ridged scapal basin, yellow
antennae, truncate, clypeal apex, subequal pronotum and scutum
(in length), blue-tinted pronotum and scutum and metapleuron with
short zone of cross-ridging.

Cladobethylus japonicus Kimsey, new species
(Fig. 9)

Holotype female. Body length 2.5 mm; face (fig. 9); scapal basin
smooth with short strip of cross-ridges on either side of broad
medial stripe; clypeus short, broadly rounded; subantennal distance
0.6 MOD; malar space 5 MOD long; face broad across genal region,
about as broad as long; midocellus 2.6 MOD from ocular margin;
ocelli arranged in nearly equilateral triangle; hindocelli separated
from ocular margin by 0.3 diameters; mesopleuron with long
parallel-sided scrobal sulcus, punctures larger than on pronotum;
metapleuron cross-ridged from hindwing base nearly to midcoxa;
propodeum posteromedial stripe rough and enclosed by strong Car-
ina; T-I smooth and impunctate; T-II-IV smooth with tiny scattered
punctures 4-6 PD apart. Body black with bluish tints on vertex and

1986]

Kimsey — New Amiseginae from Asia

159

pronotum; legs including coxae yellow; scape dark brown becoming
paler distally; pedicel and F-I-III yellow, remaining flagellomeres
brown; mandibles brown.

Male unknown.

Holotype female: JAPAN: Kyoto, 8 August 1980, H. and M.
Townes (Gainesville).

Discussion. This species has several unusual features. The eyes
have very long dense setulae, the hindocelli are very close to the
ocular margins and the malar space is also very long. In addition,
the lower face is quite broad, the scapal basin has only narrow
stripes of cross-ridging, the pronotum is longer than the scutum, the
metapleuron is cross-ridged from wing base to caxa, and only the
pedicel and F-I-III are yellow, the rest of the antenna is brown.

Isegama malaysiana Kimsey, new species
(Fig. 17)

Holotype female. Body length 3 mm. Face (fig. 17) polished with
sparse small punctures, 3-5 PD apart; eyes without distinct setulae;
scapal basin shallow with faint cross-ridges; lower face strongly
converging; eyes bulging farthest below middle; gena bulging along
lower third of eye; malar space 3.2 MOD long; clypeus short, suban-
tennal distance 0.4 MOD; midocellus 2 MOD from ocular margin;
ocelli arranged in broad triangle; hindocellus nearly touching ocular
margin; vertex strongly convex; pronotum flattened, with medial
groove and pit before lateral lobe, subequal to scutal length; meso-
pleuron with moderate punctures about 1 PD apart on anterior half,
posterior half impunctate and polished, scrobal sulcus straight, nar-
row and parallel-sided, oblique mesopleural carina well-developed;
metanotum 0.8X scutellar length, medial enclosure smooth with
tiny punctures, about 1 PD apart; propodeum with dorsal enclo-
sures polished and impunctate, posterior face rugose; T-I-III covered
with close small punctures, nearly contiguous anteriorly but becom-
ing more dispersed posteriorly, with impunctate medial stripe; T-IV
covered with small punctures. Head, thorax and abdomen black,
with blue tints on head, pronotum, scutum, scutellum and medial
enclosure of metanotum; scape, pedicel and F-I-III yellow, F-IV to
apex brown; legs including coxae yellow, wings faintly brown
tinted.

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Male. Same as female except face more coarsely punctate,
punctures 0.5- 1.0 PD apart, eyes normal; F-I 3X as long as broad;
terga more closely and coarsely punctate, without clearly indicated
impunctate medial stripe; head and thorax without metallic tints;
entire antennae reddish brown; femora dark brown becoming paler
distally; coxae black.

Holotype female. MALAYSIA: Pasoh Forest Res., Negri S., 1 1
August 1979, forest gap, P. and M. Becker (Gainesville). Paratype
male, same data as type, except 29 February 1979.

Discussion. The face of malaysiana most closely resembles that of
meaculpa Krombein due to the bulging eyes and strongly converg-
ing lower face. However, it differs significantly from meaculpa and
aridula Krombein based on characteristics of the female, including
the head only slightly wider than long, the scape, pedicel, F-I-II, and
the legs entirely yellow, and in both sexes the thoracic dorsum with
punctures relatively shallow and well-separated, the pronotum and
scutum subequal in length, and the metanotum and propodeum
without rugulae between the major ribbing.

Kryptosega Kimsey, new genus

Diagnosis. Malar space with vertical sulcus; occipital carina
well-developed, at least dorsally; scapal basin shallow, with some
cross-ridging; male flagellum elongate and cylindrical; pronotum
with shallow, occasionally faint, posteromedial groove and pit
before lateral lobe, 0.8-0. 9X scutal length (fig. 11); mesopleuron
with scrobal sulcus indicated by broad dorsally carinate groove or
nearly absent, without oblique mesopleural carina; scutum with
notauli deep posteriorly and obsolescent anteriorly, parapsides
present; metanotum with poorly defined, punctate medial area,
0.8X as long as scutellum; propodeum rounded laterally and poste-
riorly, with relatively long dorsal surface; hindcoxa without dorso-
basal carina; tarsal claw with large medial tooth; male fully winged,
forewing (fig. 19), stigma broad and elongate, without indication of
Rl, RS extended by evenly curved dark streak, medial vein arising
before cu-a; terga sharp-edged laterally and finely punctate.

Female unknown.

Type: Kryptosega anomala Kimsey.

Etymology: Krypto — hidden, sega — taken from Amisega Cameron.

1986]

Kimsey — New Amiseginae from Asia

161

18. Ma.
cuneifacialis

Figs. 10, 12, 15, 17. Front view of face. Figs. 13, 16. Lateral view of head.
Figs. 11,14. Dorsal view of thorax. Figs. 18, 19. Forewing.

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Discussion. This genus does not appear to be closely related to
any of the other Amiseginae. Kryptosega lacks most of the derived
characteristics found in other genera, except that it has no hind-
coxal carina and no indication of a lateral propodeal angle. Further
study is necessary to determine the relationships of Kryptosega.

Kryptosega anomala Kimsey, new species
(Figs. 10, 11, 13, 19)

Holotype male. Body length 3 mm. Face (figs. 10, 13); scapal
basin flattened and impunctate; frons with punctures 0.5- 1.0 PD
apart; malar space 4.4 MOD; head about as long as wide; midocel-
lus 3.6 MOD from ocular margin; ocelli arranged in broad triangle;
hindocellus separated from ocular margin by 0.3 diameter; clypeus
long and rounded apically; subantennal distance 1.8 MOD; F-I
length 4.3X breadth; F-II 3.6X as long as broad; pronotal, scutal
and scutellar punctures 0.2-0. 5 PD apart; pronotum and scutum
subequal in length; scutum with notauli absent anteriorly; meso-
pleuron with broad irregularly margined depression extending from
near pronotal lobe to scrobe, punctures 0.2- 1.0 PD apart; metano-
tum 0.8X as long as scutellum, medial enclosure with large nearly
contiguous punctures; propodeal dorsal surface with irregular longi-
tudinal carinae, posterior surface with extensive transverse carinae;
terga with sparse small punctures 2-3 PD apart. Head and thorax
black without metallic tints; abdomen black, except reddish brown
basally; scape and pedicel brown; flagellum black; coxae whitish,
legs otherwise pale brown becoming darker on tarsi; mandibles yel-
lowish becoming red apically.

Holotype male - NEW GUINEA: Mt. Kainde, 13 February- 12
March 1979, 2300 m, J. Sedlacek (Gainesville). Paratypes: 5 males,
same data as type except also collected 18 January- 14 February
1979 and 19 December 1978-8 January 1979.

Discussion. K. anomala can be distinguished from kaindeana
by the coarser punctation, longer malar space and subantennal dis-
tance, and nonmetallic color.

Kryptosega kaindeana Kimsey, new species
(Fig. 12)

Holotype male. Body length 2.5 mm. Face (fig. 12); scapal basin
impunctate and highly polished; frons with punctures shallow,

1986]

Kimsey — New Amiseginae from Asia

163

0.5- 1 .0 PD apart; malar space 2.3 MOD; midocellus 2.6 MOD from
ocular margin; ocelli arranged in nearly equilateral triangle; hindo-
cellus 1 diameter from ocular margin; head 1.2X as wide as long;
clypeus short, broadly rounded; subantennal distance 0.9 MOD; F-I
length 4.3X breadth; F-II 3.5X as long as broad; pronotal, scutal
and scutellar punctures shallow and 0.5- 1.0 PD apart; pronotum
0.8X scutal length; scutum with notauli complete; mesopleuron with
short depression near pronotal lobe, broadly separated from scrobe,
punctures large, 0.5-1 PD apart; metanotum 0.9X as long as scutel-
lum, medial enclosure punctures shallow and nearly contiguous;
propodeum strongly bulging posteromedially, dorsal surface irregu-
larly rugose, posterior surface smooth; terga with sparse tiny punc-
tures 2-4 PD apart. Head and thorax black with bronze tints
dorsally; abdomen black, except reddish brown basally and faint
bluish tints dorsally; antenna dark brown; mandibles whitish
basally, red apically; coxae whitish, legs otherwise pale brown
except hindtibial apex and tarsomeres dark brown.

Holotype male - NEW GUINEA: Mt. Kainde, 19 December
1978-18 January 1979, 2300 m, J. Sedlacek (Gainesville). One par-
atype male same data as type, except 18 January-14 February 1979.

Discussion. Unlike anomala, kaindeana is bronze colored dor-
sally. In addition, the integument appears glossy and is less coarsely
punctate and the face, particularly the malar space and subantennal
distance, are shorter.

Magdalium Kimsey, new genus

Diagnosis. Malar space with vertical sulcus; vertex with im-
punctate medial stripe extending from midocellus to occiput; occipi-
tal carina absent; scapal basin shallow, coarsely cross-ridged; male
flagellum broad, F-V-XI bulging medially (fig. 15); female flagellum
short and broad, flattened on one side; pronotum long and flat, with
oblong pits posteromedially and on lateral lobe, 0.6X combined
lengths of scutum, scutellum and metanotum (fig. 14); mesopleuron
with short oblique mesopleural carina, and scrobal sulcus long and
parallel-sided; scutum with notauli deep and straight, parapsides
faint; metanotum as long as scutellum, medial enclosure differently
sculptured than laterally; propodeum with long dorsal surface and
abruptly declivous posterior, lateral angles short and blunt; hind-
coxa with dorsobasal carina; tarsal claw with large perpendicular,

164

Psyche

[Vol. 93

medial tooth; male forewing (fig. 18), stigma long and slender, R1
not indicated, RS extended by evenly curved streak, medial vein
arising at cu-a; terga sharp-edged laterally, densely punctate except
T-I-II with impunctate medial welt.

Type: Magdalium cuneifacialis Kimsey.

Etymology: Magdalium = cylindrical figure (Latin, neuter)

Discussion. These are relatively large amisegines, which most
closely resemble Isegama, based on having a scrobal sulcus and
oblique mesopleural carina, short broad male flagellomeres and the
forewing medial vein arising at cu-a. However, Magdalium can be
distinguished by the odd lobular male flagellomeres, long flattened
pronotum, the absence of a well-defined occipital carina and the
long compressed body shape. Also, Magdalium has an impunctate
stripe on the vertex extending between the midocellus and the occi-
put much as in Cladobethylus.

Magdalium cuneifacialis Kimsey, new species
(Figs. 14-16, 18)

Holotype male. Body length 5 mm. Face (figs. 15, 16); scapal
basin with polished medial stripe and coarse cross-ridges laterally,
punctures about 1 PD apart; malar space 4 MOD long, with vertical
sulcus; head as wide as long; occipital carina present dorsally; mido-
cellus 2.5 MOD from ocular margin; ocelli arranged in broad tri-
angle; vertex with impunctate medial stripe from midocellus to
occiput; hindocellus 1 diameter from ocular margin; pronotum long
and flat, 0.6X combined lengths of scutum, scutellum and metano-
tum along midline, with large pit posteromedially and on lateral
lobe; mesopleuron with subalar fossa, short oblique mesopleural
carina and scrobal sulcus long and parallel-sided; notal punctures
0.2-0. 5 PD, larger on head and pronotum than scutum; scutum with
notauli deep and straight; metanotum 0.9X as long as scutellum;
propodeum with short blunt lateral angles; T-I-II punctures dense
and nearly contiguous, except impunctate medial welt; T-III-V with
posterior band of punctures. Head, thorax and abdomen black;
scape, pedicel and F-I-IV red; F-V-XI dark brown; forefemur dark
brown, reddish apically, midleg, foretibia and tarsi red, hindleg all
dark brown; entire body with long erect reddish setae.

Female. Same as male, except clypeus shorter; F-I 1.9-2.0X as
long as broad; F-II 0.7X as long as broad; scape, pedicel and basal
half of F-I red; rest of flagellum dark brown.

1986]

Kimsey — New Amiseginae from Asia

165

Holotype male: MALAYSIA: Pasoh Forest Res., Negri S., 17
April 1980, P. and M. Becker (Gainesville). Paratypes: 9 males, 4
females, same data as type, except differing dates, from 19 August
1978 to 29 May 1980.

Summary

Three new species of Cladobethylus, 1 new Atoposega, 1 new
Isegama and 3 new genera, Magdalium ( cuneifacialis ), Kryptosega
( anomala and kaindeana ) and Bupon {pasohanus ), are described.
Most of this material was collected in the Pasoh Forest Reserve in
Malaysia. The others, including Cladobethylus aquilus and both
species of Kryptosega are from New Guinea, and C. japonicus is
from Japan.

Acknowledgments

This study was made possible by Henry Townes, and numerous
fruitful discussions of Amiseginae with Karl V. Krombein, and was
supported by NSF Research Grant No. BSR-8407392.

CAMBRIDGE ENTOMOLOGICAL CLUB

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of each month October through May at 7:30 p.m. in Room 154,
Biological Laboratories, Divinity Avenue, Cambridge. Entomolo-
gists visiting the vicinity are cordially invited to attend.

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16 Divinity Avenue
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ISSN 0033 2615

PSYCHE

A JOURNAL OF ENTOMOLOGY

founded in 1874 by the Cambridge Entomological Club

Vol. 93 1986 Nos. 3-4

CONTENTS

The choice of web-monitoring sites by a green Miagrammopes species (Ara-

neae: Uloboridae). Brent D. Opell 167

Natal nest distribution and pleometrosis in the desert leaf-cutter ant, Acro-
myrmex versicolor (Pergande) (Hymenoptera: Formicidae). Steven W.

Rissing, Robert A. Johnson, and Gregory B. Pollock 177

Revision of the Onocosmoecus unicolor group (Trichoptera: Limnephilidae,

Dicosmoecinae). Glenn B. Wiggins and John S. Richardson 187

Population fluidity in Leptothorax longispinosus (Hymenoptera: Formici-
dae). Joan M. Herbers and Carol W. Tucker 217

Geographic variation in the cave beetle, Neaphaenops tellkampfi (Coleoptera:

Carabidae). Thomas C. Kane and George D. Brunner 231

Biosystematic revision of Epimyrma kraussei, E. vandeli, and E. foreli
(Hymenoptera: Formicidae). Alfred Buschinger, Karl Fischer, Hans-

Peter Guthy, Karla Jessen, and Ursula Winter 253

Male biology in the queenless ponerine ant, Ophthalmopone berthoudi

(Hymenoptera: Formicidae). Christian Peeters and Robin Crewe 277

Nearctic species of the new wolf-spider genus, Gladicosa (Araneae: Lycosi-

dae). Allen R. Brady 285

Nesting associations of wasps and ants on lowland Peruvian ant-plants.

Edward Allen Heere, Donald M. Windsor, and Robin B. Forster 321

Winter prey collection at a perennial colony of Paravespula vulgaris (L.)

(Hymenoptera: Vespidae). Parker Gambino 331

Young larvae of Eciton (Hymenoptera: Formicidae: Dorylinae). George C.

Wheeler and Jeanette Wheeler 341

Spatial distribution of castes within colonies of the termite, Incisitermes

schwarzi. Peter Luykx, Jack Michel, and Jeanette K. Luykx 351

A new species of Orthea, a neotropical myodochine^genus j*vith an unusual-

Blattella asahinai introduced into Florida (Blattaria: Blattellidae). Louis M.

Roth 371

Substitute names for the extinct genera Cycloptera Martynov.^ jft4fecojpt^aj|00 7

and Parelcana Carpenter (Orthoptera). Frank M. Carperiief 375

Index to Volume 93 377

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1986-1987

President James M. Carpenter

Vice-President Edward Armstrong

Secretary David Maddison

Treasurer Frank M. Carpenter

Executive Committee Heather Hermann

W. David Winter

EDITORIAL BOARD OF PSYCHE

F. M. Carpenter, (Editor), Fisher Professor of Natural History,
Emeritus, Harvard University

W. L. Brown, Jr., Professor of Entomology, Cornell University and
Associate in Entomology, Museum of Comparative Zoology
B. K. HOLLDOBLER, Alexander Agassiz Professor of Zoology, Harvard
University

H. W. Levi, Alexander Agassiz Professor of Zoology, Harvard University
M. D. BOWERS, Assistant Professor of Biology, Harvard University
E. O. WILSON, Baird Professor of Science, Harvard University
J. M. CARPENTER, Assistant Professor of Biology, Harvard University

Checks and remittances should be addressed to Treasurer, Cambridge
Entomological Club, 16 Divinity Avenue, Cambridge, Mass. 02138.

IMPORTANT NOTICE TO CONTRIBUTORS
Manuscripts intended for publication should be addressed to Professor F. M.
Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. 02138.

Psyche, vol. 93, no. 1-2, for 1986, was mailed October 15, 1986

The Lexington Press, Inc., Lexington, Massachusetts

PSYCHE

Vol. 93

1986

Nos. 3-4

THE CHOICE OF WEB-MONITORING SITES BY A GREEN
MIAG RAM MOPES SPECIES (ARANEAE: ULOBORIDAE)*

By Brent D. Opell
Department of Biology

Virginia Polytechnic Institute and State University
Blacksburg, Virginia 24061

Introduction

The varied and effective predatory strategies of spiders have
drawn more attention than have their antipredator adaptations to
threats from birds, wasps, damselflies, and other spiders (Bristowe
1941, Dorris 1970, Blanke 1972, Jackson & Blest 1982a, b). As the
majority of their predators are visual hunters, it is not surprising
that many spiders employ either protective resemblance or eucrypsis
(as defined by Robinson 1969a) to escape detection. In general,
protective resemblance seems to be more common among spiders
that use a capture web and eucrypsis among hunting spiders that
employ sit-and-wait tactics. Protective resemblance usually involves
anatomical modification and is frequently enhanced by postural
specializations (Robinson 1969b). Both of these antipredator adap-
tations are often enhanced by a spider’s color.

In order for protective resemblance and eucrypsis to be effective,
animals employing these strategies should select a background
whose color or texture closely matches their own. Such background
selection has been demonstrated for adult moths (Kettlewell 1955,
Sargent 1966, 1984; Sargent & Keiper 1969, Malcom & Hanks
1973), butterfly pupae (West & Hazel 1979), and grasshoppers
(Giles 1982). Several studies have demonstrated the protective

* Manuscript received by the editor May 16, 1986

167

168

Psyche

[Vol. 93

benefits of correct background selection in both immature and adult
insects (Hazel & West 1979, Erichsen et al. 1980, West & Hazel
1982, Sims & Shapiro 1983a, b). This study investigates background
selection by spiders of the genus Miagrammopes.

Members of the tropical genus Miagrammopes spin reduced cap-
ture webs consisting of a horizontal thread which may either be
sticky or non-sticky and may have one or several vertical or diago-
nal sticky (cribellar) capture threads extending from it (Lubin et al.
1978, Opell 1984). These spiders monitor their reduced webs from
one of the attachment points, where their postures and body form
make them cryptic. Brown species that spin their webs among twigs
and vines resemble thorns or broken twig bases, whereas green spe-
cies that spin webs on moss-covered vegetation resemble extending
moss phyllidia (Fig. 1). After subduing and wrapping prey, these
spiders return to one of their web’s attachment points and resume
their typical cryptic posture while feeding.

Most individuals of the green Miagrammopes species I observed
in Costa Rica monitored their webs from moss-covered twigs. This
occurred despite the fact that some webs were also anchored to bare
twigs. In order to test the hypothesis that members of this species
select moss-covered twigs over bare twigs as web-monitoring sites, I
conducted a series of choice experiments.

Materials and Methods

The species used in this study is an undescribed member of the
Miagrammopes aspinatus species group (Opell 1984). Voucher spec-
imens are deposited in Harvard University’s Museum of Compara-
tive Zoology. Spiders used in this study were collected from stands
of abandoned cacao (Theobroma cacao) at the Organization for
Tropical Studies’ La Selva research station located near the town of
Puerto Viejo de Sarapiqui, Heredia Province, Costa Rica. Prior to
their release onto experimental frames, these spiders were kept for
two to four days in small, cotton stoppered glass vials. During
this time, their carapace and leg lengths were measured with a
micrometer-equipped dissecting microscope. Because adult males
do not spin capture webs, only immatures and adult females were
used in this study.

Experimental frames (Fig. 2) were constructed of 2 mm diameter
hardwood applicator sticks glued together with epoxy and bound

1986]

Opell — Web-monitoring by Miagrammopes

169

Figure 1 . Adult female Miagrammopes sp. feeding on a small beetle held with the
pedipalps while monitoring an attachment line of the capture web. Setal tufts at the
distal end of the extended first legs make them resemble the moss to which the web is
attached. Scale bar represents 2 mm.

by thread. To one set of opposite vertical elements were wired moss-
covered cacao twigs and to the other set, bare cacao twigs. All twigs
were taken from the same tree and had a diameter of about 7 mm.
Two of the four frames employed bare twigs that had no evidence of
moss cover and two twigs whose moss covering was removed with-
out damage to the bark. Frames hung 98 cm apart along a taut,
north-south suspension line. To account for the possible influence
of air currents, frames were oriented so that the moss-covered twigs
occupied alternate sectors (East and West sectors, North and South
sectors, etc.).

From 26 June until 6 July 1985, these frames hung in the aban-
doned cacao plantation from which specimens were collected. From
7-15 July 1985, these study frames were transferred to a roofed
enclosure (cabina) whose screened north, east, and south walls were
covered with light colored curtains to exclude direct sunlight. Here,
frame orientation and spacing were identical to that described
above. In this enclosure, spiders were exposed only to natural light.
Each frame’s bare and moss-covered twigs were watered daily at
about 8:30 and 13:00. At the end of the study, moss on the twigs
showed no signs of thinning or turning brown and bare twigs
showed no signs of moss growth.

170

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[Vol. 93

Figure 2. Diagram of the frame used in this study, showing a spider monitoring
its web from the front, moss-covered sector.

Frames and suspension lines were cleaned of all visible silk
strands before spiders were released at 16:00 onto the top center of
each frame. Frames were checked the following morning at 8:00 and
the presence of webs and position of spiders recorded. A capture
web was defined as a web with sticky (cribellar) prey capture silk. In
contrast with the non-sticky, single-line resting web, the capture web
usually consisted of multiple, diverging threads. Spiders always
hung near one of the web’s attachment points (Fig. 1) and it was
noted whether this was a moss-covered or bare site. During the first
three days and last day of the study only a single specimen was re-
leased onto each frame. On other days, one large and one small
specimen were released on each frame. This was done to compensate

1986] Opell — Web-monitoring by Miagrammopes 171

for what initially promised to be a high percentage of spiders leaving
the frames. The size difference made it possible to distinguish indi-
viduals on each frame and to record which had made a web. After
web production and position were recorded specimens were col-
lected and released in the forest. A plot of first femur length against
carapace length for the specimens used in this study plus the values
of 61 additional specimens was used to assign the instar values to
spiders. Chi-square tests were used to evaluate the results of this
study.

Results

Earlier instars were more commonly found than later instars and,
therefore, are represented by a larger sample size. Of the 21 capture
webs constructed in the forest, nine were spun by third instars, six
by fourth instars, four by fifth instars, one by an adult female, and
one by a specimen of uncertain age. Of the 28 capture webs spun in
the enclosure, four were spun by third instars, 14 by fourth instars,
six by fifth instars, and four by adult females. Significantly more
(0.05 > p > 0.025) spiders produced capture webs in the enclosure
than in the forest (Table I). All capture webs made within frames
were monitored from a moss-covered twig. In the enclosure, six
specimens constructed their capture webs outside the frame and
monitored them from wires used to attach frames to the support
line. In most of these cases, the spider’s web was not anchored to a
moss-covered twig and the highest attachment point was the favored
monitoring site. When these six capture webs are compared with the
other 22 indoor webs monitored from moss, moss is still the favored
site (0.025 >p>0.01).

In neither habitat was there a significant difference in the moss-
covered frame sectors from which webs were monitored (Table I).
However, these results were more clear-cut in the enclosure (0.975 >
p > 0.90) than in the forest (0.50 > p > 0.10). In the latter setting,
moss-covered twigs on East and South frame sectors appear to be
favored. This may be explained by stronger and/or more unidirec-
tional wind currents in the latter setting. On seven occasions I used a
web dusting device to expel a cloud of corn starch into the air of the
forest site. On three instances the wind was blowing to the east, on
three to the southeast, and one to the southwest. These observations
suggest that a spider’s dragline had a greater chance of being carried

1986] Opell — Web-monitoring by Miagrammopes 173

to a frame’s East and South sectors and that, when moss-covered,
the twig attached to this sector would be favored over the opposite
moss-covered twig as a web-monitoring site.

Discussion

This species’ preference for moss as a web-monitoring site en-
hances its protective resemblance. The exposed first legs of these
green species have a tuft of green tibial setae that look like the small
phyllidia of an extending moss plant (Fig. 1). Although these setal
tufts are found in brown Miagrammopes species, they are more
prominent in green species, where they become disproportionately
larger in subadult and adult individuals. During the night Miagram-
mopes often abandon their typical day-time position adjacent to a
twig and hang on the monitoring line a centimeter or more from its
attachment point to a twig.

Choice of a web-monitoring site by Miagrammopes is facilitated
by the fact that most of their simple, irregular webs have no single
attachment point from which they must be monitored. A few webs
have a particular thread that probably serves as an optimal monitor-
ing line by virtue of its single attachment point to several diverging
lines. However, most newly constructed Miagrammopes webs con-
sist of an approximately horizontal thread with one or several inde-
pendently diverging vertical or diagonal threads. Either end of this
horizontal thread could serve as a monitoring site. I have seen dis-
turbed Miagrammopes run to the opposite end of their horizontal
threads and begin monitoring their webs from this new position.

The greater number of missing individuals noted in the forest
than in the enclosure experiments probably resulted from a higher
rate of spiders ballooning from the forest frames. Three factors
suggest that this difference is not due to predators removing spiders
that chose bare twigs as web-monitoring sites. First, release and
observation times were chosen so that most of site selection and web
construction took place at night when threats from visually hunting
predators were lowest. Second, during this study, no predatory
insects were seen on the experimental frames or their supporting
lines. Third, none of the spiders in the predator-free enclosure chose
bare twigs as web-monitoring sites.

This study does not address the mechanism by which individuals
select moss-covered twigs. Experimental studies on cryptic insects

174

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[Vol. 93

show that two methods may be used in selecting a matching back-
ground. Gillis (1982) showed that in one grasshopper species indi-
viduals select backgrounds whose color matches that of their
circumocular regions. By contrast, Sargent (1968) found that back-
ground selection in some moths was hereditary and was unaffected
by painting their circumocular scales. Color vision has not been
demonstrated in Miagrammopes. However, the eyes of the species
used in this study are well developed and have low f-numbers, indi-
cating that they are effective in low light intensities (Opell and Cush-
ing, in press). Tactile or moisture properties of the moss may also be
important cues for its choice as a web-monitoring site. Unlike
striped moths that must assume the proper orientation in order to
take full advantage of their cryptic markings (Sargent 1969), the
webs of Miagrammopes assure that they will assume the proper
attitude after they have selected the correct background.

Summary

Members of the spider genus Miagrammopes construct simple
capture webs consisting of only a few threads and assume a stick-
like posture as they actively monitor these webs. A green Costa
Rican species showed a statistically significant preference for moss-
covered twigs as web-monitoring sites. This choice was observed in
both a forest setting and a screened enclosure, and occurred on
experimental frames which required spiders to attach their webs to
both bare and moss-covered twigs.

Acknowledgements

I am grateful to the Organization for Tropical Studies for permit-
ting me to use its facilities and to David and Deborah Clark, co-
directors of the organization’s La Selva research station, for their
help. This study was supported by National Science Foundation
grant BSR-8407979 to the author.

References

Blanke, R.

1972. Untersuchungen zur Okophysiologie und Okethologie von Cyrto-
phora citricola Forskal (Araneae, Araneidae) in Andalusien. Forma et
Functio, 5: 125-206.

1986] Opell — Web-monitoring by Miagrammopes 175

Bristowe, W. S.

1941. The Comity of Spiders. Ray Soc., No. 128. London.

Dorris, P. F.

1970. Spiders collected from mud-dauber nests in in Mississippi. J. Kansas
Ent. Soc., 43: 10-11.

Erichsen, J. T., J. R. Krebs, and A. I. Houston.

1980. Optimal foraging and cryptic prey. J. Anim. Ecol., 49: 271-276.

Gillis, J. E.

1982. Substrate color-matching cues in the cryptic grasshopper Circotettix
rabula rabula. Anim. Behav., 30: 113-116.

Hazel, W. N. and D. A. West.

1979. Environmental control of pupal colour in swallow-tail butterflies (Lepi-
doptera: Papilioninae): Battus philenor (L) and Papilio polyxenes Fabr.
Ecol. Entomol., 4: 393-400.

Jackson, R. R. and A. D. Blest.

1982a. The biology of Portia fimbriata, a web-building jumping spider (Ara-
neae, Salticidae) from Queensland: utilisation of webs and predatory
versatility. J. Zool. (London), 196: 255-293.

1982b. The distances at which a primitive jumping spider, Portia fimbriata,
makes visual discriminations. J. Exp. Biol., 87: 441-445.

Kettlewell, H. B. D.

1955. Recognition of appropriate background by the pale and black phases of
Lepidoptera. Nature, 175: 943-944.

Lubin, Y. D., W. G. Eberhard, and G. G. Montgomery.

1978. Webs of Miagrammopes (Araneae: Uloboridae) in the Neotropics.
Psyche, 85: 1-23.

Malcom, W. M. and J. P. Hanks.

1973. Landing-site selection searching behavior in the micro-lepidopteran
Agonopteryx pulvipennella. Anim. Behav., 21: 45-48.

Opell, B. D.

1984. Phylogenetic review of the genus Miagrammopes sensu lato (Araneae:
Uloboridae). J. Arachnol., 12: 229-240.

Opell, B. D. and P. E. Cushing.

in press. Visual fields of orb-web and single-line-web spiders of the family
Uloboridae. Zoomorphology.

Robinson, M. H.

1969a. Defense against visually hunting predators. Evol. Biol., 3: 225-259.

1969b. The defensive behaviour of some orthopteroid insects from Panama.
Trans. R. Soc. London, 121: 281-303.

Sargent, T. D.

1966. Background selections of geometrid and noctuid moths. Science, 154:
1674-1675.

1968. Cryptic moths: effects on background selection of painting circumocular
scales. Science, 159: 100-101.

1969. Behavioural adaptations of cryptic moths. III. Resting attitudes of two
bark-like species, Melanolophia canadaria and Catocala uttronia. Anim.
Behav., 17: 670-672.

176

Psyche

[Vol. 93

1984. Melanism in Phigalia titea (Lepidoptera: Geometridae) in southern New
England: A response to forest disturbance? J. N. Y. Entomol. Soc., 93:
1113-1120.

Sargent, T. D. and R. R. Keiper.

1969. Behavioral adaptations of cryptic moths, I. Preliminary studies on
bark-like species. J. Lepid. Soc., 23: 1-9.

Sims, S. R. and A. M. Shapiro.

1983a. Pupal color dimorphism in California Battus philenor (Papilionidae):
Morphology factors and selective advantage. J. Lepid. Soc., 37:
236-243.

1983b. Pupal dimorphism in California Battus philenor (L): Pupation sites,
environmental control, and diapause linkage. Ecol. Entomol., 8: 95-104.

West, D. A. and W. N. Hazel.

1979. Natural pupation sites of swallowtail butterflies (Lepidoptera: Papilio-
ninae): Papilio polyxenes Fabr., P. glaucus L. and Battus philenor (L.).
Ecol. Entomol., 4: 387-392.

1982. An experimental test of natural selection for pupation site in swallowtail
butterflies. Evolution, 36: 152-159.

NATAL NEST DISTRIBUTION AND PLEOMETROSIS
IN THE DESERT LEAF-CUTTER ANT
ACROMYRMEX VERSICOLOR (PERGANDE)
(HYMENOPTERA: FORMICIDAE)

By Steven W. Rissing,* Robert A. Johnson,*
and Gregory B. Pollock**

While most ant colonies are started by single queens, colony
foundation by groups of queens, pleometrosis, also occurs (Wilson
1971, Holldobler and Wilson 1977). Several extensively studied,
highly pleometrotic species are notably similar with respect to
important aspects of colony ontogeny and population dynamics.
Myrmecocystus mimicus, Solenopsis invicta and Veromessor per-
gandei queens found colonies mutualistically without respect to
relatedness (Bartz and Holldobler 1982, Tschinkel and Howard
1983, Pollock and Rissing 1985, Rissing and Pollock 1986). Further,
while adult colonies of these species are highly territorial (Holl-
dobler 1976a, 1981; Wilson et al. 1971; Went et al. 1972, Wheeler
and Rissing 1975), natal colonies are clumped with brood raiding
and subsequent worker defection from brood-raided colonies occur-
ring (references cited above for M. mimicus and S. invicta, for V.
pergandei: Rissing and Pollock, in press). Given such frequently
deleterious natal colony interactions, adaptive value of habitat
selection by founding queens resulting in clumping of natal nests is
unclear. Natal nests of M. mimicus are generally clumped in areas
devoid of adult nests (Bartz and Holldobler 1982), yet still occur
near such nests (B. Holldobler, pers. comm.), and queens of S.
invicta show some preference for microtopographic features
(Tschinkel and Howard 1983). Here we present data relating habitat
selection and clumping of natal nests of the highly pleometrotic leaf-
cutter ant Acromyrmex versicolor (Pergande) directly to survival of
founding queens. The only other report regarding any aspect of
colony initiation in this species is a description of mating flights
following summer rains in the Sonoran Desert by Wheeler (1917).

♦Department of Zoology, Arizona State University, Tempe, AZ 85287
♦♦School of Social Science; University of California, Irvine; Irvine, CA 92717
Manuscript received by the editor April 6, 1986.

177

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A major flight of A. versicolor occurred on 19 September 1985 on
a study area in North Scottsdale, AZ 5.6 km north of Maricopa
County along Pima Rd, approximately 3.2 km west of the McDow-
ell Mountains. The habitat in this area is typical of the Sonoran
Desert with Larrea tridentata and Franseria dumosa dominant
shrubs and Olneya tesota and Cercidium microphyllum dominant
trees along numerous shallow washes in the gravel/ sandy soil. A
major storm front produced rain throughout the region the previous
day; 2.6 cm of rain was recorded at the Arizona State Laboratory of
Climatology located 32 km south of the study area.

Habitat choice by A. versicolor queens was examined by running
a transect 20 m long and 2 m wide from the base of 10 haphazardly
chosen trees on the study area. Transect direction was chosen
haphazardly. Distance of each starting nest from base of tree was
recorded and standardized into “canopy units” by dividing by dis-
tance from base of tree to outer canopy edge along each transect.
This standardization was necessitated by variance in tree size and
canopy extent. Distance to nearest neighboring tree was measured
for 20 haphazardly chosen trees and converted to canopy units using
the larger canopy extent of each pair. Number of queens per starting
nest was determined by excavating 43 nests during this time. Addi-
tionally, 21 starting nests were excavated on 22 September on a
study area of similar habitat in South Mountain Park, Phoenix,
AZ, 38.5 km southwest of the main study area.

Effect of temperature in a starting nest on queen survivorship was
determined by placing 18 queens (from the above excavated nests)
in a large test tube plugged with cotton and containing a large ball
of cotton saturated with water to prevent desiccation. Test tubes
were placed in a darkened incubator at 20, 25,. . .45°C in random
predetermined order; subsequently an additional tube was exposed
to 42.5° C. To mimic late afternoon temperature exposures in the
field, tubes remained at their given temperature for 2 hours. Queens
incapable of righting themselves after 2 hours were considered dead.
Likely temperatures in starting nests were determined by taking soil
temperatures 5 and 10 cm below surface at the trunk base (0 canopy
units), canopy edge (1 canopy unit) and in the open (> 1 canopy
unit) on the main study area between 16:00 and 17:00 h on 24

1986] Rissing, Johnson, & Pollock — Acromyrmex 179

September and 4 October 1985. All queens excavated 1-2 days fol-
lowing the mating flight were found 5-10 cm below the soil surface.

The possible importance of relatedness in formation of queen
associations was tested according to the methods of Rissing and
Pollock (1986). Eight plastic “choice boxes” (30 X 15 X 8 cm, half
filled with sand moistened in each corner and at the midpoints along
the long sides) were established with 2 sets of 4 queens, one set
collected from each of the two study sites (38.5 km apart). In 5 boxes,
queens were color marked according to collection locale; different
patterns of the same two colors were used to avoid providing cues
for recognition. As an additional control for possible paint odor,
queens in the remaining 3 boxes were not marked. Boxes were
excavated 24 hrs later when queens had dug below the surface;
location of each queen was noted relative to the others.

Results

Acromyrmex versicolor queens strongly prefer to start nests
immediately below the outer canopy of trees; while mean inter-tree
distance was 6.10 ± 1.85 canopy units (= 21.37 ± 5.14 m between
tree bases; N = 20), the average starting nest was .87 ± .53 canopy
units (= 3.15 + 1.88 m; N = 1 15) from a tree base (Fig. 1). Although
measurements were not taken, the same distribution of starting
nests was observed at South Mountain Park. All queens examined
for temperature tolerance survived exposure to temperature up to
and including 40° C for at least 2 hr; survivorship was 0, however, at
42.5° C and above. Temperatures above 40° C were found in the
open between trees (>1 canopy units) at soil depths normally occu-
pied by newly starting colonies (X5cm = 42.0 (± 1.1)° C, N = 5;
Xjocm = 36.9 (± 1.2)° C, N = 10); temperatures this high were not
found at canopy edges (= 1 canopy unit) (X5cm = 38.2 (+ 5.5)° C, N
= 5; Xjocm — 32.8 (+ 3.7)° C, N = 9) or at tree bases (= 0 canopy
units) (x5cm = 28.91 ± 2.4), N = 2; x10cm = 27.4 (± 0.8)°C, N - 5).

Acromyrmex versicolor is highly pleometrotic; 82.5% of all
queens excavated (N = 160 queens from 64 nests) were from
pleometrotic associations (Table 1). Relatedness appears unimpor-
tant in a queen’s decision to enter a foundress association; five of the
8 “choice boxes” resulted in a single starting nest occupied by all 8
queens. The remaining 3 boxes had two starting nests each: of these
6 starting nests, 4 contained queens from both collection locales, 1

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CANOPY UNITS

Figure 1. Habitat choice by Acromyrmex versicolor queens. “Canopy units”
represent distance of a starting nest from the base of tree divided by distance from
tree base to outer extent of canopy along that transect.

contained 3 queens from one site only, and the last contained a
solitary foundress.

Discussion

The distinct habitat choice of A. versicolor queens (Fig. 1) seems
a likely response to high soil temperatures in sunlit areas during the
mid-late summer flight season of this species. Queens initiating nests
in open areas will likely experience lethal temperatures even 5 cm
below the soil surface. We routinely found queens at this depth 2-3
days following the mating flight. These nests, however, were located
under trees where potentially lethal temperatures were never
recorded. Acromyrmex versicolor queen death after 2 hours of
exposure to temperatures 40-43° C is consistent with upper temper-
ature tolerances of foragers of this species of 42-43° C (Gamboa
1976).

Queens of several other ant species display varying degrees of
habitat choice. Queens of Lasius niger and Lasius flavus both prefer
bare, sunlit soil where they establish colonies more successfully than

1986] Rissing, Johnson, & Pollock — Acromyrmex 181

Table 1. Pleometrosis in Acromyrmex versicolor.

No. queens in nest

Collection frequency

in shaded soil (Pontin 1960). Queens of the tropical leaf-cutting ant
Atta cephalotes appear capable of choosing between major habitat
types (mature evergreen woodlands as opposed to deciduous forest
or cultivated fields) (Rockwood 1973). Similar ability to choose
between major habitat types (woods versus open fields) while flying
occurs in Lasius neoniger and Solenopsis molesta (Wilson and Hunt
1966).

Preference by queens for the canopy edge (as opposed to any-
where under a tree) may represent a trade-off for shade while still
being as warm as possible for rapid development of an initial worker
force and eventual establishment of a foraging territory. This would
be consistent with the high degree of pleometrosis in this species (see
below) and with the “maxi-therm” hypothesis of Hamilton (1973).
Location of a starting nest under a tree canopy (especially O. tesota
whose branches frequently droop to the ground) would permit easy
and safe access to vegetation for initiation and growth of the fungus
garden characteristic of all leaf-cutters. Unlike most species of higher
ants, queens of Acromyrmex spp., including A. versicolor, routinely
forage for vegetation, especially at colony initiation (Weber 1972,
Gamboa 1974). Trees, including O. tesota, are commonly harvested
by A. versicolor (Gamboa 1975).

Of the several hundred adult colonies of A. versicolor we have
observed within the vicinity of Phoenix, AZ, virtually all have been
located directly under adult trees and never in the open between
trees. Acromyrmex, Atta and related genera are a largely tropical,
New World group of ants; A. versicolor is the northernmost of 24
Acromyrmex species and certainly one of the most desert-adapted
of all the leaf-cutters (Creighton 1950, Weber 1972). Habitat choice
by founding queens and location of adult nests under trees may be

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an important behavioral adaptation permitting range extension into
the Sonoran Desert. Acromyrmex versicolor queens clump around
the essential resource of favorable nest sites (tree shade with ready
access to lower canopy leaves). The mating system of this species
may also permit “tracking” of this resource. Unlike some desert
species that have massive mating swarms (e.g. Pogonomyrmex spp.:
Chapman 1957; Nagel and Rettenmeyer 1973; Holldobler 1976b, c;
Davidson 1982), A. versicolor mates in small, localized groups at or
near the ground in open areas between trees (Wheeler 1917; R. A.
Johnson, pers. obs.). This behavior mimics closely the mating
behavior of M. mimicus (M. Cazier, pers. comm.) and likely V.
pergandei (Pollock and Rissing 1985), pleometrotic species with
clumped natal nests. Whether such localized mating aggregations
have led to a highly female biased sex ratio, as appears to have
occurred in V. pergandei (Pollock and Rissing 1985), is currently
unknown for these other pleometrotic species.

Some other ant species with clumped, natal nests engage in inter-
nest brood raiding in the process of establishing natal territories
(references cited above). This may select for pleometrosis (Rissing
and Pollock, in press) which generally results in more rapid produc-
tion of a larger initial worker force (Waloff 1957; Stumper 1962;
Markin et al. 1972; Taki 1976; Mintzer 1979; Bartz and Holldobler
1982, Tschinkel and Howard 1983, Rissing and Pollock, in press).
Colonies of these species are also territorial as adults. Brood raiding
also seems likely in Atta texana, another pleometrotic desert leaf-
cutter (Mintzer and Vinson 1985), which “merges” young colonies in
the laboratory and field (Echols 1966). Adult colonies of Acromyr-
mex versicolor are territorial (Gamboa 1974). We suggest such terri-
toriality, when coupled with natal nest clumping through habitat
choice, makes brood raiding and associated forms of natal nest
competition likely for this species as well.

A final similarity between A. versicolor and other pleometrotic
ants with clumped natal nests discussed here is the apparent forma-
tion of foundress associations without respect to relatedness.
Queens collected from distant locales readily associate. Similar
observations exist for M. mimicus (Bartz and Holldobler 1982), A.
texana (Mintzer and Vinson 1985) and V. pergandei (Rissing and
Pollock 1986); electrophoretic evidence indicates S. invicta queens
also associate randomly (Ross and Fletcher 1985). This differs

1986] Rissing, Johnson, & Pollock— Acromyrmex 183

dramatically from the close relatedness of cofoundresses in primi-
tively eusocial wasps (Pfennig et al. 1983 and included references).
Given the normally claustral method of colony foundation in ants,
relatedness to potential cofoundresses should be unimportant in this
essentially mutualistic process (Rissing and Pollock 1986). Colony
foundation in leaf-cutters (including A. versicolor ), however, is not
claustral; foundresses forage (references cited above). This presents
an opportunity to extend and examine the dynamics of the mutual-
istic process of colony foundation by unrelated females. Work in
this area is currently planned.

Summary

Queens of the desert leaf-cutter ant, Acromyrmex versicolor
exhibit distinct habitat choice during colony foundation; almost all
natal nests are located directly under the canopy edge of large trees.
Soil temperatures in these sites are conducive to queen survivorship
during the first several days of colony initiation while those in open
areas between trees are high enough to result in queen death. This
habitat choice results in clumping of many natal nests under indi-
vidual trees implying strong natal colony competition. Indeed, as
with several other ant species exhibiting such competition, starting
colonies are frequently pleometrotic; 82.5% of all queens excavated
were from such multiple foundress associations. As with other
pleometrotic ant species, mating aggregations of A. versicolor are
small and localized, and relatedness appears unimportant in a
queen’s decision to enter a foundress association.

Acknowledgments

G. E. Walsberg provided advice regarding queen temperature tol-
erance tests.

Literature Cited
Bartz, S. H. and B. Holldobler

1982. Colony founding in Myrmecocystus mimicus Wheeler (Hymenoptera:
Formicidae) and the evolution of foundress associations. Behav. Ecol.
Sociobiol. 10: 137-147.

Chapman, J. A.

1957. A further consideration of summit ant swarms. Canadian Entomol. 89:
389-395.

184

Psyche

[Vol. 93

Creighton, W. S.

1950. The ants of North America. Bull. Mus. Comp. Zool. 104: 1-585.

Davidson, D. W.

1982. Sexual selection in harvester ants (Hymenoptera: Formicidae: Pogono-
myrmex). Behav. Ecol. Sociobiol. 10: 245-250.

Echols, H. P.

1966. Compatibility of separate nests of Texas leaf-cutting ants. J. Econ. Ent.
59: 1299-1300.

Gamboa, G. J.

1974. Surface behavior of the leaf-cutter ant Acromyrmex versicolor versicol-
or Pergande (Hymenoptera: Formicidae). Masters Thesis; Arizona St.
Univ., Tempe, Arizona.

1975. Foraging and leaf-cutting of the desert gardening ant Acromyrmex versi-
color versicolor (Pergande) (Hymenoptera: Formicidae). Oecologia
(Berl.) 20: 103-110.

1976. Effects of temperature on the surface activity of the desert leaf-cutter
ant, Acromyrmex versicolor versicolor (Pergande) (Hymenoptera: For-
micidae). Amer. Midi. Natur. 95: 485-491.

Hamilton, W. J.

1973. Life’s Color Code. McGraw-Hill, New York, N.Y.

HOlldobler, B.

1976a. Tournaments and slavery in a desert ant. Science 192: 912-914.

1976b. The behavioral ecology of mating in harvester ants (Hymenoptera: For-
micidae: Pogonomyrmex). Behav. Ecol. Sociobiol. 1: 405-423.

1976c. Recruitment behavior, home range orientation and territoriality in har-
vester ants, Pogonomyrmex. Behav. Ecol. Sociobiol. 1: 3-44.

1981. Foraging and spatiotemporal territories in the honey ant Mrymecocys-
tus mimicus Wheeler (Hymenoptera: Formicidae). Behav. Ecol. Socio-
biol. 9: 301-314.

HOlldobler, B. and E. O. Wilson.

1977. The number of queens: An important trait in ant evolution. Naturwis-
senschaften 64: 8-15.

Markin, G. P., H. L. Collins, and J. H. Dillier.

1972. Colony founding by queens of the red imported fire ant, Solenopsis
invicta. Ann. Entomol. Soc. Am. 65: 123-124.

Mintzer, A.

1979. Colony founding and pleometrosis in Camponotus (Hymenoptera:
Formicidae). Pan-Pacific Entomol. 55: 81-89.

Mintzer, A. and S. B. Vinson.

1985. Cooperative colony foundation by females of the leafcutting ant Atta
texana in the laboratory. J. New York Entomol. Soc. 93: 1047-1051.

Nagel, H. G. and C. W. Rettenmeyer.

1973. Nuptial flights, reproductive behavior and colony founding of the west-
ern harvester ant, Pogonomrymex occidentalis (Hymenoptera: Formici-
dae). J. Kansas Entomol. Soc. 45: 82-101.

Pfennig, D. D., G. J. Gamboa, H. K. Reeve, J. S. Reeve, and I. D. Ferguson.

1983. The mechanism of nestmate discrimination in social wasps ( Polistes ,
Hymenoptera: Vespidae). Behav. Ecol. Sociobiol. 13: 299-305.

1986]

Rissing, Johnson, & Pollock — Acromyrmex

185

Pollock, G. B. and S. W. Rissing.

1985. Mating season and colony foundation of the seed-harvester ant, Vero-
messor pergandei. Psych 92: 125-134.

Pontin, A. J.

1960. Field experiments on colony foundation by Lasius niger (L.) and L.
flavus (F.) (Hym., Formicidae). Ins. Soc. 7: 227-230.

Rissing, S. W. and G. B. Pollock.

1986. Social interaction among pleometrotic queens of Veromessor pergandei
(Hymenoptera: Formicidae) during colony foundation. Anim. Behav.,
34: 226-233.

Queen agression, pleometrotic advantage and brood raiding in the ant
Veromessor pergandei (Hymenoptera: Formicidae). Anim. Behav., in
press.

Rockwood, L. L.

1973. Distribution, density and dispersion of two species of Atta (Hymenop-
tera: Formicidae) in Guanacaste Province, Costa Rica. J. Anim. Ecol.
42: 803-817.

Ross, K. G. and D. J. C. Fletcher.

1985. Comparative study of genetic and social structure in two forms of the
fire ant Solenopsis invicta (Hymenoptera: Formicidae). Behav. Ecol.
Sociobiol. 17: 349-356.

Stumper, R.

1962. Sur un effet de groupe chez les femelles de Camponotus vagus (Scopoli).
Ins. Soc. 9: 329-333.

Taki, A.

1976. Colony founding of Messor aciculatum (Fr. Smith) (Hymenoptera:
Formicidae) by single and grouped queens. Physiol. Ecol. Japan 17:
503-512.

Tschinkel, W. R. and D. F. Howard.

1983. Colony founding by pleometrosis in the fire ant, Solenopsis invicta.
Behav. Ecol. Sociobiol. 12: 103-113.

Waloff, N.

1957. The effect of the number of queens of the ant Lasius flavus (Fab.)
(Hym., Formicidae) on their survival and on the rate of development of
the first brood. Ins. Soc. 4: 391-408.

Weber, N. A.

1972. Gardening ants the Attines. Mem. Amer. Phil. Soc. 92: 1-146.

Went, F. W., G. C. Wheeler, and J. Wheeler.

1972. Feeding and digestion in some ants ( Veromessor and Manica). Bio-
science 22: 82-88.

Wheeler, J. and S. W. Rissing.

1975. Natural history of Veromessor pergandei. II. Behavior. Pan-Pacific
Entomol. 51: 303-314.

Wheeler, W. M.

1917. Notes on the marriage flights of some Sonoran ants. Psyche 24: 177-180.
Wilson, E. O.

1971. The Insect Societies. Cambridge, Massachusetts: Belknap Press of Har-
vard University Press.

186

Psyche

[Vol. 93

Wilson, E. O. and G. L. Hunt.

1966. Habitat selection by the queens of two field-dwelling species of ant.
Ecology 47: 485-487.

Wilson, N. L., J. H. Dillier, and G. P. Markin.

1971. Foraging territories of imported fire ants. Ann. Entomol. Soc. Amer. 64:
660-665.

REVISION OF THE ONOCOSMOECUS UNICOLOR GROUP
(TRICHOPTERA: LIMNEPHILIDAE, DICOSMOECINAE)

By Glenn B. Wiggins1 and John S. Richardson2
Introduction

The genus Onocosmoecus, by current definition, comprises the
unicolor group and the frontalis group (Schmid 1980, occidentalis
group = unicolor group; Wiggins 1977). From a separate study of
generic relationships within the Dicosmoecinae (G. B. Wiggins & O.
S. Flint, in prep.), it is clear that Onocosmoecus in this broad sense
is not monophyletic. The frontalis group will be considered in a
subsequent paper, but in the interim the two western North Ameri-
can species of which it is composed, O. frontalis (Banks) and O.
schmidi (Wiggins), remain nominally under Onocosmoecus. Thus,
in final analysis, this study of the unicolor group will constitute a
revision of the genus Onocosmoecus s.s., and the generic name is
used here in that restricted sense.

Among the genera of the limnephilid subfamily Dicosmoecinae,
Onocosmoecus s.s. is one of the most widespread, represented
across the whole of northern North America from Newfoundland to
Alaska, south in the western mountains to California, and across
the Bering Strait to Kamchatka. They are rather large caddisflies,
not often found in abundance but by no means rare. Larvae occur in
cool lotic habitats, and also in the littoral zone of cool lakes, where
they are detritivorous. Seven species have been assigned to the genus
in the past but reservations concerning their validity have been
expressed by several authors (e.g., Schmid 1955, 1980; Flint 1960;
Wiggins 1977). Because no analysis of types or of long series of
specimens has been undertaken, identity of the putative species has
always been doubtful. The purpose of the present study was to
undertake that analysis.

'Department of Entomology, Royal Ontario Museum, 100 Queen’s Park, Toronto,
Ontario, Canada MSS 2C6.

department of Zoology, University of British Columbia, Vancouver, B.C. Canada.
Manuscript received by the editor June 17. 1986.

187

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Extensive collections in the Department of Entomolgy, Royal
Ontario Museum (ROM) provided the main basis for this study and
were supplemented by material borrowed from other collections;
deposition of all other specimens examined is given below. On the
distribution map for O. unicolor (Fig. 14), not all records in the
central part of the range are plotted. Since complete listing of all
records for O. unicolor is too voluminous for inclusion here, an
abbreviated citation is used; localities are listed under state or pro-
vince and any other identifier, followed by the place of deposition of
the material, and the range for adult flight records is given for each
state or province. A complete listing of all records compiled is dep-
osited in the Library of the Royal Ontario Museum. Records for
larvae are included; instars are designated as LV (Larval instar V),
LIV, LIII, P (Pupa), PP (Prepupa). Life history data for O. unicolor
are grouped into weekly intervals for plotting (Fig. 10), with each
dot representing a collection comprising one or more individuals at
a given stage. Larvae were identified to the third instar, although
some characters are not as well developed as in the final instar.

Observations on food are based on analysis of the contents of the
entire gut from 10 LV, following the method of Cummins (1973).
Food data were recorded on a percentage area basis using an eye-
piece grid, and were classified under four categories: animal frag-
ments, vascular plant pieces and filamentous algae, diatoms, and
fine particulate organic material (FPOM) unidentifiable as to
origin.

Location of specimens examined

CAS California Academy of Sciences, San Francisco
CNC Canadian National Collection, Biosystematics Research
Institute, Agriculture Canada, Ottawa
DGD D. G. Denning, Moraga, California

DJB D. J. Burdick, Department of Biology, California State Uni-
versity, Fresno

INHS Illinois Natural History Survey, Champaign
LACM Los Angeles County Museum

MCZ Museum of Comparative Zoology, Harvard University,
Cambridge

1986] Wiggins & Richardson — Onocosmoecus 189

NHA N. H. Anderson, Department of Entomology, Oregon State
University, Corvallis

Oswood M. W. Oswood, Division of Life Sciences, University of
Alaska, Fairbanks

SDS S. D. Smith, Central Washington University, Ellensburg
UA Strickland Museum, University of Alberta, Edmonton
UBC Spencer Entomological Museum, University of British
Columbia

USNM National Museum of Natural History, Smithsonian Insti-
tution, Washington, D.C.

Vienna Naturhistorisches Museum, Vienna

Z.I. USSR Zoological Institute, Academy of Sciences, Leningrad

Genus Onocosmoecus Banks

Dicosmoecus (Onocosmoecus) Banks 1943, p. 357; type-species by original designa-
tion D. (O.) tristis Banks 1900.

Onocosmoecus: Schmid 1955, p. 37.

Onocosmoecus: Flint 1960, p. 19.

Onocosmoecus: Wiggins 1977, p. 268.

Onocosmoecus: Schmid 1980, p. 83.

Originally recognized as a subgenus of Dicosmoecus (Banks 1943),
Onocosmoecus was later elevated to full generic status by Schmid
(1955) on the basis of characters of adults. Larval characters added
to the generic diagnosis by Flint (1960) were augmented by Wiggins
(1977).

Description. Adults (Fig. 1) over-all light to medium brown
colour, legs uniformly light brown; fore wings yellow-brown with
variable markings, corneous spots in cells R4 and M variably pig-
mented from dark brown to colourless, variable darkish pigmented
areas around these spots and along apical and costal margins, rang-
ing from complete absence to the condition where most of the wing
is medium brown; these corneous points and surrounding pig-
mented areas sometimes show a range of expression in a series from
a single locality; hind wings paler and without markings. Venation
similar to Dicosmoecus except discoidal cell of forewing not more
than three times longer than basal radial sector (petiole). Length of
fore wing: male 14.5-22 mm; female 15-23 mm. Tibial spurs 1, 3, 4.
Head and thorax with sparse brownish and pale setae; setal warts
approximately same colour as surrounding cuticle; pleural setal

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[Vol. 93

Fig. 1. Onocosmoecus unicolor (Banks), males, a, British Columbia, Vancouver Is.; b, Michigan, Houghton Co.; c, Kurile
Isl, Chishima; d, Utah, Provo R.

1986] Wiggins & Richardson — Onocosmoecus 191

warts with pale setae, not as dense or long as in Dicosmoecus spp.
(except D. obscuripennis Banks); mesepisternum lacking second
setal wart.

Male genitalia (Figs. 2, 4, 11, 12). Segment IX broader laterally
than in Dicosmoecus , sternum not extended posteromesally. Seg-
ment X with external branches broad, flattened, longer than basal
segment of inferior appendages, fused to segment IX close to mid-
dorsal line dorsad of internal branches; internal branches variably
fused together dorsally, inferior branches lacking or occasionally
represented by small process, or well developed and prominent;
subanal plate cleft mesally, truncate apically. Phallus bearing pair of
stout spines ventrally at base of aedeagus; parameres partially and
variably fused with aedeagus, each expanded into membranous
apex bearing 1-6 stout spines, the arrangement, number and length
of these spines extremely variable and in more than 50 per cent of
specimens examined spines differ on each paramere of a single
individual.

Female genitalia { Figs. 3, 5, 6, 7, 13). Highly variable in structural
detail. Segment VIII variably sclerotized ventromesally. Segment
IX with massive rounded tergal lobes extending ventrolaterally to
sternum IX which is here interpreted as reduced to a very small lobe
or fold on each side of flattened vulval lobe; vulval lobe subdivided
into three parts apically but continuous basally, median part varia-
ble in shape, lateral parts concave mesally and variable in shape.
Segment X elongate, forming anal tube, open ventrally, variably
cleft dorsomesally; in ventral aspect, basal shoulders of segment X
highly variable in shape even within single series, frequently
extended into angulate ledge or tooth at each side.

Larva. (Fig. 8). Broad, light coloured median band extending
from coronal suture, over pronotum and mesonotum; sclerites of
head, pronotum and mesonotum with dense, minute spines. Prono-
tum lacking stout spines along anterior margin, but with row of
12-16 long, black setae just behind anterior margin, pair closest to
mid-dorsal line distinctly shorter, space between these setae and
next seta on either side narrower than that between remaining setae
of row (character valid at least to LIII); pronotum and mesonotum
with sparse, short yellowish and longer dark setae. Metanotal setae
confined to setal areas, variable: sal 4-15, sa2 3-13, sa3 9-26; metep-
imera each with approximately 8-15 setae. Setae on or near ventral

Figs. 2-3. Onocosmoecus unicolor (Banks). 2, Male genitalia (specimen from
Ontario, Durham Co.): a, lateral; b, dorsal; c, ventral; d, phallus, lateral and ventral.
3, Female genitalia (specimen from Ontario, Durham Co.): a, ventral; b, detail of
vulval lobe; c, lateral, (anal op., anal opening; gen. op., genital opening; IXt & IXs,
tergum and sternum of segment IX; inf. app., inferior appendage; ext. br. X, int. br.
X, external and internal branches of segment X; subanal pi., subanal plate).

1986]

Wiggins & Richardson — Onocosmoecus

193

edge of femora: profemur 1-3, mesofemur 4-11, metafemur 6-13;
tibiae with a single pair of stout spur-like setae; trochanteral brush
present on all legs, density variable. Femora flattened and com-
pressed, ventral edges blade-like. Abdominal segment VI with 1-4
setae posterodorsally on each side of median line; VII with 1-5,
usually 4 setae in this position; VIII with dorsal transverse row of
approximately 16-24 setae; dorsal sclerite on segment IX with 12-15
setae; lateral abdominal gills present on segments IV and V; abdom-
inal gills: dorsal, II 1-2, 3-4; III 3, 3-4; IV 3, 3-4; V 2-3, 2-3; VI 2-3,
2-3; VII 2, 2-3; VIII 0-2; lateral, II 0, 2-3; III 2-3, 2-3; IV 2-3, 2; V

1- 2; ventral, II 2, 4; III 3, 4; IV 3, 3-4; V 2-3, 4; VI 2-3, 3-4; VII 2,

2- 3. Length of larva up to 25 mm.

Case (Fig. 9). Constructed of fragments of leaves, wood and
bark, walls rather thin and flexible; length of case up to 27 mm.

Pupa. Generally as in Dicosmoecus with dorsal hook plates on
segments III-VII, dorsal sclerites on segment I with pronounced
median notch, and setal tufts present on first two antennal seg-
ments; dorsum of segment VIII with approximately 30 setae, dor-
sum IX with approximately 14; anal processes slightly curved.

Onocosmoecus unicolor (Banks)

Anabolia unicolor Banks 1897, p. 27; holotype $, Washington, Mus. Comp. Zool.
Harvard.

Asynarchus tristis Banks 1900, p. 254; cotypes IS, 2$, Colorado, Mus. Comp. Zool.
Harvard. New Synonymy.

Dicosmoecus coloradensis Ulmer 1905, p. 64, figs. 14-16; cotypes 2 S, 1$, Colorado,
Naturhistorisches Mus., Vienna. New Synonymy.

Anabolia quadrinotatus Banks 1908 (Anabolia 4-notata), p. 62, fig. 14; holotype S,
Newfoundland, Mus. Comp. Zool. Harvard. New Synonymy.

Dicosmoecus flavus Martynov 1914, p. 253, cotypes 2£, Kamchatka, Zool. Inst.,
Leningrad. New Synonymy.

Dicosmoecus (Onocosmoecus) occidentis Banks 1943, p. 362, figs. 104, 116, 124,
125, 128, 132, 136; holotype S, Idaho, Mus. Comp. Zool. Harvard. New
Synonymy.

Dicosmoecus (Onocosmoecus) alascensis Banks 1943, p. 363, figs. 105, 123, 129;
holotype (5, Alaska, Mus. Comp. Zool. Harvard. New Synonymy.

In 1943 Banks reviewed the characters used to distinguish the four
Nearctic species then assigned to D. (Onocosmoecus), and at the
same time described two additional species, D. (O.) occidentis and
D. (O.) alascensis. After studying more than 1000 adult specimens,

194 Psyche [Vol. 93

we conclude for reasons outlined below, that all of these names are
best treated as synonyms of the original species O. unicolor; also
included in the synonymy is the Palaearctic O. flavus.

In distinguishing species within the unicolor complex, Banks
(1943) utilized characters derived from colour of the fore wings and
genitalic morphology. The corneous points on the fore wings of
most Trichoptera in cells R4 and M (thyridial cell) are usually
darkly pigmented in species of Onocosmoecus and contrast strongly
with the light to medium brown fore wings. Around these points the
membrane often has indefinite darkened areas, and the extent of
these “clouds” was used by Banks as a diagnostic character (Fig. 1).
Although the darkened areas show some differences among type
series, we found that variation prevented their use as effective diag-
nostic characters. We have been unable to find in genitalic struc-
tures of either males or females throughout the unicolor complex
discrete or discontinuous character states signifying genetic groups
and taxonomic species. Differences in shape of the branches of
segment X or segments of the inferior appendages to which Banks
(1943) also referred seem valid for a few male specimens but blend
into a seemingly continuously variable range when more series are
studied. Particular importance as diagnostic characters was given by
Banks to the number and arrangement of spines on the parameres.
We found inordinate variability in these spines, ranging from one to
six on each paramere throughout the range of the unicolor complex
and frequently with a range in number exhibited within a series
from one locality; size and arrangement of the spines was equally
variable. Frequently on the two parameres of a single individual the
spines differed in both number and arrangement, sometimes exhibit-
ing conditions said to be diagnostic for two of the putative species.

Diagnosis of the females was based mainly on characters of the
shape of the tapered posterior extremity of segment X (Banks’
sheath of the ovipositor) and presence of a basolateral tooth or
ledge, and shape of the three parts of the vulval lobe. Using these
features Banks characterized the females in rather general terms but
not with precise diagnoses. We found, as with the males, that
because of many intermediate conditions in the characters proposed
we were unable to establish discrete groups for females within the
unicolor complex.

We have been cognizant of the possibility that species might be
definable within the unicolor complex on the basis of other

1986]

Wiggins & Richardson — Onocosmoecus

195

Figs. 4-6. Onocosmoecus unicolor (Banks). 4, Male genitalia (specimen from
Alaska, Admiralty Is.): a, dorsal; b, ventral. 5, Female genitalia (specimen from
Alaska, Admiralty Is.), ventral. 6, Female genitalia (specimen from Oregon, Baker
Co.), ventral.

characters, including those from other body structures, but we have
not been able to recognize discontinuities in any other characters.
Thus we conclude from our study of this material that the entire
unicolor complex is best treated as a single, variable taxonomic
species. Conclusions from study of the type specimens of the species
placed in synonymy follow.

O. unicolor (Banks). We have examined the holotype female
(Skokomish R., Washington) in the Museum of Comparative Zool-
ogy. The wings are torn and the apical lobes of segment X broken.
The two corneous points on the fore wing are only lightly pig-
mented, and the surrounding membrane only slightly darker than

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the rest of the wing. Banks (1943) stated that the apical lobes of
segment X are longer in O. unicolor than in the other species; in
females of the unicolor complex that we have examined these lobes
are elongate but variable, and discontinuously longer in none. In the
holotype there is a distinct tooth or ledge at the base of each lobe of
segment X on the lateral margin (e.g. Fig. 6); expression of this
character also shows continuous variability in our material and the
ledge is lacking in most specimens (e.g. Fig. 3a). The median and
lateral vulval lobes of the holotype taper to rounded points, which
blend continuously with a range of conditions in our material.

The male has not been clearly identified in the literature. The
illustration of male genitalia labeled as unicolor by Ross (1938, fig.
48) was given only the status of the “supposed male oi ‘unicolor"' by
Banks (1943); and although no locality data were given for the
specimen illustrated by Ross, Banks (1943: 364) stated that it came
from Inyo Co., California. Banks himself (1943) referred to speci-
mens from Banff and Alaska that “may be males of this species,”
offering as a diagnostic character that the third and fourth spines of
the paramere are not widely separated. Our material shows such a
very wide range of variation in arrangement of the spines of the
parameres that this character cannot be regarded as distinctive.

O. tristis (Banks). We have examined the three specimens
(South Park, Colorado) in the type series from the Museum of
Comparative Zoology. From these specimens Ross (1938) desig-
nated a lectotype male (17 Aug. 1899) and lectoallotype female (20
Aug. 1899); the remaining female (20 Aug. 1899) is identical to the
lectoallotype. Ross (1938) placed O. tristis in synonymy with O.
unicolor, but Banks (1943) maintained that the two were distinct
species. Although the females in the type series were characterized
by a pronounced basolateral tooth or ledge on segment X (Banks
1943), there seems little difference between these specimens and
what remains of this character in the holotype of O. unicolor (see
above). The apices of the posterior lobes of segment X are closely
appressed in both females of the type series, and all three parts of
the vulval lobe are truncate. This latter character contrasts with
somewhat more rounded lobes in the holotype of O. unicolor, but
we have many specimens showing intermediate conditions. The
male in the type series of O. tristis was distinguished by narrower
external branches of segment X (superior appendages of Banks), but

1986]

Wiggins & Richardson — Onocosmoecus

197

Fig. 7. Onocosmoecus unicolor (Banks), female genitalia (specimen from Kam-
chatka, U.S.S.R.; Syntype of O.flavus (Martynov)): a, ventral; b, lateral.

we find little distinction in this character and considerable variation
in our material generally (cf. Figs. 2 and 4). The five spines of the
parameres, offered as a diagnostic character for O. tristis by Banks,
have little value in view of the wide variability in number, size and
arrangement in our material of the unicolor complex. The three
specimens in the type series show slightly different degrees of pig-
mentation of the corneous points of the fore wing.

Although Ross (1938) designated both lectotype male and iectoal-
lotype female for O. tristis, there is no male specimen bearing a
lectotype label, and it must be concluded either that the specimen
was not labelled, or that the label or labelled specimen has been lost.
Since among the three remaining specimens of the type series there
is only one male, that specimen is here designated lectotype, an
assignment which would of course lapse should Ross’ lectotype be
found.

O. coloradensis (Ulmer). In the original description Ulmer
offered no diagnostic characters for separating O. coloradensis from
closely related forms, but he later commented in a re-description of
O.flavus (Ulmer 1927: 6) that his O. coloradensis was the same as
Asynarchus tristis Banks, the genital appendages resembling in turn
those of O. flavus Martynov. Diagnosis on the basis of only two
spines on each paramere was later proposed by Banks (1943), who

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Figs. 8-9. Onocosmoecus unicolor (Banks), larva (specimen from British Colum-
bia). 8a, larva with detail of mesofemur, lateral; b, abdominal segments VII, VIII, IX,
dorsal; c, head and thorax, dorsal. 9, case, detail of posterior end.

1986] Wiggins & Richardson — Onocosmoecus 199

cited an illustration by Ross (1938, fig. 48) as an example. It is not
clear what basis there was for this character; no reference was made
to it in Ulmer’s original description (no genitalic preparations had
been made from the type series), and Ross’ figure actually shows a
third small apical spine on the paramere. Origin of the specimen on
which Ross’ figure 48 was based was not given, although Banks
(1943: 364) stated that it came from Inyo Co., California; further-
more, although designated as O. unicolor by Ross, the specimen was
not accepted by Banks as the male of that species (see above), evi-
dently because he regarded it as O. coloradensis.

We have examined the three co-types (2<5, 1?, S. Colorado, 1879)
in the collection of the Naturhistorisches Museum, Vienna. The
parameres of one male each have three spines and those of the other,
four spines. We find no other features of these males that are dis-
tinctive. In the female of the type series, the apical lobes of segment
X are rather long and slender, lacking the basal ledge or tooth of the
holotype of O. unicolor or the females in the type series of O. tristis.
The two corneous points in the fore wing are dark in all specimens,
which in the males particularly are surrounded by a relatively large
dark area.

O. quadrinotatus (Banks). We examined the holotype male
(Grand Lake, Newfoundland, 28 July 1906) in the Museum of
Comparative Zoology. This is the only name based on material
from eastern North America, and was distinguished from the west-
ern forms by uniformly dark fore wings (Banks 1943). The holotype
displays this character, but in some eastern populations there is a
tendency for slighty darkening around the spot in cell R4 (Fig. lb).
Moreover, some specimens from western North America also have
uniformly dark fore wings, e.g. Fig. Id. Some of these have several
spines on the parameres (e.g., California, Nevada Co., Sage-
hen Cr., 4 Aug. 1985, 1(5, ROM), but in others the spines are
reduced to one or two (Utah, Summit Co., E. Fork Bear R., ca. 2
mi. above confluence with Bear R., 4-5 Aug. 1985, 7 $, ROM;
Idaho, Teton Co., Darby Cr., 6-7 Aug. 1985, 6(5, ROM). The holo-
type male has five and six spines respectively on the two parameres,
distinguished by the basal spine being little longer than the others
and not reaching the tip of the paramere (Banks 1943). Within the
eastern part of North America where no western species has ever
been recorded in the literature, we found spines of the parameres

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ranging from three to six, with the basal spine in some extending to
the end of the paramere; and within a single series (Province of
Quebec, Wacouno R., n. Sept. lies, 10 Aug. 1973, ROM) all
conditions from three to six are represented.

No precise diagnosis was offered for the female by Banks, but
only the general characters of rather short apical lobes (Banks’
sheath of the ovipositor) and absence of a basal ledge on segment X,
and a broad median vulval lobe (Fig. 3). Our sample of females
from eastern populations comprises only six specimens (Ont., P.Q.,
N.H., Mich.), but genitalic structures differ considerably among
them: shape of the vulval lobes, and on segment X, the length and
taper of the apical lobes and development of the basolateral ledge.
These variations concern the same characters proposed by Banks
for diagnosis of the western species of Onocosmoecus, and we find
no other basis for identification of O. quadrinotatus as a separate
species.

O. occidentis (Banks). We examined the holotype male (Wal-
lace, Idaho, 1 October) in the Museum of Comparative Zoology.
Diagnosis was based solely on genitalic characters. In the male the
internal branch of segment X (Banks’ superior plate) was said to be
broadened toward the base and to have a median separation extend-
ing to the basal fourth; our examination of the holotype reveals no
distinctive broadening in the shape of these combined internal
branches and the median separation extends no more than half the
length, which is generally characteristic of males of the unicolor
complex. Spines of each paramere are four in number as stated, but
the arrangement attributed to them holds true only for one para-
mere of the holotype, spines of the other being quite different. The
female was distinguished by characters of segment X — short apical
lobes with slightly divergent tips and lacking the basolateral tooth
or ledge; over the range of characters in O. unicolor s.L. none of
these characters is unique as described, and we find nothing that
would serve to distinguish this species.

O. alascensis (Banks). We have examined the holotype male (1
Aug. 1917) and single male paratype (29 July 1917), both from
Iditarod, Alaska, from the collection of the Museum of Compara-
tive Zoology. Among the diagnostic characters proposed by Banks
(1943) was four spines on the parameres, which the holotype has,
but the paratype has three and five spines respectively on the two

1986] Wiggins & Richardson — Onocosmoecus 201

parameres. The external branches of segment X (superior appen-
dages of Banks) are slightly narrowed at the base, but this feature is
variable and appears not to be of diagnostic value. The two corne-
ous points on the fore wings are darkly pigmented and each is
surrounded by a fairly well defined dark area. Within the material of
the unicolor complex that we have examined none of these charac-
ters is distinctive, and we find no reliable basis for identifying this
species.

O. flavus (Martynov). Recognition of this Palaearctic species
was somewhat irregular in that the description of the female
appeared as Dicosmoecus sp. (sp.n.?) (Martynov 1913: 477), with
the name proposed later (Martynov 1914: 253). To the original
description, Martynov (1913: 478) added the comment: “This spe-
cies resembles D. unicolor Banks from Washington Territory. But
having seen no specimens of the last named species, and the struc-
ture of its genital appendages being entirely unknown, I cannot
identify my specimens with D. unicolor Judging by the illustra-
tions, the female appears to have been described again as Dicosmo-
ecus sp. (Martynov 1925, figs. 1, 2). The male was described and
illustrated by Ulmer (1927) along with the female; Ulmer mentioned
the surprising similarity between flavus and the North American
coloradensis [= unicolor ] which he had described earlier. We have
examined from the Zoological Institute, Academy of Sciences,
Leningrad, one of the two female syntypes (Pushino, Kamchatka
R., 19 July 1908) and a male evidently identified by Martynov; and
in the ROM are additional specimens from two localities in Kam-
chatka (Dalneje Lake, 1(5, 19, and Ponomarskaya R., 1(5), and
from the Kurile Islands (Chishima, 1(5) (Fig. lc). The syntype
female (Fig. 7) fits readily into the range exhibited by our Nearctic
material, and we found no unique genitalic characters; the dorsal
lobes of segment X lack a basolateral tooth and the median vulval
lobe is narrow and well separated from the lateral lobes. In genitalic
characters this flavus syntype is very close to females from Washing-
ton (Olympic National Park, 29 June-1 July 1969, ROM #690148)
and from the Yukon (Dempster Hwy., km. 72, 1 Aug 1979, ROM
#791 191b); other Washington females (Minotaur Cr., Chelan Co.,
Sept. -Oct. 1976, S. D. Smith coll.) have several similar characters.
By contrast, in the Dalneje Lake female, segment X has a basolat-
eral tooth and the median vulval lobe is broad with its sides largely

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representing collection of 1 or more individuals at 1 site.

1986] Wiggins & Richardson — Onocosmoecus 203

fused to the lateral lobes, demonstrating a tendency for variation
similar to that which is so widespread among Nearctic specimens.
All four male specimens are generally consistent in genitalic charac-
ters with the external branches of segment X expanding rather
broadly at mid-length and tapering toward a rounded apex, but this
condition also occurs widely in Nearctic material; two of the speci-
mens have four spines on each paramere, but the Dalneje Lake male
shows three and five spines respectively on each paramere, and the
Chishima male five and six spines. The fore wings of the syntype
female have dark corneous points surrounded by faint darkish areas
of moderate size, similar to the type specimens of the Nearctic O.
occidentis, alascensis, and coloradensis; and these darkish areas are
somewhat variable in size in the other specimens. The few specimens
we have seen are smaller [length of fore wing male 14.5-16 mm (n =
4), female 15-17.5 (n = 2)] than most of our North American spec-
imens, although specimens of that size are represented in our mate-
rial. Finding no characters to separate these representatives of O.
flavus from the Nearctic populations of O. unicolor , we extend our
interpretation of O. unicolor as a widespread and highly variable
species to include the Palaearctic O. flavus.

Other variants. One of the extreme variants encountered occurs
in Alaska (Admiralty Island, Young Bay, 23 July 1981, 1<5, 1?,
ROM). The male of this series (Fig. 4) shows both pronounced
narrowing at the base of the external branch of segment X and
broadening toward the apex, as well as a strong tooth on the mesal
edge of the basal segment of the inferior appendage (Fig. 4b). In the
female (Fig. 5), segment X forms a slender tubular ovipositor lack-
ing any dorsomedian subdivision, and the basal shoulders of X are
not produced as a ledge. By contrast, in a female from Oregon
(Baker Co., Pine Cr., 14 July 1967, 1<5, 1?, ROM), segment X has
the form of a slender ovipositor (Fig. 6), but the base of X is
strongly produced as a sharp dentate ledge. While these are repre-
sentative of the extreme variation, we found intermediates between
them and less extreme genital structures. In a single series from
Oregon (Lane Co., 12 mi. SE Eugene, 22 Sept. 1968, 2(5, 2?, ROM)
segment X in ventral aspect of one female forms an elongate ovi-
positor similar to that in Figure 5, but in the other female the ovi-
positor is extremely wide; in one of these females the lateral vulval
lobes are enlarged apically into a thick truncate knob, very unlike
the more usual flattened condition in Figure 3b.

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Diagnosis for adults of O. unicolor (Banks) s A. Fore wings rang-
ing in colour from light yellow brown to dark brown; length of fore
wing: male 14.5-18.5 mm; female 15-21 mm.

Male genitalia (Figs. 2, 4). Segment IX not unusually short;
inferior appendages variable in shape of segments, ventromesal
angle of basal segment in ventral aspect ranging from obtuse (Fig.
2c) to sharply pointed (Fig. 4b). Segment X with external branches
tending to be orientated in an oblique to horizontal plane, usually
narrowed basally and broader toward the apex; internal branches
fused together into a flattened, somewhat pointed median lobe vari-
ably cleft at the apex; inferior branches usually absent, occasionally
represented by a small protuberance or angulate vertical lobe. Phal-
lus with parameres variably fused to aedeagus, ranging from little
separation (Fig. 2d) to almost complete separation (as in Fig. lid);
spines at apex of parameres extremely variable, ranging from 1 to 6,
usually straight and singlepointed.

Female genitalia (Figs. 3, 5, 6, 7). Segment IX with enlarged
tergal lobes uniformly bulbous; sternum IX reduced to a small
sclerotized lobe at each side of the vulval lobe. Segment X in ventral
aspect tapered and tubular, broadly open ventrally, dorsally entire
or with a narrow median cleft, base of X extended into a lateral
shoulder in ventral aspect, variable in shape and frequently dentate.

Biology. Larvae of O. unicolor live in slow water and pool areas
of cool rivers and streams, and also in the littoral zone of cool lakes.
There appears to be little preference in substrate since larvae occur
in stony streams and organic sediments of lake margins. Larvae
usually burrow into bottom sediments for pupation, fixing the case
to some larger object such as a rock. Collection records plotted by
week for specimens examined (Fig. 10) are interpreted as a univol-
tine life cycle. Most adults emerge in the period 15 July- 15 Sep-
tember. Early larval development proceeds quickly, third instars
appearing at least by early September, fourth instars by mid-
September, with fourth and fifth instars overwintering. In contrast
to Dicosmoecus (Wiggins & Richardson 1982, figs. 33, 34), no dia-
pausing fifth instar larvae were found in O. unicolor. Pupae were
collected from June to the middle of October. These data are similar
to those from an intensive study of a population in Marion Lake,
B.C. (Winterbourn 1971), except that most larvae overwintered
there as instars III and IV; egg masses (4.5-5 mm diam., approx. 150
eggs each) were found 9-24 September.

1986] Wiggins & Richardson — Onocosmoecus 205

Larvae are shredders, vascular plant pieces and filamentous algae
combined accounting for over 71% of the total gut content in mate-
rial (10 LV) we sampled. Animal fragments averaged 9.7% but in
one individual accounted for 97% of the gut content. Diatoms were
present in small numbers in most guts, averaging 12.2%, although
one specimen contained approximately 77 percent diatoms. Fine
particulate organic matter averaged 6.9%. Our data contrast
strongly with those of Winterbourn (1971) who reported only sedi-
ment and animal fragments in the guts of this species in a lake
habitat.

Head widths for the last three instars have been established as
follows (n = 204): LV, 1.62 mm (range 1.25-2.0); LIV, 1.12 mm
(1.025-1.125); LIII, 0.725 mm.

Range. (Fig. 14). As defined here, O. unicolor is transcontin-
ental through northern North America, extending throughout the
western mountains and into eastern Asia. In North America the
species is recorded from Alaska, Alberta, British Columbia, Cali-
fornia, Colorado, Idaho, Maine, Manitoba, Massachusetts, Michi-
gan, Montana, Nevada, Newfoundland, New Hampshire, New
Mexico, New York, Northwest Territories, Nova Scotia, Ontario,
Oregon, Quebec, Saskatchewan, South Dakota, Utah, Vermont,
Washington, Wisconsin, Wyoming, and Yukon.

Material examined and other records. ALASKA. 24 June-27
September. Mile 140, Hwy. 3 (CNC). Admiralty Is. (ROM).
Anchorage (ROM). Angel Cr. (ROM). Bear Cr. (CNC). Byers Cr.
(ROM). Chatanika R. (Oswood). Chena R. (Oswood). Chichagof
Is. (ROM). Chilkoot R. (ROM). Circle (ROM). Delta (CNC).
Eklutna Lk. (INHS). Etolin Is. (ROM). Fairbanks (CNC). Gle-
nallen (ROM). Gulkana R. (ROM). Haines (ROM). Hood Bay Cr.
(ROM). Iditarod (MCZ, USNM). Juneau (ROM). Kenai Peninsula
(ROM). Kodiak Is. (ROM). Lk. Iliamna (ROM). Lowe (Oswood).
Lower Summit Lk. (ROM). Moon Lk. (ROM). Palmar (USNM).
Parks Hwy., mp. 128.5 (ROM). Port Heiden (ROM). Portage
(CNC). Prince of Wales Is. (DGD). Reflection Lk. (ROM). Sadle-
rochit Spring (USNM). Squirrel Cr. Cpgrd. (CAS, DGD). Steese
Hwy., ml. 35-97.2 (ROM). Tolsona R. (ROM). Trapper Cr. (CNC).
Turner Lk. (ROM). Ugak Bay (ROM). Umnak Is. (ROM). Upper
Gulkana R. (INHS). Wasilla (INHS). Wrench Cr. (ROM).
ALBERTA. 16 July-12 October. Banff (INHS, ROM, UA,
USNM). Calgary (INHS). Canmore (ROM, UA). Coleman (UA).

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Crowsnest R. (UA). Cypress Hills Prov. Pk. (ROM). Dungarvan
Cr. (ROM). Edson (ROM, UA). Fairview (ROM). Fawcett (UA,
USNM). Ft. McMurray (USNM). Ft. Vermilion (ROM). Galwey
Brook (ROM). Gorge Cr. (UA). Hinton (UA). House R. (ROM).
Jasper Nat. Pk. (ROM, UA). Kananaskis (UA). LaBiche R.
(ROM). Longview (UA). Lundbreck Falls (ROM, UA). McLeod R.
(UA). Nojack (ROM, UA). Nordegg (ROM, UA). N. Ram R.
(ROM). Red Deer Crossing (ROM). Sheep R. (UA). Ware Cr.
(UA). Waterton Nat. Pk. (ROM, UA). Whitecourt (ROM). Wild-
horse Camp (ROM). Yara Cr. (ROM). BRITISH COLUMBIA. 2
July- 14 November. Atlin (CNC). Babine R. (INHS). Beaverdell
(CNC). Cassiar Jet. (CNC). Clinton (ROM). Creston (CNC). Cultus
Lk. (CNC, INHS, ROM). D’Arcy (ROM). E. C. Manning Prov.
Pk. (ROM). Edgewood (INHS). Fernie (INHS, ROM). Fraser Lk.
(CNC). Galena Bay (CNC). Glacier (ROM). Golden (ROM). Haney
(ROM). Harrison Lk. (CNC, INHS). Highland R. Prov. Pk.
(ROM). Invermere (ROM). Jesmond (CNC). Kamloops (ROM).
Knutsford (ROM). Langley (ROM). Lillooet (CNC, USNM). Little
Fort (CNC). Lower Post (CNC). McBride (CNC). Miledge Cr.
(CNC). Mt. Robson Prov. Pk. (ROM). New Denver (CNC). Nicola
(CNC). Pemberton (CNC). Princeton (CNC). Prophet R. Prov. Pk.
(ROM). Queen Charlotte Islands (USNM). Revelstoke (USNM).
Rolls (INHS). Rosebery (CNC). Salmon Arm (INHS). Sandon
(USNM). Sicamous (CNC). Squamish (CNC). Stanley (CNC).
Summerland (CNC). Terrace (CNC, INHS, USNM). Topley
(CNC). Trutch (CNC). Valemount (CNC). Vancouver (INHS).
Vancouver Is. (CNC, ROM). Vavenby (USNM). Walhachin (ROM).
Wycliffe (ROM). Yellowhead Pass (ROM). CALIFORNIA. 23
July.- 11 October. Alpine Co. (INHS). Fresno Co. (DJB). Inyo Co.
(DJB, INHS). Modoc Co. (CAS, CNC, USNM). Napa Co. (ROM).
Nevada Co. (ROM). Placer Co. (DGD, LACM). Plumas Co.
(DJB). Santa Cruz Co. (INHS). Sequoia Nat. Pk. (INHS). Siskiyou
Co. (CAS, USNM). Trinity Co. (CNC). Yosemite Nat. Pk.
(LACM). COLORADO. 2 August- 1 October. Cameron Pass
(INHS). Chaffee Co. (CAS). Custer Co. (USNM). El Paso Co.
(INHS). Jefferson Co. (USNM). Larimer Co. (INHS, ROM). Park
Co. (USNM). Routt Co. (CAS). Saquache Co. (ROM). S. Colo-
rado (Vienna). S. Park (MCZ). IDAHO. 5 July-1 October. Ban-
nock Co. (ROM). Bonner Co. (CAS, ROM). Idaho Co. (ROM,

1986]

Wiggins & Richardson — Onocosmoecus

207

USNM). Latah Co. (ROM). Teton Co. (ROM). Valley Co. (CAS,
ROM). Wallace (MCZ). MAINE. 4 August- 15 September. Cum-
berland Co. (USNM). Oxford Co. (INHS). Piscataquis Co.
(USNM). MANITOBA. 27 August. Flin Flon (ROM). God’s R.
(ROM). Hayes R. (INHS). MASSACHUSETTS. No adults. Berk-
shire Co. (USNM). MICHIGAN. 28 August-29 August. Emmet
Co. (ROM). Houghton Co. (ROM). Lake Co. (fide Flint 1960).
MONTANA. 7 August-28 September. Carbon Co. (ROM). Cas-
cade Co. (ROM). Flathead Co. (ROM). Gallatin Co. (GNC). Glac-
ier Nat. Pk. (DGD, ROM, USNM). Missoula Co. (INHS, ROM).
Ravalli Co. (ROM). NEVADA. 31 July. Washoe Co. (USNM).
NEWFOUNDLAND. Grand Lake, 28 July, (MCZ). Cartwright
(Labrador) 2 August (ROM). NEW HAMPSHIRE. 4 August- 10
September. Coos Co. (INHS, ROM, USNM). NEW MEXICO. 4
September. Rio Arriba Co. (INHS). NEW YORK. 7 September.
Ulster Co. (INHS). NORTHWEST TERRITORIES. 6 July-24
August. Aklavik (CNC, ROM). Great Slave Lk. (UA). Norman
Wells (ROM). NOVA SCOTIA. 12 August, Baddeck (fide Banks
1943). ONTARIO. 9 August- 18 September. Algoma Dist. (ROM).
Belfountain (ROM). Cochrane Dist. (ROM). Durham Co. (ROM,
USNM). Kenora Dist. (ROM). Lk. Superior (ROM). Midland
(ROM). Oro Station (ROM). Rainy R. Dist. (ROM). Thunder Bay
Dist. (ROM). Wellington Co. (ROM). OREGON. 21 June-13
November. Baker Co. (ROM, USNM). Benton Co. (INHS, NHA,
ROM). Blue Mtns. (ROM). Clackamas Co. (ROM). Clatsop Co.
(INHS, ROM, USNM). Crook Cr. (ROM). Deschutes Co. (NHA,
ROM). Douglas Co. (ROM). Grant Co. (ROM). Harney Co.
(ROM). Hood River Co. (ROM). Jefferson Co. (ROM). Klamath
Co. (NHA). Lake Co. (DGD, ROM). Lane Co. (ROM). Lincoln
Co. (DGD, ROM). Linn Co. (ROM). Umatilla Co. (ROM). Union
Co. (NHA, SDS). Wallowa Co. (INHS, NHA, ROM). Wasco Co.
(ROM). Wheeler Co. (ROM). Yamhill Co. (INHS). QUEBEC. 29
June-23 September. Brebeuf (ROM). Cascapedia (INHS). Harring-
ton (ROM). Matamek R. (fide Williams and Williams 1979). Mt.
Lyall (INHS). Wacouno R. (ROM). Other records fide Roy and
Harper 1979. SASKATCHEWAN. 22 August-2 September. N.
Battleford (ROM). Pierceland (ROM). Prince Albert (INHS,
ROM). SOUTH DAKOTA. No adults. Lawrence Co. (ROM).
U.S.S.R. 19 July-17 September. Kamchatka (ROM). Kurile Islands,

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Chishima (ROM). Pushino (Z.I. USSR). UTAH. 23 July-16 Sep-
tember. Cache Co. (INHS, ROM). Carbon Co. (USNM). Daggett
Co. (ROM). Garfield Co. (ROM). San Juan Co. (USNM). Summit
Co. (CAS, ROM, USNM). Wasatch Co. (INHS, ROM). Washing-
ton Co. (USNM). VERMONT. 11 September-23 September. Ben-
nington Co. (ROM). Windham Co. (DGD). WASHINGTON. 1
June-9 October. Chelan Co. (ROM, SDS, USNM). Jefferson Co.
(ROM, USNM). King Co. (ROM). Kittitas Co. (ROM, SDS,
USNM). Mt. Rainier Nat. Pk. (ROM). Okanogan Co. (USNM).
Pacific Co. (ROM). Snohomish Co. (MCZ). Whatcom Co. (CNC,
ROM, USNM). Whitman Co. (INHS). Yakima Co. (ROM). WIS-
CONSIN. (fide Longridge and Hilsenhoff 1972). WYOMING. 6
August- 1 September. Albany Co. (ROM). Carbon Co. (ROM).
Teton Co. (INHS). Uinta Co. (USNM). Yellowstone Nat. Pk.
(INHS). YUKON. 26 June-29 August. Alaska Hwy. At Aishihik R.
(CAS) and at Koidern R. (ROM). Bearfeed Cr. (ROM). Blackstone
(ROM). Burwash Landing (CNC). Champagne (CNC). Christmas
Cr. (ROM). Clear Cr. (ROM). Dawson (CNC). Dempster Hwy.,
kmp 72, 140.5 (ROM). Dezadeash Lk. (ROM). Eagle Plain (ROM).
Eagle R. (ROM). Engineer Cr. (ROM). Flat Cr. (ROM). George’s
Gorge (CNC). Haines Jet. (CNC). Haines Rd., kmp 175 (ROM).
Klondike Hwy., kmp 476, 562, 572, 626 (ROM). Kluane (UBC).
Lake Laberge (UBC). Lapie R. Canyon (ROM). Lower Rancheria
R. (ROM). Mayo Rd., kmp 14 (ROM). McQuesten R. (ROM).
Money Cr. (ROM). Pelly Crossing (ROM). Pine Cr. (ROM). Quiet
Lk. (ROM). Rancheria (UBC). Rose Lk. (UBC). South Canol Rd.,
kmp 22, 39.5, 154, 172 (ROM). Sulphur Lk. (ROM). Tagish (UBC).
Takhanne R. (ROM). Tatchun Cr. (ROM). Teslin (CNC). Watson
Lk. Cpgrd. (ROM). Whitehorse (CNC). Willow Cr. (CNC, ROM).

Onocosmoecus sequoiae n.sp.

Figs. 11-13

Almost all of the several hundred adult specimens examined fall
within the bounds of continuous variation described above in the O.
unicolor complex, except some from a few localities for the most
part in the Sierra Nevada Mountains of California. Because these
specimens show clear structural differences from O. unicolor as
defined above, and because intermediate forms have not been

1986]

Wiggins & Richardson — Onocosmoecus

209

Figs. 1 1-13. Onocosmoecus sequoiae n.sp. 1 1, Holotype male, Tulare Co., Cali-
fornia, genitalia: a, lateral; b, dorsal; c, ventral; d, phallus. 12, Variant male, Shasta
Co., California, phallus. 13, Allotype female, Tulare Co., California, genitalia: a,
ventral; b, lateral, (ext. br. X, int. br. X., inf. br. X, external, internal, & inferior
branches of segment X; subanal pi. X, subanal plate of segment X; IXt, tergum of
segment IX; X, segment X).

210 Psyche [Vol. 93

found, we consider them to represent a distinct and previously
unrecognized species.

Adult. Similar to O. unicolor in general body characters and
venation, but distinguished by characters of the male and female
genitalia as outlined in the key to species. Colour more similar to
yellowish variants of O. unicolor than to the darker brown speci-
mens; dark markings on the fore wings around the corneous spot in
cell R4 and around the thyridium variable. Length of the fore wing:
$ 18-20 mm; $ 20-21 mm.

Male genitalia (Fig. 11). Segment X with external branches in
lateral aspect broad at the base, usually somewhat tapered toward
the apex, orientation mainly in a vertical plane; internal branches in
dorsal aspect with a double-edged median crest; subanal plate in
dorsal aspect extending beyond the periphery of the internal
branches; inferior branch of X prominent as a flattened tongue
between the external branch and the subanal plate. Phallus with
parameres entirely separate from the aedeagus except at the basal
articulation, spines fewer than in O. unicolor and bent, proximal
spines cusped with small accessory points, apex of each paramere a
prominent membranous lobe.

Female genitalia (Fig. 13). Segment IX with tergum consisting
of enlarged lateral lobes as in O. unicolor, but the lobes concave
ventrolaterally; sternum of IX somewhat folded and less sclerotized
than in O. unicolor. Segment X in ventral aspect shorter and
broader than in O. unicolor, and more widely divided dorsally,
prominent basal shoulder lacking. Vulval lobe in ventral aspect with
median portion wider and more pointed than is usually so in O.
unicolor.

Larva. Unknown for this species, and consequently Onocos-
moecus larvae from the general range of O. sequoiae cannot yet be
assigned to species.

Types. Holotype male (pinned): CALIFORNIA, Tulare Co.,
Salmon Cr., trib. Kern R., Horsemeadow Campground, Sequoia
National Forest, approx. 7000 ft., 10 mi. NE Kernville, 7 August
1985, black light trap, R. W. Wisseman; Allotype female (pinned),
same data as holotype; Paratypes 48(5 17?, same data as holotype,
specimens pinned and in alcohol. These specimens are deposited in
the Department of Entomology, Royal Ontario Museum, Toronto.

1986]

Wiggins & Richardson — Onocosmoecus

211

Fig. 14. Nearctic distribution of Onocosmoecus spp.

212

Psyche

[Vol. 93

Additional Paratypes. CALIFORNIA: El Dorado Co., Tahoe Par-
adise, 5-13 August 1985, 4 <3, W. H. Tyson, DGD; 1 mi. SW Meyers,
13 July 1984, 1$, W. H. Tyson, DGD. Madera Co., Red’s Meadow,
16 August 1941, 3 S 19, M. V. Hood, LACM; Nelder Cr. Camp,
4600 ft., 25 August 1973, 19, W. H. Tyson, USNM; Central Camp,
5500 ft., 30 July 1983, 3<J, J. Larson, DGD. Siskiyou Co., Shadow
Cr., 7 mi. E Cecilville, 5 September 1968, 19, USNM. Tulare Co.,
Salmon Cr. at Horsemeadow Campground, Sequoia National
Forest, 31 July 1965, 2$, W. P. Vann, DGD; Johnsondale,
Aug. -Sept. 1985, many $$ 99, uvl, D. J. Burdick, CAS, DJB,
USNM, ROM.

From extensive u.v.l. collections made by D. J. Burdick, we have
been able to examine long series of adults of O. sequoiae. There is
no evidence of intergradation between O. sequoiae and O. unicolor,
and both species were represented in two of the series examined:
Madera Co., Lewis Cr., 16-22 September 1983; El Dorado Co., 1
mi. SW Meyers, 30 August 1984 (specimens in collection of D. J.
Burdick). In two male specimens of O. unicolor from Fresno Co.
(Friant) the inferior branch of segment X was an angulate vertical
plate, but not the flattened tongue of O. sequoiae; and the para-
meres of these specimens were typical of O. unicolor.

Range and Habitat. Adults of this species have been collected
in the vicinity of streams mainly in the Sierra Nevada Mountains of
California in El Dorado, Inyo, Madera, Plumas, Shasta, Siskiyou,
and Tulare Counties (Fig. 14). In the absence of information on
larvae of O. sequoiae, any difference in habitat between the two
species remains unknown.

Variation. Another form, provisionally considered a variant of
O. sequoiae, has been found in collections from Shasta County
(Castle Cr., approx. 3 mi. w. Hwy. 1-5, 9 August 1985, 18(5 79, uvl,
ROM; Indian Cr., Castle Crags State Park, 9 August 1985, 1<J, uvl,
ROM; Hat Cr., 25 June 1947, 1$, CAS) and Plumas County
(Thompson Cr., 0.6 mi. above Thompson Meadows, s.w. Quincy,
16-17 July 1985, 1<5, uvl, ROM). These specimens are larger than
most O. unicolor and typical O. sequoiae (length of fore wing: $
21-22 mm; 923 mm), but the principal difference is in the parameres
of the males (Fig. 12) where the proximal spine is a long, stout
straight process with a cluster of short denticles at the apex. The
distal spines on the parameres are reduced in size and nearly

1986]

Wiggins & Richardson — Onocosmoecus

213

straight, and the apical membranous lobe is also reduced. The fused
internal branches of segment X are more flattened than in the typi-
cal form and the median crest less distinct; the external branches of
X tend to be straight-sided and less tapered than in the typical form,
but are enlarged apically in one specimen in the Shasta County
series. Females of this variant are similar to the typical form.

None of these variant specimens is included in the type material of
O. sequoiae, and in the continued absence of intermediates, they
could be considered as representing a distinct species.

Key to adults of Onocosmoecus s.s. species

1 Males 2

Females 3

2(1) In lateral aspect, inferior branches of segment X large and well
developed into a flattened tongue between the external branch
and subanal plate, external branches of X largely orientated in
a vertical plane, frequently broadest at the base and tapering
apically (Fig. 11a). Known only from the Sierra Nevada

Mountains of eastern California (Fig. 14) sequoiae

In lateral aspect, inferior branches of segment X usually lack-
ing, occasionally present but very small, external branches of X
oriented more horizontally, frequently narrow at the base and
broadened apically (Fig. 2a). Widely distributed in North

America including California (Fig. 14) and also far eastern

USSR unicolor

3(1) Segment IX with terga enlarged and bulbous, segment X in
ventral aspect a tapered tube, open ventrally but closed dor-
sally for the most part or with a narrow apical cleft (Figs. 3, 5,

6, 7) unicolor

Segment IX with terga enlarged but concave ventrolaterally,
segment X in ventral aspect tubular but tapered apically little if
at all, and cleft both dorsally and ventrally in a broad V-shape
(Fig. 13) sequoiae

Acknowledgments

This study was completed under financial support (to GBW) from
the Natural Sciences and Engineering Research Council of Canada.
Field studies were supported by previous grants (to GBW) from the

214

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[Vol. 93

National Research Council of Canada (A5707), the U.S. National
Science Foundation (G22135) and the Canadian National Sports-
men’s Show. For the loan of type specimens and of general collec-
tions, we acknowledge the assistance of N. E. Woodley, A. F.
Newton and N. D. Stone, Museum of Comparative Zoology, Har-
vard University; A. Kaltenbach, Naturhistorisches Museum Wien;
O. S. Flint, United States National Museum of Natural History; D.

G. Denning, Moraga, California; D. J. Burdick, California State
University, Fresno; J. D. Unzicker, Illinois Natural History Survey;
S. D. Smith, Central Washington State College; A. P. Nimmo,
University of Alberta; N. H. Anderson and R. W. Wisseman,
Oregon State University; I. M. Levanidova, Institute of Biology and
Pedology, Vladivostok; M. W. Oswood, University of Alaska; F.
Schmid, Biosystematics Research Institute, Ottawa; L. Zhiltzova,
Zoological Institute, USSR Academy of Sciences, Leningrad; W. J.
Pulawski, California Academy of Sciences, San Francisco. For
assistance with field collections we are indebted to G. W. Courtney,

H. E. Frania, E. R. Fuller, R. Jaagumagi, L. H. Kohalmi, B. D.
Marshall, C. R. Parker, R. S. Scott, I. M. Smith, R. N. Vineyard,
R. W. Wisseman, and T. Yamamoto.

Line drawings were prepared by Anker Odum and Zile Zichmanis
of the Royal Ontario Museum. Photographs were made by Brian
Boyle, ROM Photography. Susan Pasch, Shakilah Mehrunnisa and
E. R. Fuller assisted with the preparation of the manuscript.

Summary

From analysis of type specimens of the seven putative species of
the Onocosmoecus unicolor group and of extensive collections from
many localities in North America, six names ( Asynarchus tristis
Banks, Dicosmoecus coloradensis Ulmer, Anabolia quadrinotatus
Banks, Dicosmoecus (O.) occidentis Banks, Dicosmoecus (O.) alas-
censis Banks all from North America, and Dicosmoecus flavus Mar-
tynov from Kamchatka) are proposed as junior subjective synonyms
of Onocosmoecus unicolor (Banks). Other variables are discussed
and it is concluded that existing evidence shows O. unicolor to be a
highly variable and widespread species ranging through northern
and montane North America to eastern Asia. A new species Ono-
cosmoecus sequoiae is recognized from several localities, mainly in

1986]

215

Wiggins & Richardson — Onocosmoecus

the Sierra Nevada Mountains of California. These two species con
titute Onocosmoecus s.s.; geographic distribution is summarized
and observations on biology are included.

References

Banks, N. 1897. New North American neuropteroid insects. Trans. Am. ent. Soc.
24:21-31.

1900. New genera and species of Nearctic neuropteroid insects. Trans.

Am. ent. Soc. 26: 239-259.

1908. Some Trichoptera, and allied insects, from Newfoundland. Psyche

15:61-67.

1943. Notes and descriptions of Nearctic Trichoptera. Bull. Mus. comp.

Zool. 92: 341-369.

Cummins, K. W. 1973. Trophic relations of aquatic insects. Ann. Rev. Entomol.
18: 183-206.

Flint, O. S. 1960. Taxonomy and biology of Nearctic limnephilid larvae (Tri-
choptera), with special reference to species in eastern United States. Entomolog-
ica am. 40: 1-120.

Longridge, J. L. and W. L. Hilsenhoff. 1972. Trichoptera (Caddisflies) In
Aquatic Insects of the Pine Popple River, Wisconsin by W. L. Hilsenhoff et al.
Wisconsin Dept. Nat. Resources, Tech. Bull. 54, pp. 1-44.

Martynov, A. V. 1913. Trichoptera of the Kamtshatka expedition. Russk. ent.
Obozr. 13: 476-481.

1914. Les Trichopteres de la Siberie et des regions adjacentes. IV-e

partie. Sousfam. Limnophilinae (fam. Limnophilidae). Ezheg. zool. Muz. 19:
173-285.

1925. Trichoptera recueillis au Kamtshatka par l’expedition de Mr. Th.

Riabusinskij en 1908-1909. Ezheg. zool. Muz. 26: 10-26.

Ross, H. H. 1938. Lectotypes of North American caddis flies in the Museum of
Comparative Zoology. Psyche 45: 1-61.

Roy, D. and P. P. Harper. 1979. Liste preliminaire des Trichopteres (insectes)
du Quebec. Ann. Soc. ent. Quebec 24: 148-171.

Schmid, F. 1955. Contribution a l’etude des Limnophilidae (Trichoptera).
Mitt, schweiz. ent. Ges. 28.

1980. Genera des Trichopteres du Canada et des Etats adjacents. Les

Insectes et Arachnides du Canada, Partie 7. Can. Agric. Publ. 1692. 296 pp.
Ulmer, G. 1905. Neue und wenig bekannte aussereuropaischeTrichopteren,
hauptsachlich aus dem Wiener Museum. Annin naturh. Mus. Wien 20: 59-98.

1927. Entomologische Ergebnisse der schwedischen Kamtchatka-Expedi-

tion 1920-1922. 11. Trichopteren und Ephemeropteren. Ark. Zool. 19: 1-17.
Wiggins, G. B. 1977. Larvae of the North American Caddisfly Genera (Trichop-
tera). University of Toronto Press, Toronto and Buffalo. 401 pp.

Wiggins, G. B. and J. S. Richardson. 1982. Revision and synopsis of the cad-
disfly genus Dicosmoecus (Trichoptera: Limnephilidae; Dicosmoecinae). Aquat.
Insects 4: 181-217.

216 Psyche [Vol. 93

Williams, N. E. and D. D. Williams. 1979. Distribution and feeding records of
the caddisflies (Trichoptera) of the Matamek River region, Quebec. Can. J.
Zool. 57: 2402-2412.

Winterbourn, M. J. 1971. The life histories and trophic relationships of the
Trichoptera of Marion Lake, British Columbia. Can. J. Zool. 49: 623-635.

POPULATION FLUIDITY IN LEPTOTHORAX
LONGISPINOSUS (HYMENOPTERA:FORMICIDAE)*

By Joan M. Herbers and Carol W. Tucker

Department of Zoology, University of Vermont
Burlington VT 05405

Introduction

Although social insect colonies are commonly conceived as stable
entities in time and in space, considerable information exists to
demonstrate that population fluidity can be pronounced. Data on
ants show that workers can be exchanged between nests (Kan-
nowski 1959; Scherba 1965; Chauvin and Leconte 1965; Alloway et
al 1982; Del Rio Pesado and Alloway 1983; MacKay and MacKay
1983); a colony can undergo budding (Scherba 1958; Talbot 1961;
Brian 1965; Cherix et al 1980; Stuart 1985; Pamilo et al. 1985); and
entire nests can move from one site to another (Van Pelt 1976;
Smallwood and Culver 1979; Smallwood 1982; Droual 1984;
Herbers 1985). These observations lead to the conclusion that in
some species the colony is not a fixed entity, but rather a shifting
collection influenced by ecological contingencies.

That a given colony can occupy more than one physical nest site,
a condition known as polydomy, deserves particular attention
(Fletcher and Ross 1985). Evolutionary dynamics under conditions
of colony fractionation are poorly understood, even though the
consequences for eusocial evolution may be profound. There is sur-
prisingly little information to document and measure the extent of
population fluidity for any species, a gap we help to fill in this paper.

Recent work demonstrates that some species of leptothoracine
ants are polydomous (Alloway et al 1982; Del Rio Pesado and
Alloway 1983; Stuart 1985). These inconspicuous temperate species
are well-suited for detailed studies of polydomy because they are
small and easy to culture. Here we quantify nest fission, fusion,
migration, and other features of polydomy for Leptothorax longis-
pinosus kept under semi-natural conditions in the laboratory. While

* Manuscript received by the editor May 12, 1986

217

218

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[Vol. 93

a complete understanding of population fluidity must be predicated
on work conducted in the field, our results provide insight into the
evolutionary ecology of this ant.

Population structure in L. longispinosus

Many ants of the genus Leptothorax are polygynous (Buschinger
1968, 1974), and L. longispinosus is no exception (Talbot 1957;
Headley 1943; Alloway et al 1982). Previous work on the E. N.
Huyck Preserve (Albany County, New York) showed the popula-
tion to be facultatively polygynous: some nests contain no queen,
others have one, and still others have multiple queens (Herbers

1984) . Moreover, there was a strong winter-summer dichotomy in
queen distribution. Many nests in summer are queenless, whereas in
winter such groups are rare (Herbers 1986a); similarly, the average
number of queens per nest is lower in summer. Finally, nests are
considerably more spread out in summer than in winter (Herbers

1985) . These results are best explained as correlates of a seasonal
shift in spatial structure: colony fractionation in summer and con-
densation for overwintering. It appears that, for the most part,
overwintering nests are independent colonies that become poly-
domous in summer when they fractionate to occupy several nest
sites (Herbers 1986a). This cyclic polydomy hypothesis is supported
by behavioral evidence reported below.

Methods

Nests of L. longispinosus were excavated from the New York site
in late October 1983, when they exhibited spatial relationships and a
distribution of queens among nests that is typical of winter. Each
nest was returned to the laboratory and removed from its stick,
acorn, or root. The ants were then resettled into glass tubes 10 cm
long and 4 mm in diameter. Each nest was put into a separate box
and incubated at 4°C for overwintering. In March the temperature
and light-dark cycles were slowly incremented to match outside
conditions. On May 8, 1984 the conditions were stabilized at 14
hours of light. On that date, we positioned 17 nests on 4 artificial
forest floors to duplicate their spatial positions in nature the pre-
vious fall (Figure 1). Observations and censuses were then con-
ducted until August 27, 1984, when the experiments were terminated.

1986] Herbers Tucker — Leptothorax longispinosus 219

The artificial forest floors were lm X lm in size. Each had a red
glass base upon which autoclaved pine needles, leaf fragments and
other debris typical of the habitat were scattered. The sides of the
floor were coated with petroleum jelly to prevent worker escapes,
and the entire structure was enclosed in mosquito netting to restrict
alate fights. Lights above and below the red glass base provided
illumination. The temperature was maintained at 18-20°C and rela-
tive humidity at 60-90%. Periodically water was sprinkled on the
floor to simulate rainfall. In addition to placing nests on the floor
according to where they had been collected, we supplied additional
tubes so that each floor had a total of 10 nesting sites. Nests were
supplied with water ad libidum and solid food (both frozen fruitflies
and a formula based on Bhatkar and Whitcomb’s (1970) recipe)
three times weekly. Detailed observations of behavior were taken
for the first 3 weeks (2 hours of continuous observation daily from
9:00-11:00 as well as periodic checks), after which the intensity of
observation was reduced to 2 hours per week. Nests were censused
daily for the first two weeks and weekly for the rest of the period.

Results

The initial contents of nests are given in Table 1. Four nests on 1 A
(all polygynous), three nests on IB (one queenless, one monogy-
nous, one polygynous) and five nests on 2B (two queenless, one
monogynous, two polygynous) were positioned to duplicate their
natural locations with respect to each another (Figure 1).

Direct observations of the ants showed that initially aggression
was common: workers engaged in fighting behavior, wherein two
workers would interlock mandibles, attempt to sting each other,
push or pull by the mouthparts, and so on. These encounters some-
times resulted in death of one or both participants. Not all interac-
tions were aggressive, however; workers were observed to carry
other workers, brood, and in one case a queen outside the nest.
Several occasions of tandem running (which usually precedes a col-
ony migration (Moglich 1978) were observed. In addition to inter-
acting with other ants, workers were often observed to explore,
forage, and manipulate pieces of detritus and food.

Particularly striking was exploration of the empty tubes which
represented potential new nesting sites. This exploratory behavior is

220

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1A 2A

YC •

• YD
• WH

• BD

BD •

• YD

RC •

Fig. 1 . Spatial relationships of nests placed on floors (each 1 m X 1 m). Additional
nesting tubes were supplied to give a total of 10 on each floor.

apparent from censuses when one or two workers were observed
within a tube (cf. Table 2). Sometimes this exploration was followed
by immigration, but more often there was no apparent result.

The time scale within which population changes occurred is given
in Figure 2. Fighting between workers was most frequent in the first
three weeks of the season, and virtually nonexistent after 8 weeks.
Similarly, observations of workers carrying other workers were
clustered in the first few weeks of the experiments. Exploration of
new nesting sites was quite high initially, then fell off by the second

1986] Berbers & Tucker — Leptothorax longispinosus 221

Table 1. Occupants of nests positioned on forest floors in early May

Marker

Queens

Workers

Eggs

Larvae

Floor 1A

178

Floor IB

Floor 2A

Floor 2B

1 1

101

week. Moreover, the five observations of tandem running behavior
were restricted to the first 2 weeks. Workers and/or brood moved
between existing nests primarily within the first four weeks. By
mid-June there was little activity on the floors other than routine
foraging.

The first month of census data for nests on floor 2B are given in
Table 2. From these data we can infer the following: a group of
workers moved from RC to GD on May 9. On the 10th, a queen and
some workers moved from GD to RC, and the fusion of GD and
RC continued over the following three days. On May 28 the YC nest
split, with 47 workers moving to GD and 35 remaining behind. At
about this time members of the RC site started to explore OH; this
tentative exploration continued for about two weeks more. Thus a
great deal of information about population fluidity can be gleaned
from census data alone.

The census data also showed striking differences in activity
between the first few weeks and the rest of the summer. Wholescale
migration, fission into subunits, and fusion of nests occurred most
often early in the experiments (Figure 3). Of three migration events,
two occurred in the first two weeks. Of four fission events, two

OBSERVATIONS PER HOUR

222 Psyche [Vol. 93

WEEK OF OBSERVATION

Fig. 2. Frequencies of certain events associated with population changes. The
time scale is irregular to indicate how activity dropped off in summer.

1986] Herbers & Tucker — Leptothorax longispinosus 223

occurred in the second week, and one in the fourth week. Of two
fusions, one occurred on the third day of observation. The remain-
ing events occurred in late July, and involved ants only on Table 1 A
(Figure 4): one nest moved and split within the next week; one of
those subunits was apparently joined by a second nest immediately
thereafter. Thus, although the two nests had not interacted in any
discernible way prior to the end of July, they demonstrated a
remarkable fluidity after being in place for eight weeks.

Fission rates may be a function of nest size (Stuart 1985). Nests
that underwent fractionation tended to have more queens than
those which failed to subdivide during this study (average ranks of
10.4 and 8.6, respectively), but this difference was not significant
(Mann- Whitney U-test; P > .05). Similarly, nests that underwent
fission tended to have more workers (R = 12.8) than those which
failed to subdivide (R = 7.9), but again the differences were not
significant (U = 40.6, P = .07). Although the small number of
fissions reduced the power of our analysis, nonetheless out results
are consistent with Stuart’s observations.

Most nests in this experiment reared sexuals. Since it is extremely
difficult to mimic the naturally-occurring reproductive flights of this
species in the laboratory and thus our observations of reproductive
behavior may not be indicative of natural activity, we give only a
brief account: males eclosed in late July, and after staying in the
nesting tubes for a week or so, they started to emerge onto the forest
floor. There they explored and took a few preliminary hops before
returning to their natal nests (at which point they were not always
allowed reentry). By late August, all males left their nests perman-
ently, and many were dead. Female alates, however, were much
more reclusive, and came outside the nest rather infrequently. As a
rule, these females were reaccepted into their natal nests readily. In
only two cases was a gyne from one nest accepted into a second nest;
thus acceptance of non-natal new queens may be rare in nature as
well. These observations suggest that polygyny develops in L. lon-
gispinosus nests primarily when daughters rejoin their nest of origin.

Discussion

Like all laboratory studies, our work can be criticized on the
grounds that behavior of disturbed nests in seminatural conditions

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1986] Herbers & Tucker — Leptothorax longispinosus 225

bears no resemblance to field behavior. While sensitive to this
argument, we nonetheless maintain that our data can be extended to
evolutionary and ecological considerations. There were no gross
differences between lab and field behavior; indeed a striking feature
of these Leptothorax ants is how readily they adapt to laboratory
conditions (e.g. Wilson 1975). Comparable studies of polydomy in
these tiny ants cannot be conducted in the field. Given that this
species adjusts well to captivity and no other avenue of investigation
is possible currently, we proceed to interpret results of our labora-
tory studies.

When the ants were first introduced to the artificial forest floor,
they not only encountered a new environment that required explo-
ration, but also met members of other nests. Thus the effects of
exploring new habitat and encountering new ants were initially
confounded in this study. However, we argue that, within a week,
the behavior of these ants came to reflect what might be observed in
the field. Leptothorax workers seem to become familiar with their
surroundings quickly; certainly when these ants are placed into a
new nest box the initial intense exploration wanes within 2-3 days.
Moreover, the ants would have encountered each other in nature
under spring conditions, just as they did in the lab. Therefore, while
the effects of exploring a new habitat cannot be separated out, we
feel they are relatively inconsequential after the first week of our
observations.

The most striking aspect of this study was how critical spring
activity is in determining a population structure that remains rela-
tively stable throughout the rest of the summer. The vast majority of
aggressive encounters (which may result in intraspecific dulosis
(Alloway 1980)), exploration of new nesting sites, apparent
recruitment of nestmates (tandem running), and colony subdivision
occurred within three weeks of the arrival of “spring”. In fact, very
little behavior of interest to this study was observed after June 15.
Most ants emerging from the nest in summer were apparently
searching for food or water; when two individuals met, they usually
antennated briefly and then went their separate ways. This pattern is
consistent with their natural history. The only field observations of
queens walking alone and of workers carrying other workers or
brood have been recorded in early May. Extra-nest worker activity
from June- August appears restricted to individual foraging (Herbers,
pers. obs.).

226

Psyche

[Vol. 93

FISSIONS S

FUSIONS

MIGRATIONS • •

MAT JUN JUL AUG SEP
MONTH

Figure 3. Large-scale events occurred infrequently over the season and were
clustered in early spring.

These behavioral observations accord well with inferences made
about population structure that are based on spatial distributions of
nests and queen distribution among nests (Herbers 1986a). If the
cyclic polydomy hypothesis is correct, then the nests used in this
study, which had been collected in late fall, had already undergone
colony coalition for overwintering. The units that were set out on
the floors, then, were presumably functional, independent colonies.
When ants from different colonies came back into contact after
overwintering, they re-established dominance relations through
aggressive encounters and perhaps staked out territories. Likewise,
under spring conditions, colonies fractionated to occupy empty
nesting sites. After a period of fusions, fissions, brood exchange,
and the like, a spatial pattern was achieved that was largely main-
tained throughout the rest of the summer. We expect that, had we
been able to expose the floors to more autumn-like conditions, we
would have observed nest fusions and colony condensations to
increase.

This seasonal cycle makes comparisons to other studies difficult.
Alloway et al (1982) reported that fusion resulted in each of three

1986] Herbers & Tucker — Leptothorax longispinosus 227

SITE MARKER 7/23 7/ 30 8/6

Figure 4. Schema of changes on floor 1 A in late July.

separate experiments where 2 nests of L. longispinosus were posi-
tioned naturally on floors. They also examined the closely related L.
ambiguus, for which fusions occurred in 16 of 21 replicates. Their
experiments were apparently conducted on nests collected from
spring through mid-summer, which probably included parts of
polydomous colonies. That we observed only 2 fusions in a com-
parable study may reflect the fact that we placed functional colonies
on our floors; a lower fusion rate would be expected for entire
colonies than for subunits of polydomous colonies.

In contrast to fusion events, reports of spontaneous polydomy
show rough similarity between species. Stuart (1985) found that 12
of 57 nests of L. curvispinosus underwent fission in the laboratory,
events that were dispersed throughout the season. Our fission rate (4
events for 17 nests) is quite comparable, although we observed spon-
taneous polydomy primarily in spring. Thus fission events may not
be as strongly seasonal as our results imply.

The above data are entirely consistent with the cyclic polydomy
hypothesis, since activities associated with colony fractionation
(brood transport, tandem running, fissions, worker exchange)
occurred mainly in early spring. The fluid nature of this L. longispi-
nosus population is quite evident, and can help to explain summer-
winter differences in queen and worker distribution (Herbers
1986a). The causes of cyclic polydomy are obscure at present. Col-
ony fission during spring and summer may serve to alleviate compe-
tition for food (Herbers 1985), but nest coalition in fall is more

228

Psyche

[Vol. 93

difficult to explain. Condensation for overwintering might serve
important social functions. Alternatively, a tantalizing suggestion
based on laboratory data is that nest survivorship in winter is a
function of resident queen number (Herbers 1986b). If the same
relation holds in nature, then nests subunits may have higher survi-
vorship in concert than they would alone. Whatever the proximate
and ultimate causes, the seasonal cycle in polydomy deserves closer
scrutiny.

Acknowledgments

This study was supported by grants from the National Academy
of Sciences and the National Science Foundation (DEB 82-02361
and BSR-85 16639).

References

Alloway, T. M. 1980. The origins of slavery in leptothoracine ants (Hymenop-
tera: Formicidae). Amer. Nat. 115: 247-261.

Alloway, T. M., A. Buschinger, M. Talbot, R. Stuart, and C. Thomas. 1982.
Polygyny and polydomy in three North American species of the ant genus
Leptothorax Mayr (Hymenoptera: Formicidae). Psyche 89: 249-274.

Bhatkar, A. & W. H. Whitcomb. 1970. Artificial diet for rearing various species
of ants. Fla. Ent. 53: 271-232.

Brian, M. V. 1965. Social insect populations. London: Academic Press.
Buschinger, A. 1968. Monogynie und Polygynie bei Arten der Gattung Lepto-
thorax Mayr. Ins. Soc. 15: 217-226.

Buschinger, A. 1974. Monogynie und Polygynie in Insektsozietaten. In: G. H.
Schmidt, ed. Sozialpolymorphismus bei Insekten. Stuttgart: Wissenschaftliche
Verlagsgesellschaft.

Chauvin, R. and J. LeComte. 1965. Evolution des Echanges entre differentes
colonies-filies de Formica polyctena. Mesuree a l’aide des radio-isotopes. Ins.
Soc. 12: 197-200.

Cherix, D., P. Werner, and F. Catzeflis. 1980. Organisation spatial d’un sys-
teme polycalique chez Formica (Coptoformica) exsecta Nyl. (Hymenoptera:
Formicidae). Bull. Soc. Ent. Suisse 53: 163-172.

Del Rio Pesado, M. G. and T. M. Alloway. 1983. Polydomy in the slave-
making ant Harpagoxenus americanus (Emery) (Hymenoptera: Formicidae)
Psyche 90: 151-162.

Droual, R. 1984. Anti-predator behaviour in the ant Pheidole desertorum: the
importance of multiple nests. Anim. Beh. 32: 1054-1058.

Fletcher, D. J. C. and K. G. Ross. 1985. Regulation of reproduction in eusocial
hymenoptera. Ann. Rev. Entomol. 30: 319-343.

Headley, A. E. 1943. Population studies of two species of ants, Leptothorax
longispinosus Roger and Leptothorax curvispinosus Mayr. Ann. Ent. Soc.
Amer. 36: 743-753.

1986] Herbers & Tucker — Leptothorax longispinosus 229

Herbers, J. M. 1984. Queen-worker conflict and eusocial evolution in a polygy-
nous ant species. Evol. 38: 631-643.

Herbers, J. M. 1985. Seasonal structuring of a northeastern ant community. Ins.
Soc. 32: 224-240.

Herbers, J. M. 1986a. Nest site competition and facultative polygyny in Lepto-
thorax longispinosus. Beh. Ecol. Sociobiol. 19: 115-122.

Herbers, J. M. 1986b. Effects of ecological parameters on queen number in Lep-
tothorax longispinosus (Hymenoptera: Formicidae). J. Kans. Ent. Soc. (in
press)

Kannowski, P. 1959. The use of radioactive phosphorus in the study of colony
distribution of the ant Lasius minitus. Ecol. 40: 162-165.

MacKay, W. and E. MacKay. 1983. Analysis of internest movement in Formica
haemorrhoidalis Emery. Southwest. Natur. 28: 295.

MOglich, M. 1978. Social organization of nest emigration in Leptothorax
(Hym., Form.) Ins. Soc. 25: 205-225.

Pamilo, P., R. H. Crozier, and J. Fraser. 1985. Inter-nest interactions, nest
autonomy, and reproductive specialization in an Australian arid-zone ant, Rhy-
tidoponera sp. 12. Psyche 92: 217-236.

Scherba, G. 1958. Reproduction, nest orientation, and population structure of
an aggregation of mound nests of Formica ulkei Emery (Formicidae) Ins. Soc. 2:
201-213.

Scherba, G. 1965. Analysis of inter-nest movement by workers of the ant For-
mica opaciventris Emery. Anim. Behav. 12: 508.

Smallwood, J. 1982. Nest relocations in ants. Ins. Soc. 29: 138-147.

Smallwood, J. and D. C. Culver. 1979. Colony movements of some North
American ants. J. Anim. Ecol. 48: 373-382.

Stuart, R. J. 1985. Spontaneous polydomy in laboratory colonies of the ant
Leptothorax curvispinosus Mayr (Hymenoptera: Formicidae). Psyche 92:
71-81.

Talbot, M. 1957. Population studies of the slave-making ant Leptothorax dulo-
ticus and its slave Leptothorax curvispinosus. Ecology 38: 449-456.

Talbot, M. 1961. Mounds of the ant Formica ulkei at the Edwin S. George
Reserve, Livingston County, Michigan. Ecol. 42: 202-205.

Van Pelt, A. 1976. Nest relocation in the ant Pogonomyrmex barbatus. Ann.
Ent. Soc. Amer. 69: 493.

Wilson, E. O. 1975. Leptothorax duloticus and the beginnings of slavery in ants.
Evol. 29: 108-119.

GEOGRAPHIC VARIATION IN THE CAVE BEETLE
NEAPHAENOPS TELLKAMPFI
(COLEOPTERA: CARABIDAE)

By Thomas C. Kane1 and George D. Brunner

Department of Biological Sciences, University of Cincinnati
Cincinnati, OH 45221 USA

Introduction

More than 200 species of cave limited (i.e., troglobitic) trechine
carabid beetles are known from caves of the eastern United States
(Barr, 1979b, 1981). These species are generally considered to be
derived from ancestral surface species which were widespread dur-
ing the cold, moist climates associated with glacial maxima (Barr,
1968). Subsequent warming and drying of these regions, as glaciers
retreated, led ultimately to the extirpation of surface populations,
with only some of the cave limited stocks surviving. Available evi-
dence suggests that for trechines cave isolation is irreversible (Barr,
1968, 1979a). Therefore, geographic spread of and gene flow in
troglobitic trechines will be restricted to subterranean routes (Barr,
1968). The interconnectivity of caves and the presence of geological
barriers (e.g., noncavernous strata and large rivers) become impor-
tant factors in determining the geographic extent of and degrees of
gene flow within these troglobitic taxa.

In extensive and highly continuous limestone cave systems, such
as those of the Mississippian plateaus, interpretation of evolution-
ary relationships between closely similar taxa becomes especially
complicated (Barr, 1979b). One question which arises is whether
such taxa represent multiple isolations of a common surface dwell-
ing ancestor or are the product of more recent divergence in a
common troglobitic ancestor. Even when the latter scenario appears
to be the case, divergence may only involve subtle, although gener-
ally consistent, differences in minor morphological characters.
Thus, inferences about such factors as the amount of gene flow, if

'Author to whom all editorial correspondence and reprint requests should be
addressed.

Manuscript received by the editor March 25, 1986.

231

232

Psyche

[Vol. 93

any, still occurring among the taxa, the relative degree of differenti-
ation between the various taxa, and the manner in which the present
geographic pattern has been produced may be strengthened by the
availability of genetic data such as those obtained through gel elec-
trophoresis (Barr, 1979b; Turanchik and Kane, 1979).

As Barr (1979b) has indicated, the large geographic distribution
and abundance of Neaphaenops tellkampfi populations present an
excellent opportunity to assess the extent of gene flow between local
populations of a troglobitic trechine using both morphological and
electrophoretic data. Among the many species of troglobitic tre-
chine carabid beetles in the United States, Neaphaenops tellkampfi
is noteworthy for having the most extensive geographic range and
being one of the most abundant species of the group (Barr, 1979b,
1981). The species is distributed (Fig. 1) from just south of the Ohio
River in the north to its southern limit near the Tennessee border, in
the highly cavernous Mississippian limestones of the Pennyroyal
Plateau in west central Kentucky (Barr, 1979b). The western extent
of its range is delimited by the noncavernous Big Clifty sandstone,
and the eastern and southeastern limits of the range correspond
roughly with the contact with the Salem and Warsaw limestones
(Barr, 1979b).

Neaphaenops tellkampfi , like other cave trechines, is an impor-
tant predator in terrestrial cave communities (Barr and Kuehne,
1971; Kane and Poulson, 1976). Unlike other troglobitic trechines in
the Pennyroyal Plateau, however, N. tellkampfi has evolved special-
ized behaviors which allow it to prey on the eggs and early instar
nymphs of the common cave “cricket” Hadenoecus subterraneus
(Orthoptera: Rhaphidophoridae), resources which are energy rich
and seasonally abundant (Kane and Poulson, 1976; Hubbell and
Norton, 1978). This predator-prey interaction has evolved to the
extent that no N. tellkampfi populations occur outside the range of
H. subterraneus (Hubbell and Norton, 1978). In fact, Barr (1979b)
has suggested that at least part of the eastern limits of the N. tell-
kampfi range may be determined by the absence of H. subterraneus
further east, rather than to the presence of any extrinsic geological
barrier.

Using morphological and geological criteria, Barr (1979b) has
recognized four subspecies of N. tellkampfi. The nominate subspe-
cies, N. t. tellkampfi, on which most of the ecological studies dis-

1986]

Kane & Brunner — Neaphaenops tellkampfi

233

Figure 1. Map of west central Kentucky showing locations sampled for Nea-
phaenops tellkampfi in this study. Taxonomic designations of populations at these
sites (after Barr, 1979b) are as follows: N. t. henroti: BL; CW; SS; T , N. t. meridionalis:
H; OS; ST; N. t. tellkampfi: B; BH; GO; HA; LB; P; RB; WH; N. t. viator: C; CB; S;
N. t. meridionalis X N. t. tellkampfi hybrid: F.

cussed previously have been done, is distributed in the central
portion of the range to include the caves of Mammoth Cave
National Park. Neaphaenops t. meridionalis, the southern subspe-
cies, is limited to the north by the noncavernous sandstones near the
Barren River. However, two populations are known in the south-
eastern part of the range which are morphologically intermediate
between nominate tellkampfi and meridionalis for six of nine diag-
nostic characters, suggesting a narrow zone of hybridization
between the two subspecies. Barr (1979b) points out, however, that
despite the limited gene flow, meridionalis is morphologically the
most distinct of the four subspecies. Morphological evidence (Barr,
1979b) suggests a broad zone of hybridization between nominate
tellkampfi and the eastern subspecies N. t. viator, with gradual
intergradation between the two subspecies over approximately an

234

Psyche

[Vol. 93

eight km. distance. The eastern extent of the viator range is delim-
ited by the contact of the St. Louis/ Salem and Warsaw limestones
and, perhaps more directly, by the absence of H. subterraneus
further east (Barr, 1979b). As is the case with nominate tellkampfi,
populations of viator are known from caves on both the north and
south sides of the Green River. The northern limits of the viator
range are set in large part by a sandstone ridge and extensive fault
zone across Hart County. This geological feature also appears to be
a complete barrier to gene flow between the northern subspecies N .
t. henroti and either nominate tellkampfi or viator to the south (Fig.
1) (Barr, 1979b). Despite the absence of any known hybrid popula-
tions, tellkampfi and henroti are the most similar subspecies
morphologically, and henroti also shows a large degree of morpho-
logical affinity with viator as well (Barr, 1979b).

Previous studies using gel electrophoresis (Giuseffi et al., 1978;
Turanchik and Kane, 1979) have shown that genetic variability in
local populations of N. t. tellkampfi approach those observed in
similar surface dwelling invertebrates. These results, coupled with
similar subsequent findings in other species (e.g., Dickson et al.,
1979), suggest that cave adaptation does not necessarily result in a
reduction in genetic variation. Further, genetic similarity values (I)
(Nei, 1972) among eight local populations of nominate tellkampfi
fall in the range (i.e., 0.90-1.00 (Turanchik and Kane, 1979)) com-
monly reported for populations of continuously distributed surface
dwelling species. These results substantiate the contention that con-
tinuous limestone expanses can act as underground dispersal high-
ways for cave limited species (Barr, 1968).

The purpose of the present study was to examine electrophoreti-
cally several local populations of each of the other three subspecies
of N. tellkampfi. We were interested in determining how infrasub-
specific variation in these subspecies compared with that of nomi-
nate tellkampfi. Further, we wished to use these electrophoretic data
to quantitatively assess relationships among subspecies and also to
gain some insight to how the present distributional pattern of the
species has been produced. In these regards, Barr’s (1979b) morpho-
logical and biogeographic work provides a model against which the
electrophoretic data can be examined.

1986] Kane & Brunner — Neaphaenops tellkampfi 235

Methods

Electrophoretic data gathered from a total of 1 8 populations (Fig.
1) of Neaphaenops tellkampfi were analyzed in this study. All of the
electrophoretic data for ten of these populations were gathered dur-
ing the course of the present study, between 1980 and 1983. These
ten populations include three each of N. t. henroti (BL, CW and
T/SS; Fig. 1), N. t. meridionals (H, OS and ST; Fig. 1) and N. t.
viator (C, CB and S; Fig. 1) as recognized by Barr (1979b). The
tenth population (F; Fig. 1) is a purported meridionals X tellkampfi
hybrid on morphological grounds (Barr, 1979b). Most, but not all,
of the electrophoretic data on the eight populations of N. t. tell-
kampfi (B, BH, GO, HA, LB, P, RB and WH; Fig. 1) were col-
lected in 1977-78 and reported by Turanchik and Kane (1979).
Modifications of and additions to the nominate tellkampfi data set
will be discussed in appropriate sections below. All 18 of the popula-
tions sampled, with the exception of the SS and T sites of henroti,
represent a single cave location. During the course of the study
permission to sample the SS site was rescinded before a sample
adequate for complete electrophoretic survey could be obtained.
Subsequently the nearby T site was located but it harbored a much
smaller henroti population and failed to yield a large enough sample
to obtain data on all electrophoretic loci. Pooling of the data from
the two sites, which appears to be justified by their geographic
proximity, did produce a complete set of electrophoretic data.

Beetles were maintained alive at 5°C or frozen at -80° C prior to
electrophoresis. All electrophoresis was conducted on vertical
polyacrylamide slab gels using an Ortec Model 4200 Electrophoresis
System or a Hoefer Scientific SE600 System. Sample preparation
and run procedures used in this study were similar to those dis-
cussed by Giuseffi et al. (1978) and Turanchik and Kane (1979).
Each animal provided enough homogenate for two assays.

Six enzyme systems provided a total of seven consistently scora-
ble loci. These included: alkaline phosphatase (ALP) (1); esterase
(EST) (1); malate dehydrogenase (MDH) (2); phosphoglucomutase
(PGM) (1); phosphoglucose isomerase (PGI) (1); and, xanthine
dehydrogenase (XDH) (1). In addition a general protein (GP) stain
revealed two sets of consistently scorable bands which are also

236

Psyche

[Vol. 93

included in the data. The more complete data of this study suggested
interpretational changes at two loci from those reported by Turan-
chik and Kane (1979). The present data show that the ALP bands
are properly interpreted as a single variable locus rather than as two
separate loci. Also, we have chosen a more conservative interpreta-
tion of the XDH data. Electrophoretic analysis of XDH in N. tell-
kampfi populations produces a single band per beetle with slight
differences in mobility between some individuals. Initially these data
appeared to be consistent with data reported by Singh et al. (1976)
for a variable XDH locus in Drosophila pseudoobscura. However,
application of additional techniques which Singh et al. (1976) used
to reveal multiple bands in D. pseudoobscura heterozygotes, failed
to reveal any multiple banded N. tellkampfi individuals at the XDH
locus. More recently, Finnerty and Johnson (1979) have shown that
data such as these may be the result of post-translational modifica-
tion of an enzyme encoded by a monomorphic locus. We have
chosen this interpretation of the XDH locus in the present study.
PGM was not assayed in previous studies of N. tellkampfi (Giuseffi
et al., 1978; Turanchik and Kane, 1979) and therefore populations
of N. t. tellkampfi were re-collected and surveyed for this enzyme.
The majority of the data analysis was accomplished using a Fortran
77 version of the BIOSYS-1 program developed by Swofford and
Selander (1981).

Results

Of the nine putative genetic loci examined in this study, five were
polymorphic and the remaining four were monomorphic with the
same variant fixed in all populations of the four taxa (Table 1).
Genetic variability in N. tellkampfi populations has been estimated
as the proportion of polymorphic loci per population (P) and the
average frequency of heterozygous loci per individual (H) (Table 2).
The average N. tellkampfi population is polymorphic at approxi-
mately 30% of its loci and the average individual in such a popula-
tion is heterozygous at 9.4% of its loci (Table 2). These values are
somewhat lower than those reported previously by Turanchik and
Kane (1979) as a result of the addition of another invariant locus
(PGM) and the more conservative interpretation of the XDH locus.
However, these values of P and H still approach values typically
reported for many surface invertebrates (Selander, 1976). Therefore,

1986]

Kane & Brunner — Neaphaenops tellkampfi

237

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these data continue to support the contention that cave isolation
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ability for a species (Barr, 1968; Giuseffi et al., 1978; Dickson et al.,
1979; Turanchik and Kane, 1979).

Estimates of P and H by subspecies (Table 2) indicate no differ-
ences in genetic variability between the four taxa. Of the five polymor-
phic loci examined, two, EST and PGI, are diagnostic of subspecific
differentiation (Table 1). Three variants have been detected at each
of these loci, with all meridionalis populations fixed for a slow
migrating electromorph at both loci, all viator populations fixed for
electromorphs of intermediate mobility at each locus, and all hen-
roti and nominate tellkampfi populations being fixed for the fast
migrating electromorphs at both loci. The only local population
that is polymorphic at these two loci is population F (Fig. 1 and
Table 1). This population, which is morphologically intermediate
between meridionalis and tellkampfi and a purported hybrid of the
two subspecies (Barr, 1979b), contains both the slow and fast elec-
tromorphs at both the EST and PGI loci. The fact that these elec-
tromorphs are alternatively fixed in meridionalis and tellkampfi
populations respectively provides biochemical evidence of the
hybrid nature of this population. By contrast, all three of the viator
populations are fixed for the intermediate mobility electromorphs at
both the EST and PGI loci even though two of these populations, C
and S, lie in the zone of morphological intergradation between tell-
kampfi and viator { Barr, 1979).

Rogers’ (1972) estimate of genetic similarity (S) was used for
pairwise comparisons of the 18 populations (Table 3). Rogers’ dis-
tance values were used in a UPGMA clustering procedure to pro-
duce a biochemical dendrogram (Fig. 2). Infrasubspecific genetic
identities are all greater than 0.90. This includes some populations,
such as the C population of viator, separated from other popula-
tions of the same subspecies by shallow rivers such as the Green
River. This finding is consistent with earlier work (Turanchik and
Kane, 1979) on populations BH, RB and B of nominate tellkampfi
and with findings on at least one other cave limited species in the
same area (Laing et al., 1976), and serves to reconfirm the fact that
rivers per se are not necessarily dispersal barriers for cave limited
forms. Genetic differentiation between subspecies is substantial in
some cases (Fig. 2). Neaphaenops t. meridionalis and N. t. viator

1986]

Kane & Brunner — Neaphaenops tellkampfi

239

Table 2. Genetic variability in four subspecies of Neaphaenops tellkampfi.
P = average proportion of polymorphic loci per population; H = average proportion
of heterozygous loci per individual.

Subspecies

Site

OBS.

EXP

Avg. Alleles/
Locus

henroti

0.333

0.088

0.091

1.145

0.333

0.117

0.110

1.182

T/SS

0.222

0.091

0.081

1.187

AVG

0.296

0.099

0.094

1.171

meridionalis

0.333

0.133

0.124

1.216

0.333

0.119

0.113

1.201

0.111

0.040

0.058

1.109

AVG

0.259

0.097

0.098

1.175

tellkampfi

0.333

0.058

0.079

1.116

0.333

0.077

0.094

1.132

0.222

0.088

0.086

1.153

0.333

0.107

0.11 1

1.166

0.222

0.070

0.067

1.119

0.333

0.076

0.092

1.152

0.333

0.073

0.112

1.184

0.333

0.076

0.079

1.114

AVG

0.305

0.078

0.090

1.142

viator

0.333

0.093

0.075

1.132

0.222

0.101

0.102

1.171

0.333

0.101

0.104

1.177

AVG

0.296

0.098

0.094

1.160

mer. X tell.

0.444

0.192

0.167

1.265

hybrid

Neaphaenops tellkampfi

AVG

0.302

0.094

0.097

1.162

OVERALL

0.556

show levels of similarity to each other and to the other two subspe-
cies in the range of 0.69-0.78 (Fig. 2). Genetic similarity between
henroti and nominate tellkampfi (S > 0.96; Table 3) is as great as
similarity values among local populations within a subspecies.
Although these two subspecies are the most similar of the four

240

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[Vol. 93

Table 3. Rogers’ (1972) coefficients of genetic similarity (S) for comparisons of
four subspecies of Neaphaenops tellkampfi. Values shown are averages of pairwise
comparisons of appropriate populations. Values in parentheses are the ranges of
similarity values appropriate to each comparison.

Subspecies

No. of
Pops.

N. t. h.

Subspecies

N. t. m. N. t. t.

N. t. v.

henroti

0.975

(0.971-0.978)

meridionalis

0.748

(0.732-0.766)

0.956

(0.942-0.983)

tellkampfi

0.963

(0.928-0.982)

0.730

(0.689-0.763)

0.963

(0.917-0.995)

viator

0.741

(0.714-0.758)

0.740

(0.727-0.766)

0.737

(0.713-0.765)

0.963

(0.945-0.984)

mer. X tell.
hybrid

0.878

(0.863-0.886)

0.833

(0.828-0.836)

0.865

(0.844-0.883)

0.770

(0.750-0.781)

morphologically, this large a genetic similarity is somewhat unex-
pected given the presence of the Hart Co. Ridge, an apparent geo-
logical barrier between these two subspecies.

Genetic differentiation within and between subspecies was exam-
ined using F-statistics (Wright, 1978) and a Chi-square contingency
analysis of heterogeneity (Workman and Niswander, 1970). Allo-
zyme phenotype frequencies for the 18 populations were used to
calculate genetic differentiation (i.e., F-statistics) in a hierarchichal
manner (Wright, 1978). The two hierarchical levels are subspecies
within species and local populations within subspecies. Since the
hybrid F population could not be unequivocally assigned to either
tellkampfi or meridionalis, it was considered as a fifth “subspecies”
at that level of the hierarchy. Three loci (ALP; GPT-1; MDH-2) are
variable in some or all local populations of each subspecies. Signifi-
cant heterogeneity in gene frequencies (Chi-square) was observed
among N. t. tellkampfi populations at the ALP and MDH-2 loci but
not at the GPT-1 locus (Table 4). Significant heterogeneity in gene
frequencies at the GPT-1 locus was observed among local popula-
tions of viator and among local populations of meridionalis, but no
differentiation was observed among local populations of either sub-
species at the ALP or MDH-2 loci (Table 4). No heterogeneity in
gene frequency was observed among henroti populations at any of

1986]

Kane & Brunner — Neaphaenops tellkampfi

241

0.40 0.33 0.27 0.20 0.13 0.07 0.00

ROGERS' DISTANCE

Figure 2. UPGMA dendrogram of 18 populations of Neaphaenops tellkampfi,
generated from Rogers’ genetic distance values for nine biochemical loci.

the three variable loci. The slightly greater differentiation observed
among tellkampfi populations may be due to the fact that this sub-
species has a somewhat larger geographic range than any of the
other three subspecies, or simply to the fact that more populations
(8) were examined for nominate tellkampfi than for any of the other
three subspecies.

Whereas genetic differentiation between infrasubspecific popula-
tions is slight to moderate, differentiation between subspecies is very
great (Table 5). At the level of subspecies, variation is observed at
the EST and PGI loci in addition to the three loci discussed above.
Significant heterogeneity in allele frequency between subspecies was
observed at all five loci (Table 5) and overall genetic differentiation
is very great (Fst = 0.528), with the EST and PGI loci essentially
fixed for alternative alleles in three of the four subspecies.

242 Psyche [Vol. 93

Table 4. F-statistics and heterogeneity chi-square values for four subspecies of N.
tellkampfi.

Fit

F,s

fst

SUBSPECIES

ALP LOCUS

henroti

-0.007

-0.013

0.006

0.510ns

meridionalis

0.098

0.083

0.016

1.690ns

tellkampfi

0.217

0.150

0.080

16.416*

viator

-0.183

-0.201

0.015

1.291ns

GPT-1 LOCUS

henroti

-0.166

-0.207

0.034

5.613ns

meridonalis

-0.004

-0.154

0.130

12.145***

tellkampfi

-0.083

-0.157

0.064

13.810ns

viator

0.193

0.105

0.098

12.465***

MDH-2 LOCUS

henroti

-0.007

-0.089

0.006

1.958ns

meridionalis

-0.055

-0.100

0.042

4.549ns

tellkampfi

0.209

0.176

0.041

17.605*

viator

-0.034

-0.058

0.022

2.539ns

ns = P > 0.05; * = P < 0.05; *** = P > 0.005

Fit = correlation between uniting gametes relative to the gametes of the total
population

F,s — average correlation over subdivisions of uniting gametes relative to those of
their own subdivision

Fst = correlation of random gametes within subdivisions relative to gametes of the
total population

Slatkin (1981) has proposed a method to estimate overall gene
flow in natural populations in a qualitative manner from gene fre-
quency data. Using computer simulation, Slatkin (1981) has demon-
strated a dependence between gene flow and the conditional average
frequency of an allele, p(i) where:
d = number of demes sampled
i = number of demes in which the allele occurs
p = average frequency of the alleles in those demes
Caccone (1985) used Slatkin’s technique to assess gene flow in
several species of cave animals, based on her own data for H. sub -
terraneus, the data of Laing et al. (1976) for the scavenger beetle
Ptomaphagus hirtus and Turanchik and Kane’s (1979) data for the

1986]

Kane & Brunner — Neaphaenops tellkampfi

243

Table 5. Hierarchichal F-statistics and heterogeneity chi-square analyis of allelic
frequencies between subspecies of Neaphaenops tellkampfi

Locus

fct

Fcs

Fst

ALP

0.022

0.023

-0.001

16.224**

EST

0.958

0.000

0.958

1564.120***

GPT-1

0.081

0.074

0.007

22.227**

MDH-2

0.044

0.035

0.009

17.900**

PGI

0.988

0.000

0.988

1841.171***

TOTAL

0.546

0.038

0.528

** = PC0.01; *** = P <0.005

FCt — correlation of random gametes in local populations relative to the gametes of
the total population

Fcs = average correlation over subspecies of uniting gametes relative to those of
their own subspecies

Fst = correlation of random gametes within subspecies relative to gametes of the
total population

subspecies N. t. tellkampfi. Thus, an analysis of gene flow in all four
N. tellkampfi subspecies is appropriate since both H. subterraneus
and P. hirtus are sympatric with N. tellkampfi. Further, the range of
H. subterraneus examined by Caccone (1985) is more comparable to
that of N. tellkampfi (s.l.) than simply to that of nominate
tellkampfi.

The Slatkin analysis suggests that N. tellkampfi may be qualita-
tively described as a species in which gene flow level is low. Alleles
with low incidence values (i/d) have high conditional frequencies (p)
(Fig. 3). Caccone (1985) showed that P. hirtus is also a species with
low gene flow levels. By contrast, PI. subterraneus is seen to be a
species with intermediate gene flow levels (Caccone, 1985). As indi-
cated earlier, the range of H. subterraneus is larger than and
includes the entire range of N. tellkampfi. Unlike N. tellkampfi and
P. hirtus, however, H. subterraneus is troglophilic (facultative cave
dweller) and thus is capable of some dispersal on the surface in
addition to the subterranean routes available to troglobites. Analy-
sis of the eight nominate tellkampfi populations indicates a high
level of gene flow within this subspecies (Fig. 3) despite some het-
erogeneity in gene frequencies among these populations (Table 4).
The overall pattern of gene flow is generally consistent with the
pattern of genetic differentiation obtained from the F-statistics.

244

Psyche

Discussion

[Vol. 93

The patterns of variation described here for N. tellkampfi provide
a basis for understanding some of the factors which cause genetic
differentiation in cave limited species. Barr (1979b) suggested that
three different patterns of gene flow were indicated by the morpho-
logical and geological data on the four subspecies. These include: (1)
no gene flow ( henroti with either tellkampfi or viator ); (2) very
limited gene flow ( meridionalis with tellkampfi ); and, (3) moderate
gene flow (tellkampfi with viator). Initially the biochemical data
seem to support only pattern (2) with population F clearly contain-
ing meridionalis X tellkampfi hybrids and with other meridionalis
and tellkampfi populations examined in this study showing no bio-
chemical evidence of hybridization. Thus, the morphological data
(Barr, 1979b) and now the biochemical data suggest that hybridiza-
tion is restricted to a very narrow geographic area.

The allozyme data directly support only part of pattern (1). The rela-
tively large genetic distance between henroti and viator (D = 0.289)
and the lack of any biochemical, as well as morphological (Barr,
1979b), evidence of hybridization support the assertion that
the Hart Co. Ridge is acting as a complete barrier to gene flow
between these two subspecies. The large genetic similarity between
henroti and tellkampfi (S > 0.96) does not lend support to the
conclusion that these two subspecies are also extrinsically isolated
from each other. However, allozyme studies on the scavenger beetle
P. hirtus (Laing et al., 1976) show that a population north of the
Hart Co. Ridge has a genetic similarity (I) of approximately 0.75
with two populations south of the Ridge in caves GO and RB, which
are also occupied by nominate tellkampfi. Further, the Hart Co.
Ridge coincides with the southern range limit of Orconectes inermis
(Decapoda: Astacidae) and the northern range limit of O. pelluci-
dus, two species of troglobitic crayfish whose ranges are almost
completely separate (Hobbs and Barr, 1972). Thus the evidence for
the Hart Co. Ridge as a dispersal barrier is overwhelming.

The close genetic similarity between henroti and tellkampfi is
consistent with Barr’s (1979b) supposition that all four subspecies of
N. tellkampfi are descended from a common ancestral stock that
became isolated in caves in the southern portion of the present
range. Barr argues that henroti was derived from a peripheral popu-
lation of nominate tellkampfi which penetrated north of the Hart

1986]

Kane & Brunner— Neaphaenops tellkampfi

245

i/d

Figure 3. Conditional allele frequencies (p(i)) as a function of their incidence
(i/d) in four taxa of cave-dwelling organisms. Three qualitative patterns of gene flow
are inferred: low gene flow: Neaphaenops tellkampfi (filled circles) and Ptomaphagus
hirtus [open circles (data from Laing et al., 1976)]; intermediate gene flow: Hadenoe-
cus subterraneus [triangles (data from Caccone, 1985)]; and, high gene flow: Nea-
phaenops tellkampfi tellkampfi (squares).

246

Psyche

[Vol. 93

Co. Ridge through some of the scattered cave systems known in the
area. The close biochemical similarity of henroti and tellkampfi
support this view over the alternative hypothesis that henroti
represents a separate isolation of the surface dwelling ancestral spe-
cies. Furthermore, Barr (1979b) notes that henroti has apparently
not extended its range as far northward and westward as the geolog-
ical evidence and the distribution of Hadenoecus subterraneus
would suggest is possible. This observation, coupled with the evi-
dence of high genetic similarity between henroti and tellkampfi, is
supportive of a southern origin for N. tellkampfi with the range of
henroti representing the most recent northward dispersal.

The allozyme data fail to demonstrate a broad zone of hybridiza-
tion between tellkampfi and viator (pattern (3) above). Moreover,
inclusion of additional information fails to explain the discrepancy
between the biochemical distinctness of the two taxa, on the one
hand, and the independent evidence for a broad zone of hybrid-
ization on the other. The lack of any geological barrier between
tellkampfi and viator and the large degree of morphological inter-
gradation between the two taxa (Barr, 1979b) give great support to
the hypothesis of hybridization. Two of the viator populations
examined in this study (i.e., C and S) lie within the zone of morpho-
logical intergradation, making the lack of biochemical hybridization
even more puzzling.

Genetic differentiation in N. tellkampfi occurs primarily between
subspecies, with high genetic similarity (S > 0.90) and only slight
(Fst < 0.05) to moderate (0.05 < Fst <0.15) genetic differentiation
among infrasubspecific populations. Culver (1982) reanalyzed
Laing et al.’s (1976) data on P. hirtus and found that the average
between area Nei index for P. hirtus populations in the ranges of
different N. tellkampfi subspecies was I = 0.794. The average I
between N. tellkampfi subspecies from the present study is 0.791.
Further, analysis based on conditional allele frequencies indicates
that gene flow level in both species can be qualitatively described as
low. Interestingly the two species differ greatly in their ecological
and demographic characteristics (Kane, 1982) and a substantial
amount of evidence suggests that N. tellkampfi has a longer evolu-
tionary history of cave isolation than does P. hirtus (Laing et al.,
1976; Barr, 1979b).

Caccone (1985) suggests that gene flow levels and degree of
genetic differentiation in cave species may be influenced by their

1986] Kane & Brunner — Neaphaenops tellkampfi 247

degree of dependence on the cave environment. Troglobitic species
such as N. tellkampfi and P. hirtus, which are restricted to subterra-
nean routes of dispersal, might be expected to show lower gene flow
levels and greater genetic differentiation than cave dwelling species
which are still capable of some dispersal on the surface. Although its
distribution is restricted to cave regions, H. subterraneus emerges
from caves on warm humid evenings to feed. Thus, the intermediate
levels of gene flow inferred for H. subterraneus, as opposed to low
levels for the two troglobites, may result from limited surface dis-
persal. Morphological evidence (Hubbell and Norton, 1978) also
suggests a lesser degree of geographic differentiation in H. suberra-
neus than in N. tellkampfi over approximately the same area. Mor-
phological differences occur between southwestern populations of
H. subterraneus (i.e., in the range of N. t. meridionalis ) and those to
the north. However, there is no significant morphological differen-
tiation among the northern populations of H. subterraneus (Hub-
bell and Norton, 1978), whereas in the same region N. tellkampfi is
morphologically differentiated into three distinct subspecies (i.e.,
henroti, tellkampfi and viator). Trogloxenes show less cave depend-
ence than troglophiles. Such species often use caves only sporadi-
cally and only for shelter. Unfortunately no genetic data are
available for trogloxenes which are partially or wholly sympatric
with the species described above. Caccone (1985) does report genetic
data for Euhadenoecus puteanus, a relative of H. subterraneus,
which is a forest dweller and a sporadic trogloxene over a range
from southern New York to Georgia. She finds relatively high levels
of gene flow between five cave populations of E. puteanus which is
at least consistent with the expectations for a trogloxene.

Although degree of cave dependence appears to play a major role
in determining the degree of gene flow and genetic differentiation
over the geographic range of cave dwelling species, ecological differ-
ences between species may also influence their genetic characteris-
tics. Neaphaenops tellkampfi and P. hirtus are both troglobites and
show similar biogeographic patterns of genetic differentiation.
However, ecologically the two species are dissimilar. Whereas N.
tellkampfi is a specialized predator which tends to establish large
permanent populations (Kane and Ryan, 1983), P. hirtus is more
opportunistic. Local populations may develop on small isolated
patches of organic matter such as carrion or feces from reproduc-
tion by a few founders (Peck, 1973) and such populations are often

248

Psyche

[Vol. 93

ephemeral. Thus, stochastic events may have a greater influence on
the genetic characteristics of local P. hirtus populations than on
those of N. tellkampfi. In fact, genetic variability in local P. hirtus
populations (P = 0.154; H = 0.048 (Laing et al., 1976)) appears to
be about half that of local N. tellkampfi populations (P = 0.302;
H = 0.094). Further, the average Nei index between local P. hirtus
populations in the range of N. t. tellkampfi is I = 0.874 (Culver,
1982), whereas the average I between local nominate tellkampfi
populations is 0.981. Thus, if ecological differences influence genetic
patterns of similarly cave dependent species, the effects appear to be
manifested at the level of local populations.

Acknowledgments

We would particularly like to thank Dr. Thomas C. Barr, Jr. for
valuable discussions throughout the study, including a critical
review of the manuscript. In addition, Dr. Barr kindly provided us
with locations for several of the cave sites and in one case (ST site)
provided us with material for electrophoresis. We would also like to
acknowledge and thank Curtis Meininger for field assistance and
useful discussions and Kevie Vulinec for drafting the figures. The
National Part Service, Mammoth Cave National Park, KY, kindly
provided access to several of the sites used in this study. This
research was partially supported by a National Speleological
Society Research Grant to GDB and by grants from the American
Philosophical Society (Penrose Fund No. 8718) and the National
Science Foundation (DEB-8202273) to TCK.

Summary

An understanding of patterns of geographic variation is impor-
tant in interpreting evolutionary relationships between closely sim-
ilar taxa and in inferring levels of gene flow between geographic
populations. For obligate cave dwelling (i.e., troglobitic) species,
dispersal and gene flow are restricted to subterranean routes. Thus,
the interconnectivity of caves and the presence of geological barriers
become important factors in determining the geographical distribu-
tion and the degree of gene flow among populations of troglobitic
species.

Neaphaenops tellkampfi , a troglobitic trechine beetle, has the
most extensive geographic range and is one of the most abundant of

1986] Kane & Brunner — Neaphaenops tellkampfi 249

the approximately 200 species of cave trechines in the eastern United
States. Four morphological subspecies of N. tellkampfi have
been described over its range in west central Kentucky. In the present
study, electrophoretic data were collected on a total of 18 popula-
tions to include all four subspecies. These data support the hypothe-
sis that N. tellkampfi has been derived from a single isolation of a
surface dwelling ancestor. The present distribution has apparently
resulted from a northward movement of the troglobitic stock
through subterranean routes. Morphological (i.e., subspecific) dif-
ferentiation appears to be directly related to the presence of partial
and/or complete geological barriers to dispersal in certain portions
of the range.

Comparison of genetic data on N. tellkampfi with those on other
sympatric cave dwelling species suggests that level of gene flow and
degree of genetic differentiation may be related to the degree of cave
dependence of such species. Troglobites show lower levels of gene
flow and greater genetic differentiation over their geographic ranges
than do more facultative cave dwellers (e.g., troglophiles and tro-
gloxenes) in which intermediate to high levels of gene flow have
been reported. Ecological differences between species with similar
degrees of cave dependence do not appear to produce differences in
genetic patterns on a biogeographic scale. There is some evidence to
suggest, however, that ecological differences between such species
may affect genetic variability and genetic distance at the level of
local populations.

References

Barr, T. C. 1968. Cave ecology and the evolution of troglobites. In: T. Dob-
zhansky, M. K. Hecht and W. C. Steere (eds.) Evolutionary Biology, Plenum
Press, New York pp. 35-102.

Barr, T. C. 1979a. Revision of Appalachian Trechus (Coleoptera: Carabidae).
Brimleyana 2: 29-75.

Barr, T. C. 1979b. The taxonomy, distribution, and affinities of Neaphaenops
with notes on associated species of Pseudanophthalmus (Coleoptera: Carabi-
dae). American Museum Novitates: No. 2682 pp. 20.

Barr, T. C. 1981. The cavernicolous carabid beetles of North America. Proceed-
ings of the Eighth International Congress of Speleolology 1: 343-344.

Barr, T. C. and R. A. Kuehne. 1971. Ecological studies in the Mammoth Cave
ecosystem of Kentucky. II. The ecosystem. Annales de Speleolologie 26: 47-96.

Caccone, A. 1985. Gene flow in cave arthropods: a qualitative and quantitative
approach. Evolution 39: 1223-1235.

250

Psyche

[Vol. 93

Culver, D. C. 1982. Cave life: evolution and ecology. Harvard University Press,
Cambridge, Mass.

Dickson, G. W., J. C. Patton, J. L. Holsinger and J. C. Avise. 1979. Genetic
variability in cave-dwelling and deep-sea organisms with emphasis on Cran-
gonyx antennatus (Crustacea: Amphipoda) in Virginia. Brimleyana 2: 1 19-130.

Finnerty, V. and G. Johnson. 1979. Post-translational modification as a poten-
tial explanation of high levels of enzyme polymorphism: xanthine dehydroge-
nase and aldehyde oxidase in Drosophila melanogaster. Genetics 91: 695-722.

Giuseffi, S., T. C. Kane and W. F. Duggleby. 1978. Genetic variability in the
Kentucky cave beetle Neaphaenops tellkampfi (Coleoptera: Carabidae). Evolu-
tion 32: 679-681.

Hobbs, H. H., Jr. and T. C. Barr. 1972. Origins and affinities of the troglobitic
crayfishes of North America (Decapoda: Astacidae). II. Genus Orconectes.
Smithsonian Contributions to Zoology: No. 105 pp. 84.

Hubbell, T. H. and R. M. Norton. 1978. The systematics and biology of the
cave crickets of the North American Tribe Hadenoecini (Orthoptera Saltatoria:
Ensifera: Rhaphidophoridae: Dolichopodinae). Miscellaneous Publications
Museum of Zoology, University of Michigan, Ann Arbor: No. 156 pp. 124.

Kane, T. C. 1982. Genetic patterns and population structure in cave animals. In:
D. Mossakowski and G. Roth, eds. Environmental adaptation and evolution.
Gustav Fischer, Stuttgart, West Germany, pp. 131-149.

Kane, T. C. and T. L. Poulson. 1976. Foraging by cave beetles: spatial and
temporal heterogeneity of prey. Ecology 57: 793- 800.

Kane, T. C. and T. Ryan. 1983. Population ecology of carabid cave beetles.
Oecologia (Berlin). 60: 46-55.

Laing, C. D., G. R. Carmody and S. B. Peck. 1976. Population genetics and
evolutionary biology of the cave beetle Ptomaphagus hirtus. Evolution 30:
484-498.

Nei, M. 1972. Genetic distance between populations. American Naturalist 106:
283-292.

Peck, S. B. 1973. A systematic revision and the evolutionary biology of the
Ptomaphagus (Adelops) beetles of North America (Coleoptera: Leiodidae), with
emphasis on cave-inhabiting species. Bulletin of the Museum of Comparative
Zoology (Harvard). 145: 29-162.

Rogers, J. S. 1972. Measures of genetic similarity and genetic distance. Studies in
genetics, University of Texas Publications 7213: 145-153.

Selander, R. K. 1976. Genic variation in natural populations. In: F. J. Ayala
(ed.) Molecular Evolution, Sinauer, Sunderland, Mass. pp. 21-45.

Singh, R. S., R. C. Lewontin and A. A. Felton. 1976. Genetic heterozygosity
within electrophoretic “alleles” of xanthine dehydrogenase in Drosophila pseu-
doobscura. Genetics 84: 609-629.

Sl atkin, M. 1981. Estimating levels of gene flow in natural populations. Genet-
ics 99: 323-335.

Swofford, D. L. AND R. B. Selander. 1981. BIOSYS-1: a FORTRAN program
for the comprehensive analysis of electrophoretic data in population genetics
and systematics. The Journal of Heredity 72: 281-283.

1986] Kane & Brunner — Neaphaenops tellkampfi 251

Turanchik, E. J. and T. C. Kane. 1979. Ecological genetics of the cave beetle
Neaphaenops tellkampfi (Coleoptera: Carabidae). Oecologia (Berlin) 44: 63-67.

Workman, P. L. and J. D. Niswander. 1970. Population studies on southwest-
ern Indian tribes. II. Local genetic differentiation in the Papago. American
Journal of Human Genetics 22: 24-49.

Wright, S. 1978. Evolution and the genetics of populations, vol. 4. Variability
within and among natural populations. University of Chicago Press, Chicago.

BIOSYSTEMATIC REVISION OF EPIMYRMA KRAUSSEI,
E. VAN DELI, AND E. FORELI
(HYMENOPTERA: FORMICIDAE)

By Alfred Buschinger1, Karl Fischer1, Hans-Peter Guthy1,
Karla Jessen1, and Ursula Winter2

Introduction

The myrmicine genus Epimyrma Emery 1915 presently comprises
11 described species, inhabiting central and southern Europe and
North Africa. They all are living as social parasites together with
host species of the genus Leptothorax (subgenera Myrafant Smith
1950 and Temnothorax Mayr 1861), some as active slavemakers,
e.g. E. ravouxi (Andre 1896) (Winter 1979), others as “degenerate
slavemakers” (£. kraussei Emery 1915 (Buschinger & Winter 1982)),
and E. Corsica (Emery 1895) as a workerless permanent parasite
(Buschinger & Winter 1985).

The taxonomy of the genus is not yet completely consolidated.
Thus, in the most recent revision, Kutter (1973) comes to the con-
clusion that E. kraussei, E. vandeli Santschi 1927, and E. foreli
Menozzi 1921, are so similar that a future comparison of larger
series presumably would reveal their synonymy. It is the object of
this paper to provide evidence for the accuracy of Kutter’s predic-
tion. E. kraussei was described by 2 5$ and 1 ? (Emery 1915) from
Sorgono, Sardegna. Menozzi (1921) established E. foreli on the
basis of 4 colonies from the vicinity of Sambiase di Calabria, S ’Italy,
and E. vandeli was described after 6 colonies collected by A. Vandel
near Miramont-de-Quercy and Touffailles, Dept. Tarn-et-Garonne,
in S’France (Santschi 1927, Vandel 1927). The most distinctive
characters of the 3 species were slightly different shapes of the peti-
oli, different grades of coloration from light, yellow-brown in E.
foreli to a nearly black in E. vandeli, and the lack of $ $ in the latter
as opposed to E. foreli and E. kraussei.

’FB Biologie, Institut fur Zoologie, der Technischen Hochschule, Schnittspahnstr.
3, D-6100 Darmstadt, FRG

2FB 2 (Biologie), Universitat Bremen, Postfach 330440, D-2800 Bremen, FRG
Manuscript received by the editor December 23, 1985.

253

254

Psyche

[Vol. 93

During the past years, we have collected E. kraussei from numer-
ous localities in the mediterranean area, including the type localities
of E. vandeli and E. foreli. We were studying their populations, <$9
9 -production in the lab and in the field, their reproductive behav-
ior, colony foundation behavior, and karotypes. Crossbreeding of
several populations including E. vandeli and E. foreli was possible.
All observations pointed towards a synonymy of the 3 species.
Finally, the types were examined, and morphological studies includ-
ing the $$ of the 3 species were carried out. This considerable body
of evidence now clearly demonstrates that E. foreli and E. vandeli
represent but junior synonyms of E. kraussei.

Material Collected and Range of
Epimyrma kraussei

A total of 337 colonies of E. kraussei (including E.v. and E.f)
have been collected between 1975 and 1984 (table 1). Populations
are numbered for an easier identification in the following text. Fig. 1
may provide a visual impression of the range of E. kraussei; it also
contains a few additional localities from the literature, mainly those
from North Africa (Cagniant 1968). Nests usually are found in cre-
vices between flat stones, most easily in old dry walls of terraced
vineyards and olive orchards, but also in rocky slopes underneath
shrubs (Buschinger & Winter 1983). Colonies are small and can thus
be aspirated almost completely. In the type locality of E. vandeli, we
did not find the species in the exact sites of Vandel; however, we
could collect a sample of 1 1 colonies near Lauzerte, only 5 km W of
the original site, in the limestone slopes of the Barguelonne valley
(table 1, no 5). E. foreli had been found near Sambiase di Calabria,
in moss covering the bark of olive trees (Menozzi 1921). We tried in
vain to find Leptothoracini in such sites, presumably because the
ants have been decimated there by pesticide treatment of the trees.
However, in several localities around Sambiase (table 1, no 19), we
found 22 colonies of a yellowish Epimyrma with Temnothorax
hosts, again in terrace walls. We are convinced that they represent
members of the same population as that studied by Menozzi. Unfor-
tunately, the search for E. kraussei in its type locality, Sorgono in
Sardegna, Italy, in April 1985, remained unsuccessful. Even the host
species was quite rare in this area. From the map (Fig. 1) we may
conclude that both the type localities of E. vandeli and E. foreli are
situated well within the area of E. kraussei.

1986] Buschinger et al. — Revision of Epimyrma 255

Table 1. Localities and numbers of colonies collected of Epimyrma kraussei
Emery 1915 (no 5a: Type locality of E. vandeli Santschi 1927, no 19: Type locality of
E. foreli Menozzi 1921).

population

no.

locality

n colonies

1981/07/14-30

Calpe (Spain, E’coast)

2 a

1981/03/30

Banyuls (S’France)

1984/04/03

Puig de Pani (NE’Spain)

1984/04/03

Selva de Mar (NE’Spain)

1984/04/03

Faro de Sarnella (NE’Spain)

3 a

1984/04/05

Pont de Bar/Seo de Urgel
(Span. Pyrenees)

1984/04/05

Tremp/Tolva (Span. Pyr.)

1984/04/06

Ainsa (Span. Pyr.)

1984/04/06

Broto (Span. Pyr.)

1984/04/10

Chapelle St. Pons (S Bouleterne,
French Pyrenees)

5 a

1981/03/31

1981/04/01

Lauzerte/Quercy (S’France)

1978/08/10

Cabrespine/ Aude (S’France)

1981/03/23

La Couronne/ Bouches-du-Rhone
(S’France)

1981/04/02

Nyons/ Drome (S’France)

1984/04/11

Suze-la-Rousse/ Vaucluse (S’France)

1983/05/07-08

Ste. Maxime, Puget Ville/Alpes
Maritimes (S’France)

1982/03/25

Venaco/ Haute Corse (France)

1983/05/03-06

Alassio. Albenga, Ranzo, Toirano
Ventimiglia/ Prov. Imperia and
Savona (N ’Italy)

1975/05/29

Aosta (N’ltaly)

12 a

1978/05/02

Ossuccio/Lago di Como (N’ltaly)

1978/10/14

1980/10/13

Biolo/ Valtelino (N’ltaly)

1980/10/12

Lovere (Lago d’Iseo, N’ltaly)

1979/04/09

1980/05/05-06

1980/10/11

Tignale (Lago di Garda, N’ltaly)

113

256

Psyche

[Vol. 93

Table 1, continued.

population

no.

locality

n colonies

1981/03/26

1982/10/12

1974/06/15

Salorno (Adige, N’ltaly)

1981/09/23-26

Krk (Dalmatia, Yugoslavia)

1983/09/29

Pag (Dalmatia, Yugoslavia)

1978/08/22

Nacionalni park Paklenica (Dalmatia,
Yugoslavia)

1982/10/03-10

Gizzeria, Rogliano, near Sambiase
(Calabria, S’ltaly)

337

Morphological Studies

Comparison of the type material of E. kraussei, E. vandeli, and E.
foreli with new material

The types of E.v. and E.f are deposited in the Naturhistorisches
Museum Basel, Switzerland. We could study 1$ E. foreli, and 1? E.
vandeli, both from the type series. The Museo Civico di Storia
Naturale “Giacomo Doria” in Genova, Italy, has provided us with
the types (1$, lj) of E. kraussei.

With a close examination of these types we could only confirm
the similarity of all 3 “species” as was already stated by Kutter
(1973). We therefore refrain from a detailed presentation of mea-
surements and structures compared. We also did not find any con-
stant differences between the types and specimens from our newly
collected material, with respect to size, shape of petioli, head and
thorax, length of body hairs etc.; just the coloration was slightly
variable between different populations. Thus, the population from
Calabria (E. foreli), and one from Spain (pop. no. 3) exhibit a quite
light, yellowish brown coloration of $ and $. Other E.k. popula-
tions appear brownish, whereas a dark brown or nearly black is
typical for 9 E. vandeli (pop. no. 5), for a colony from La Couronne
(no. 6), and for population no. 9 from Corsica. Young 9 9 are
darker in coloration than old queens, and callow 9 2 usually exhibit
some darker spots in the thorax, and a yellow base of the gaster,

1986]

Buschinger et al. — Revision of Epimyrma

257

Fig. 1. Distribution of Epimyrma kraussei Emery 1915. •: Our collecting sites
listed in Table 1. No. 5: Type locality of E. vandeli Santschi 1927; no. 19: Type
locality of E.foreli Menozzi 1921. A and C: Localities of E. vandeli in N’Africa cf.
Cagniant (1968), B: Locality of E. kraussei cf. Cagniant (1968); D: Type locality of E.
kraussei in Sardegna.

whereas the coloration in old queens is usually uniform. This age-
dependent color variation is also typical for E. ravouxi (Andre 1896)
(Buschinger 1982).

Male genitalia, wing venation, and shape of 9$ petioli

We studied wing venation and genitalia of E.k. $$, and the out-
lines of the 9 and $ petioli of specimens from Tignale and Biolo
(Italian Alps), Calpe (Spanish Mediterranean coast), Calabria
(S’ltaly, E. foreli), and Lauzerte (S’France, E. vandeli). The same
characters were investigated in E. ravouxi from several distant pop-
ulations [Taubertal: Bavaria (D), Swiss Valley (CH), S’France, Cor-
sica (F)], in order to compare their variation within and between the
species. E.r. is clearly distinct from E.k. (Buschinger & Winter 1983,
Winter and Buschinger 1983), and thus may serve as a reference
species. Males preserved in alcohol were dissected, and permanent

258

Psyche

[Vol. 93

Fig. 2: Sagittae of Epimyrma a: Pop. no. 19 (E. foreli)\ b: Pop. no. 14 (E.

kraussei ); c: E. ravouxi from Corsica: d: Pop. no. 5a (E. vandeli).

mounds were made of the subgenital plate, the sagittae, and volsel-
lae with laciniae, as well as the forewings and antennae. The outlines
of 2 and 2 petioli were drawn and superimposed following a
slightly modified method of Wehner (1983). As far as possible we
always studied 10 (522 from each of the populations mentioned
above.

Male genitalia

Table 2 reveals that the numbers of sagittal teeth (Fig. 2) vary
both within E.k. and E.r., but with higher mean values in E.k.,
including the populations of E.v. and E.f.

The volsellae and laciniae (Fig. 3, table 3) exhibit a high confor-
mity in E.k. and the two populations of E.v. and E.f., in that the
cuspis (tip of lacinia) rarely reaches, and never overlaps the digitus
(terminology following Bitsch 1979). In E. ravouxi, on the contrary,

1986]

Buschinger et al. — Revision of Epimyrma

259

Table 2. Numbers of sagittal teeth in SS of Epimyrma kraussei Emery 1915
(= E. vandeli Santschi 1927, = E. foreli Menozzi 1921), and of E. ravouxi Andre
1896) from different populations.

species / population

min

n teeth

max

n sagittae
checked

E. kraussei

no 14 Tignale

13.9

no 12b Biolo

13.6

no 1 Calpe

14.3

no 19 Calabria {E.f.)

14.4

no 5a Lauzerte (E.v.)

13.7

E. ravouxi

Bavaria (D)

11.5

Nyons (F)

12.6

Corsica (F)

10.3

Swiss Valley (CH)

12.4

the cuspis usually overlaps or at least reaches the digitus, with very
few exceptions.

The subgenital plates did not differ between populations or
species.

Male wing venation

Wing venation in Epimyrma 92 is quite variable (Andre 1896,
Kutter 1973). In $ forewings the radial cell is short and open, the
cubital cell long and usually closed, the discoidal cell may be closed,
open, or nearly lacking, and the recurrens can be complete, incom-
plete, or absent. Reductions of wing venation need not be symmetri-
cal in the two forewings of a specimen. We compared mainly the
shape of the discoidal cells, which exhibits sizable differences
between the species, but varies also within E.k. and E.r. considera-
bly (Fig. 4).

Thus, table 4 shows the numbers of wings with open or closed
discoidal cell. This character apparently is not appropriate for a
differentiation of species or populations. A slightly better distinc-
tion is possible with the shape of the discoidal cell (table 4). In E.
ravouxi this cell is near to quadratic, with a slightly shorter anterior
border. This is also true for a good deal of the N’ltalian and the
Spanish populations of E.k., but already in these populations, and
more in the Calabrian (no 19, E.f) and the Lauzerte (no 5, E.v.)

260

Psyche

[Vol. 93

0, 2 mm

Fig. 3: Volsellae and laciniae of Epimyrma a: Population no. 14 ( E . kraus-

sei)\ b: Pop. no. 19 ( E.foreli)\ c: Pop. no. 5a (E. vandeli)\ d: E. ravouxi from Corsica.

populations the anterior border becomes shorter until the discoidal
cell is triangular.

Shape of the petioli in $$ and 99

In several publications (e.g. Menozzi 1931, Sadil 1953) the pro-
files of 9 and 9 petioli and postpetioli were used as the most impor-
tant characters for the determination of Epimyrma species. Kutter
(1973), however, clearly demonstrated with 92 from a single E.
ravouxi colony that these profiles may vary to such an extent that
they are useless for species discrimination.

Nevertheless, we again studied this character, using a slightly
modified method of Wehner (1983). The outlines of the petioli of 10
99 and 10 $9 (exceptions: Population 12b: 5 99 > and population
5a: 399) Per population were drawn with the aid of a Wild M5

1986]

Buschinger et al. — Revision of Epimyrma

261

Table 3. Morphological comparison of the shape of volsella and lacinia in SS of
Epimyrma kraussei Emery 1915 (= E. vandeli Santschi 1927, = E. for eli Menozzi
1921), and of E. ravouxi (Andre 1896) from different populations

species/ population

n volsellae and laciniae where
cuspis

antrum reaches c. overlaps

open digitus digitus

checked

E. kraussei

no 14 Tignale

no 12b Biolo

no 1 Calpe

no 19 Calabria {E.f.)

no 5a Lauzerte {E.v.)

E. ravouxi

Bavaria (D)

Nyons (F)

Corsica (F)

Swiss Valley (CH)

dissecting microscope and a drawing tube at about X 88. The draw-
ings then were superimposed in such a way that they all were of the
same size and overlapped to a maximal degree (Fig. 5). However,
sizes and profiles of the petioli are varying within each population
so much that a clear distinction of populations by this character is
impossible. Even between E. kraussei and E. ravouxi we could not
find any reliable differences in the petiolar outlines. The character,
therefore, is useless for taxonomical purposes in the Epimyrma spe-
cies investigated, and it can neither support nor contradict a syn-
onymization of E.f and E.v. with E. kraussei.

Karyology

Karyotypes were studied using the air-drying technique of Imai et
al. (1977). Usually we made preparations from testes of $ pupae,
and a few from cerebral ganglia of prepupae. E. kraussei from sev-
eral populations (pop. no. 1, 5b, 6, 7, 9, 12a) and E. vandeli (pop.
no. 5a) were checked, whereas no preparations of E. foreli could be
made.

A total of 215 metaphase cells of 16 E. kraussei-$ pupae from 8
colonies of 6 different localities showed 10 chromosomes each (Fig.
6). 6 cells had 9 chromosomes, and 5 cells had the diploid number of

262

Psyche

[Vol. 93

Fig. 4: Forewings of Epimyrma SS- Left: Variation of wing venation of E.

ravouxi from 3 populations; reduction of the subrectangular discoidal cell, a: Corsica,
b: Swiss Valley, c: Taubertal, Bavaria. Right: Variation of wing venation within
one E. Araussez-population (no. 1, Calpe, Spain). d,e,f: Reduction of the sub-
triangular discoidal cell.

2n = 20. 9 cells of 2 additional, apparently $, prepupae contained 20
chromosomes, 2 others had 17 and 15, respectively. Chromosome
numbers of less than the haploid (n = 10) or diploid (2n = 20)
number are probably due to loss of chromosomes during prepara-
tion. Single diploid cells in haploid $ $ were occasionally found in
other species, too (e.g. Hauschteck 1962, 1965).

In 2 E. vandeli $ pupae from a colony from the type locality (pop.
5a), 26 and 15 cells, respectively, were checked. They all had 10
chromosomes each.

The karyotypes of E. kraussei and E. vandeli with n = 10 chromo-
somes are apparently identical. They consist of 6 small to medium-
sized metacentrics, 3 medium-sized submetacentrics and 1 large
subtelocentric. E.k. and E.v. share this karotype with all the species
of this genus so far studied [E. bernardi Espadaler 1982, E. Corsica
(Emery 1895), E. ravouxi (Andre 1896) and E. stumperi (Kutter

venation in forewings of SS of Epimyrma kraussei Emery 1915 (= E. vandeli Santschi 1927, - E. foreli

1986]

Buschinger et al.— Revision of Epimyrma

263

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[Vol. 93

1950)], and with Myrmoxenus gordiagini Ruszky 1902, a species
very closely related to Epimyrma (Buschinger et al. 1983, Fischer
unpubl.). No host species of Epimyrma and no other Leptothora-
cine species having this particular karyotype could yet be found.
Thus, we may suppose that E. foreli as well has the karyotype of the
genus, and no arguments for or against the synonymization of the 3
snecies in auestion can be derived from our karyological studies.

Biological data

Host specificity

The host species of E. kraussei in all populations investigated,
including those ascribed to E. vandeli and E. foreli, is invariably
Leptothorax ( Temnothorax ) recedens (Nylander 1856). All other
Epimyrma species have different host species belonging to the sub-
genus Myrafant (Kutter 1973, Espadaler 1982, Buschinger & Winter
1985), and no other Epimyrma species has ever been found with
Temnothorax hosts. In or close to the localities where we have
collected E. kraussei (table 1) we usually found several other Lepto-
thorax species, particularly often L. (Myrafant) unifasciatus (La-
treille 1798), which then was parasitized by the slavemaking ants, E.
ravouxi or Chalepoxenus sp., but never by E. kraussei. Host speci-
ficity, is thus apparently a good character for species discrimination
in the genus Epimyrma, and the joint use of Temnothorax by E.v.,
E.f, and E.k. is an argument for their synonymization.

Population Data

Reproductive biology and colony foundation

Epimyrma species, as far as is known, may differ considerably
with respect to their sex ratios. Thus, E. ravouxi has a sex ratio of
about 1.5 (5/2); in E. kraussei from population no. 14 (Tignale) this
ratio is about 0.3 in field colonies; and 0.2 in laboratory culture
(Winter & Buschinger 1983), and in E. Corsica it is 0.08 (Buschinger
& Winter 1985). Sex ratios correspond well with the reproductive
biology of the species concerned: E. ravouxi is characterized by
extranidal mating, whereas E. kraussei (pop. no. 14 Tignale) and E.
Corsica mate inside the mother nests and thus continually inbreed.
The inseminated, dealate 22 of E.k. and E.c. remain in the mother

1986]

Buschinger et al. — Revision of Epimyrma

265

/\A

/v\

Fig. 5: Shape of petiolus and postpetiolus in Epimyrma $$and a,b,c:

kraussei from populations no. 1 (a), 12b (b), 14 (c); d: no. 19 ( E.foreli)\ e: no. 5a
vandeli). Usually the drawings of 10 specimens (b$: 5, e$: 3) were superimposed
following the method of Wehner 1983.

266

Psyche

[Vol. 93

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1986] Buschinger et al. — Revision of Epimyrma 267

nests over winter, and colony foundation through invading of a host
colony occurs in spring. E.r. young queens, on the other hand, begin
with colony foundation immediately after swarming, in late
summer.

In most of the populations of E.k., E.f and E.v., we found
evidence of a reproductive biology identical to that of E.k. pop. no
14 (Tignale), where we first have observed this kind of behavior
(Winter & Buschinger 1983). 3 of the 1 1 colonies of E.v., which were
collected on 31 March and 1st April, contained young Epimyrma-
22 still engaged with throttling the host colony queens. The £./.-
population, on the other hand, was studied in fall, October 3-10,
and most of the colonies contained dealate young 22, a few alate
ones, and some $$. Reproductive behavior, thus, is identical in
E.k., E.v. and E.f, with intranidal mating and colony foundation in
spring. So far as it could be checked, also the production of sexuals
and the sex ratios are quite similar (table 5), the sex ratios indicating
a generally high 2-bias.

Epimyrma worker-numbers

Slave-making ant species are characterized by the presence of a
comparatively high number of 22 in their nests, apart from
incipient colonies. In the genus Epimyrma, we found a considerable
variation of 2 numbers in different species, dependent upon their
respective type of parasitism. Thus, E. ravouxi, an active slave-
maker, has up to 77 £.-22 (mean 24.9) in a nest, whereas the
“degenerate slavemaker”, E. kraussei, had an average of only 3.5 and
a maximum of 10 £-22 (Buschinger & Winter 1983). £. Corsica
(Emery 1895) has lost the 2 -caste completely (Buschinger & Winter
1985). £ vandeli was originally said to be workerless, whereas 22
had been described of £ kraussei and £. foreli. We therefore
censused the £-22 m most of our field-collected colonies, and also
the 2 -production of a representative number of colonies in
laboratory culture.

In table 6 we compare the Epimyrma 2 -numbers of 4 larger
populations including 2 ascribed to £. kraussei (no 14 and 2a), and
the populations no 5a (E.v.) and no 19 (E.f), and of 5 local
populations of £. kraussei from the Spanish Pyrenees with nests
always found in close vicinity.

268 Psyche [Voi. 93

l \ 5 * a

$ if. * ' < * Hitt

Fig. 6: Karyotypes of a: Epimyrma kraussei from pop. no. 5b (Aude, S’France)

and b: Epimyrma ravouxi from pop. no. 12a (Lago di Como, N’ltaly).

Most striking is the fact, that our 1 1 field colonies of E. vandeli
did not contain any E.-$$. This corresponds to the original de-
scription of 6 colonies without 5$ (Vandel 1927). In laboratory
culture, however, we obtained a few $$ from colonies of this
population (see below).

Workerlessness is also found in a certain amount of colonies in
most populations of E. kraussei. In part, this is due to the fact the
newly founded colonies do not yet contain £.-$$, and most of our
collecting was done in spring during the time of colony foundation.
Therefore, it is not surprising that the population of E.foreli is the
only one where all colonies contained at least one E. $ : The sample
was entirely collected in the fall. On the contrary, our material from
the type locality of E. vandeli was collected in spring, and in 3 of the
11 colonies the E.-Q was still engaged in throttling the Temno-
thorax queen. A few more colonies may as well have been incipient
ones, where the host queen had already been eliminated. Further-
more, 3 colonies in the laboratory produced unusually high
amounts of E.-$$, and when dissected, the queens proved to be
poorly inseminated, having very few sperm cells in their receptacula.
The lack of £.-$$ in our sample is thus at least in part explained by
these facts.

The highly variable average and median values of $ -numbers as
well as the maximum values in other populations are also very
remarkable. In some populations, like that of E.foreli, but also at

1986]

Buschinger et al. — Revision of Epimyrma

269

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Psyche

[Vol. 93

Banyuls (no 2a) and along the Spanish Pyrenees (no 2c, 3b, 3d), we
found a few colonies with 15 to more than 20 £.-$$, which would
be sufficient for an effective slave-raiding. As was suggested for E.
kraussei from Tignale (Buschinger & Winter 1983), however, we
believe that slave-raids do occur only exceptionally, if at all, in the
other populations now studied: Most colonies comprise but very
few E.-QQ, and colonies with higher E.-Q -numbers on average do
not contain more host species workers than those with few or no

E. -QQ. From table 6 we may conclude that Epimyrma kraussei has
established numerous local populations in which the reduction of
$ -numbers has occurred to highly variable degrees. The population
ascribed to E. vandeli then would be close to one end of the scale
which is complete loss of the £ -caste like in E. Corsica (Buschinger
& Winter 1985), and E.foreli is among the populations with highest

F. -$ -numbers. It must be stated, however, that a geographical vari-
ation of 5 -numbers, e.g., in the sense of a cline, is lacking: Popula-
tions with low $ -numbers have been found in S ’France (no 5a, E.
vandeli) and in N’Spain (no 3a), and high $ -numbers occur close to
the latter locality (no 3b) as well as in S ’Italy (no 19, E. foreli).

In laboratory culture the $ -production of Epimyrma colonies
roughly corresponds to the field data. Table 7 provides a compari-
son of 5 -production in colonies from 3 populations. Most impor-
tant is the fact that appeared in 2 of the 5 laboratory-kept

colonies from population no 5a (Lauzerte, E. vandeli).

Worker numbers, thus, are not contradictory to a synonymiza-
tion of E.v. and E.f. with E. kraussei.

Crossbreeding experiments

Intranidal mating is an excellent condition for experimental
crossbreeding of sexuals from different populations and even spe-
cies. Colonies are kept in nearly natural annual cycles with a long
hibernation of about 6 months at 10° C a “spring” and “fall” phase
in daily temperature rhythms of 10°C (12h, dark) and 20° C (12h,
light) for 2 weeks each, and a summer phase of 15°C (lOh, dark) and
25° C (14h, light) for 2 weeks, followed by 2 months of 17°C (lOh,
dark) and 28° C ( 14h, light), and again 2 weeks of 1 5° C/ 25° C when
pupation decreases. For details of formicaries, feeding etc. see
Buschinger (1974). All $ pupae from colonies of 2 populations or
species are exchanged. Further $ pupae arising newly from the

1986]

Buschinger et al. — Revision of Epimyrma

271

Table 7. Worker-production in colonies of Epimyrma kraussei Emery 1915 from
3 populations, in the first summer after collecting (including the populations of
E.foreli Menozzi 1921 and E. vandeli Santschi 1927).

population

n colonies

total

Epimyrma $$ produced
mean median

range

no 14 Tignale (E.k.)

0.52

0- 2

no 19 Calabria (E.f.)

2.0

0-10

no 5a Lauzerte (E.v.)

1.0

0- 4

remaining brood are either removed or exchanged. Usually the
foreign pupae are easily accepted, and also the sexuals hatching
from them. After dealation of the young 2? a few of them are
dissected for control of insemination. In the following spring the 9$
leave the nest chambers and can be placed with host colonies, where
they found their own colonies. The first sexual offspring usually
develops from rapid brood in the year of colony foundation (Winter
& Buschinger 1983).

It must be said, however, that the rate of successful colony foun-
dations is generally low, both with cross-mated 2? and those having
normally mated with brothers, Quite often this is due to insufficient
insemination, and perhaps to not yet optimal laboratory conditions.
We therefore present only a preliminary survey of successful cross-
breedings (table 8) without giving data on numbers of replicates or
numbers of offspring produced. These experiments are being
continued.

Table 8 clearly reveals that crossbreeding between different E.k.
populations, and also between E.k. and E.v. or E.f, is possible. This
result, however, can only weakly support our supposition of the
synonymy of the 3 species, since we also succeeded in crossbreeding
E.k. with E. Corsica, and with E. bernardi, both of which are
morphologically and biologically distinct good and species.

Discussion and Conclusion

The meaning of the morphological and biological characters stud-
ied in E. kraussei, E. vandeli and E. foreli, has been discussed with
reference to the question of synonymy of the 3 species already in the
respective sections. We found no morphological characters which
would allow a clear distinction between them. The karyotype is
apparently homologous in all Epimyrma species. The 3 species

272

Psyche

[Vol. 93

investigated have a common host species, Leptothorax (T.) reced-
ens, which is not parasitized by any other Epimyrma species. The
numbers of Epimyrma-Q 5 are variable, but low in all the 3 species,
which therefore should represent “degenerate slavemakers” as was
already stated for E. kraussei (Buschinger & Winter 1983). Field
data and laboratory breeding results indicate that the 3 species have
a highly 9-biased sexual production, intranidal mating and in-
breeding, that the young 99 overwinter in their mother nests and
invade own host colonies in spring. Crossbreeding experiments
reveal that a strict genetical isolation is lacking. The 3 original sam-
ples, comprising only few specimens, were apparently described as
separate species mainly because they were found in quite distant
localities, and because the variability of their slight morphological
differences could not be evaluated then.

We therefore synonymize E. vandeli Santschi 1927 and E. foreli
Menozzi 1921 with E. kraussei Emery 1915.

Population structure and reproductive biology in this species,
however, are highly remarkable (Winter & Buschinger 1983). The
inbreeding system with young queens spreading on foot, and thus
over only short distances, must result in an extremely restricted gene
flow, even if a rare mating of sexuals from neighboring colonies
might occur. The populations from different continents (northern
Africa, southern Europe) and islands (Sardegna, Corsica), but also
from more neighboring localities (southern France, northern
Spain), must have been isolated for a very long time. This isolation,
in our opinion, is responsible for the differences in coloration,
morphology of wings and genitalia, and worker numbers, which we
observed in certain populations. The replacement of one of these
characters by another one can only occur through interdemic
selection, through supplantation of a local population by another
one which is somewhat more effective. Since E. kraussei, however,
does not inhabit large, continuous habitats, but instead forms
numerous small, patchily distributed populations, this process must
be slow and rare. The reduction of worker numbers in favor of a
higher 9 production should be highly adaptive in this species. Since,
however, the genetical basis for this evolution cannot spread, e.g.,
through flying we may speculate that different demes just have
reached different degrees of worker reduction. Crossbreeding ex-
periments have been started in order to find out whether or not

1986]

Buschinger et al.— Revision of Epimyrma

273

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274

Psyche

[Vol. 93

worker number in E. kraussei populations is genetically determined.
If so, we may predict that somewhere in the range of E. kraussei,
populations will be found with high $ -numbers, and still actively
slave-raiding, and other perhaps truly workerless demes. The
evolution from outbreeding and slave-raiding towards intranidal
mating and reduction of worker numbers and slave-making
behavior, is an apparently widespread trait in the genus Epimyrma.
Intranidal mating has been found also in E. bernardi and in E.
Corsica, two species which are morphologically clearly separated
from E. kraussei. Whereas E. bernardi “still” produces a consider-
able amount of $$, E. Corsica has lost this caste completely
(Buschinger & Winter 1985). Future studies will be necessary to find
out whether worker reduction in Epimyrma is developing in several
species or species groups independently, in parallel evolution, or
whether the species with different worker numbers form a series of
descent. The present study of E. kraussei evidently favors the first
alternative.

Summary

Epimyrma vandeli Santschi 1927 and E. foreli Menozzi 1921 are
junior synonyms of E. kraussei Emery 1915. A comparison was
made of the type specimens and of newly collected material from the
type localities of E.v. and E.f, and from numerous populations of
E.k.. No reliable morphological differences could be found, despite
a certain variation in $ genitalia, wing venation and body colora-
tion of different populations. Karyotypes are homologous in all
Epimyrma species and populations yet studied. The host species is
Leptothorax ( Temnothorax ) recedens (Nylander 1856) in all E.k.
populations including E.v. and E.f., whereas all other Epimyrma
species have different host species. Epimyrma $ -numbers vary
between populations, E.v. having a particularly low, and E.f. quite a
high one, both, however, remaining within the range of the other
E.k. populations. Sexual production is similar in all populations
with a remarkably low (^-production. In all populations studied,
sexuals mate within the mother nests, and inseminated, dealate
young 99 remain there over winter until they leave for colony foun-
dation in spring. E.v. and E.f. could be successfully crossbred with
E.k., and sexuals from several E.k. populations among each other.
Differences between E.k. populations presumably are due to their

1986] Buschinger et al. — Revision of Epimyrma 275

quasi-clonal structure with very restricted or lacking gene flow
between colonies and demes.

Acknowledgements

This work was supported by grants of the Deutsche Forschungs-
gemeinschaft.

References

Andr6, E. 1896: Description d’une nouvelle fourmi de France. - Bull. Soc. Ent

France 2: 367-368.

Bitsch, J. 1979: Morphologie abdominale des Insectes. In: Grasse, P.-P., Traite

de Zoologie, t. VIII, fasc. II, 290-578; Masson, Paris.

Buschinger, A. 1974: Experimente und Beobachtungen zur Griindung und Ent-

wicklung neuer Sozietaten der sklavenhaltenden Ameise Harpagoxenus sublae-
vis (Nyl.). - Ins. soc. 21: 381-406.

Buschinger, A. 1982: Epimyrma goesswaldi Menozzi 1931 = Epimyrma ravouxi

(Andre 1896) - Morphologischer und biologischer Nachweis der Synonymie
(Hym., Formicidae). - Zool. Anz. 208: 352-358.

Buschinger, A. & Winter, U. 1983: Population studies of the dulotic ant, Epi-

myrma ravouxi, and the degenerate slavemaker, E. kraussei (Hymenoptera:
Formicidae). - Entomol. Gener. 8: 251-266.

Buschinger, A. & Winter, U. 1985: Life history and male morphology of the

workerless parasitic ant Epimyrma Corsica (Hymenoptera: Formicidae). -
Entomol. Gener. 10: 65-75.

Buschinger, A., Winter, U. & Faber, W. 1983: The biology of Myrmoxenus

gordiagini Ruzsky, a slave-making ant (Hymenoptera, Formicidae). - Psyche 90:
335-342.

Cagniant, H. 1968: Du nouveau sur la repartition des Epimyrma d’Algerie

(Hymenopteres - Formicidae - Myrmicinae). - Bull. Soc. Hist. Nat. Toulouse
104: 427-429.

Emery, C. 1915: Contributo alia conoscenza delle formiche delle isole italiane.

-Ann. Mus. Civ. Stor. Nat. Genova, Ser. 3a, 46: 244-270.

Espadaler, X. 1983: Epimyrma bernardi n.sp., a new parasitic ant. - Spixiana 5:

1-6.

Hauschteck, E. 1962: Die Chromosomen einiger in der Schweiz vorkommender

Ameisenarten. - Vierteljahresschr. Naturf. Ges. Zurich 107: 213-220.

Hauschteck, E. 1965: Halbe haploide Chromosomenzahl im Hoden von Myr-

mica sulcinodis Nyl. (Formicidae). - Experientia 21: 323-325.

Imai, H. T., Crozier, R. H. & Taylor, R. W. 1977: Karyotype evolution in

Australian ants. - Chromosoma 59: 341-393.

Kutter, H. 1950. Gber zwei neue Ameisen. - Mitt. Schweiz. Ent. Ges. 23:
337-346.

Kutter, H. 1973: Beitrag zur Losung taxonomischer Probleme in der Gattung

Epimyrma (Hymenoptera Formicidae). Mitt. Schweiz. Ent. Ges. 46: 281-289.

Psyche

276

[Vol. 93

Menozzi, C. 1921: Formiche dei dintorni di Sambiase di Calabria. - Boll. Lab.

Zool. gener. Agr. Portici 15: 24-32.

Menozzi, C. 1931: Revisione del genere Epimyrma Em. (Hymen. Formicidae) e

descrizione di una specie inedita di questo genere. - Mem. Soc. Ent. Ital. 10:
36-53.

Sadil, J. 1953: Epimyrma zaleskyi nov. spec. (Hym., Formicoidea). - Casop.

Cesk. Spol. ent. 50: 188-196.

Santschi, F. 1927: Notes myrmecologiques. I. Sur quelques nouvelles fourmis de

France. - Bull. Soc. Ent. France 1927: 126-127.

Vandel, A. 1927: Observations sur les moeurs d’une Fourmi parasite: Epimyrma

vandeli Santschi. - Bull. Soc. Ent. France 1927: 289-295.

Wehner, R. 1983: Taxonomie, Funktionsmorphologie und Zoogeographie der

saharischen Wilstenameise Cataglyphis fortis (Forel 1902) stat. nov. (Insecta:
Hymenoptera: Formicidae). - Senckenbergiana biol. 64: 89-132.

Winter, U. 1979: Epimyrma goesswaldi Menozzi, eine sklavenhaltende Ameise.

-Naturwissenschaften 66: 581.

Winter, U. & Buschinger, A. 1983: The reproductive biology of a slavemaker

ant, Epimyrma ravouxi, and a degenerate slavemaker, E. kraussei (Hymenop-
tera: Formicidae). - Entomol. Gener. 9: 1-15.

MALE BIOLOGY IN THE QUEENLESS PONERINE ANT
OPHTHALMOPONE BERTHOUDI
(HYMENOPTERA: FORMICIDAE)

By Christian Peeters* and Robin Crewe

Department of Zoology, University of the Witwatersrand,
Johannesburg 2001, South Africa

Introduction

Various ponerine ants exhibit significant modifications in their
pattern of male dispersal, and this is associated with changes in the
queenright social structure. In some species the queen caste has
become permanently wingless (= ergatoid queens), and in others it
has been replaced by mated laying workers (= gamergates; Peeters
and Crewe, 1984). Thus, male nuptial flights take on new character-
istics since they have to locate flightless sexual partners. Data on
male behavior are only available for a few of the ponerine species
without a queen caste, but generally males disperse individually and
orientate to foreign nests, around which mating then occurs. Brown
(1953) observed low-flying males entering nests in two species of
Rhytidoponera. Mating can occur outside the nest entrances (e.g. in
R. chalybaea ; Ward, 1981), or inside the nest (e.g. in Diacamma
rugosum\ Wheeler and Chapman, 1922).

Ophthalmopone berthoudi Forel is permanently queenless, and
details of its reproductive system and polydomous organization
appear elsewhere (Peeters and Crewe, 1985, MS). This paper deals
with the pattern of male behavior in the field and the characteristics
of male production in a breeding system made up exclusively of
laying workers.

Methods

Colonies of Ophthalmopone berthoudi were studied in one local-
ity in Mkuzi Game Reserve (north-eastern Natal, South Africa),
during 1981-1983. Observations were made throughout the year,

♦Present address: School of Zoology, University of New South Wales, P.O. Box 1,
Kensington N.S.W., Australia 2033.

Manuscript received by the editor May 20, 1986.

277

278

Psyche

[Vol. 93

but the ones specifically reported in this paper were made during the
period of male activity (January-April). Male behavior was usually
studied near colonies under intensive study (Peeters, 1984). In such
colonies the location of all the nest entrances was known (colonies
are polydomous), and all the workers active outside the nests had
been color-marked with individual codes. In addition, a few males
were marked on the thorax and then released. The presence of males
was determined by observation of their activity outside nests and by
examination of the contents of excavated nests.

Results

Dates of male activity

Normal winged males are produced in this species and were found
inside most nests excavated during January-April (Table 1). This
limited period of male production was confirmed by finding male
pupae during January-April only. A subjective impression is that
the number of males present above ground reached a peak in Feb-
ruary. Excavations also revealed that males are present in every
nest of a colony. However, nests collected in the same month could
contain different numbers of males (Table 1). During January and
February, a few males were seen to be carried between the nests of a
colony. This carrying did not follow any organized pattern, and
occurred together with the recruitment of workers and brood. Many
of the cocoons that were transferred between nests during that
period contained male pupae (A sample of cocoons then found in
the nests yielded 70 male pupae and 248 worker pupae).

The investment in male production does not appear exceptional
in this queenless species; a colony (464 workers) with five nests
excavated in February 1982 yielded 60 males (Table 1), and this is in
addition to those that had already departed as well as pupae.

Dispersal behavior

Every day during a three-week observation period in January-
February 1982, a few males (1-8) left from each of six nests under
intensive observation. Departing males left the nests, often
climbed up low vegetation and flew off. Once on the wing, they
could no longer be followed. The time of departure (9H00 to 12H00)
often coincided with the period when workers were no longer active
on the surface because of high soil temperatures. Male exit times
appeared not to be affected by cloudy or cooler weather.

1986] Peeters & Crewe — Male biology in Ophthalmopone 279

Table 1. Size of the male population in nests excavated during January-April.
Males were not present at other times of the year.

Date of
excavation

Number of nests Number of males found in each nest
excavated (together with number of adult workers)

January 1981

4 tt

5(145), 0(84), 6(142), 10(227)

January 1982

1 7(> 1 40)

January 1983

3 ft

3(116), 9(222), 6(121)

February 1982

5 +

7(20), 9(80), 19(168), 13(72), 12(124)

March 1982

2 +

0(119), 0(77)

April 1981

3 +

0(318), 0(75), 1(445)

April 1983

2 +

0(106), 2(121)

# from 2 colonies
+ from same colony

Evidence that males remain in their natal nests until they are
physiologically ready to mate was adduced from the following
observations. An adult male that was painted while being carried
between two nests, remained inside the second nest for nine days
before it left and flew off. Dissection of males collected during
excavations revealed that there was little or no sperm in the vasa
deferentia and ejaculatory ducts of many of them.

After the initial dispersal flight, males alighted on the ground and
appeared to search for nests haphazardly. They walked quickly with
frequent changes of direction, and investigated little holes and
depressions in the soil. They regularly climbed up short grass stems
or low vegetation from which they flew off, often for only a short
distance. This behavior was interspersed with ground searches. On a
number of occasions, males were observed either landing very close
to nest entrances, or walking straight towards occupied nests shortly
after landing. Five marked males were observed outside one nest on
two successive days, indicating that after locating a foreign nest,

Behavior around the entrances of foreign nests

During the period of their activity, males were observed waiting
immobile outside nest entrances, either on the ground or on top of
short grass stems. The working assumption was that such males did
not originate from these nests, because they always flew away from
their natal nests. Some nests frequently had many males in their
vicinity, while other nearby nests seldom had any around them.

Males usually investigated entrance holes with their antennae and
hesitantly walked in; some ran out immediately afterwards. Individ-
ual males were repeatedly evicted from nests by workers (in nests

280

Psyche

[Vol. 93

under intensive study, these were often marked workers, i.e. active
on the surface). Males were held by their legs, wings or antennae,
and resisted fiercely; some managed to struggle free. After releasing
the males in the vicinity of the nests (30 cm -1 m away), the workers
ran back into the entrance holes. The uninjured males cleaned their
antennae and then immediately attempted to enter the nests again.
On some occasions a number of workers cooperated in the eviction
of foreign males, and some workers also chased males when they
came across them outside the nests. Eviction did not always follow
a male’s entrance, and some marked males remained underground
for at least 15 minutes.

Discussion

In Ophthalmopone berthoudi copulation was never observed
above ground, and it is inferred that it occurs exclusively inside
foreign conspecific nests. This is an unusual situation in ants, who
usually mate some distance from the nests. However, copulation can
take place in the immediate vicinity of nests in queenright and
queenless ponerines, and in socially parasitic myrmicines (e.g. Har-
pagoxenus ; Buschinger and Alio way, 1979). In Rhytidoponera
chalybaea, in which colonies have either a queen or gamergates,
large numbers of workers and males mill around nest entrances,
and males make repeated attempts to mate with workers (Ward,
1981). However, males also enter nests and may mate with workers
there. In the queenless R. metallica, workers attract males by the
release of a pygidial gland pheromone; this distinct behavior (‘sex-
ual calling’) occurs outside the nest entrances (Holldobler and Has-
kins, 1977). The pygidial gland has been found in O. berthoudi
(Villet et al., 1984), and we speculate that if young workers release
this sex pheromone, they only do so inside the nests and hence
encounter males underground. Sexual calling was never observed inf
the field or in the laboratory.

Direct data are not available on the activities of males inside
foreign nests, and the occurrence of mating is inferred from the large
proportion of inseminated workers in nests excavated after the
period of male activity (Peeters and Crewe, 1985). The existence of
many gamergates in some nests (up to 108) strongly suggests that
males copulate more than once; otherwise, such nests would need to
be visited by larger numbers of males than we observed entering any

1986] Peeters & Crewe — Male biology in Ophthalmopone 281

nest. The substantial variations in the percentages of gamergates
present in different nests at any one time of the year (Peeters and
Crewe, 1985) suggest that the number of male visits to a particular
nest is irregular. Some nests may be located more often than others,
and consequently varying numbers of young workers become
mated. In polydomous colonies such as these, gamergates can be
transferred between nests and, hence, a colony should survive from
year to year as long as one of its nests is visited by males.

The exit of males from their natal nests is not coordinated, and
they disperse over a period of a few weeks. This is different to the
situation in queenright species where the emergence of all the male
and female reproductives is synchronized in time (e.g. in Campono-
tus herculeanus, through the release of a mandibular gland phero-
mone by the males; see Holldobler and Bartz, 1985). Dispersal is
then often associated with the initiation of new nests, which must
occur during optimal environmental conditions (e.g. after rain). In
contrast, copulation in O. berthoudi is not followed by independent
colony foundation by the mated workers, because colonies repro-
duce by fission (Peeters, 1984). Thus it is no longer selectively
advantageous for males to disperse simultaneously in response to a
specific environmental cue. However, males continue to be pro-
duced only during a short period of the year. Unmated workers
show no ovarian activity in O. berthoudi, and haploid eggs are laid
exclusively by gamergates (Peeters and Crewe, 1985). Egg fertiliza-
tion is thus a voluntary act by the mated workers, and males are
produced following the first summer rains. Sperm exhaustion is
unlikely since individual gamergates lay relatively few eggs during
their lifetime. It is not known whether all the gamergates in a nest
produce haploid eggs; the inter-nest transfer of male adults and
pupae would make this hard to determine.

The importance of chemical attractants during nest location
remains unclear. In Leptogenys ocellifera, a ponerine with ergatoid
queens, dispersing males search for the chemical trails that lead
from the nests into the surroundings (Maschwitz and Muhlenberg,
1975), and males of Megaponera foe tens follow trails laid by
workers during raids on termite nests (Longhurst and Howse, 1979).
This is impossible in O. berthoudi because continuous trails are not
laid. There is evidence that discrete scent marks are deposited on the
substrate by inexperienced foragers (Peeters and Crewe, MS), but
this may be of no use to males. It is conceivable that the pygidial

282

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[Vol. 93

gland secretions also work as a long-distance attractant. In addition
to signalling sexual receptiveness to the males inside the nests, these
volatile secretions (which are produced by many workers) may dif-
fuse out of the nests and be perceived by searching males.

Males of O. berthoudi need to enter foreign nests in order to find
sexual partners. The colony units have distinct identities (Peeters,
1984), and alien males are recognized as different by workers, which
then attempt to remove them from the nest; similar hostility is also
displayed in R. chalybaea (Ward, 1981). This aggression contrasts
with the acceptance of alien males by workers in ponerine species
with ergatoid queens, e.g. males in Leptogenys and Megaponera
were not attacked following their entry into foreign colonies
(Wheeler, 1900; Longhurst and Howse, 1979). In the queenless
Dinoponera gigantea, Overal (1980) observed a male being carried
into a nest by a forager. Carrying of males in O. berthoudi was
always between the nests of a single polydomous colony and is thus
not equivalent to the observations made by Overal. Access by males
to foreign nests may be facilitated by the fact that the older workers
that perform activities on the surface and are responsible for the
evictions, are usually not active during the daily peaks of male
activity. The younger workers confined inside the nests are those
likely to become mated (Peeters and Crewe, 1985), and these
probably do not behave aggressively towards foreign males.

If the queenright ancestors of this species exhibited the typical
formicid pattern of reproduction, then male and female reproduc-
tives would have been produced seasonally. With the change to
worker reproduction, the sexually-attractive workers do not dis-
perse from their nests prior to mating, and mating is no longer
coupled with colony foundation, hence the times of male activity no
longer need to be synchronized with female activity periods or with
appropriate environmental conditions for colony foundation. This
relaxation of the selective pressures on the timing of male dispersal
has resulted in an extended mating period. Nonetheless, male activ-
ity remains seasonal. This has no adaptive significance in O. ber-
thoudi, because young workers that can be mated occur throughout
the year. However it has the effect of ensuring that an adequate
number of infertile workers are present in the colonies.

1986] Peeters & Crewe — Male biology in Ophthalmopone 283
Acknowledgments

We are grateful to R. H. Crozier for comments on the manu-
script. We thank the Natal Parks Board for permission to work in
Mkuzi Reserve, and Peter Goodman for his hospitality. This work
was supported by grants from the University of the Witwatersrand
and the Council for Scientific and Industrial Research.

References

Brown, W. L.

1953. Characters and synonymies among the genera of ants. Part I. Breviora,
Mus. Comp. Zool., 11: 1-13.

Buschinger, A. and T. M. Alloway

1979. Sexual behaviour in the slave-making ant, Harpagoxenus canadensis
M. R. Smith, and sexual pheromone experiments with H. canadensis, H.
americanus (Emery), and H. sublaevis (Nylander) (Hymenoptera; For-
micidae). Z. Tierpsychol., 49: 113-119.

HOlldobler, B. and C. P. Haskins

1977. Sexual calling behavior in primitive ants. Science, 195: 793-794.

HOlldobler, B. and S. H. Bartz

1985. Sociobiology of reproduction in ants. In: Experimental Behavioral
Ecology and Sociobiology (eds. B. Holldobler, M. Lindauer) Gustav
Fischer Verlag, Stuttgart pp. 237-257.

Longhurst, C. and P. Howse

1979. Some aspects of the biology of the males of Megaponera foetens.
Insectes Soc., 26: 85-91.

Maschwitz, U. and M. Muhlenberg

1975. Zur Jagdstrategie einiger orientalischer Leptogenys- Arten (Formicidae:
Ponerinae). Oecol., 20, 65-83.

OVERAL, W. L.

1980. Observations on colony founding and migration of Dinoponera gigan-
tea. J. Ga Entomol. Soc., 15: 466-469.

Peeters, C. P.

1984. Social organization, breeding biology and the process of reproductive
differentiation in Ophthalmopone berthoudi Forel, a ponerine ant.
Unpublished Ph.D. Thesis, University of the Witwatersrand Johannes-
burg, South Africa.

Peeters, C. and R. Crewe

1984. Insemination controls the reproductive division of labour in a ponerine
ant. Naturwiss., 71: 50-51.

1985. Worker reproduction in the ponerine ant Ophthalmopone berthoudi: an
alternative form of eusocial organization. Behav. Ecol. Sociobiol., 18:
29-37.

284

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[Vol. 93

MS. Foraging and recruitment in ponerine ants: solitary hunting in the
queenless Ophthalmopone berthoudi (Hymenoptera: Formicidae).
Submitted.

VlLLET, M., C. PEETERS AND R. CREWE

1984. The occurrence of a pygidial gland in four genera of ponerine ants
(Hymenoptera: Formicidae). J. Georgia Entomol. Soc., 19: 413-416.

Ward, P. S.

1981. Ecology and life history of the Rhytidoponera impressa group (Hyme-
noptera: Formicidae) II. Colony origin, seasonal cycles, and reproduc-
tion. Psyche, 88: 109-126.

Wheeler, W. M.

1900. A study of some Texan Ponerinae. Biol. Bull., 2: 1-31.

Wheeler, W. and J. Chapman

1922. The mating of Diacamma. Psyche, 29: 203-211.

NEARCTIC SPECIES OF THE NEW WOLF SPIDER
GENUS GLADICOSA (ARANEAE: LYCOSIDAE)*

By Allen R. Brady

Department of Biology, Hope College
Holland, Michigan 49423

This is the second paper in a projected series of systematic studies
of the Nearctic Lycosidae described primarily in the genus Lycosa.
Over 50 species of medium to large size wolf spiders from the Nearc-
tic Region have been placed in this genus. However, recent studies
indicate that several distinct genera are included under Lycosa.
Matters have been complicated at the generic level by C. F. Roewer
(1954) who listed 44 new genera of Lycosinae in the Katalog der
Araneae. They are nomina nuda, lacking descriptions. Later Roewer
(1959, 1960) defined these 44 genera, thus validating the names, and
added seven more new ones to the Lycosinae as well. These genera
were established primarily on the basis of differences in the number
of posterior cheliceral teeth and eye arrangement (particularly eyes
of the anterior row). Investigations of North American Lycosidae
(Brady 1962, 1972, 1979) indicate that the number of posterior che-
liceral teeth is an unreliable character in delimiting genera. Recent
studies indicate that color patterns on the dorsal surface of the
carapace, length of legs relative to body size, and particularly the
structure of the male and female genitalia are most reliable in
determining generic relationships. Certain features of the eye
arrangement, as well as information about habitat, behavior, and
life history are also useful. In the final analysis, it is the unique
combination of all these features that should be employed to
distinguish genera.

Gladicosa gen. nov.

Lycosa (part) Walckenaer 1837: 338. Emerton 1885: 485. Marx 1890: 562; 1892: 160.
Stone 1890: 423, 426. Montgomery 1902: 538, 546, 566; 1904: 277-280; 1905:
174; 1909: 514. Banks 1901: 184; 1910: 55, 57; 1911: 454. Chamberlin 1904: 147;
1908: 225, 226, 265; 1924: 28. Petrunkevitch 1911: 560. Comstock 1913: 631, 639;

* Manuscript received by the editor July 15, 1986

285

286

Psyche

[Vol. 93

1940: 644, 650. Bishop and Crosby 1926: 207. Wood 1926: 174. Crosby and
Bishop 1928: 1067. Elliott 1930: 5; 1932: 423. Worley and Pickwell 1931: 91, 93.
Chickering 1932: 351. Gertsch 1934: 7, 8; 1949: 82. Gertsch and Wallace 1935:
20-22; 1937: 10. Kaston 1935: 191; 1936: 103, 114; 1938: 184; 1948: 322, 328;
1981: 322, 328. Allard 1936: 67. Jones 1936: 69. Chamberlin and Ivie 1944: 142,
144. Bonnet 1957: 2607, 2635, 2645. Fitch 1963: 108-109. Whitcomb, Exline,
Hunter 1963: 656. Whitcomb and Bell 1964: 45. Dorris 1965: 408; 1968: 36.
Drew 1967: 194. Harrison 1969: 14-16. Bultman, Uetz, Brady 1982: 26.
Leimonia (part) Simon 1864: 352.

Trochosa (part) Montgomery 1904: 301, 305. Chamberlin and Ivie 1942: 35.

Avicosa (part) Roewer 1954: 236.

Hogna (part) Roewer 1954: 258.

Scaptocosa (part) Roewer 1954: 293.

Varacosa (part) Roewer 1954: 306.

Alopecosa (part) Bonnet 1955: 248.

Type species. Gladicosa gulosa (Walckenaer)

Etymology. The generic name is a combination of gladius
(Latin for sword) referring to the unique sword-shaped embolus of
the male palpus, and cosa derived from the generic name Lycosa. It
is considered feminine.

Diagnosis. Gladicosa may be distinguished from other lycosid
genera by the following combination of characters: (1) the swordlike
or bladelike form of the embolus (em) and its clockwise orientation
in ventral view of the left palpus of the male (Fig. 33), (2) the modi-
fication of the terminal apophysis (ta), which is also broadly flattened
and parallels (and partly supports) the embolus (Figs. 33, 34), (3) the
rectangular or wedge shape of the transverse piece (tp) of the scape
of the epigynum, together with its white pearlescent appearance, in
whole or part (Fig. 10) and (4) the dorsal color pattern illustrated in
Figures 1-5 and described below.

Description. Total length 7.8 to 18.8 mm. Carapace length 4.2
to 8.3 mm; width 3. 1 to 6.4 mm. Carapace viewed dorsally, narrow-
ing at level of PLE row, smoothly convex along lateral margins,
with posterior margin concave; viewed laterally essentially the same
height from eye region to posterior declivity (highest point is poste-
rior cephalic region in front of dorsal groove with the carapace
sloping very slightly anteriorly). Dorsal groove long and distinct.
Dorsal color pattern with light uneven submarginal stripes and wide
median light colored stripe, narrow between ALE, widening until
just anterior to dorsal groove (where it is usually constricted),
becoming wider again parallel to groove, and then narrowing as it

1986]

Brady — Ne arc tic Gladicosa

287

follows thoracic declivity to posterior edge of carapace. Black mark-
ings framing median stripe at posterior declivity. Dark areas of
carapace brown to dark brown and black. Light stripes pale yellow
to yellow-orange (Figs. 1-5).

Anterior median eyes (AME) slightly larger than anterior lateral
eyes (ALE). Anterior eye row much narrower than posterior median
eye row (PME), with dorsal tangent slightly procurved. Posterior
lateral eye row (PLE) much the widest (see Tables 1-6).

Chelicerae dark reddish brown to black; anterior and posterior
margin each with three teeth, the anterior triad crowded more
closely together.

Legs when compared to body dimensions relatively longer than in
Trochosa; without distinct annulations; yellow, yellow-orange to
golden brown in color. Order of leg length IV-I-II-III. Tibial spina-
tion in female: leg I, 2-2-2 ventral, 1-0 or 1-1 prolateral; leg II 2-2-2
ventral, 1-1 prolateral; leg III 2-2-2 ventral, 1-1 prolateral, 1-1 retro-
lateral, 1-1 dorsal; leg IV 2-2-2 ventral, 1-1 prolateral, 1-1 retrolat-
eral, 1-1 dorsal. Tibial spination in the male is the same with the
addition on leg I of 1-1 retrolateral and leg II 1-1 retrolateral.

Dorsal abdominal pattern variable according to size and hirsute-
ness, but generally with anterio-lateral black markings aligned with
those on carapace, cardiac area well marked, and often with pattern
of chevrons as indicated in Figures 1-5. Dark colors on dorsum of
abdomen brown to black, lighter colors cream to tan or beige. Ven-
ter of abdomen cream to light brown in gulosa, huberti, and euepi-
gynata; dark brown to black in pulchra and bellamyi. Region
anterior to epigastric furrow of contrasting darker or lighter color
respectively.

Male palpus with stridulatory file situated retrolaterally at tip of
tibia. Cymbium with cluster of macrosetae at tip, and with stridula-
tory scraper retrolaterally at base. Male palpal sclerites as seen in
ventral view: Palea (pa) concave, largely hidden by embolus, visible
along retrolateral margin. Embolus (em) blade-like, tapering to a
point, with clockwise orientation (from left to right) in left palpus,
which is opposite to that of most Lycosinae. Conductor (co) con-
cave, with cuplike portion containing tips of the terminal apophysis
(ta) and the embolus. Terminal apophysis large, flattened and paral-
leling embolus, with its tip serving partly as a conductor. Median
apophysis (ma) with a flattened ridge extending retrolaterally and

Psyche

288

[Vol. 93

coming to a point near margin of cymbium (cy); heavily sclerotized
spur directed medially (Figs. 30, 33, 34).

Epigynum of female with scape shovel-shaped with elongate lon-
gitudinal piece (lp) (handle) and rectangular or trapezoidal trans-
verse piece (tp) (blade). The transverse piece is unusual in being
wholly or partly translucent white or pearlescent in appearance (Fig.
10). Spermathecae (s) smooth and round to ovoid (Fig. 7), rarely
elongate ovoid (Fig. 15); usually their diameter apart.

Methods

The techniques and methods employed in the study of Gladicosa
were essentially the same as for Trochosa (Brady 1979) and are
described there. Color descriptions are based upon appearance of
specimens in alcohol illuminated by microscope lamp. Measure-
ments are listed in millimeters, but for Gladicosa the mean and
standard error (SEM) are listed instead of the mean and range as in
the previous paper. Methods and techniques of measurement are
described in the paper on Trochosa (Brady 1979). Under Records
specific localities are given for uncommon species and the peripheral
range for common species, otherwise localities of specimens exam-
ined are indicated by counties.

Acknowledgments

This study was made possible by the loan of large numbers of
specimens from the Museum of Comparative Zoology, Cambridge,
Massachusetts, the American Museum of Natural History, New
York City, and the Canadian National Collection, Ottawa, Canada.

I wish to thank sincerely the curators of those collections, Dr. H. W.
Levi, Dr. N. J. Platnick, and Dr. C. D. Dondale respectively for the
use of these materials. The loan of type specimens from the Museum
of Comparative Zoology, the American Museum and the Phila-
delphia Academy of Natural History is gratefully acknowledged.
Thanks are offered to Mr. Donald Azum for loan of the latter.

I am indebted to the following individuals and institutions for
making available regional collections that provided a much better
picture of geographical distribution and clarified the relationships
of certain populations: Dr. Richard Brown and Ms. Pat Miller of
the Entomological Museum, Mississippi State University; Mr. Tim

1986]

Brady — N ear c tic Gladicosa

289

Lockley, Deta State Research Center, USDA, Stoneville, Missis-
sippi; Dr. G. B. Edwards and Dr. H. K. Wallace of the University of
Florida, Gainesville; and Dr. Andrew Penniman, Defiance College,
Defiance, Ohio.

Special thanks are extended to Dr. C. D. Dondale and Dr. H. W.
Levi who consented to review the manuscript and offered construc-
tive criticism and friendly advice. I am also grateful to Mr. F. R.
Wanless for sending specimens of Lycosa pulchra Keyserling from
the L. Koch collection maintained in the British Museum (Natural
History). A note of special appreciation to Ms. Amy Youatt, who
helped with general sorting, compilation of locality data, and prep-
aration of distribution maps.

National Science Foundation grant DEB-7803561 assisted in
defraying expenses of the investigation. A summer grant from the
faculty development program at Hope College (1980) helped to
initiate this project.

Key to Females

la Transverse piece (tp) of scape of epigynum rectangular, about

equal in length and width (Figs. 6-14) 2

lb Transverse piece (tp) of scape of epigynum irregular in shape
(Figs. 15-17) or, if rectangular, much wider than long

(Figs. 18-26) 3

2a Transverse piece entirely pearlescent in appearance. Longitudi-
nal piece (lp) lacking indentations where it joins transverse

piece (Figs. 6-9) gulosa

2b Transverse piece only partly pearlescent white. Longitudinal
piece (lp) with indentations at posterior end where it joins

transverse piece (Figs. 10-14) pulchra

3a Transverse piece irregular in shape and broadly joined by longi-
tudinal piece (Figs. 15-17) euepigynata

3b Transverse piece somewhat rectangular, much wider than long

and narrowly joined by longitudinal piece 4

4a Width of transverse piece greater than length of longitudinal
piece. Longitudinal piece about the same width throughout

its length (Figs. 18-20) huberti

4b Width of transverse piece equal to or less than length of trans-
verse piece. Longitudinal piece wider anteriorly, narrowing

posteriorly (Figs. 21-26) bellamyi

290

Psyche

Key to Males

[Vol. 93

la Both embolus (em) and terminal apophysis (ta) bladelike,
paralleling one another with each separate and drawn out to

a point (Figs. 27, 28, 35-42) 3

lb Embolus bladelike, but terminal apophysis not resembling it;
the two not as distinctly separated as above (Figs. 29-34,

43-46) 2

2a Relatively small species. Total length 7.8 to 11.0 mm (Figs.

29-34). Not reported from central Texas bellamyi

2b Relatively large species. Total length 10.4 to 13.9 mm (Figs.

43-46). Distribution central Texas euepigynata

3a Embolus with relatively short, pointed tip (Figs. 27, 28)

. huberti

3b Embolus with longer drawn out tip that is curved at end 4

4a Tip of embolus pointed; median apophysis (ma) with large ret-

rolateral spur (Figs. 35-36) gulosa

4b Tip of embolus flattened; median apophysis (ma) with small
retrolateral spur (Figs. 37-42) pulchra

Gladicosa gulosa (Walckenaer), comb. nov.

Figures 5, 6-9, 35, 36. Map 1.

Lycosa gulosa Walckenaer, 1837: 338. Male holotype from North America, de-
stroyed. Marx 1890: 562. Chamberlin 1908: 225, 226, 265, pi. 21, figs. 4, 7, S $.
Montgomery 1909: 514. Petrunkevitch 1911: 560. Comstock 1913: 631, 639, figs.
720 g-h, 9 S', 1940: 644, 650, figs. 720 g-h, 9 S- Bishop and Crosby 1926: 207.
Wood 1926: 174. Crosby and Bishop 1928: 1067. Elliott 1930: 5; 1932: 423.
Worley and Pickwell 1931: 91, 93. Chickering 1932: 351. Gertsch 1934: 7; 1949:
82. Gertsch and Wallace 1935: 20. Kaston 1935: 191; 1936: 103, 114; 1938: 184;
1948: 322, 328, pi. 57, figs. 1 106-1109, 9 S', 1981: 322, 328, figs. 1 106-1109, 9 S-
Allard 1936: 67. Fitch 1963: 108-109, fig. 46. Whitcomb, Exline, Hunter 1963:
656. Whitcomb and Bell 1964: 45. Dorris 1965: 408; 1968: 36. Drew 1967: 194.
Harrison 1969: 14-16. Bultman, Uetz, Brady 1982: 26.

Leimonia gulosa: Simon 1 864: 352.

Lycosa kochi: Emerton 1885: 485, pi. 46, figs. 6-6c, 9 S', 1902: 74, figs. 179, 180, 9-
Stone 1890: 423, 426, pi. 15, fig. 3. Marx 1892: 160. Gertsch and Wallace 1935:
21, figs. 39, 42, S $• Not Lycosa kochi Keyserling.

Lycosa helluo: Banks 1901: 184 (part).

Lycosa nigraurata Montgomery, 1902: 538, 546, pi. 30, fig. 53, S ■ Male holotype
from Medford, Burlington Co., New Jersey (N.J. Stone), examined. Synony-
mized with Lycosa purcelli Montgomery by Montgomery 1904: 305.

Lycosa purcelli Montgomery, 1902: 538, 566, pi. 30, figs. 30, 31, 9 S • Female
syntype from Philadelphia, Philadelphia Co., Pennsylvania, May, 1888, and

1986] Brady — Nearctic Gladicosa 291

male syntype from Point Pleasant, Ocean Co., New Jersey, 30 April 1889 (N.J.

Stone), examined. Synonymized with Lycosa kochi: Emerton by Gertsch and

Wallace 1935: 21.

Trochosa purcelli, Montgomery, 1904: 301, 305.

Lycosa pulchra: Chamberlin 1904: 147 (part); Banks 1910: 57 (part).

Varacosa gulosa: Roewer 1954: 306.

Alopecosa gulosa: Bonnet 1955: 248.

Discussion. The nomenclatural history of G. gulosa is complex.
Walckenaer’s (1837) seven-line description without figures is not
diagnostic for this species. The locality given is North America, and
that doesn’t help. To complicate matters, Emerton (1885) misidenti-
fied this species as Tarentula kochi Keyserling and transferred it to
the genus Lycosa. Gertsch and Wallace (1935) discussed the syste-
matic and nomenclatural problems associated with G. gulosa and
suggested using the name Lycosa kochi Emerton for this species
since Emerton (1885) had placed the species in a different genus.
However, according to Article 49 of the International Code of Zoo-
logical Nomenclature (1985): “A previously established species-
group name wrongly used to denote a species-group taxon because
of misidentification cannot be used for that taxon even if it and the
taxon to which the name correctly applies are in, or are later
assigned to, different genera, except when a previous misidentifca-
tion is deliberately used in fixing the type species of a new nominal
genus.” Bonnet (1955) points out that the name nigraurata or pure-
celli of Montgomery should have been used for the species. Mont-
gomery (1904) himself synonymized nigraurata with purcelli and
the name purcelli has been used only by Montgomery (1902, 1904).
The name gulosa, on the other hand, has been employed numerous
times since Gertsch and Wallace’s (1935) invocation of kochi, and
even by Gertsch (1949) in his book American Spiders. It therefore
seems best to retain the name gulosa for this species to promote
stability of nomenclature by preserving a long accepted name in its
accustomed meaning.

Color. Females. Face yellow or yellow-orange, to pale golden
brown. Eye region darker with nacelles black. Chelicerae yellowish
brown to dark reddish brown, almost black at distal ends. Condyles
yellow or orange, to golden brown.

Carapace light brown to brown, with broad yellow to yellow-
orange median stripe. Narrow irregular submarginal yellow stripes
suffused with brown. Posterior declivity with black patches as in
Figure 3.

292

Psyche

[Vol. 93

Fig. 1 . Gladicosa huberti (Chamberlin), female from Bar M Ranch near Boston,
Thomas Co., Georgia, 2 Mar. 1973. Fig. 2. Gladicosa bellamyi (Gertsch and Wal-
lace), female from 2 mi. N of Stoneville, Washington Co., Mississippi, 9-11 May
1983. Fig. 3. Gladicosa pulchra (Keyserling), female from Gainesville, Alachua
Co., Florida, 14 June 1935. Fig. 4. Gladicosa gulosa (Walckenaer), female from 4
mi. S of New Richmond, Allegan Co., Michigan, 16 Sept. 1974. Fig. 5. Gladicosa
euepigynata (Montgomery), Camp Verde, Kerr Co., Texas, Dec. 1939.

1986] Brady — Nearctic Gladicosa 293

Dorsum of abdomen light brown to brown with pair of black
anterior-lateral patches as in Figure 5. Anterior cream to yellow
spots mark depressions of internal muscle attachments. Cardiac
area faintly indicated. Venter of abdomen cream or light beige to
pale yellowish brown. Few scattered darker spots. Overlaid with
fine coat of white hair.

Legs yellow or pale yellow-orange to yellowish brown, darker
distally. Femora with dusky bands on dorsal and lateral surfaces.
Ventral surface lighter yellow.

Labium and endites brownish orange to brown with distal ends
yellow to cream. Sternum yellow to light golden brown.

Color. Males. Face yellow to yellow-orange, darker brownish
in eye region. Chelicerae with basal areas yellow to orange-yellow,
darker brown to reddish brown distally. Condyles orange-yellow to
orange. Cymbia of palpi dark brown.

Carapace brown with a broad median yellow stripe and irregular
yellowish submarginal stripes obscured by thicker clothing of white
hair.

Dorsum of abdomen beige to light brown with black markings
along sides beginning anteriorly and continuing posteriorly. Black
markings often more prominent than in female. Posterior of dorsum
without distinct chevrons as in other species. Venter of abdomen
pale yellow to beige, clothed with white hair which is more abun-
dant laterally.

Legs yellow to brownish yellow. Darker dorsally without dusky
markings on femora as in female.

Labium and endites orange-yellow to orange-brown with distal
ends lighter yellow to beige. Sternum orange to orange-brown.

Measurements. Ten females and ten males from Allegan Co.,
Michigan. See Table 1.

Diagnosis. Gladicosa gulosa is closest to G. pulehra in size and
coloration. The markings of pulehra offer greater contrast, and
chevrons are usually visible on the dorsum of the female abdomen
(compare Fig. 5 with Fig. 4). The epigyna of the females and the
palpi of the males also resemble one another in appearance, but are
distinctly different when compared in detail. The epigynum of
gulosa has the transverse piece entirely pearlescent white, whereas
pulehra has some white, but nearly always shows darker brown
sclerotized areas on the transverse piece (compare Figs. 6, 8, 9 with
Figs. 10, 11, 13, 14). In gulosa the embolus is pointed at the end,
whereas that of pulehra is somewhat spatulate in shape (compare
Figs. 35, 36 with Figs. 37, 38).

294

Psyche

[Vol. 93

Table 1. Measurements of ten females and ten males of Gladicosa gulosa from
Allegan Co., Michigan.

Females:

Mean SEM

Ant. Eye Row

.985 ± .023

Femur I

4.26 ± .06

PME

1.218 ± .016

Pat. -Tibia I

5.50 ± .09

PLE

1.623 ± .020

Meta. I

3.13 + .05

POQ

1.138 ± .015

Tarsus I

1.86 + .03

Car. Width

4.36 ± .08

Total I

14.74 + .22

Car. Length

5.88 ± .09

Femur IV

4.92 ± .08

Body Length

13.18 +.49

Pat. -Tibia IV

5.77 ± .08

Pat. -Tibia II

4.96 +.09

Meta. IV

5.40 ± .07

Pat. -Tibia III

4.37 +.08

Tarsus IV

2.34 ± .02

Total IV

18.44 + .24

Males:

Mean SEM

Ant. Eye Row

.900 ± .025

Femur I

4.13 ± .06

PME

1.141 ± .021

Pat. -Tibia

5.46 ± .09

PLE

1.503 ± .028

Meta. I

3.46 ± .06

POQ

1.049+ .018

Tarsus I

1.89 ± .03

Car. Width

4.14 +.06

Total I

14.93 ± .23

Car. Length

5.50 +.12

Femur IV

4.63 ± .08

Body Length

11.46 +.30

Pat. -Tibia IV

5.50+ .10

Pat. -Tibia II

4.79 +.08

Meta. IV

5.27 ± .08

Pat. -Tibia III

4.18 +.07

Tarsus IV

2.33 ± .05

Total IV »

17.73 ± .30

Natural History. Kaston (1948) reports gulosa running over
dead leaves on forest floors in Connecticut. I have found it in leaf
litter of deciduous woods in Michigan. Here it is found in more
open Oak woodlands as opposed to the shaded floor of Beech-
Maple forests. In Michigan and New England gulosa usually
matures late in the fall, overwinters as an adult, and mates in early
spring. Kaston (1936) made the following observations of courtship
behavior in the species:

Immediately upon coming in contact with the female, or
within 3 minutes thereof, the male begins to drum his palps
rapidly against the floor of the cage. These drumming move-
ments are made so rapidly that a distinct purring or humming
sound can be heard. The palps are used alternately and are
raised only a very short distance during the process. The body
is held at an angle so that the posterior end of the abdomen
almost touches the floor. As a consequence when the male
begins to twitch his abdomen in a vertical plane the tip strikes

1986]

Brady — Near c tic Gladicosa

295

Map 1. Distribution of G. gulosa.

the floor. However, I could not detect any sounds made by this
part of the body. It is highly probable that the vibrations set up
in the substratum by the tapping movements of the palps and
abdomen are perceived by the female. This may exert an excit-
ing influence on her in a manner analogous to that which
occurs in web-building species, where the male tweaks the
threads of the female’s snare.

The male now moves slowly toward the female without
courting. When near her he reaches over to touch her. At first
she may jump at him and chase him away. Later, if she is
receptive she allows him to stroke her legs or abdomen. After
this contact with the female the male resumes his courtship
movements. Later on, if the male gets more excited he begins to
raise his forelegs off the floor about 1 or 2 mm, and lower them
quickly. During this process the legs quiver violently.

After 13 minutes of this courting one male began to mount
the female, but before he could get into the final copulatory
position, she ran away from him. Another male had courted
only seven minutes when the female allowed him to mount. The
position is the usual one for Lycosids, the male using his palps

296

Psyche

[Vol. 93

alternately during the 10 minutes the act lasted. This duration
time may not be the usual one for the species, however, for one
pair were observed in the field, when collected, which were
already in copula and remained so for about another half hour.
The sound produced during courtship was also reported by
Allard (1936). Observations were made on a collecting trip in the
Bull Run Mountains of Virginia during late April. He described the
sound as a distinct purring produced by drumming rapidly upon dry
leaf surfaces. He reports:

The creatures were very wary, but with care I was able to
examine their movements critically from a distance of only a
few inches. When the spider moved and made its sounds, the
fore part of the body quivered perceptibly and the palpi, too,
executed gentle up and down movements. The quivering
movements brought the chelicerae directly in contact with the
dry leaf surface, and the latter alone appeared to be responsible
for the rather loud sounds I had heard.

According to Allard these tapping sounds could be heard a distance
of 10 feet or more.

Rovner (1975) investigated sound production in three species of
Schizocosa and six species of Lycosa, including gulosa. Previous
investigators, as with gulosa above, had regarded such sounds as
being solely percussive, generally produced by a tapping or scraping
of the palps or the chelicerae against the substratum. High-speed

Figs. 6-9. Gladicosa gulosa (Walckenaer) 6-7. Female from 4 mi. S of New
Richmond, Allegan Co., Michigan, 16 Sept. 1974. 6. Epigynum. 7. Internal geni-

talia. 8. Epigynum of female from Pepperell, Middlesex Co., Massachusetts, Apr.
1973. 9. Epigynum of female from Cove Creek Valley, 15 mi. S of Prairie Grove,

Washington Co., Arkansas.

Figs. 10-14. Gladicosa pulchra (Keyserling). 10. Epigynum of female from
Stone Co., Mississippi, 21 Dec. 1964. 1 1. Epigynum of syntype from North Ameri-

ca. 12-13. Female from Gainesville, Alachua Co., Florida, 14 June 1935.
12. Internal genitalia. 13. Epigynum. 14. Epigynum of holotype of Lycosa inso-
pita Montgomery [= Gladicosa pulchra (Keyserling)] from Austin, Travis Co.,
Texas.

Figs. 15-17. Gladicosa euepigynata (Montgomery). 15-16. Female from Camp
Verde, Kerr Co., Texas, Dec. 1939. 15. Internal genitalia. 16. Epigynum. 17.

Epigynum of holotype of Lycosa euepigynata Montgomery [= Gladicosa euepigy-
nata (Keyserling)] from Austin, Travis Co., Texas. Ip, longitudinal piece of scape; s,
seminal receptacle; tp, trnasverse piece of scape.

1986]

Brady — Nearctic Gladicosa

297

298

Psyche

[Vol. 93

film analysis by Rovner (1975) revealed the prescence of a
stridulatory organ at the tibio-tarsal joint. This apparatus consists
of a file on the distal end of the tibia and a scraper at the base of the
palpal cymbium. Further examination revealed a group of stout
spines or macrosetae at the tip of the palpal tarsus. These spines
apparently aid in coupling the tarsus to the substratum. Thus, the
sound produced by gulosa ©and other lycosids is not generated
simply by drumming, but involves a rapid oscillation at the
tibio-tarsal joint facilitated by macrosetae that anchor the palpus to
the substratum.

Kaston (1948) reports seeing mature females of gulosa from Sep-
tember, through winter, to June suggesting that some may live for
two years. Egg sacs appear in early April and are produced until late
May. Egg sacs vary from 6-10 mm in diameter and egg counts range
from 118-274, each egg about 1 mm in diameter.

Distribution. From southern Canada in the northeast to eastern
Texas in the southwest. Not recorded from Florida and a single
specimen from Colorado (Map 1).

Records. CANADA. Nova Scotia. Bridgewater; Kentville.
Quebec. Ft. Coulonge; King Mtn., Gatineau National Park; Ste.
Rose. Ontario. Arnprior; Belleville; Chatterton; Haliburton; Mar-
mora; Mazinaw Lake; Ottawa; Pelee Island; Port Credit; Rondeau
Provincial Park; Simcoe; Toronto.

UNITED STATES. Maine. Androscoggin Co.: Poland Spring,
15 June 1904, 9 (J. H. Emerton); York Co.: Wells, 12 Aug. 1933 (W.
Ivie). New Hampshire. Belknap; Carroll; Cheshire; Hillsboro; Sulli-
van. Vermont. Caledonia; Windham; Windsor. Massachusetts.
Barnstable; Berkshire; Essex; Franklin; Hampden; Middlesex; Nor-
folk; Worcester. Connecticut. Providence; Fairfield; Litchfield;
Middlesex; New Haven; Windham. New York. Allegany; Cat-
taraugus; Courtland; Essex; Fulton; Monroe; Nassau; Oneida;
Onondaga; Queens; Richmond; Rockland; Steuben; Suffolk; Sul-
livan; Tompkins; Westchester; Wyoming. New Jersey. Bergen;
Burlington; Camden; Mercer; Ocean; Union. Pennsylvania. Butler;
Cambria; Carbon; Mifflin; Philadelphia; Pike; Venango; Westmore-
land. Ohio. Champaign; Columbiana; Hocking; Knox; Ottawa;
Washington. Maryland. Anne Arundel; Baltimore City; Mont-
gomery. District of Columbia. Washington. West Virginia. Poca-
hontas. Virginia. Fairfax; Falls Church (Indep. City); King William;

1986]

Brady — Ne arc tic Gladicosa

299

Montgomery; Prince Edward; Richmond (Indep. City); Rocking-
ham; Shenandoah. Kentucky. Breathitt; Wolfe. Tennessee. Sevier.
North Carolina. Beaufort; Buncombe; Chatham; Cherokee; Dur-
ham; Hartnett; Haywood; Henderson; Jones; Lee; Macon; Onslow;
Orange; Swain; Transylvania; Wake. Georgia. Rabun. Alabama.
Bibb; Butler; Lee. Mississippi. Forrest; George; Hinds; Jackson;
Perry. Louisiana. Caddo; Grant. Michigan. Allegan; Barry; Cal-
houn; Charlevoix; Cheboygan; Clare; Iosco; Jackson; Lake; Liv-
ingston; Midland; Oakland; Ontonagon; Osceola; Ottawa; Ros-
common; Washtenaw; Wexford. Indiana. Jackson; LaPorte; Parke;
Vermillion. Wisconsin. Adams; Buffalo; Chippewa; Dane; Ozaukee;
Polk; Rusk; Sauk; Sheboygan; Vernon; Velas; Washburn; Wau-
shara. Illinois. Champaign; Cook; Ogle; Piatt; Shaunee. Minnesota.
Hennepin; Ramsey. Missouri. Boone; Greene; St. Charles; St.
Louis City. Arkansas. Carroll; Lawrence; Montgomery; Polk;
Washington. South Dakota. Lincoln Co.: Newton Hills St. Pk., 6
mi. SSE of Canton, 9 June 1957, 9 (T. J. Cohn). Nebraska. Jeffer-
son Co.: Fairbury, 1 May 1957, 9 (W. F. Rapp, Jr.); Lancaster Co.:
Lincoln, 1941, 552 (M. J. Harbaugh); Saline Co.: Crete, 12 Sept.
1948, 9 (J- & W. Rapp). Kansas. Cowley Co.: Winfield, 92;
Kingman Co.: Kingman Co. St. Pk. near Calista, 13 Oct. 1963,
(5:399 (J. & W. Ivie); Riley Co.: Manhattan, (559 (N. Banks), Apr.
1903, 9 (T. H. Sheffer). Oklahoma. Canadian Co.: Yukon, 10 Sept.,
355;22 (N. M. Newport); Cleveland Co.: Norman, 552 (J- H.
Emerton); Creek Co.: Drumright, 26 Feb. 1927, 9 (Byers). Texas.
Dallas Co.: Dallas, 28 Jan. 1954, $ (E. E. Gilbert), White Rock
Creek, 13 Dec. 1934, 9 (N. E. Vickery & S. Jones); Grayson Co.: 6
mi. N of Denison, 20 Oct. 1963, 9 (K. W. Haller); Jasper Co.:
Jasper, 26 Jan. 1962, 55 (High School Sci. Club); Wichita Co.:
Burkburnett, 12 Oct. 1964, 5:499 (K. W. Haller). Colorado.
Bluebell Canvon near Boulder, 23 Oct. 1944, 9 (R. E. Gregg).

Gladicosa pulchra (Keyserling), comb. nov.

Figures 4, 10-14, 37-42. Map 2.

Tarentula pulchra Keyserling, 1877: 628, pi. 7, figs. 13, 14, $9- Syntypes ((59 ) from
“North America,” L. Koch collection, deposited in the British Museum (Natural
History), examined. Banks 1893: 124.

Lycosa pulchra: Montgomery 1904: 277. Banks 1910: 57; 1911: 454. Banks, Newport,
and Bird 1932: 31. Gertsch 1934: 8. Gertsch and Wallace 1935: 21, figs. 38, 41,
59. Jones 1936: 69.

300

Psyche

[Vol. 93

Lycosa gulosa: Chamberlin 1908: 265 (part).

Lycosa insopita Montgomery, 1904: 278, 280, figs. 3, 4, <$$. Syntypes ($$ ) from
Austin, Travis Co., Texas, deposited in the American Museum of Natural His-
tory, examined; 1905: 174; 1909: 514. Petrunkevitch 1911: 560. First synonymy
with Lycosa pulchra by Gertsch 1934.

Scaptocosa pulchra: Roewer 1954: 293.

Alopecosa pulchra: Bonnet 1955: 256.

Discussion. Montgomery (1904) described this species under
Lycosa insopita. He apparently did not have the Keyserling syn-
types for comparison. Gertsch (1934) was the first to recognize the
synonymy.

Color. The range of color in G. pulchra is greater than that of
G. gulosa. I have noted light forms and dark forms of pulchra.
These do not represent a genetic polymorphism but are the extremes
in a color continuum. There is no discernible correlation between
geographic locality and color pattern among the specimens exam-
ined. The darker forms are much more numerous than the light
colored ones. The range of color is indicated in the following
descriptions.

Color. Female. Face orange-brown to dark reddish brown.
Chelicerae dark reddish brown to black with condyles lighter
orange-brown.

Carapace dark brown to a dark reddish brown with a broad
median yellow stripe suffused with white hair. Irregular lighter
submarginal yellow stripes similarly clothed with white hair. Pattern
as in Figure 4.

Dorsum of abdomen brown to brown mottled with black.
Anterio-lateral areas black, blending with similar black areas on
cephalothorax. Five pairs of white spots (in well-marked specimens)
beginning in cardiac area and continuing posteriad. White spots
connected by dark brown chevrons as in Figure 4. Cardiac area
darker brown, outlined by lighter brown or yellowish.

Venter of abdomen dark brown to almost black posterior to epi-
gastric furrow. Yellowish anterior to furrow.

Legs light brown with darker black annulations on femora to
dark reddish brown without distinct annulations.

Labium and endites light brown to black with pale yellowish
distal ends. Sternum yellow brown (golden), dark reddish brown to
black.

1986]

Brady — N ear c tic Gladicosa

301

Figs. 18-20. Gladicosa huberti (Chamberlin). 18-19. Female from Bar M
Ranch near Boston, Thomas Co., Georgia, 2 Mar. 1973. 18. Internal genitalia. 19.

Epigynum. 20. Epigynum of female from Welaka Reserve, Putnam Co., Florida, 1 1
Nov. 1972.,

Figs. 21-26. Gladicosa bellamyi (Gertsch and Wallace). 21-22. Holotype
female from Liberty Co., Florida, 12 Apr. 1935. 21. Internal genitalia. 22. Epigy-

num. 23. Epigynum of holotype of Trochosa cherokee Chamberlin and Ivie,
[= Gladicosa bellamyi (Gertsch and Wallace)]. Ft. Gibson, Muskogee Co.,
Oklahoma, 21 July 1937. 24-26. Females from 2 mi. N of Stoneville, Washington

Co., Mississippi. Internal genitalia. 25. Epigynum. 26. Epigynum.

302

Psyche

[Vol. 93

Color . Male. Face yellow-orange to orange-brown. Dark in
ocular area. Chelicerae brownish orange to dark reddish brown.
Cymbia of palpi yellow-orange to dark reddish brown.

Carapace orange-brown to dark orange-brown with broad yellow
to pale orange median stripe overlaid with white hair. Irregular
submarginal stripes of same color, sometimes indistinct.

Dorsum of abdomen with median area light to medium brown,
bordered by black. Five pairs of white spots beginning in cardiac
area and continuing posteriad. Spots joined by black chevrons.
Cardiac area brown, enclosed by lighter pale brown to yellow-
brown. Pattern similar to female. Venter of abdomen brown to
black posterior to epigastric furrow. Light brown to pale yellow or
cream anterior to furrow.

Labium and endites yellow-orange to orange with distal ends
cream. Sternum yellow-orange to orange.

Measurements. Ten females and ten males from Florida. See
Table 2.

Diagnosis. Gladicosa pulchra is closest to G. gulosa in size, col-
oration, and genitalic structure. Gladicosa pulchra is a larger species

1986]

Brady — Ne arctic Gladicosa

303

than gulosa (compare Table 1 with Table 2) and is usually darker in
color with a more distinct pattern (compare Fig. 4 with Fig. 3). In
most specimens of pulchra the venter of the abdomen is dark brown
to black behind the epigastric furrow, while that of gulosa is
yellowish to light brown. Differences between female and male
genitalia of these two species are noted under gulosa and in the keys.

Natural History. Little is known of the habitat or behavior of
pulchra. I’ve collected this species in Florida from the trunks of
deciduous trees where their color blends well with the bark sub-
strate. G. B. Edwards (personal communication) has collected spec-
imens from similar microhabitats in Florida. Pat Miller (personal
communication) reported collecting both male and female pulchra
from the trunks of pine trees at night in Perry, Florida, on
December 5, 1982. Montgomery (1904) reported finding pulchra
near Austin, Texas, in drier habitats than gulosa and less
abundantly. He noted that the females live under stones where they
make a shallow horizontal burrow lined with silk. Whether this
behavior is consistent throughout the life cycle or represents a
temporary adjustment to molting or egg laying is a question to be
answered. Gladicosa pulchra is not the abundant inhabitant of
deciduous leaf litter, as are gulosa and huberti. Of the species
investigated pulchra is the most variable in coloration of the body
and structure of the epigynum. It is possible that more than
one species is represented in this complex.

Roble (1986) reported rearing Mantispa viridis from a Gladicosa
pulchra egg sac. It is the first record of a lycosid spider serving as a
host of M. viridis. When the spider died, its egg sac was opened and
a mantispid cocoon and 95 surviving spiderlings were found. This
corroborates an earlier observation of high spiderling survival
within a mantispid-infested egg sac of Lycosa rabida.

Distribution. From Long Island, New York, along the East
Coast to Texas in the southwest. Limited in its northern range
inland to the southern parts of Kansas and Missouri and northern
Kentucky. More abundant in the southeastern United States (Map
2).

Records. UNITED STATES. New York. Suffolk Co.: Coram,
Long Island, 19 Oct. 1934, $ (E. L. Bell). New Jersey. Cape May
Co.: Cape May, 29 Sept. 1945, $ (C. & M. Goodnight). Virginia.
Alexandria (Indep. City); Falls Church (Indep. City); Fairfax. Ken-
tucky. Woodford Co.: Kentucky River, 16 Sept. 1920, $. Tennessee.

304 Psyche [Vol. 93

Table 2. Measurements of ten females and ten males of Gladicosa pulchra from
Florida.

Females:

Mean SEM

Ant. Eye Row

1.304 ± .028

Femur I

5.46 + .12

PME

1.734 ± .040

Pat. -Tibia I

7.23 + .16

PLE

2.284 ± .052

Meta. I

4.23 ± .10

POQ

1.622 ± .036

Tarsus I

2.18 ± .05

Car. Width

5.50 ±.16

Total I

19.09 + .43

Car. Length

7.23 +.19

Femur IV

5.93 + .14

Body Length

15.89 +.56

Pat. -Tibia IV

7.36+ .18

Pat. -Tibia II

6.75 +.16

Meta. IV

6.75 + .19

Pat. -Tibia III

5.88 +.14

Tarsus IV

2.70 + .07

Total IV

22.73 + .51

Males:

Mean SEM

Ant. Eye Row

1.176+ .022

Femur I

5.79 + .11

PME

1.604 ± .032

Pat. -Tibia I

7.88 ± .19

PLE

2.044 ± .050

Meta. I

5.46 + .15

POQ

1.514+ .032

Tarsus I

2.47 + .06

Car. Width

4.94 +.14

Total I

21.59 + .50

Car. Length

6.54 +.18

Femur IV

6.19 + .11

Body Length

12.35 +.33

Pat. -Tibia IV

7.71 + .16

Pat. -Tibia 11

12.02 + .17

Meta. IV

7.83 + .18

Pat. -Tibia III

6.09 +.16

Tarsus IV

2.95 + .09

Total IV

24.69 + .51

Knox Co.: Knoxville, 8 Oct., $ (W. B. Cartwright). North Carolina.
Alamance; Durham; Moore; Wake. Georgia. Floyd; Screven. Flor-
ida. Alachua; Baker; Citrus; Gadsden; Lake; Leon; Levy; Liberty;
Marion; Oklaloosa; Putnam; Polk; Sarasota; Taylor, Volusia. Ala-
bama. Baldwin; Butler; Lee; Mobile. Mississippi. Forrest; Jackson;
Marshall; Noxubee; Oktibbeha; Pike; Stone. Louisiana. Caddo;
Evangeline; Madison. Missouri. Pulaski Co.: Richland, 20 Apr.
1962, $ (W. Ivie). Arkansas. Lawrence; Montgomery; Sharp;
Washington. Kansas. Bourbon Co.: Redfield, 14 Oct. 1963, $ (J. &
W. Ivie). Texas. Bandera Co.: Dec. 1939, 322 (D. & S. Mulaik);
Comal Co.: Hancock, 27 May 1948, 2 with egg case (I. J. Ander-
son); DeWitt Co.: 16.4 mi. SE of Cuero, 23 Dec. 1955 (W. McAlis-
ter); Hale Co.: Wimberley, 1948, 2 (Exline coll.); Harris Co.: Clear
Lake near Seabrook, Sept. 1959, 2 (J- C. Bequaert); Kerr Co.:
Raven Ranch, Dec. 1939, $$ : 1022 (D. & S. Mulaik); Smith Co.:
Tyler St. Pk., 12 Mar. 1982, 2 (S. M. Roble); Travis Co.: Austin,
522 (T. H. Montgomery).

1986]

Brady — Nearctic Gladicosa

305

Gladicosa huberti, comb. nov.

Figures 1, 18-20, 27, 28. Map 3.

Lycosa huberti Chamberlin, 1924: 28, pi. 6, fig. 44, $. Female holotype from Tali-
sheek, St. Tammany Par., Louisiana, 4 March 1920 (H. E. Hubert), deposited in
the Museum of Comparative Zoology, examined. Gertsch and Wallace 1935: 22,
figs. 40, 43, (5$. Chamberlin and Ivie 1944: 144. Bonnet 1957: 2645.

Scaptocosa huberti: Roewer 1954: 293.

Discussion. Gladicosa huberti together with G. pulchra were
placed in the genus Scaptocosa by Roewer (1954) with Lycosa mis-
souriensis (Banks) [= Geolycosa\ as the type species. Five other
North American species now considered to be in Geolycosa and one
species of Schizocosa were included in Scaptocosa as well. It is not
clear what distinguishes this odd assemblage.

Color. Females. Face orange-brown to reddish brown with
eye nacelles black. Chelicerae dark reddish brown (mahogany) to
black. Condyles orange-brown.

Carapace orange-brown to reddish brown with broad median
pale orange stripe from PME to posterior edge. Lighter irregular
submarginal stripes less distinct than median. Pattern as in Figure 1 .

Dorsum of abdomen brown to dark brown with cardiac area
outlined in black. Chevrons faintly indicated along posterior half
with white spots marking their lateral edges. Anterior lateral edges
of dorsum darker as in Figure 1. Venter pale yellow-orange to
darker brown. Lateral areas darker in pale-colored individuals, con-
colorous brown in others.

Legs yellow-orange to orange-brown, without darker annulations.

Labium and endites orange-brown to dark reddish brown, with
distal ends yellowish to cream. Sternum yellow-orange to light
orange-brown.

Color. Males. Face dark orange-brown to very dark reddish
brown, eye region black. Chelicerae dark reddish brown to black.
Condyles lighter. Cymbia of palpi dark red-brown.

Carapace orange-brown to darker reddish brown with light
orange broad median stripe from eye region to posterior edge. Light-
er, irregular submarginal stripes, not so distinct as median one.

Dorsum of abdomen medium to dark brown with cardiac area
lighter, outlined by black line which is enclosed in turn with lighter
color extending laterally. Anterior lateral areas marked by black
color, which extends more posteriad than in female. Venter of

307

1986] Brady — Ne arc tic Gladicosa

abdomen orange-brown to dark brown. Central area somewhat
lighter.

Legs yellow-orange to orange-brown, somewhat lighter ventrally,
without darker bands.

Labium and endites yellow-orange to dark reddish brown, with
distal ends pale yellow to cream. Sternum yellow to reddish
orange-brown.

Measurements. Ten females and ten males from Georgia and
Florida.

Diagnosis. Gladicosa huberti is closest to G. bellamyi in body
size and shape of the epigynum, but resembles G. gulosa in colora-
tion and structure of the male palpus. Gladicosa huberti is lighter in
color than bellamyi and smaller in size than gulosa. It may be dis-
tinguished from either of these species by comparing the epigynum
(Figs. 19, 20) to bellamyi (Figs. 22, 23, 25, 26) or gulosa (Figs. 6,
8, 9) and the palpus (Figs. 27, 28) to bellamyi (Figs. 29-34) ox gulosa
(Figs. 35, 36).

Natural History. Nothing concerning the natural history of this
species is reported in the literature. I have collected it in leaf litter
near the edge of woods in Georgia and in a marshy area near the
edge of a pond beneath a pine tree canopy in Florida. The great
majority of the adult specimens were collected from February
through April (see Records below).

Distribution. Southeastern United States (Map 3).

Records. South Carolina. Jasper Co.: Ridgeland, 28 Mar.-6
Apr. 1975, ? with egg case (D. Brody). Georgia. Chatham Co.: 8 mi.

Figs. 27-28. Gladicosa huberti (Chamberlin), left palpus of male from Bar M
Ranch near Boston, Thomas Co., Georgia, 2 Mar. 1973. 27. Retrolateral

view. 28. Ventral view.

Figs. 29-34. Gladicosa bellamyi (Gertsch and Wallace). 29-30. Male from
Sharon Woods Metropolitan Park, Columbus, Franklin Co., Ohio 1-8 May 1973.
29. Left palpus, retrolateral view. 30. Left palpus, ventral view. 3 1 -34. Males from
2 mi. N of Stoneville, Washington Co., Mississippi 9-31 May 1983. 31. Ventral

view. 32. Retrolateral view. 33. Ventral view. 34. Retrolateral view.

Figs. 35-36. Gladicosa gulosa (Walckenaer), left palpus of male from 4 mi. S of
New Richmond, Allegan Co., Michigan, 16 Sept. 1974. 35. Retrolateral view.

36. Ventral view.

Figs. 37-38. Gladicosa pulchra (Keyserling), left palpus of male syntype of
Lycosa pulchra Keyserling from North America. 37. Ventral view. 38. Retrolat-
eral view, co, conductor; cy, cymbium; em, embolus; ma, median apophysis; pa,
palea; ta, terminal apophysis.

308

Psyche

[Vol. 93

Map 3. Distribution of G. huberti, euepigynata, and Bellamyi.

W of Savannah, 5 Apr. 1943, $, 3 mi. SE of Savannah, 8 Apr. 1943,

2 (W. Ivie); Chattahoochee Co.: Fort Benning, 31 Oct. 1943, 2 (D.
C. Beck); Screven Co.: 1 mi. N of Sylvania, 9 Apr. 1943, 5, 2 mi. N
of Sylvania, 11 Apr. 1943, 29, 7 mi. N of Sylvania, 2 (W. Ivie);
Thomas Co.: Bar M Lodge near Boston, 2 Mar. 1973, 5599°°
(A. R. Brady).

Florida . Alachua Co.: 6 Apr. 1935, 2 with egg case, 26 Nov.

1936, 52, 2-3 Feb. 1937, 455:422:0, 18 Feb. 1937, 522, 6 Mar.

1937, 2, 23-27 Apr. 1937, 722, 12 June 1937, 322, 20 Nov. 1938, 2
(H. K. Wallace); Calhoun Co.: Blountstown, 28 Apr. 1935, 2 (H. K.
Wallace); Columbia Co.: 27 Apr. 1935, 5:422 (H. K. Wallace),

3 Feb. 1938, 2 (Beck); Levy Co.: 20 Apr. 1935, 2 (H. K. Wallace);
Putnam Co.: Welaka Reserve, 11 Nov. 1972, 355:99 (A. R.
Brady).

Mississippi. Forrest Co.: Camp Shelby near Hattiesburg, Oct.-
Nov. 1943, 2 (C. D. Michener); George Co.: Lucedale, Mar. 1930,
22 (Dietrich).

Gladicosa bellamyi (Gertsch and Wallace) comb. nov.
Figures 2, 21-26, 29-34. Map 3.

Lycosa bellamyi Gertsch and Wallace, 1937: 10, fig. 14, $. Female holotype from
Liberty Co., Florida, 12 April 1935 (H. K. Wallace) deposited in the American
Museum of Natural History, examined. Chamberlin and Ivie 1944: 142. Bonnet
1957: 2635.

1986]

Brady — Ne arc tic Gladicosa

309

Trochosa cherokee Chamberlin and Ivie, 1942: 35, fig. 76, $. Female holotype from
Fort Gibson, Muskogee Co., Oklahoma, 21 July 1937 (Standish-Kaiser) depos-
ited in the American Museum of Natural History, examined. NEW SYNONYM.
Avicosa bellamyi: Roewer 1954: 236.

Discussion. Gladicosa bellamyi was placed in the new genus
Avicosa by Roewer (1954) with Avicosa avida (Walckenaer) [=
Schizocosa ] as the type species. Two other North American species
now placed in Schizocosa ( minnesotensis and wasatchensis =
mccookx) as well as Lycosa ceratiola and Tarentula pictilis (now
Alopecosa pictilis ) were also included in this new genus. Avicosa is
certainly an artificial conglomeration without systematic foundation.

Table 3. Measurements of ten females and ten males of Gladicosa huberti from
Georgia and Florida.

Females:

Mean SEM

Ant. Eye Row

.959 + .013

Femur I

3.63 ± .09

PME

1.138 ± .018

Pat. -Tibia I

4.68 ± .11

PLE

1.478 ± .025

Meta. I

2.56 ± .09

POQ

1.048 ± .019

Tarsus I

1.57 ± .03

Car. Width

3.84 +.12

Total I

12.48 ± .32

Car. Length

5.09 +.11

Femur IV

4.12 ± .09

Body Length

11.18 +.46

Pat. -Tibia IV

5.03 ± .10

Pat. -Tibia II

4.27 +.11

Meta. IV

4.53 ± .09

Pat. -Tibia III

3.68 +.08

Tarsus IV

2.03 ± .03

Total IV

15.70 ± .31

Males:

Mean SEM

Ant. Eye Row

.861 ± .006

Femur I

3.62 ± .05

PME

1.061 ± .009

Pat. -Tibia

4.85 ± .06

PLE

1.364 ± .011

Meta. I

3.11 ± .04

POQ

.966 ± .010

Tarsus I

1.71 ± .03

Car. Width

3.60 ± .04

Total I

13.30+ .17

Car. Length

4.81 +.07

Femur IV

4.14 ± .06

Body Length

8.98 +.17

Pat. -Tibia IV

4.96 ± .07

Pat. -Tibia II

4.32 ± .06

Meta. IV

4.61 ± .07

Pat. -Tibia III

3.67 ± .05

Tarsus IV

2.03 ± .03

Total IV

15.73 ± .22

Color. Females. Face orange-brown to dark reddish brown.
Chelicerae dark reddish brown to black. Condyles lighter yellowish.

Carapace dark brown to dark reddish brown with broad median
yellow-orange to pale brownish orange stripe from PME to poste-
rior declivity as in Figure 2. Indistinct submarginal stripes of same
color.

310 Psyche [Vol. 93

Dorsum of abdomen pale yellow-brown to medium brown, often
with darker brown cardiac mark and darker chevrons posteriorly as
in Figure 2. Slight indication of black counter-shading anterio-
laterally. Venter of abdomen dark brown posterior to epigastric
furrow; median area sometimes mottled with light orange-brown.
Lighter yellowish anterior to furrow.

Legs brown to dark brown dorsally. Pale yellowish brown to
golden brown ventrally. Legs without distinct bands.

Labium and endites dark reddish brown to orange-brown with
distal ends lighter golden to yellow.

Color. Males. Face dark red-brown. Eye region black. Che-
licerae dark brown to black with inner distal margins lighter orange-
brown. Condyles lighter orange to yellow. Cymbia of palpi brown
to dark brown.

Carapace dark reddish brown overlaid with fine black hair. Broad
median pale yellow-orange to orange-brown stripe from PME to
posterior edge.

Dorsum of abdomen beige to light brown. Black countershading
in anterio-lateral areas, extending posteriorly farther than in female.
Indistinct chevrons posteriorly. In some specimens the median longi-
tudinal area of the dorsum is pale yellow to cream with darker
brown at edges and along sides. Venter of abdomen dark brown to
black posterior to epigastric furrow, lighter yellowish brown ante-
riorly. Lateral areas often somewhat lighter in color.

Legs orange-brown to dark brown dorsally, paler golden to yel-
lowish brown ventrally. Without darker bands. Tibia and metatar-
sus I black, tarsus yellow.

Table 4. Measurements of ten females and ten males of Gladicosa bellamyi from
Ohio.

Females:

Mean SEM

Ant. Eye Row

.891 ± .016

Femur I

3.39 ± .08

PME

1.135 ± .018

Pat. -Tibia I

4.46+ .11

PLE

1.478 ± .024

Meta. I

2.47 ± .05

POQ

1.065 ± .015

Tarsus I

1.53 ± .03

Car. Width

3.66 ± .08

Total I

11.85 + .26

Car. Length

4.86 +.09

Femur IV

3.95 ± .09

Body Length

10.43 ± .27

Pat. -Tibia IV

4.86+ .11

Pat. -Tibia II

4.05 ± .09

Meta. IV

4.51 ± .10

Pat. -Tibia III

3.49 +.08

Tarsus IV

1.96 + .04

Total IV

15.28 ± .33

1986]

Brady — Nearctic Gladicosa

311

Males:

Mean SEM

Ant. Eye Row

.839 ± .014

Femur I

3.31 ± .06

PME

1.071 ± .013

Pat. -Tibia I

4.59 ± .06

PLE

1.369 ± .019

Meta. I

2.79 ± .04

POQ

.993 ± .014

Tarsus I

1.62 ± .03

Car. Width

3.34 ± .05

Total I

12.30 ± .15

Car. Length

4.40 ± .06

Femur IV

3.73 ± .06

Body Length

8.56 ±.14

Pat. -Tibia IV

4.59 ± .06

Pat. -Tibia II

3.97 ±.05

Meta. IV

4.38 ± .07

Pat. -Tibia III

3.40 ± .04

Tarsus IV
Total IV

1.98 ± .04
14.68 ± .17

Labium and endites orange-brown to dark brown with distal ends
lighter yellow to golden. Sternum light orange-brown to darker
reddish brown.

Measurements. Ten females and ten males from Ohio, and ten
females from Mississippi. See Tables 4 and 5.

Table 5. Measurements of ten females of Gladicosa bellamyi from Mississippi.

Mean SEM

Ant. Eye Row

.925 ± .013

Femur I

4.23 ± .07

PME

1.216 ± .01 1

Pat. -Tibia I

5.73 ± .10

PLE

1.553 ± .021

Meta. I

3.54 ± .09

POQ

1.121 ± .009

Tarsus 1

1.95 ± .03

Car. Width

4.15 ±.08

Total I

15.44 ± .26

Car. Length

5.32 ±.09

Femur IV

4.82 ± .08

Body Length

9.94 ±.19

Pat. -Tibia IV

5.91 ± .11

Pat. -Tibia II

5.04 ±.07

Meta. IV

5.61 ± .10

Pat. -Tibia III

4.33 ±.07

Tarsus IV

2.43 ± .04

Total IV

18.75 ± .31

Diagnosis. Gladicosa bellamyi is closest to G. huberti in body
size and in shape of the epigynum (compare Figs. 22, 23, 25, 26 with
Figs. 19, 20). It is more darkly colored than huberti and the light sub-
marginal stripes on the carapace are narrower. Gladicosa bellamyi
can be easily distinguished from huberti by the structure of the male
palpi (compare Figs. 29-34 with Figs. 27, 28). Other than the type
specimens of Lycosa bellamyi and Trochosa cherokee, this species is
represented by specimens taken in pitfall traps near Stoneville, Mis-
sissippi and Columbus, Ohio. The males from Mississippi, which

312

Psyche

[Vol. 93

are the predominant sex in these collections, are distinctly larger
than the Ohio males as indicated by the Measurements, but the
similarity of coloration, genitalic structure, and anatomical propor-
tions led me to think that only one species is represented. The
southern populations are simply larger in size.

Natural History. Andrew Penniman (personal communication)
collected this species in some abundance by using pitfall traps in a
wooded area in central Ohio. The collecting period extended from
24 April to 28 August 1973 and the relative abundance of the sexes
taken in these traps is indicated in the records below. Four females
with egg cases were collected from 29 May- 12 June. The egg cases
contained 53, 56, 91, and 106 eggs. Tim Lockley (personal commun-
ication) also captured this species in pitfall traps placed at the edge
of a deciduous woods in Mississippi. Most of these specimens were
males as indicated in the records below. A single female with egg
case was collected between 3-6 June 1983.

Distribution. Ohio southeastward to western Florida and south-
westward to Oklahoma (Map 3).

Records. Ohio. Franklin Co.: Sharon Woods Metropolitan
Park, Columbus, 24 April- 1 May, 19<3<3:?; 1-8 May, 28(3(3:3??;
8-15 May, 23(3(3:10??; 15-22 May, 8??; 22-29 May, ??;
29 May-5 June, <3(3:7??; 5-12 June, <3(3:3??; 19-26 June,
?; 26 June-3 July, 3??; 10-17 July, ?; 21-28 Aug. 1973, <3
(A. J. Penniman). Florida. Liberty Co.: 12 April 1935, ?
(H. K. Wallace). Mississippi. Washington Co.: 2 mi. N of Stoneville,
21-25 April, <3; 27-29 April, <3; 29 April-2 May, 9<3<3; 2-4 May,
3(3(3; 4-6 May, 4<3<3; 6-9 May, 4<3<3; 9-11 May, <3(3?;

13-16 May, 5<3<3; 16-18 May, 23-25 May, <3; 25-31 May,
<3<3; 3-6 June, <3?; 1-6 July 1983, ? (T. C. Lockley). Oklahoma.
Muskogee Co.: Fort Gibson, 21 July 1937, ? (Standish-Kaiser).

Gladicosa euepigynata (Montgomery) comb. nov.

Figures 3, 15-17, 43-46. Map 3.

Lycosa euepigynata Montgomery, 1904: 277, 279, pi. 28, figs. 1, 2, <5‘$- Holotype
female from Austin, Travis Co., Texas (T. H. Montgomery) deposited in the
American Museum of Natural History, examined. Montgomery 1909: 514.
Banks 1910: 55. Gertsch 1934: 8. Gertsch and Wallace 1935: 22, figs. 44, 45, <?<$.
Bonnet 1957: 2607.

1986] Brady — Nearctic Gladicosa 313

Lycosa gulosa: Chamberlin 1908: 265 (in part). Petrunkevitch 1911: 560 (in part).

Not Lycosa gulosa (Walckenaer).

Hogna euepigynata: Roewer 1954: 258.

Discussion. Chamberlin (1908) synonymized G. euepigynata
with G. gulosa commenting upon the variation in size and color of
gulosa. Montgomery (1909) rightfully defended his designation of
euepigynata as a distinct species.

Color. Females. Face with sides orange-yellow, eye region
brown. Chelicerae dark reddish brown, darker distally.

Carapace brown with broad, irregular median stripe of orange-
yellow to yellow. Irregular submarginal stripes of orange-yellow,
intersected by black lines radiating from thoracic area. Pattern illus-
trated in Figure 3.

Dorsum of abdomen mottled with beige, spots of white, and dark
brown along the edges. Faint indications of chevron markings pos-
teriorly as in Figure 3. A series of five white spots marking edges of
chevrons. Venter of abdomen pale cream to yellow.

Legs yellow-gold to brownish orange. Pale ventrally with dorsal
surfaces of femora marked by three irregular dark brown bands.

Labium reddish brown with distal end yellow. Endites orange-
brown to reddish brown with distal ends yellow. Sternum orange-
brown to reddish brown.

Color. Males. Face yellow to brownish yellow, eye region
brown. Cymbia of palpi brown.

Carapace brown with broad median yellow stripe and irregular
submarginal stripes of same color, producing a pattern very similar
to that of female (Fig. 3).

Dorsum of abdomen with mottled pattern of light and dark
brown overlaid with white hair. White hairs forming five paired
spots beginning in cardiac area and continuing posteriad. Cardiac
area outlined with dark brown. Overall pattern as in female (Fig. 3).
Venter of abdomen cream to pale brown or beige.

Legs yellow to golden brown, darker on dorsal surface. Each
femur with three dark brown irregular bands that are more distinct
on dorsal surfaces.

Labium yellow to gold. Endites brown, with distal ends yellow.
Sternum golden yellow.

Measurements. Ten females and ten males from Texas. See
Table 6.

314

Psyche

[Vol. 93

Figs. 39-42. Gladicosa pulchra (Keyserling). Left palpus of male syntype of
Lycosa insopita Montgomery [= Gladicosa pulchra (Keyserling)] from Austin, Tra-
vis Co., Texas. 39. Retrolateral view. 40. Ventral view. 41-42. Left palpus of
male from Gainesville, Florida, 14 June 1935. 41. Ventral view. 42. Retrolateral

view.

Figs. 43-46. Gladicosa euepigynata (Montgomery). 43-44. Left palpus of male
syntype of Lycosa euepigynata Montgomery from Austin, Travis Co., Texas.
43. Retrolateral view. 44. Ventral view. 45-46. Left palpus of male from Camp
Verde, Kerr Co., Texas, Dec. 1939. 45. Ventral view. 46. Retrolateral view.

315

1986] Brady — Near die Gladicosa

Table 6. Measurements of ten females and ten males of Gladicosa euepigynata

from Texas.

Females: Mean SEM

Mean SEM

Ant. Eye Row
PME
PLE
POQ

Car. Width
Car. Length
Body Length
Pat. -Tibia II
Pat. -Tibia III

Males:

Ant. Eye Row
PME
PLE
POQ

Car. Width
Car. Length
Body Length
Pat. -Tibia II
Pat. -Tibia III

1.268 + .016
1.692 + .016
2.124 ± .024
1.610+ .014
5.32 +.08
7.21 +.12
16.88 +.35
6.54 ± .09
5.76 ± .07

Mean SEM

1.152+ .018
1.580+ .018
1 .964 ± .028

I. 466 ± .016
4.84 +.11
6.60 +.17

II. 91 +.28
6.24 ± .08
5.59 ± .08

Femur I
Pat. -Tibia I
Meta. I
Tarsus I
Total I
Femur IV
Pat. -Tibia IV
Meta. IV
Tarsus IV
Total IV

Femur I
Pat. -Tibia
Meta. I
Tarsus I
Total I
Femur IV
Pat. -Tibia IV
Meta. IV
Tarsus IV
Total IV

5.39 ± .06
7.06 + .10
4.15 ± .05

2.40 ± .02
19.05 ± .21

6.19 ± .08
7.51 ± .08

6.86 ± .07
2.93 ± .02

23.48 ± .22

Mean SEM

5.19 ± .10

6.87 ± .07
4.54 ± .07
2.44 ± .04

18.97 ± .25
5.91 ± .09
7.20 + .10
6.82 ± .07
2.84 ± .05
22.76 ± .29

Diagnosis . Gladicosa euepigynata is closest to G. pulchra in size
and coloration (compare Fig. 3 with Fig. 4). The epigynum of
euepigynata (Figs. 15-17) and the palpus (Figs. 43-46) distinguish it
from pulchra and all other species of Gladicosa.

Natural History. Montgomery (1904) reported this species as
being abundant near Austin, Texas. There he found it under stones
near water. Males were most numerous in January.

Distribution. South central Texas (Map 3).

Records. Texas. Bandera Co.: 2 mi. N of Medina, Dec. 1939,
39 (S. & D. Mulaik); Hays Co.: 15 Apr. 1939, 399 (D. & S.
Mulaik); Kerr Co.: Camp Verde, Dec. 1939, 5:392> Raven Ranch,
Dec. 1939, 92, Turtle Creek, Dec. 1939, 32 (D. & S. Mulaik);
Kendall Co.: Dec. 1939, 9 (D. & S. Mulaik); Tom Green Co.: San
Angelo, Dec. 1939, 9 (S. Mulaik); Travis Co.: Austin, 1333;2322
(R. V. Chamberlin).

316 Psyche [Voi. 93

References Cited

Allard, H. A.

1936. The drumming spider ( Lycosa gulosa Walckenaer). Proc. Biol. Soc.
Washington, 49: 67-68.

Banks, N.

1893. Notes on spiders. Jour. N. Y. Ent. Soc., 1: 123-134.

1901. Notes on some spiders of Walckenaer, Koch and others. Jour. N. Y. Ent.
Soc., 9: 182-189.

1910. Catalogue of Nearctic spiders. Bull. U.S. Nat. Mus., 72: 1-80.

1911. Some Arachnida from North Carolina. Proc. Acad. Nat. Sci. Phila., 65:
676-687.

Banks, N., N. M. Newport and R. D. Bird.

1932. Oklahoma spiders. Publ. Univ. Okla., Biol. Surv., 4(1): 7-49.

Bishop, S. C. and C. R. Crosby

1926. Notes on the spiders of the southeastern United States with descriptions
of new species. Jour. El. Mitch. Sci. Soc., 41(3-4): 163-212.

Bonnet, P.

1955. Bibliographia Araneorum. Toulouse, 2(1): 1-918.

1957. Bibliographia Araneorum. Toulouse, 2(3): 1927-3026.

Brady, A. R.

1962. The spider genus Sosippus in North America, Mexico, and Central
America (Araneae, Lycosidae). Psyche, 69(3): 129-164.

1972. Geographic variation and speciation in the Sosippus floridanus species
group (Araneae: Lycosidae). Psyche, 79(1-2): 27-48.

1979. Nearctic species of the wolf spider genus Trochosa (Araneae: Lycosidae).
Psyche, 86(2-3): 167-212.

Bultman, T. L., G. W. Uetz and A. R. Brady

1982. A comparison of cursorial spider communities along a successional gra-
dient. Jl. Arachn., 10: 23-33.

Chamberlin, R. V.

1904. Notes on generic characters in the Lycosidae. Canad. Ent., 36: 145-148,
173-178.

1908. Revision of North American spiders of the family Lycosidae. Proc.
Acad. Nat. Sci. Philad., 60: 158-318.

1924. Descriptions of new American and Chinese spiders, with notes on other
Chinese species. Proc. U.S. Nat. Mus., 63(13): 1-38.

Chamberlin, R. V. and W. Ivie

1942. A hundred new species of American spiders. Bull. Univ. Utah., 32(13):
1-117.

1944. Spiders of the Georgia region of North America. Bull. Univ. Utah, 35(9):
1-267.

Chickering, A. M.

1932. Notes and studies on Arachnida. II. Araneae from the Douglas Lake
Region, Michigan I. Pap. Mich. Acad. Sci., 15: 349-355.

Comstock, J. H.

1913. The spider book. Doubleday & Page, Garden City, New York, 721 pp.

1986] Brady — Near c tic Gladicosa 317

1940. The spider book. Rev. and ed. by W. J. Gertsch. Comstock, Ithaca, 729

pp.

Crosby, C. R. and S. C. Bishop

1928. Araneae in a list of the insects of New York. Cornell Univ. Agr. Exper.
Sta. Mem., 101: 1034-1074.

Dorris, P. R.

1965. A list of spiders collected in northern Mississippi. Trans. Amer. Micros.
Soc., 84(3): 407-408.

1968. A preliminary study of the spiders of Clark County, Arkansas compared
with a five year study of Missisippi spiders. Ark. Acad. Sci. Proc., 22:
33-37.

Drew, L. C.

1967. Spiders of Beaver Island, Michigan. Publ. Mus. Mich. St. Univ., Biol.
Ser., 3(3): 153-208.

Elliott, F. R.

1930. An ecological study of spiders of the Beech-Maple forest. Ohio Jour.
Sci., 30(1): 1-22.

1932. Revision of and additions to the list of Araneae (spiders) of Indiana.
Proc. Indiana Acad. Sci., 41: 419-430.

Emerton, J. H.

1885. New England Lycosidae. Trans. Conn. Acad. Arts Sci., 6: 481-505.

1902. The common spiders of the United States. Boston, 225 pp.

Fitch, H. S.

1963. Spiders of the University of Kansas Natural History Reservation and
Rockefeller Experimental Tract. Univ. Kan. Mus. Nat. Hist., Misc.
Pub., 33: 1-202.

Gertsch, W. J.

1934. Notes on American Lycosidae. Amer. Mus. Nov., 693: 1-25.

1949. American spiders. D. Van Nostrand Co., Inc., Princeton, New Jersey,
285 pp.

Gertsch, W. J. and H. K. Wallace

1935. Further notes on American Lycosidae. Amer. Mus. Nov., 794: 1-22.

1937. New American Lycosidae with notes on other species. Amer. Mus. Nov.,
919: 1-22.

Harrison, J. B.

1969. Acoustic behavior of the wolf spider Lycosa gulosa. Animal Behavior,
17: 14-16.

Jones, S. E.

1936. The Araneida of Dallas County: Preliminary note. Field Lab., 4(2):
68-70.

Kaston, B. J.

1935. The slit sense organs of spiders. Jour. Morph., 58: 189-207.

1936. The senses involved in the courtship of some vagabond spiders. Ent.
Amer., 16(2): 97-167.

1938. New spiders from New England with notes on other species. Bull. Brook-
lyn Ent. Soc., 33(4): 173-191.

1948. Spiders of Connecticut. Bull. Conn. St. Geol. Nat. Hist. Surv., 70:
1-874.

318 Psyche [Vol. 93

1981. Spiders of Connecticut. Rev. ed. Bull. Conn. St. Geol. Nat. Hist. Surv.,
70: 1-1020.

Keyserling, E. G.

1877. Ueber amerikanische Spinnenarten der Unterordnung Citigradae. Verh.
Zool.-Bot. Ges. Wien, 26: 609-708.

Marx, G.

1890. Catalogue of the described Araneae of temperate North America. Proc.
U.S. Nat. Mus., 12: 497-594.

1892. A list of the Araneae of the District of Columbia. Proc. Ent. Soc. Wash.,
2(2): 148-161.

Montgomery, T. H.

1902. Descriptions of Lycosidae and Oxyopidae of Philadelphia and its vicin-
ity. Proc. Acad. Nat. Sci. Phila., 54: 535-592.

1904. Descriptions of North American Araneae of the families Lycosidae and
Pisauridae. Proc. Acad. Nat. Sci. Phila., 56: 261-323.

1905. The spermatogenesis of Syrbula and Lycosa, with general considerations
upon chromosome reduction and the heterochromosomes. Proc. Acad.
Nat. Sci. Phila., 57: 162-205.

1909. Remarks on Prof. Chamberlin’s revision of North American Lycosidae.
Proc. Acad. Nat. Sci. Philad., 60: 513-515.

Petrunkevitch, A.

1911. A synonymic index-catalogue of spiders of North, Central and South
America with all adjacent islands, Greenland, Bermuda, West Indies,
Terra del Fuego, Galapagos, etc. Bull. Amer. Mus. Nat. Hist., 29: 1-791.
Roble, S. M.

1986. A new spider host association for Mantispa viridis (Neuroptera, Manti-
spidae). Jl. Arach., 14: 135-136.

Roewer, C. F.

1954. Katalog der Araneae. Band 2, Institut Royal des Sciences Naturelles de
Belgique, Bruxelles, 1: 1-923.

1959, 1960. Exploration du Parc National de l’U pemba. Araneae, Lycosifor-
mia II (Lycosidae). Institut des Parcs Nationaux du Congo Beige, Brux-
elles. 1-518 (1958), 519-1040 (1959).

Rovner, J. S.

1975. Sound production by wolf spiders: a substratum-coupled stridulatory
mechanism. Science, 190: 1309-1310.

Simon, E.

1864. Histoire naturelle des Araignees (Araneides). Paris. 540 pp.

Stone, W.

1890. Pennsylvanian and New Jersey spiders of the family Lycosidae. Proc.
Acad. Nat. Sci. Phila. 1890(3): 420-434.

Walckenaer, C. A.

1837. Histoire naturelle des insectes: apteres. Tome I. Paris, 549 pp.
Whitcomb, W. H.

1967. Wolf and lynx spider life histories. Univ. Arkansas Div. Agr., Dept.
Ent., Term Rept. NSF Grants, 141 pp.

1986]

319

Brady — Nearctic Gladicosa

Whitcomb, W. H. and K. Bell

1964. Predaceous insects, spiders, and mit^s of Arkansas cotton fields. Agr.
Exp. Sta., Univ. Ark. Bull., 690: 3-84.

Whitcomb, W. H., H. Exline and R. C. Hunter

1963. Spiders of the Arkansas cotton field. Ann. Ent. Soc. Amer., 56: 653-660.
Wood, F. D.

1926. Autotomy in Arachnida. Jour. Morph. 42: 143-195.

Worley, L. G. and G. B. Pickwell

1931. The spiders of Nebraska. Univ. Stud. Nebraska, 27(1-4): 1-129.

NESTING ASSOCIATIONS OF WASPS AND ANTS
ON LOWLAND PERUVIAN ANT-PLANTS

By Edward Allen Herre,12 Donald M. Windsor,1
and Robin B. Foster1

Introduction

Neotropical vespid wasps are known to form nesting associations
with other species of wasps and ants. For instance, Mischocyttarus
immarginatus nests primarily in association with the larger and
more aggressive colonies of certain polybiine wasps in the savannas
of northwestern Costa Rica (Windsor 1972, 1973). Examples of
wasp species which form interspecific nesting associations with ants
include Polybia rejecta and Synoeca chalybea, whose nests are usu-
ally associated with carton building Azteca spp. ant colonies
throughout the neotropics (Vesey-Fitzgerald 1938, Richards 1945,
DMW, RBF, EAH personal observations).

Often the ants with which wasps nest are involved in more or less
specific associations with host ant-plants. In addition to Azteca
spp., Polybia rejecta nests can be found in ant acacias which support
healthy colonies of Pseudomyrmex ferruginea (DMW, RBF per-
sonal observations). Zikan (1949) has reported that several Mischo-
cyttarus species nest on the ant plant Cordia nodosa (Boraginaceae)
inhabited by Azteca spp. and an unidentified species of myrmecace-
ous Melastomataceae. Richards (1945) reported collecting at least
one nest of M. metoecus and M. decimus from C. nodosa in
Guyana.

Why do these nesting associations exist? Windsor (1972, 1973)
demonstrated that Mischocyttarus immarginatus nests associated
with nests of other, more aggressive wasps species survive longer
and produce more brood. It appears that such nests suffer less dam-
age from birds which destroy nests and rob brood. Richards (1951)
suggested that nesting with ants such as Azteca may be one of
relatively few possible defenses available to tropical wasps against

'Smithsonian Tropical Research Institute, Box 2072, Balboa, Republic of Panama
department of Biology, University of Iowa, Iowa City, Iowa 52242, U.S.A.
Manuscript received by the editor March 20, 1986

321

322

Psyche

[Vol. 93

the organized raids of army ants (Ecitonini). Below we describe
numerous species of vespid wasps which form nesting associations
with Allomerus octoarticulatus ants inhabiting the plant Tococa
guianensis Aublet (Melastomataceae) and with Pheidole spp. ants
inhabiting Maieta poeppigii Mart, ex Triana (Melastomataceae)
and show that by nesting on these plants the wasps escape army ant
raids.

Study Area

The study site is in Loreto, Peru, at Estacion Biologica Callicebus
which is located 3-5 km. south of the village of Mishana on the Rio
Nanay in Loreto, Peru. The village is approximately 30 km. east of
the confluence of the Nanay with the Amazon River near Iquitos.
The Estacion consists of a forest camp and an extensive trail net-
work through the apparently uncut and non-indundated forest. The
forest grows on low hills composed of a mosaic of white sand and
dark brown sand and is drained by tea-colored streams. These sedi-
ments are derived from the ancient Guiana and Brazilian Shields
and have been eroded and redeposited following the Andean uplift.
White sand areas, though common in the Rio Negro drainage, are
infrequent in the Western Amazon (see Kinsey & Gentry, 1979). The
brown sand soils support a flora typical of much of the non-
indundated Peruvian Amazon. The white sand soils have a distinc-
tive flora which shows strong affinities to the flora of the Guiana
Highlands. The latter areas also have a shorter forest canopy (20 m.
vs. 30-35 m.); fewer lianas, straighter, thinner, and less-branched
understory trees and shrubs; and a thick mat of roots over the white
sand. The observations presented below were collected during 8
short visits (4 by EAH in August 1978, October 1978, in December
1979, and in June 1983; 1 by DMW in November 1978; and 3 by
RBF in August 1974, 1978, and 1980).

Observations and Results

Understory ant-plants are common at Mishana, especially on the
brown sand soils, and are represented by a diversity of families:
(Melastomataceae) Tococa (3 spp.), Maieta (2 spp.); (Chrysobala-
naceae) Hirtella (2 spp.); (Boraginaceae) Cordia nodosa ; (Rubia-
ceae) Duroia hirsuta. Of these, only Tococa guianensis is abundant
on the white sand areas. This 2-4 m. treelet is most common on

1986] Heere, Windsor, and Foster — Wasps and ants 323

upper slopes and hilltops, primarily in gaps formed by treefalls. The
population is polymorphic for a bright red-purple color on the
undersides of the leaves. The petioles have a large, bilobed, hollow
expansion (formicarium) with a pair of openings onto the under-
sides of the leaf blade (see figure 1). Maieta poeppigii, in
contrast, is an arching shrub less than 1 m. tall most common on
brown sand soils, primarily on lower slopes and streambanks. The
formicarium of Maieta consists of a pair of raised, hollow chambers
on either side of the midrib at the base of the leafblade.

Although we found colonies of at least eight different species of
ants inhabiting different Tococa guianensis individuals (eg. Azteca
spp., Dolichoderus spp., Pseudomyrmex spp., Crematogaster spp.,
and Gnamtogenys spp.), the majority of the plants we encountered
were occupied by colonies of the ant Allomerus octoarticulatus
(Allomerus) (18 of 34 plants in one census). Allomerus builds a
characteristic carton tunnel of cemented debris with small holes
regularly spaced over the surface (see figure 1). These structures
envelope most of the stems, connect the formicaria, and extend
down the main stem to within 20 cm. of the ground. Most ant
activity is confined to the formicaria and these tunnels. Unlike
other species of ants which we observed on these plants, we did not
observe Allomerus foraging off the host plant either on any casually
observed plants or on focal plants watched at hourly intervals
between 5 am. and 1 am. In addition, the presence of coccids and
structures which may have been food bodies or feeding glands for
the ants (see Roth, 1970) led us to believe that Allomerus derives all
its nutrition either directly or indirectly from the host T. guianensis,
much as Pseudomyrmex satanica is supported by farming coccids
within the hollow outer twigs of Triplaris cumminghami (DMW,
personal observation in Costa Rica). Allomerus aggressively recruits
onto leaf surfaces when a plant is disturbed. However, the ants do
not harm wasp broods although they will swarm all over the wasp
nest.

Maieta poeppigii plants were overwhelmingly occupied by Phei-
dole spp. ants (94 of 101 plants). Unlike Allomerus, the Pheidole
spp. do not build tunnels, although they do characteristically store
debris in one of the two paired chambers at the base of each leaf.
The Pheidole spp. ants are not particularly aggressive. Occasional
minor workers can be found outside the formicaria. Major workers
and minor workers emerge from the formicaria in large numbers

324

Psyche

[Vol. 93

Figure 1. Nests and adults of the wasp Mischocyttarus inSolitis shown beneath
the leaves of Tococa guianensis plants inhabited by Allomerus octoarticulatus ants.
Notice the separation of the cells on the multiple pedicels. Also notice the characteris-
tic carton tunneling and formicaria used by the Allomerus ants

1986] Heere, Windsor, and Foster — Wasps and ants 325

only if the leaves are violently shaken or if the formicaria are
directly disturbed. The Pheidole ants were not observed on the wasp
nests.

Of those wasps which construct small open nests, we found ten
Mischocyttarus species and one Polistes species in the forest under-
story. All but two of thirty-one active colonies encountered
occurred on the undersides of Tococa guianensis or Maieta poeppi-
gii leaves and only when the plants were occupied by Allomerus and
Pheidole ants, respectively (see table 1). Two of the Mischocyttarus
wasp species, M. latissimus and M. insolitis, build multi-pediceled
nests arranged in rows along the midrib of the leaf. The cells are
fused in the nests of M. latissimus while the nests of M. insolitus
consist of separate clusters of one to four brood cells with each
cluster supported by its own pedicle (see photograph 1). The net
result is the subdivision and separation of the broodcells which
comprise the nests of M. insolitus. The other species of Mischocyt-
tarus build nests more typical of the genus; a cluster containing all
cells supported by a single pedicel. All of the Mischocyttarus. spe-
cies are extremely timid, flying away from their nests at the slightest
disturbance and making no attempt at brood defense. In addition,
two colonies (one each of Polybia signata and Polybia spp.) out of
six total colonies of socially complex, aggressively swarming Poly-
biinae wasps were found attached to limbs of T. guianensis.

A small number of Tococa plants supported a disproportionate
number of wasp colonies and this was most obvious with the nests
of Mischocyttarus insolitis. In a census of 43 T. guianensis plants
with Allomerus ants, five plants were the host for single Mischocyt-
tarus nests while seven plants had two or more colonies. In a survey
of 1 16 Maieta poeppigii plants with Pheidole spp. ants, one plant
had three nests and two plans each had one.

Several observations and manipulations we performed indicate
that by nesting on these myrmecacious melastomes the wasps avoid
nest plundering by army ants. While following the raiding swarms
of Eciton burchelli and Eciton rapax we noticed that these ants
never ran on to either Tococa guianensis or Maieta poeppigii plants.
The avoidance of these two plants contrasted sharply with the army
ants’ rapid climbing and investigating most other plants in their
path.

326 Psyche [Voi. 93

Table 1. A list of wasp species collected at Mishana, 1-5 November 1978. The
number of nests found on each type of host plant ( Maieta , Tococa, or other) is
indicated. Brachygastra melania was previously only known from Bolivia.

Species with open nests:

Nest

Found

Maieta Tococa other

Mischocyttarus synoecus Rich.

Yes

M. lecointei Ducke

4 2 1

M. pallidus Zikan

M. insolitus Zikan

M. latissimus Rich.

M. decimus Rich.

M. sp. near mirificus Zikan

M. carbonarius Sauss.

M. silvieola Zikan

M. sp. near interruptus Rich.

Polistes rufiventris Ducke

P. pacificus

Species with closed nests:

(found in clearing near river)

Angiopolybia pallens Lep.
A. paraensis Spirola

morph ruficornis Ducke

Apoica thoracica R. du Buyss.

Brachygastra bilineolata Spinola

B. buyssoni Ducke

B. melania Richards

B. moebiana Sauss.

B. myersi Bequaert

Polybia signata Ducke

P. sp. near fastidiosuscula Sauss

P. sp.

1 1

P. rejecta F.

P. liliaceae F.

Protopolybia acutiscutis Cameron

Pseudopolybia vespiceps Sauss.

Stelopolybia angulicollis Spinola

Synoeca surinama Lep.

S. virginea F.

1986] Heere, Windsor, and Foster — Wasps and ants 327

The perceived avoidance was substantiated when we moved a
twig that the Eciton ants were using as a bridge against a stem of M.
poeppigii. The army ants stopped when they came in contact with
the stem and although ants from the rear continued moving forward
until there was a great tangled mass of ants at the front, no ants
crawled onto the stem. Next, we placed stems of Tococa guianensis
and Maieta poeppigii with intact leaves and formicaria across active
Eciton trails and found that the trails were quickly rerouted around
the plants. Similar responses were not obtained when we placed
other plant species or Tococa guianensis without Allomerus inhab-
itants across the path of the army ants. Further, in three instances,
we observed army ants passing by T. guianensis plants with Allome-
rus ants and active wasps nests. We removed two T. guianensis
leaves minus formicaria with attached wasp nests, placed them on
twigs at the same height off the ground as they had been on the
plants, and put the twigs in front of the Eciton raiding swarms. In
both instances the army ants swiftly scaled the twigs and seized the
wasp brood.

Discussion

Predatory ants pose a particularly important threat to the nests
and broods of tropical wasps (Jeanne 1972, Litte 1977). In discuss-
ing this problem in his revision of the genus Mischocyttarus,
Richards (1945) states, “A number of species have entered into some
sort of association with ants and have thereby found safety by firmly
grasping the nettle.” Clearly the wasps nesting on these plants
benefit by having a neutral border maintained for them. With access
to the sole connection to the terrestrial world guarded by Allomerus
or Pheidole ants, there is little or no risk that hostile army ant
species will come plundering down the pedicel. In this light the unus-
ual (for Mischocyttarus wasps) nest architecture of M. insolitis
becomes more comprehensible.

As Jeanne (1979) demonstrated, building a highly subdivided nest
composed of multiple combs uses materials for nest construction
very inefficiently and requires a much higher expenditure of time
and energy per cell than does the nest architecture more characteris-
tic of polistine wasps. However, a highly subdivided nest no longer
provides as concentrated a target for a bird which plunders by
knocking down whole nests and then leisurely eating the brood (eg.

328

Psyche

[Vol. 93

Windsor 1972). More passes are needed and the return in food per
time and effort is less. Further, a subdivided nest is less vulnerable
to being entirely wiped out by nest parasites which can move from
cell to cell (eg. tineid moth larvae described by Jeanne 1979).

There appears to be no obvious benefit that the ants derive from
the presence of the timid Mischocyttarus wasps. Why do the ants
tolerate the presence of these wasps? A review of the ant species with
which various vespid wasps are reported to form nesting associa-
tions shows that with the exception of some Azteca species, the ants
all appear to be nutritionally supported by their host plants. Appar-
ently the ants either cannot eat the wasp brood or do not recognize
the wasp brood as a potential meal. Further, in the case of some
ant-wasp associations such as that between Azteca spp. and Polybia
rejecta, the wasps have been reported to benefit the associated ant
colony by discouraging anteaters (R. Silberglied, personal commun-
ication). The Tococa guianensis plants on which the aggressive Poly-
bia and Polistes wasps nested were difficult to approach without
being stung. It is likely that the presence of these wasps reduces
damage to the host plant and, consequently, the ant colony caused
by mammals. Therefore, the Mischocyttarus wasps, while not being
a detriment to the ants, may simply be taking good advantage of a
tolerance that the ants have developed to more beneficial species of
symbiotic wasps.

Acknowledgments

We wish to thank Don Francisco Pizarro for generous hospitality
and essential logistic help during our various visits to Casaria
Mishana, Dr. William Brown for kindly indentifying all ant species
mentioned, Dr. J. J. Wurdack for identifying the Tococa and
Maieta species, and, especially, the late O. W. Richards for identify-
ing the wasp species and encouraging this work with his enthusiasm
and expertise. D. E. Wheeler, D. M. Feener, L. Johnson and the
Iowa Writing Seminar Group provided helpful comments on the
manuscript. This work was supported by The Smithsonian Tropical
Research Institute (DMW, EAH), The Harris Foundation (EAH),
and The University of Iowa’s Teaching and Research Fellowship
Program (EAH).

1986] Heere, Windsor, and Foster— Wasps and ants 329

Summary

Twelve species of vespid wasps were found nesting on two species
of melastomataceous ant plants in a mixed lowland forest near
Iquitos, Peru. Although eight different species of ants inhabited
different individual plants of Tococa guianensis (Melastomataceae),
wasps only nested on those plants inhabited by the ant Allomerus
octoarticulatus. Nests were also found on Maieta poeppigii
(Melastomaceae) inhabited by Pheidole spp. Several Mischocyttarus
species exhibited nest architectures atypical of the group. Observa-
tions and manipulations indicate that by nesting on these ant plants
inhabited by those particular ants the wasps avoid nest plundering
by army ants.

References

Jeanne, R. L.

1972. Social Biology of the Neotropical Wasp Mischocyttarus drewsenii. Bull.

Mus. Comp. Zool., Harvard Univ. 144: 63-150.

1979. Construction and Utilization of Multiple Combs in Polistes canadensis
in Relation to the Biology of a Predaceous Moth. Behav. Ecol. Socio-
biol. 4, 293-310.

Kinsey, W. G. and A. H. Gentry

1979. Habitat utilization in two species of Callicebus. in Primate Ecology:
Problem Oriented Field Studies. R. W. Sussman ed. Wiley and Sons.
New York.

Litte, M.

1977. Behavioral ecology of the Social Wasp Mischocyttarus mexicanus.
Behav. Ecol. Sociobiol. 2: 229-246.

Richards, O. W.

1945. A Review of the Genus Mischocyttarus de Saussure. Trans. Roy.
Entomol. Soc. London. 95: 295-462.

Richards, O. W. and M. J. Richards.

1951. Observations on the Social Wasps of South America (Hymenoptera,
Vespidae). Trans. Roy. Entomol. Soc. London. 102: 1-170.

Roth, I.

1976. Estructura Interna de los Domacios Foliares en Tococa (Melastomeae).
Acta Biol. Venez., 9(2): 221-258.

Vesey-Fitzgerald, D.

1938. Social Wasps (Hym. Vespidae) from Trinidad, with a note on the genus
Trypoxylon Latreille. Trans. Roy. Entomol. Soc. London. 87: 181-191.
Windsor, D. M.

1972. Nesting Association between two Neotropical Polybiine Wasps (Hyme-
noptera, Vespidae). Biotropica 4: 1-3.

330

Psyche

[Vol. 93

1973. Birds as Predators on the Brood of Polybia Wasps (Hymenoptera: Ves-
pidae: Polistinae) in a Costa Rican Deciduous Forest. Briotropica 8(2):
111-116.

ZlKAN, J. F.

1949. O Genero Mischocyttarus Saussure (Hymenoptera, Vespidae), com a
decriciao de 82 especies novas. VBoln. Parq. nac. Itatiaia 1: 1-251.

WINTER PREY COLLECTION AT A PERENNIAL COLONY
OF PARAVESPULA VULGARIS (L.)
(HYMENOPTERA: VESPIDAE)

By Parker Gambino
Department of Entomological Sciences
University of California
Berkeley, California, U.S.A. 94720

Introduction

Diet is a fundamental aspect of an organism’s biology. In euso-
cial vespid wasps the food intake of a mature colony, including
nutrition of immatures, is determined by the foraging behavior of
workers. Yellowjackets of the genus Paravespula Bliithgen meet
the protein requirements of the colony by capturing live arthropods
and collecting flesh from dead animals. By enabling these species to
utilize a broader resource base, scavenging likely contributed to the
evolution in this genus of a colony cycle characterized by higher
worker populations and greater longevity than in Vespula Thomp-
son, a closely related genus in which only live prey is taken (Mac-
Donald et al., 1976).

Prey collection by freely foraging Paravespula colonies has been
described in detail by Kleinhout (1958), Kemper and Dohring
(1962), Broekhuizen and Hordijk (1968), and Archer (1977).
Numerous shorter lists of prey are available, (cf. Spradbery (1973)
for a literature review). Broekhuizen and Hordijk (1968) investi-
gated the response of P. vulgaris (L.) to artificial manipulations of
prey densities in trees, while MacDonald et al. (1974) offered var-
ious prey items in screen-enclosed foraging areas. Heinrich (1984)
gave a good account of general foraging behavior of individual
workers and Free (1970) investigated handling of honeybee prey by
workers.

Paravespula species undergo an annual monogynous cycle over
most of their range, but in mild-weathered areas, perennial polygy-
nous colonies sometimes develop (Spradbery, 1973). These colonies,
characterized by enormous populations of workers, occur especially

* Revised manuscript received by the editor July 7, 1986.

331

332

Psyche

[Vol. 93

in areas recently invaded by Paravespula. Although perennation
and polygyny represent significant deviations from the typical Para-
vespula pattern, the sporadic appearance of such colonies has hin-
dered study. Published prey studies have addressed only annual
colonies, thus the discovery of a perennial Paravespula colony at the
University of California provided an opportunity to study its winter
diet.

Materials and Methods

I first noticed the colony of Paravespula vulgaris (L.) on October
10, 1984. Typical annual colonies of this native species initiated in
April or May usually begin to decline in the fall. The high level of
activity (about 300 worker sorties per minute) indicated that this
colony had been functional since at least spring 1984, and suggested
that it might persist for another year. In fact, the colony remained
vigorous through a second summer, with a final observation of
external worker activity on February 6, 1986.

The colony was located about 25 m north of Callaghan Hall ticket
kiosk on the Berkeley campus of the University of California, at
elevation 75 m. The immediate surrounding area is a mixed stand of
Monterey pine ( Pinus radiata D. Don) and coast live oak ( Quercus
agrifolia Nee) over a grass ground cover dominated by Ehrharta
erecta Lam. Strawberry Creek, flowing basically east to west, passes
within 40 meters.

The subterranean nest was under a fallen log about 1 meter in
diameter, which supported a lush growth of ivy ( Hedera helix L.).
Active entrance holes were at ground level on both sides of the log.
The log was well shaded, although the west entrance received some
direct afternoon sun.

To facilitate sampling, I constructed devices to restrict yellow-
jacket access to the nest at each entrance. To sample from the east
entrance, I sealed it and netted the returning foragers as they
hovered near it. After separating prey from the workers by shaking
the net, I either allowed workers to fly from the net or anaesthetized
them with carbon dioxide and removed them. A typical 40 minute
net-sampling session involved approximately 40 sweeps of the net.

Beginning April 5, 1985, I used a modified funnel trap to collect
from the west entrance. This passive method was more efficient at
collecting foragers returning with prey. The trap was left in place

1986] Gambino — Winter prey of Paravespula 333

approximately 15 minutes per sampling session. Captured workers
were anaesthetized and shaken from the trap. Anaesthetized
workers were returned to the vicinity of the nest entrance for which
they had been bound. Items separated from the workers were
immediately transferred to 70% EtOH.

I identified sorted samples to lowest feasible taxonomic levels
with the assistance of workers at the Essig Museum, University of
California, Berkeley. An item which was recognizable as a single
prey load was counted even if it was only a fragment of an organism.
For example, a honeybee abdomen counted as one record of Apis
mellifera L.

I visited the colony to observe wasp behavior daily from January
5 to May 10, 1985, and sampled approximately weekly. Time of day
and environmental conditions during sampling varied somewhat,
but most sessions were during the early to mid afternoon of bright
sunny days.

Results

I analyzed a total of 1306 items, many of which were only frag-
ments and/or badly mauled. Precision of identification was variable
. Thus, while some relatively intact prey items could be identified to
species, other more macerated fragments of arthropods could not be
identified below phylum. Because there is no way to know which
items were captured live (predation sens, str .) and which were scav-
enged, I classified all food items as prey. No items of food made or
prepared by humans were identified.

The 914 prey items that could be identified at least to order are
summarized in Table 1 according to taxa and collection dates.1
Temporal variation of selected prey items in the colony’s diet, illus-
trated in Figure 1, reflects the sequential availability of potential
prey species, based on their life history patterns.

I began the study during the flight period of the sawfly, Xyela
radiatae Burdick, when adults were so abundant that they actually
crawled into my net during several collection sessions. Accordingly,
X. radiatae was the dominant prey item in January (79% of deter-
mined specimens). Yellowjackets commonly hunted in the short

'A more detailed list of prey is available from the author on request.

Table 1. Prey items taken from Paravespula vulgaris foragers. Totals are inclusive; each higher category includes numbers
of identified prey in lower categories within the hierarchy, if any.

DATE

January February March April May

Prey Item 1 5 10 16 22 29 4 8 15 22 17 15 22 29 5 12 20 26 3 10

334

Psyche

[Vol. 93

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1986]

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Gambino — Winter prey of Paravespula 335

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Table 1. (con’td)

336

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[Vol. 93

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1986] Gambino — Winter prey of Paravespula 337

grass near the colony, especially beneath Pinus radiata , the xyelid’s
host plant (Burdick, 1961). Workers flying close to the ground thor-
oughly scanned plant surfaces, paying special attention to areas of
contrasting colors and textures to locate and capture surface-
inhabiting arthropods, which comprised the vast majority of the
colony’s prey. I observed attempted and successful captures of X.
radiatae adults on grass blades. These sawflies, presumably newly
emerged from underground pupae, seemed especially vulnerable to
Paravespula predation.

In late February, the beetle, Byturellus grisescens (Jayne),
reached its greatest abundance in the prey samples. Although I was
unable to capture any of these beetles myself, they oviposit on oak
catkins (J. Doyen, pers. comm.). This was the first good suggestion
of the importance of oak insects in the diet of the colony. Of the
many tree species occurring on the University campus, Q. agrifolia,
a native, is one of the most common.

As the season progressed, hunting at ground level became less
frequent, and foragers shifted their attention to tree foliage, particu-
larly Q. agrifolia . Local population explosions of caterpillars (Lepi-
doptera) in late March and April, and the treehopper, Cyrtolobus
vanduzeei Goding, in May were also tracked by this colony (Fig. 1).
Again, most of the identifiable Lepidoptera and Membracidae were
of taxa known to be associated with Q. agrifolia.

Discussion

The wide taxonomic array of arthropod prey and focus on
abundant prey species shown by the observation colony are consist-
ent with known habits of the genus Paravespula. Scavenging, a
characteristic of the genus, is suspected in the collection of pieces of
earthworm and Apis mellifera, as well as some other items which
were tangled in silk strands and may have been taken from spider
webs. Collection of proteinaceous food prepared for human con-
sumption, a habit accounting for the pest status of P. vulgaris in
many areas (MacDonald et al., 1976) was not detected. Although
such food was certainly within the flight range of foraging workers,
it was not common in the immediate vicinity of the colony, and
foragers may have become conditioned to locate arthropod prey. In
general, the data from the observation colony indicate that the flesh

338

Psyche

[Vol. 93

»-

IN 3 OH 3 d

Figure 1. Temporal variation in prey composition. Percentages of identified items comprised of Xyela radiatae Burdick,
Byturellus grisescens (J ayne), Lepidoptera larvae, and Cyrtolobus vanduzeei Goding.

1986] Gambino — Winter prey of Paravespula 339

collection behavior of perennial colonies probably does not differ
substantially from that of annual colonies.

Summary

This study suggests that in coastal California natural food resour-
ces are sufficient to sustain healthy overwintering Paravespula colo-
nies. The P. vulgaris colony under study took a wide assortment of
prey, and adjusted its diet according to local abundances of prey
species. Q. agrifolia, a native tree common in the vicinity of the
colony, was the source of many of the insects comprising its diet.

Acknowledgments

The success of this study resulted from the contributions of many
co-workers in the Department of Entomological Sciences, Univer-
sity of California, Berkeley. Vernard Lewis discovered the colony,
and Tina Sterret provided technical assistance. Howell Daly, John
Doyen, Jerry Powell, Evert Schlinger, Stuart McKamey, Woodrow
Middlekauff, and Jim Whitfield assisted in identifying prey. Howell
Daly, John De Benedictis and Woodrow Middlekauff reviewed the
manuscript and offered suggestions for its improvement. Financial
support was furnished in part by the Northern California chapter of
the ARCS Foundation.

Literature Cited

Archer, M. E.

1977. The weights of forager loads of Paravespula vulgaris (Linn.) (Hymenop-
tera: Vespidae) and the relationship of load weight to forager size. Ins.
Soc. 24(1): 95-102.

Broekhuizen, V. S., AND C. Hordijk

1968. Untersuchungen iiber die Beute von Paravespula vulgaris L. (Hym.,
Vespidae) und ihre Abhangigkeit von der Beutetierdichte. Z. Ang.
Entomol. 62: 68-77.

Burdick, D. J.

1961. A taxonomic and biological study of the genus Xyela Dalman in North
America. Univ. Calif. Publ. Entomol. 17(3): 285-356.

Free, J. B.

1 970. The behavior of wasps ( Vespula germanica L. and V. vulgaris L.) when
foraging. Ins. Soc. 17(1): 1 1-20.

Psyche

[Vol. 93

340

Heinrich, B.

1984. Strategies of thermoregulation and foraging in two vespid wasps, Doli-
chovespula maculata and Vespula vulgaris. J. Comp. Physiol. B 154:
175-180.

Kemper, V. H., and E. Dohring

1962. Untersuchungen liber die Ernahrung sozialer Faltenwespen Deutsch-
lands, inbesondere von P. germanica und P. vulgaris. Z. Ang. Zoologie
49(2): 227-280.

Kleinhout, J.

1958. Het verzmelen van prooien van sociale wespen. De Levende Natuur 61:
179-182.

MacDonald, J. F., R. D. Akre, and W. B. Hill

1974. Comparative biology and behavior of Vespula atropilosa and V. pensyl-
vanica (Hymenoptera: Vespidae). Melanderia 18: 1-65.

MacDonald, J. F., R. D. Akre, and R. W. Matthews.

1976. Evaluation of yellowjacket abatement in the United States. Bull.
Entomol. Soc. Amer. 22(4): 397-401.

Spradbery, J. P.

1973. Wasps. University of Washington Press, Seattle, xvi + 408 pp.

YOUNG LARVAE OF ECITON
(HYMENOPTERA: FORMICIDAE: DORYLINAE)1

By George C. Wheeler and Jeanette Wheeler
3358 NE 58th Avenue,

Silver Springs, Florida 32688

I. Instars

In our previous studies of ant larvae we have been concerned
primarily with generic characterizations and differences based on
mature larvae. We described immature stages when available, which
wasn’t often. And even when we did, we didn’t know the instars.
Never have we had a complete larval series from egg to semipupa.
Yet many authors have stated quite glibly the number of larval
instars. At least it seems glib to us, for we consider it hard work to
establish the number of instars. To do this we require that following
specimens: a first-instar larva inside an egg ready to hatch; a second
instar larva inside a first ready to moult; a third-instar inside a
second-instar ready to moult; etc.; and finally a mature larva. How
can we prove maturity? By comparison with a semipupa, which will
reveal all characters of a mature larvae except shape. For further
confirmation one should have a worker pupa or a worker to verify
size. The identification of sexual larvae presents a further complica-
tion. If the larva is larger than a worker semipupa it is probably a
sexual or at least a queen. In most species we have not been able to
recognize younger sexual larvae.

In polymorphic species (e.g., Eciton, Atta, Acromyrmex, Cam-
ponotus) such procedures are even more difficult. How can one tell
whether a small larva is a young major or a mature minum or
whether a large larva is a half-grown major or a mature intermediate?

Two interesting papers afford a partial solution to this problem:
Tafuri (1955) on Eciton hamatum and Lappano (1958) on E. bur-
chelli

Eciton is an ideal genus for such a study: there can be no mixing
of broods; except for one all-sexual brood per year, all larvae will

1 Manuscript received by the editor June 30, 1986.

341

342

Psyche

[Vol. 93

become workers. All one needs to determine, then, is whether the
larvae in such a brood foreshadow adult polymorphism, and if so,
how? The solution depends upon the fact that at the middle of the
statary phase the queen lays during one week a single batch of 60,000
to 130,000 eggs and then no more until the next statary phase.

“In E. hamatum the adult polymorphic workers form a continu-
ous series from the smallest worker minor to the largest soldier
form. . .. Besides differences in size there are apparent qualitative
differences in this series marked primarily by the exceptional
hooked manidbles and head pattern of the major workers.” (Tafuri
1955: 32.) In the larvae, however, such differences “are not noticea-
bly apparent.” Any distinction of growth stages (i.e. nomadic days)
is impossible on the basis of body size alone, because of overlap-
ping. The larvae likewise form a smooth series from the smallest to
the largest forms. The author therefore based his determination of
larval age (in nomadic days) on the allelomorphic growth of the
imaginal leg discs.

“[It] is highly probable that the largest larvae of any stage have
developed from the eggs first to be laid and first to hatch and
represent the potential major workers of the mature brood. Sim-
ilarly, the smallests [larvae] presumably develop from the eggs last
to be laid and last to hatch and represent the potential workers
minima of the mature brood.” (Lappano 1958: 49).

From these two articles we get the impression that larval devel-
opment in Eciton is a smooth process from hatching to pupation
without any such interruptions as molts. The word “instar” is not
found in either of these articles.

So we re-examined our supply of doryline larvae and found
graded series of larvae of Eciton hamatum sent to us by the late Dr.
T. C. Schneirla (including some of the sample studied by Lappano)
from Barro Colorado Island (Panama) and a similar supply of E.
burchelli larvae collected in Trinidad by Dr. N. A. Weber.

The great advantage of the Tafuri/ Lappano method is that it
requires no technique and can be applied to either living or pre-
served material. However, after applying our tedious technique
(Wheeler and Wheeler 1960) of cleaning, staining and mounting in
balsam, we found that we had the prerequisite for identifying all
instars, except the mature larva, which we had already studied
(Wheeler and Wheeler 1984). We should warn, however, that the

343

1986] Wheeler & Wheeler — Larvae of Eciton

preparation of these immature was the most difficult we have ever
experienced.

II. Interspecific Differences

Whenever we have had two or more species in the same genus, we
have either given a complete description of each or at least men-
tioned differences. We have not been willing to go beyond that,
because we did not know the extent of intranidal or internidal or
intraspecific differences. Here at last, we have series of Eciton bur-
chelli and E. hamatum which embolden us to make a tentative
comparison. Table 1 gives a few characters which can be measured
for each instar in each species. The “spiracle diameter” which we
have not mentioned previously, is the diameter of the atrium and
not of the opening into it.

Eciton burchelli Westwood
Figure 1

Egg. About 0.3 X 0.54 mm.

First Instar Larva. Length 0.59-0.9 mm long (average 0.73
mm). Head greater in diameter than body which tapers to the poste-
rior end. Anus subterminal. Segmentation distinct. Spiracles about
0.001 mm in diameter. Entire integument sparsely spinulose, the
spinules minute and isolated. Body hairs lacking. Cranium subcircu-
lar. Antennae minute, just above midlength of cranium. Head hairs
lacking. Labrum arcuate, about 1/4 width of cranium; with a few
spinules and sensilla on and near ventral surface. Mandible with
straight apical tooth which is feebly sclerotized, remainder not
sclerotized. Maxilla broadly paraboloidal and appearing adnate;
palp represented by 7 sensilla in a loose cluster; galea represented by
2 sensilla. Labial palp represented by 3 sensilla; opening of sericter-
ies a short transverse slit.

Second Instar Larva. Length (through spiracles) 0.9- 1.5 mm
(average 1.2 mm). Head same diameter as T1 and AV, the widest
parts of the body. Spiracles about 0.003 mm in diameter. Entire
integument spinulose, the spinules moderately abundant and iso-
lated. Body hairs 0.006-0.012 mm long; few, most on Tl, fewer on
T2 and T3. Cranium subhexagonal; integument with a few spinules.
Antennae just above midlength of cranium. About 30 head hairs;

[Vol. 93

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1986] Wheeler & Wheeler — Larvae of Eciton 345

Figure 1. Eciton burchelli. BI, first instar larva; BII, second instar larva;
Bill, third instar larva; BIV, head of fourth instar larva; BV, head of fifth instar
(= mature) larva. Side views, X 38; head in anterior view, X 50. Be, egg nearly ready
to hatch, X 38. bl-b5, larvae of the five instars in side view to show relative sizes, X 9.

0.006-0.019 mm long; simple. Labrum feebly bilobed. Otherwise
similar to first instar larva.

Third Instar Larva. Length (through spiracles) 1. 5-2.9 mm.
Spiracles about 0.013 mm in diameter. Integumentary spinules more
conspicuous and in rows. Body hairs 0.037-0.075 mm long; more
numerous but largely confined to thorax. Cranium subhexagonal
and with bulging genae. Head hairs 0.036-0.075 mm long, slender
and flexuous; some dorsal hairs curved downward and a few ventral
upward; about 60 present. Labrum with transverse rows of spinules
on anterior and posterior surfaces on and adjacent to ventral sur-
face; median sulcus with about 10 sensilla on and near ventral sur-
face. Mandible with apical tooth slightly curved medially and with

346

Psyche

[Vol. 93

medial border erose. Maxilla with apex narrowly paraboloidal and
bearing rather long spinules in short transverse rows; palp repres-
ented by a cluster of 8 sensilla; galea a slight elevation with 2 sen-
silla. Anterior surface of labium with minute spinules in short
transverse rows. Otherwise similar to second instar larva.

Fourth Instar Larva. Length (through spiracles) 3.2-7 mm.
Diameter uniform. Spiracles about 0.019 mm in diameter. Integu-
ment with minute spinules in transverse rows. Body hairs 0.025-0.15
mm long; on all somites but most numerous on T1-T3 and AVIII-
AX. Head hairs 0.025-0.1 mm long; about 100; several ventral hairs
curved upward. Labrum with lateral borders sinuate. Maxillary
palp represented by a cluster of 9 sensilla; galea a short sclerotized
frustum with 2 apical sensilla. Labium with anterior surface bearing
numerous short transverse rows of spinules; opening of sericteries a
transverse slit in the bottom of a depression. Otherwise similar to
third instar larva.

Mature larva. Length (through spiracles) 5-9.2 mm. Compared
to E. hamatum in our 1984: 270.

Material studied: numerous larvae from Trinidad, courtesy of Dr.
N. A. Weber.

Eciton hamatum (Fabricius)

Fig. 2

Egg. About 0.25 X 0.5 mm.

First Instar Larva. Length 0. 5-0.9 mm. Head of same diameter
as Tl; body straight, diameter decreasing posteriorly. Spiracles
about 0.001 mm in diameter. Entire integument spinulose, the spin-
ules minute and isolated. No body hairs. Cranium transversely sub-
elliptical. Antennae above midlength of cranium. No head hairs.
Labrum cresentic. Mandibles subtriangular, with straight apical
tooth, feebly sclerotized. Maxilla with broadly paraboloidal apex,
appearing adnate; palp represented by a cluster of 6-8 sensilla; galea
represented by 2 sensilla. Labial palp represented by 3 sensilla;
opening of sericteries very short.

Second Instar Larva. Length (through spiracles) 1.3- 1.8 mm.
Body of nearly uniform diameter. Spiracles 0.006 mm in diameter.
Integument coarsely spinulose, the spinules isolated. Body hairs
0.006 mm long, simple, few, mostly on venter of TL Cranium trans-
versely subelliptical. Head hairs about 0.006 mm long, simple, about

347

1986]

Wheeler & Wheeler — Larvae of Eciton

Figure 2. Eciton hamatum. HI, first instar larva; HII, second instar larva; Hill,
third instar larva; HIV, fourth instar larva; HV, head of fifth instar (= mature) larva.
Side views, X 38; heads in anterior view, X 50. He, larva inside egg with mouth parts
breaking shell, X 38.

12. Maxillary palp represented by a cluster of 10 sensilla. Otherwise
similar to first instar larva.

Third Instar Larva. Length (through spiracles) 2. 5-3. 5 mm.
Widest at AVII. Spiracles about 0.01 mm in diameter. Body hairs
0.013-0.037 mm long, most on thorax and AI and a few of A VI- AX.
Cranium slightly broader than long. Head hairs 0.013-0.037 mm
long, about 50. Labrum feebly bilobed. Mandible with apical half
more narrowly tapered to a sharp point, apex straight. Maxillary

Psyche

348

[Vol. 93

palp a cluster of 8 sensilla. Labium with a few minute spinules
medially. Otherwise similar to second instar larva.

Fourth Instar Larva. Length (through spiracles) 3.5-4 mm.
Spiracle diameter about 0.013 mm. Body hairs 0.025-0.05 mm long,
sparse, most numerous on T1 and AIX-AX. Head hairs 0.025-0.05
mm long; some dorsal hairs curved downward, few ventral upward;
about 100. Labrum bilobed and with sinuate lateral borders; with a
few spinules medioventrally. Mandible with apical half tapering to a
narrow sharp point and slightly curved medially. Maxillary palp a
cluster of 7 sensilla. Otherwise similar to third instar larva.

Fifth Instar Larva = Mature Larva. Length (through spiracles)
4.4-12.1 mm. Spiracles about 0.025 mm in diameter. Entire integu-
ment densely and coarsely spinulose, the spinules rather long and
the rows so close together that the spinules overlap. Body hairs
moderately numerous; 0.075-0.2 mm long, longest around anus.
Cranium with entire integument spinulose, the spinules isolated or
in rows. Head hairs 0.033-0.165 mm long; about 120; some ventral
hairs curved strongly upward. Labrum with a few sensilla ventro-
medially; spinulose, the spinules minute and isolated or in short
rows, on all surfaces. Mandible with 3-4 small denticles on apical
half. Maxilla broadly paraboloidal and appearing adnate, entire
surface spinulose, the spinules isolated or in short rows; palp a
slightly elevated sclerotized cluster of 8 sensilla; galea a small
sclerotized cone with 2 apical sensilla. Labium with entire surface
spinulose, the spinules isolated or in short rows. Otherwise as in the
fourth instar larva. See our 1984: Fig. 9 on p. 271.

Material studied: numerous larvae from Barro Colorado Island,
Panama, courtesy of the late Dr. T. C. Schneirla.

Our tentative conclusions are:

1 . In each species instars may be distinguished by spiracle diame-
ter; body hair length and distribution; head hair length and number.

2. The two species are indistinguishable in the first and fifth
instars. In the second instar they may be separated by a spiracle
diameter; uniformity in length and distribution of body hairs; length
and number of head hairs. In instar three: length and distribution of
body hairs; length and number of head hairs. In the fourth instar:
spiracle diameter; distribution and uniformity of length of body
hairs.

1986]

349

Wheeler & Wheeler — Larvae of Eciton

Literature Cited

Lappano, Eleanor R.

1 958. A morphological study of larval development in polymorphic all-worker
broods of the army ant Eciton burchelli. Insectes Sociaux 5: 31-66.
Tafuri, J. F.

1955. Growth and polymorphism in the larva of the army ant ( Eciton ( E .)
hamatum Fabricius). Jour. New York Entomol. Soc. 63: 21-41.
Wheeler, G. C. and Jeanette Wheeler.

1960. Techniques for the study of ant larvae. Psyche 67: 87-94.

1984. The larvae of the army ants: a revision. J. Kansas Entomol. Soc. 57:
263-275.

SPATIAL DISTRIBUTION OF CASTES WITHIN COLONIES
OF THE TERMITE INCISITERMES SCHWARZE

By Peter Luykx, Jack Michel, Jeannette K. Luykx* 2
Introduction

In order to describe the social organization of termites with any
precision, it is essential to have quantitative information on the
spatial distribution of castes within the colony. Such information is
important not only for descriptive purposes, but also because it can
give clues to the interactions that take place within and among the
different castes.

Precise information on caste distribution within colonies is ordi-
narily not easy to obtain, because colonies are usually completely
disrupted in opening them up, and because in any case the descrip-
tion of spatial organization in large three-dimensional or dispersed
colonies in quantitative terms is difficult. But in some locations,
colonies of certain kalotermitid species offer a unique opportunity
to obtain just such data. In the Oleta River Mangrove Preserve just
north of Miami, Florida, large numbers of Incisitermes schwarzi are
found in slender, dead mangrove tree-trunks, where they form
nearly one-dimensional colonies. Because the colonies are relatively
small and are entirely above ground, and because the termites do
not forage outside the wood, whole colonies can be collected in
segments and analyzed. The results of such an analysis are the sub-
ject of this paper.

While some of the findings of this study — the association of lar-
vae with the royal pair, the aggregation of nymphs and alates — have
been noted before in a casual way in the general descriptions of
many other students of the Isoptera (e.g., Imms, 1919; Grasse,
1949), this is the first quantitative description of the spatial distribu-
tion of castes in a termite, and is worth putting on record for that
reason.

'This is contribution no. 245 from the Program in Tropical Biology, Ecology, and
Behavior, Dept, of Biology, Univ. of Miami.

2 Department of Biology, University of Miami, Coral Gables, FL 33124, U.S.A.
Manuscript received by the editor June 7, 1986

351

352

Psyche

Materials and Methods

[Vol. 93

Colonies of Incisitermes schwarzi Banks (Kalotermitidae) were
collected from the Oleta River Mangrove Preserve, North Miami
Beach, Florida, on four collecting trips carried out between the
hours of 9 a.m. and 12 noon, at low tide, during the months of
March, April and May, 1985. (In this species, the annual reproduc-
tive cycles of different colonies are not synchronized, so that differ-
ent reproductive stages may be found at any time of the year
(Luykx, 1986).) Colonies of /. schwarzi were found only in standing
trunks, not in fallen dead wood. Small, dead mangrove trees
(Laguncularia racemosa) 3-4 cm in diameter and 1-3 m tall were
selected and quickly cut into 10-12 cm segments with a chain saw.
To minimize the possibility of redistribution of the termites during
the sectioning, the tree was not touched before the first cut was
made; the first cut was made at ground level, and all subsequent cuts
were made with the tree held horizontally (to prevent the vibration
of the saw from shaking termites from one segment to a lower
segment). Complete sectioning of each tree was accomplished with
60-90 seconds of the first cut. We estimate we might have killed
about 5% of the termites in each colony with the saw.

If a dead tree had termites (about half the ones chosen did), the
segments were put into numbered plastic bags and taken back to the
laboratory for opening and analysis. Determination of the sex and
caste of each individual in each segment was usually carried out
within one day of collection. We obtained useful data on a total of 9
complete colonies.

For the purpose of this analysis, seven castes were distinguished:
larvae (the first three instars), workers (or pseudergates: later
instars, with wing buds not readily seen with the naked eye), early-
and late-stage nymphs (the last two pre-imaginal molts, with elon-
gated wing pads easily seen with the naked eye), alates (imagos),
soldiers (small and large), and reproductives (king and queen). In
the Kalotermitidae, the larvae, workers, nymphs, and alates repres-
ent a developmental series; the only truly sterile castes are the
soldiers.

Males and females occur in all castes, with typically a slight excess
of males among the soldiers and among the nymphs (Luykx, 1987).
Except for a slight statistical tendency for soldiers of one sex to be
associated with non-soldiers of the opposite sex, the sexes within the

1986] Luykx, Michel, & Luykx—Incisitermes 353

colonies were distributed essentially at random (Luykx et al., 1987),
and will not be further considered here.

Results and Conclusions

The distribution of the castes in nine pieces of wood is represented
in Fig. 1. In eight of the nine pieces a single colony was found. In
one piece two colonies were found: PL487, and a small incipient
colony consisting only of the royal pair, one soldier, one larva, and
five workers. This small colony, PL488, contained entirely within a
single short segment, does not, of course, give any information on
caste distribution, and will not be considered further.

The major portion of most of the colonies (with the exception of
PL476 and PL486) was found toward the bottom, where the wood
was less deteriorated and less fragile. (The topmost portions of the
dead trunks are often thoroughly tunneled and in a highly deterio-
rated condition, and rarely contain any termites.) In most colonies,
the king and queen were found together in the lower part of the
colony (Fig. 2). It seems likely that the royal pair might initiate the
colony at any level in suitable dead wood, but then move down into
sounder wood as the colony grows.

Larvae (the first two or three instars) were found preferentially in
the same segments as the royal pairs. As illustrated in Fig. 2, among
the segments with reproductives, 7 out of 9 had more larvae than
expected for those segments. Twelve of the 14 segments with more
larvae than expected also had a reproductive or was adjacent to one
that did. That this is a real association, and not just a common
tendency for both larvae and reproductives to be located in the
lower parts of the colony, is suggested by colony PL476, the one
colony in which the reproductives were found in a segment in the
upper part of the colony: in this colony the larvae also were concen-
trated in this same segment (Fig. 2).

The members of the different castes representing successive stages
of development — workers, early-stage nymphs, late-stage nymphs
and alates — showed successively greater degrees of aggregation. For
example, when the cumulative proportions of workers, early-stage
nymphs, and late-stage nymphs in colony PL482 are plotted separ-
ately as a function of the segment number in which they were found,
it is apparent that the workers were distributed over a wider number
of segments than the early-stage nymphs, and the early-stage

354

Psyche

[Vol. 93

Fig. 1. Segment-by-segment distribution of castes in colonies contained in 9
pieces of wood. The base line indicates the height of the wood in each case. Repres-
ented on the left side of each colony are the numbers of individual soldiers (solid
symbols) and presoldiers (open symbols) — circles, small soldiers and presoldiers;

1986]

Luykx , Michel, & Luykx — Incisitermes

355

squares, large soldiers and presoldiers. Also shown are the reproductives — +, king;
X, queen. See colony PL487 for legend for other castes. The space between successive
tick-marks at the bottom of each colony represents ten individuals. Note that the
scales for Figs. 1 A and IB are different.

356

Psyche

[Vol. 93

Fig. 2. Segment-by-segment distribution of larvae in relation to reproductives in
6 colonies (colonies PL477, PL480, and PL490 were omitted because they had too
few larvae). The size of individual segments analyzed separately is indicated by the
tick-marks on PL476. For each colony, the baseline alone indicates uninhabited
wood; low boxes indicate fewer larvae, high boxes indicate more larvae than
expected for that segment (based on the average number of larvae per segment for
that colony). Dots represent reproductives. The shaded segment represents PL488,
an incipient colony contained entirely within a single segment.

1986] Luykx, Michel, & Luykx — Incisitermes 357

nymphs over a wider number of segments than the late-stage
nymphs (Fig. 3). The variance in position of the members of a caste
can be used as a measure of the dispersion or aggregation of that
caste, and then compared with the position-variance for all the
members of all the major castes of the colony taken together. In
colony PL487, for example, the ratio of the caste variance to total
colony variance was 1.30, 0.80, 0.52, and 0.40, for the workers,
early-stage nymphs, late-stage nymphs, and alates, respectively. A
summary of all variance ratios for all seven of the colonies with
nymphs or alates is given in Fig. 4.

There was no regularity in the mean positions of the major castes
in relation to each other nor in relation to the top or bottom of the
colony. In several colonies (e.g., PL482, Fig. 3), the mean position
of the nymphs was higher than that of the workers, but just as often
the reverse was true. In colonies PL482 and PL487, the mean posi-
of the early-stage nymphs was between that of the workers and
of the late-stage nymphs, but in colony PL486 it was below that of
those two castes. Neither was the mean position of alates (in the
three colonies that had alates) consistent in relation to that of the
other major castes.

The mean position of the soldiers, however, with the exception of
those in colony PL477, was always above that for the bulk of the
colony (e.g., colony PL482, Fig. 1). This makes sense in terms of the
function of soldiers in defending the colony, for the wood in the
upper part of the colonies is generally more deteriorated than that
lower down, and presumably more susceptible to invasion by
predators.

Six of the 9 colonies— PL476, PL477, PL480, PL486, PL487, and
PL490 — had bimodal distributions (Fig. 1). There was no clear or
consistent difference between the top and bottom groups in total
numbers of termites nor in overall caste composition in any of these
colonies. (The excess numbers of nymphs and alates in the bottom
groups of PL480 and PL486 are probably a secondary effect of the
tendency of these castes to clump together.)

Discussion

The association of larvae with the reproductives has been casually
noted by many students of the Kalotermitidae, but has not been

358

l.o n

Q-

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.6 -
.4 -
.2 -

Psyche [Vol. 93

W 1

n 1

n‘ I

Fig. 3. Distribution of workers (w), early-stage nymphs (n), and late-stage
nymphs (n') in colony PL482. The mean position and standard deviation of the
distribution for each caste is indicated by the three lines at the top.

documented quantitatively until now. Certainly the larvae are
mobile enough to disperse themselves more widely. Even in the
laboratory, after the disruption of field-collected colonies in open-
ing them up and transferring them to petri dishes, the larvae are
often found later to have re-aggregated under one fragment of
wood, often in association with the reproductives. The significance

1986]

Luykx, Michel, & Luykx — Incisitermes

1.4

359

1.2 J
1.0

VC/ .8]

7Vt

.6
.4 -
.2-
.0-

•••

••

w — ► n — ► n

CASTE

Fig. 4. Aggregation of individuals with successive developmental stage: w,
workers; n, early-stage nymphs; n', late-stage nymphs; a, alates (imagos). VC/VT,
ratio of position-variance of members of a given caste to position-variance of all
castes in the colony taken together.

of the association is not entirely clear. It is usually thought (e.g., see
Wilson, 1971) that in termites the care of the youngest larvae is
assumed by older siblings — this is, after all, one of the hallmarks of
eusociality. It might seem surprising, therefore, that the larvae
remain associated with their parents even in the presence of numer-
ous older siblings. It may be that in some termite species, particu-
larly among the lower termites, the parents continue to provide
some essential nutrients to newly-hatched larvae, something that
cannot readily be provided by older siblings. Something like this has
been seen by Nalepa (1984) in family groups in Cryptocercus punc-
tulatus, a subsocial wood-eating cockroach widely regarded as a
model of termite ancestors.

360

Psyche

[Vol. 93

Alternatively, the significance of the association between larvae
and reproductives may be just the reverse: the larvae may be feeding
the royal pair. What were classified as “larvae” in this study were
approximately the first three instars. The newly hatched larvae,
lacking intestinal flagellates, cannot feed themselves and therefore
would not be expected to be able to feed other individuals either.
But by the third instar the termites possess the intestinal symbionts,
and can feed themselves. It may be that the younger instars (beyond
the first or second) are responsible for the care of the reproductives.
There is some evidence in other species of termites (reviewed by
McMahan, 1979) that it is the younger workers that are primarily
concerned with colony feeding functions, while older workers spe-
cialize in other acitivities.

These two alternatives could probably be distinguished by means
of careful observations on the behavior of larvae and reproductives
in laboratory colonies.

The aggregation of alates within the colony is interesting, and
parallels laboratory observations on groups of alates removed from
colonies. The aggregation may reflect a tendency of the alates to
accumulate near an exit hole in preparation for emergence. The data
in Fig. 4 demonstrate that the tendency to aggregate begins in the
preceding nymphal stages. Buchli (1961) described an accumulation
of late-stage nymphs and alates in the upper and peripheral regions
of nests of Reticulitermes lucifugus, but this was apparently due to
an antagonism between these stages and the main body of workers
of the nest. In /. schwarzi nymphs and alates appear to aggregate in
the main part of the colony without any mutual show of antagonism
with other nestmates.

A striking feature of six of the nine colonies (PL476, PL477,
PL480, PL486, PL487, and PL490) was a tendency for the termites
to distribute themselves in the wood in two distinct groups. In gen-
eral, the caste composition was about the same for the upper and
lower groups (the excess of nymphs or alates in the bottom group in
colonies PL480 and PL486 may simply be a secondary effect of the
tendency of these castes to clump together). It seems unlikely, given
the regular differences found in the distribution of the major castes,
that this bimodal distribution is somehow an artefact of the proce-
dure used in cutting the colony into segments with a consequent
wholesale redistribution of members of the colony. It may be that as

1986] Luykx, Michel, & Luykx — Incisitermes 361

a colony grows, the wood in the center of the colony is often used
up, and the members of the colony then spread upwards and
downwards from the center.

Acknowledgments

This work was supported by grant no. BSR-81 19692 from the
National Science Foundation. We are grateful to Dr. Steven Green
for suggestions on how to evaluate the distribution of the major
castes (Figs. 3 and 4), to Dr. Keith Waddington for comments on
the manuscript, and to Carol A. Provost for help in preparing the
figures.

Summary

Nine colonies of the dry-wood termite Incisitermes schwarzi were
rapidly cut into segments in the field, and the numbers of individu-
als of different castes in each segment analyzed in order to learn
something about the distribution of castes within natural colonies.
The main findings are that the royal pair is usually in the lower part
of the colony, associated with small larvae; the mean position of
soldiers is usually higher than the mean position for the whole col-
ony; and, relative to the pseudergates, the early-stage nymphs, late-
stage nymphs, and alates are successively more clumped or
aggregated within the colony.

Literature Cited

Buchli, H.

1961. Les relations entre la colonie maternelle et les jeunes imagos ailes de
Reticulitermes lucifugus. Vie et Milieu, 12: 627-632.

Grass£, P.-P.

1949. Ordre des isopteres ou termites. Traite de zoologie 9: 408-544. Mas-
son, Paris.

Imms, A. D.

1919. On the structure and biology of Archotermopsis, together with descrip-
tions of new species of intestinal protozoa, and general observations on
the isoptera. Phil. Trans. Roy. Soc. Lond., B., 209: 75-180.

Luykx, P.

1987. Termite colony dynamics as revealed by the sex- and caste-ratios of
whole colonies of Incisitermes schwarzi Banks (isoptera: Kalotermiti-
dae). Insectes Sociaux (in press).

362 Psyche [Vol. 93

Luykx, P., J. Michael and J. K. Luykx.

1987. The spatial distribution of the sexes in colonies of the termite
Incisitermes schwarzi Banks (Isoptera: Kalotermitidae). Insectes Sociaux
(in press).

McMahan, E. A.

1979. Temporal polyethism in termites. Sociobiology, 4: 153-168.

Nalepa, C. A.

1984. Colony composition, protozoan transfer and some life history charac-
tristics of the woodroach Cryptocercus punctulatus Scudder (Dictyop-
tera: Cryptocercidae). Behav. Ecol. Sociobiol., 14: 273-279.

Wilson, E. O.

1971. The insect societies. Harvard Univ. Press.

A NEW SPECIES OF ORTHAEA, A NEOTROPICAL

MYODOCHINE GENUS WITH AN UNUSUAL HABITAT

(HEMIPTERA: LYGAEIDAE: RHYPAROCHROMINAE)*

By B. J. Harrington

Department of Entomology, University of Wisconsin
Madison, Wisconsin 53706

The genus Orthaea, as described by Dallas (1852), was monotypic,
with O. consuta the type species, and was treated by Stal (1874) as a
subgenus of Pamera ( Say, 1832). In 1914, Van Duzee argued against
the use of the generic name Pamera, which Say (1832) had merely
employed in a faunal list with no type or original species given, and
suggested Orthaea as the valid generic name for a growing assem-
blage of myodochine species. In his subsequent catalogue of Hemip-
tera (Van Duzee, 1917) Pamera Stal ( nec Say, 1832) 1874,
Plociomerus A & S 1843, Gyndes Stal 1862, and Diplonotus Stal
1872 were listed as synonyms of Orthaea, which generally persisted
as the name employed for the group in question until Barber (1939)
synonymized it with Pachybrachius (Hahn, 1826). Harrington’s
1980 monograph of the tribe Myodochini recognized the large,
catch-all genus Pachybrachius as polyphyletic, including several
genera and representing separate lineages involving three of the four
male genitalic types for the tribe. In that study (Harrington, 1980),
the genus Orthaea, with genitalic Type IV, was resurrected from
synonymy with Pachybrachius and noted to include the type species
O. consuta and one other species, Orthaea procincta (Breddin)
(1901).

The present paper describes a new species, Orthaea alveusincola,
and provides features to distinguish it and the other two known
species from each other. Details of the habitat in which the type
series was collected are provided since this genus apparently occu-
pies a niche unique for members of the tribe Myodochini.

All measurements in the following description are in millimeters
and the Villalobos color chart (Palmer, 1962) has been used as a
standard.

* Manuscript received by the editor September 2, 1986.

363

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[Vol. 93

Orthaea alveusincola Harrington, new species
(Figure 1)

Description. Head, anterior pronotal lobe including collar, and
scutellum sooty black. Posterior pronotal lobe, background color of
clavus and corium, and majority of hemelytral membrane blackish
brown; posterior pronotal lobe subtly lighter, grading toward light
chestnut on humeral angles. A pair of small maculae on either side
of midline on anterior half of posterior pronotal lobe, anterior one-
half of corial margin of clavus, adjacent base of corium, an elongate
macula midlength along claval margin of corium, lateral corial mar-
gin except for apical corial angle, an elongate macula running just
inside and extending less than half the length membranal margin of
corium (forming a V-shape with the line-like pale lateral corial mar-
gin), and a small macula on hemelytral membrane adjacent to apical
corial angle pale, between tawny and buffy yellow. A small diffuse
area between cream and pale gray marking the posterior margin of
the hemelytral membrane medially. Antennal segment I, distal one-
fourth of segment II, distal one-half of segment III, and extreme
proximal portion and distal one-fourth of segment IV dark, fuscous
tinged with chestnut. Femora of all three pairs of legs pale cream
basally grading to between fuscous and tawny; the extent of the
dark area greatest on the forelegs, covering almost three-fourths
their length. Tibiae light tawny with distal ends fuscous. Tarsi with
segments I and II light tawny and segment III darker. Abdomen
laterally and ventrally dark chestnut, except pygophore dark tawny.

Legs, antennae, and labium smooth; antennae with short hairs
and legs and labium with sparse elongate hairs. Meso-and metatibae
also with bristles along full length. Head subshining with micro-
rugosity and numerous short, recumbent, anteriorly directed hairs.
Pronotum pruinose and with fine recumbent hairs. Collar of ante-
rior pronotal lobe and posterior pronotal lobe prominently punc-
tate; punctures present but smaller, sparse and very shallow on
anterior lobe. Scutellum pruinose, punctate, and clothed with fine
hairs. Hemelytra subshining with sparse hairs emerging from punc-
tures. Clavus with punctation in three regular rows plus an incom-
plete fourth. Corium with a regular row of punctures along claval
suture and another parallel row along cubitus; other claval puncta-
tion randomly distributed. Abdomen ventrally and laterally sub-
shining, clothed with numerous fine recumbent hairs.

1986]

Harrington — New species of Orthea

365

Fig. 1. Orthaea alveusincola Harrington, new species, holotype, dorsal

view

366

Psyche

[Vol. 93

Head barely declivent anteriorly; somewhat prolonged in postoc-
ular region, but not constricted to form a neck. Tylus not attaining
midlength on antennal segment I. Vertex flattened, slightly depressed
before ocelli. Ocelli behind hind margin of eyes. Eyes rounded.
Length of head 1.40; postocular length 0.30; width across eyes 1.10;
interocular distance 0.52. Anterior pronotal lobe rounded; anterior
margin with a distinct band-like collar. Transverse pronotal impres-
sion well demarked and complete save for a narrow, median, dull
carina. Posterior pronotal margin straight across base of scutellum.
Humeral angles truncate, rounded. Length anterior pronotal lobe
1.22; width 1.38; width transverse impression 1.28; length posterior
pronotal lobe 0.80; width across humeral angles 1.98. Length scutel-
lum 1.20; width 1.04. Hemelytra not quite attaining end of abdo-
men; rounded rim of pygophore visible posteriorly beyond heme-
lytral membrane. Lateral corial margins vaguely sinuate at level of
claval commissure. Length corium 2.06; midline distance apex
corium to apex membrane 3.04; length claval commissure 0.82; mid-
line distance apex clavus to apex corium 1.82. Labium attaining
anterior margin of metacoxal cavities. Length labial segments I
1.20, II 1.30, III 1.08, IV 0.58. Bucculae short, projecting anteriorly
around base of labium; buccular juncture broadly V-shaped and
occurring at level of antenniferous tubercles. Antennae slender and
extremely elongate; segment IV fusiform and slightly curving.
Length antennal segments I 1.32, II 2.22, III 2.14, IV 2.30. Legs
elongate, slender. Fore femur slightly incrassate with spines double
ranked, the anterior row extending proximad two-thirds the femoral
length. Middle one-half or more of fore tibia bearing a single row of
small spines. Mesofemur with a single row of spines on anterior
surface. Mesepimeron barely emergent. Metathoracic scent gland
auricle strongly elevated from pleural surface. Total length 8.98.

Holotype. Panama: La Mesa above El Valle, 13-1-1974, B. J.

Harrington and J. A. Slater. In American Museum of Natural His-
tory, New York.

Paratypes. 1$, 10?, Same data as holotype. In American
Museum of Natural History, New York; United States National
Museum of Natural History, Washington; British Museum (Natural
History), London and private collections of P. D. Ashlock, B. J.
Harrington, and J. A. Slater.

1986] Harrington — New species of Orthea 367

Variation. Female specimens lack spines on the foretibia and
mesofemur. They also have the anterior pronotal lobe smaller, less
rounded, and in a plane lower than that of the posterior pronotal
lobes.

Etymology. This species is named O. alveusincola , “river-bed
dweller”, for the surprising habitat in which the type series was
collected.

Diagnosis. O. alveusincola, consuta, and procincta can be dis-
tinguished from each other on the basis of their hemelytral color
patterns. In alveusincola the lateral corial margin is narrowly pale
complete to the subapical macula, which continues the pale area
inward along the membranal margin in a characteristic V-shape. In
both consuta and procincta the narrow pale area along the lateral
corial margin extends posteriorly only about one-half to two-thirds
the corial length stopping short of the pale subapical corial macula
and that macula is broad and transverse, extending medially to the
membranal margin instead of running at an angle as a stripe along
the membranal margin. O. consuta lacks pale markings on the cla-
vus, while both alveusincola and procincta have them, and O. con-
suta also lacks the distinctive pair of orange maculae on either side
of midline on the anterior one-half of the posterior pronotal lobe
that are present in the other two species. O. consuta and procincta
have the lateral margins of the posterior pronotal lobe broadly
marked with orange, contrasting with the dark background; in
alveusincola these margins are not so distinctly marked and only
vaguely, if at all, lighter than the background. O. procincta lacks
foretibial and mesofemoral spines in the males as well as females,
while alveusincola and consuta males have rows of spines in both
areas.

Habitat. The type series of O. alveusincola was collected among
rocks in the partially dry bed of a mountain or highland stream
(approximate elevation 750 m.) in Panama. The insects were most
abundant in hollows and around rocks where seeds of an overhang-
ing tree were concentrated. They ran rapidly, often entering the
edges of trapped pools of water, and flew readily when pursued,
indicating full macroptery consistent with the temporary nature of
the habitat. Two series of O. procincta from Peru that were exam-
ined in this study each also have labels reporting collection in asso-
ciation with a rapid stream at high elevations (500 m. and 1600 m.).

368

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[Vol. 93

One generally would not anticipate finding rhyparochromine
Lygaeidae closely associated with a stream, since their diet of seeds
would be expected to either rot or germinate on moist ground. Yet a
highland stream, which can by flash flooding wash and concentrate
seeds and then dry rapidly, would provide a very suitable habitat
with a rich concentration of a seed resource to be exploited.
Members of the genus Orthaea have apparently adapted to capital-
ize on this resource, since two of the three known species have been
collected in such a habitat.

Currently, known distributions for the genus include O. consuta
from British Guiana and Colombia, O. procincta from Ecuador,
and O. alveusincola from Panama. As Orthaea is apparently a high-
land genus in an unexpected habitat and thus not commonly col-
lected, it is quite likely that additional new neotropical species may
be found, having evolved as montane isolates.

Summary

A new species, Orthaea alveusincola, from Panama is described.
Diagnostic features are presented to distinguish it and the other two
species in the genus, O. consuta Dallas and O. procincta (Breddin).
The type locality is described and the unusual river-bed habitat of
the genus is discussed. A full dorsal view illustration of the holotype
of O. alveusincola is provided.

Acknowledgments

I thank Dr. H. Dodge Engleman of the Coco Solo Hospital,
Panama Canal Zone, who was a generous host and collecting asso-
ciate during a field trip to Panama. I appreciate the loan of speci-
mens of described species by W. R. Dolling of the British Museum
(Natural History), London and P. D. Ashlock of the Snow Entomo-
logical Museum, University of Kansas, Lawrence, KS. I thank
Jeffrey Sternberg, University of Wisconsin, Madison for the excel-
lent dorsal view illustration of the holotype. This research was sup-
ported by the College of Agricultural and Life Sciences, University
of Wisconsin, Madison (Project No. 2578).

1986]

Harrington — New species of Or the a

369

Literature Cited

Barber, H. G. 1939. Scientific survey of Porto Rico and the Virgin Islands:
Insects of Porto Rico and the Virgin Islands: Hemiptera-Heteroptera (excepting
the Miridae and Corixidae). Sci. Surv. P. Rico 14: 3: 263-441.

Breddin, G. 1901. Neue neotropische Wanzen und Zirpen. Soc. Entomol. 16: 59.

Dallas, W. S. 1852. List of the specimens of Hemipterous insects in the collec-
tion of the British Museum. Part II. London: Taylor and Francis Inc. p. 580.

Hahn, C. W. 1826. leones ad monographium Cimicum. Nurnberg: Lechner. 1:
18.

Harrington, B. J. 1980. A generic level revision and cladistic analysis of the
Myodochini of the world (Hemiptera, Lygaeidae, Rhyparochrominae). Bull.
Amer. Mus. Nat. Hist. 167:2: 45-116.

Palmer, R. S. 1962. Handbook of North American birds. Vol. I Loons through
flamingos. Yale University Press, New Haven col. pi.

Say, T. 1832. Descriptions of new species of Heteropterous Hemiptera of North
America. New Harmony, Indiana. 1831: 16.

StAl, C. 1874. Enumeratio Hemipterorum pt 4. K. svenska VetenskAkad.
Handl. 12:1: 1-186.

Van Duzee, E. P. 1914. Nomenclatural and critical notes on Hemiptera. Canad.
Entomol. 46: 377-389.

Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America North of Mexico
(excepting the Aphididae, Coccidae and Aleurodidae). University of California
Press, Berkeley, p. 183.

BLATTELLA ASAHINAI INTRODUCED INTO FLORIDA
(BLATTARIA: BLATTELLIDAE).

By Louis M. Roth*

Museum of Comparative Zoology, Harvard University
Cambridge, MA 02138

On March 3, 1986, Dr. Philip G. Koehler of the Florida Exten-
sion Service, University of Florida, sent me some cockroaches from
Lakeland, Florida, for identification. These had been submitted to
him by Mr. Ed Shower, a pest control worker, who referred to
them as German cockroaches, but pointed out that they were un-
usual because they flew readily and were common outdoors. Until
now only 2 species of Blattella occur in the United States, namely,
vaga Hebard (India, Afghanistan, Pakistan, Sri Lanka, Mexico,
and the United States [California, Arizona, Texas]), and the cos-
mopolitan germanica (Linn.), both originating from Asia (Roth,
1985).

I decided that the “unusual germanica ” could be Blattella bey-
bienkoi Roth, which is found in Sri Lanka, Andaman Islands,
Burma, Chagos Archipelago, China, India, and Thailand (Roth,
1985). However, it also agreed with specimens of Blattella asahinai
Mizukubo, described from Okinawa (Mizukubo, 1981; Asahina,
1985). I was unaware of this species when I completed my revision
of Blattella and submitted it for publication in 1982.

I sent several Lakeland specimens to Dr. Mizukubo, who con-
cluded that they are asahinai. He also made a detailed comparison
of Sri Lanka paratypes of B. beybienkoi, and asahinai from Florida
and Okinawa, and could find no significant differences between the
two species, which I am here synonymizing.

My (Roth, 1970) attempts to cross B. germanica with 6 other
species of Blattella, namely, bisignata (Brunner), lituricollis
(Walker), sauteri (Karny), roederi Roth [as sp. C], humbertiana
(Saussure) [as sp. D], and lobiventris [as sp. E], were generally
unsuccessful. B. germanica males mated only once with bisignata

♦Correspondence: 81 Brush Hill Road, Box 238, Sherborn, MA 01770
Manuscript received by the editor August 23, 1986

371

372

Psyche

[Vol. 93

Figs. 1-5. Tergal gland reservoirs on abdominal segment 8 of Blattella spp.: Figs.
1,2. B. germanica from Lakeland, Fla. Figs. 3, 4. B. asahinai from Lakeland and
Okinawa, respectively. Figs. 5-8. F, males resulting from a cross between male
asahinai and female germanica.

and a male of the latter mated once with a female germanica. No
offspring resulted from these 2 crosses. However, in the laboratory,
B. asahinai males do cross readily with germanica females producing
Fi offspring which, to date, have produced F2 nymphs. Attempts to
produce offspring from the opposite cross of germanica males and
asahinai females, have been unsuccessful (Patterson, et al., 1986);
these results suggest that the 2 species are distinct, but very closely
related.

One of the best diagnostic morphological characters for distin-
guishing asahinai from germanica is the shape (KOH preparation)

of the male tergal gland reservoirs on the eighth abdominal segment
(cp. Figs. 1 and 2 with Figs. 3 and 4). Although there is some
variation in the shapes of these sacs in germanica (see Figs. 6, 7, in
Roth, 1985), their posterior margins curve cephalad where they may
or may not join with the anterior margins. In asahinai the sacs are
connected anteriorly, but their hind margins are widely separated
(see Figs. 12B, C, 13A-F, in Roth, 1985). The Fj males resulting
from crossing male asahinai with female germanica have reservoirs
which are more typical of germanica (Figs. 5-8).

Blattella asahinai is very widely distributed. On Okinawa it is
usually found among dead leaves and litter on the ground, and
occasionally was collected by sweeping over tree blossoms (Mizu-
kubo, 1981). In Florida it is considered to be a potential pest since it
is found in large numbers in lawns, on bushes, and invades houses.
It is now being studied by members of the USDA-ARS, and the
University of Florida, Household Insects Project (Patterson et al.,
1986).

Acknowledgments

I thank the following: Dr. Takayuki Mizukubo, National Insti-
tute of Agro-Environmental Sciences, Japan, for his opinions con-
cerning B. asahinai from Florida and B. beybienkoi; Dr. Syoziro
Asahina, retired, for specimens of B. asahinai from Okinawa; Mr.
Donald Azuma, Academy of Natural Sciences of Philadelphia for
sending paratypes of B. beybienkoi to Dr. Mizukubo at my request;
Dr. R. S. Patterson, Dr. R. J. Brenner, and Dr. P. G. Koehler,
USDA-ARS, Gainesville, Florida, and the University of Florida,
for specimens and biological information of asahinai from Florida
and for Fj specimens resulting from a cross between asahinai and
germanica.

I am grateful to The American Philosophical Society for partial
support.

References

Asahina, S.

1985. Taxonomic notes on Japanese Blattaria, XIV. Descriptions of one new
and four little-known species. Cho Cho, pub. by Gen-Gen Sha,
Kitakyushu-shi, Wakamatsu, Miyamaru, Nippon, 8: 19-26.

Mizukubo, T.

1981. A revision of the genus Blattella (Blattaria: Blattellidae) of Japan. 1.
Terminology of the male genitalia and description of a new species from
Okinawa Island. Esakia, 17: 149-159.

374

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[Vol. 93

Patterson, R. S., P. G. Koehler, and R. J. Brenner.

1986. Personal communication.

Roth, L. M.

1970. Interspecific mating in Blattaria. Ann. Ent. Soc. Amer. 63: 1282-1285.
Roth, L. M.

1985. A taxonomic revision of the genus Blattella Caudell (Dictyoptera, Blat-
taria: Blattellidae). Entomol. scand. Suppl. No. 22: 1-221.

SUBSTITUTE NAMES FOR THE EXTINCT GENERA
CYCLOPTERA MARTYNOVA (MECOPTERA)

AND PARELCANA CARPENTER (ORTHOPTERA)*

By Frank M. Carptenter
Museum of Comparative Zoology
Harvard University, Cambridge, Mass. 02138

In 1958 Dr. Olga M. Martynova described a fossil mecopteron
belonging to a new genus, Cycloptera, and representing a new fam-
ily, Cyclopteridae. Since the name Cycloptera turns out to be pre-
occupied, a substitute name is needed. Dr. Martynova has asked me
to propose a replacement name, and, following her suggestion for
the name, I propose the following:

Cyclopterina, nomen novum , pro Cycloptera Martynova, 1958,
p. 85, non Audinet-Serville, 1839, p. 439. The type species, Cyclop-
tera autumnale Martynova, 1958, original designation, becomes
Cyclopterina autumnalis (Martynova), new combination. The new
generic name is derived from Cycloptera with the addition of the
feminine suffix -ina and is considered feminine. The genus is known
only from the Permian of the Kuznetz Basin, near the Tom River,
Kemerovsk Region, USSR.

The family name, Cyclopteridae Martynova, 1938, p. 84, is herein
replaced by Cyclopterinidae. Cyclopterina is the only genus known
in the family at present.

In 1966 I described a Permian orthopteron, placing it in a new
genus, Parelcana, of a new family, Parelcanidae. I have only
recently realized that the name Parelcana is preoccupied and I take
this opportunity to propose the following substitute name:

Anelcana, nomen novum pro Parelcana Carpenter, 1966, p. 84,
non Handlirsch, 1906, p. 420. The type species, Parelcana dilatata
Carpenter, 1966, original designation, becomes Anelcana dilatata
(Carpenter), new combination. The new generic name is derived
from Elcana with the prefix an (“not”). The genus is known only
from the Permian of Kansas, U.S.A.

* Research supported by National Science Foundation grant DEB 8205398, F. M.
Carpenter, Principal Investigator.

375

376

Psyche

[Vol. 93

The family name, Parelcanidae Carpenter, is herein replaced by
Anelcanidae. The genus Petrelcana Carpenter (1966), from the same
locality, is the only other genus in the family.

References

Audenit-Serville, J. G.

1839. Histoire naturelle des insectes, Orthopteres. Paris Libraraire Encyclope-
dique de Roret, p. 1-776, pi. 1-14.

Carpenter, F. M.

1966. The Lower Permian insects of Kansas. Part 11. The orders Protorthop-
tera and Orthoptera. Psyche, 73: 48-88, PI. 4-7, text-fig. 1-20.

Handlirsch, Anton

1906. Die fossilen Insekten und die Phylogenie der rezenten Formen. P. 1-640,
pi. 1-26, Engelmann (Leipzig).

Martynova, O. M.

1958. Novye nassekomye ix permskikh i mezozojskikh ot lozxgenij SSSR.
Materialy k Osnovam Paleontologii, 2: 69-94, text-fig. 1-23.

PSYCHE

A Journal of Entomology

Volume 93
1986

Editorial Board

F. M. Carpenter, Editor H. W. Levi

W. L. Brown, Jr. M. D. Bowers

E. O. Wilson J. M. Carpenter

B. K. Holldobler

Published Quarterly by the Cambridge Entomological Club
Editorial Office: Biological Laboratories
16 Divinity Avenue
Cambridge, Massachusetts, U.S.A.

The numbers of Psyche issued during the past year were mailed on the following
dates:

Vol. 92, no. 4, April 27, 1986
Vol. 93, nos. 1-2, October 10, 1986

PSYCHE

INDEX TO VOLUME 93, 1986

INDEX TO AUTHORS

Barr, Thomas C., Jr. An eyeless subterranean beetle ( Pseudanophthalmus ) from a
Kentucky coal mine (Coleoptera: Trechinae). 47

Brady, Allen R. Nearctic species of the new wolf spider genus, Gladicosa (Araneae:
Lycosidae). 285

Brunner, George D. See Kane, Thomas C.

Buschinger, Alfred, Karl Fischer, Hans-Peter Guthy. Karla Jessen, and Ursula Winter.
Biosystematic revision of Epimyrma krussei, E., vandeli, and E. foreli (Hymenoptera:
Formicidae). 253

Carpenter, Frank M. Substitute names for the extinct genera Cycloptera Martynova
(Mecoptera) and Parelcana Carpenter (Orthoptera). 375

Carpenter, James M. A synonymic generic checklist of the Eumeninae (Hymenoptera:
Vespidae). 61

Chandler, Donald S. New Pselaphidae from New Hampshire (Coleoptera). 121

Crewe, Robin. See Peeters, Christian.

Eberhard, William G. Pupation in mycetophilid flies: a correction. 117

Fischer, Karl. See Buschinger, Alfred.

Forster, Robin B. See Heere, Edward Allen

Gambino, Parker. Winter prey collection at a perennial colony of Paravespula vulgaris
(L.) (Hymenoptera: Vespidae). 331

Guthy, Hans-Peter. See Buschinger, Alfred.

Heere, Edward Allen, Donald M. Winsor, and Robin B. Forster. Nesting associations of
wasps and ants on lowland Peruvian ant-plants. 321

Harrington, B. J. A new species of Orthea, a neotropical myodochine genus with an usual
habitat (Hemiptera: Lygaeidae: Rhyparochrominae). 363

Herbers, Joan M. and Carol W. Tucker Population fluidity in Leptothorax longispinosus
(Hymeniptera: Formicidae). 217

Jessen, Karla. See Buschinger, Alfred.

Johnson, Robert A. See Rissing, Steven W.

Kane, Thomas C. and George D. Brunner. Geographic variation in the cave beetle,
Neaphaenops tellkampfi (Coleoptera; Carabidae). 231

Kimsey, Lynn Siri. New species and genera of Aniseginae from Asia (Chrysididae,
Hymenoptera). 153

379

Kronestedt, Torbjdm. A presumptive pheromone-emitting structure in wolf spiders
(Araneae: Lycosidae) 127

Levi, Herbert W. The orb-weaver genus Witica (Araneae: Araneidae). 35

Luykx, Peter, Jack Michel, and Jeanette K. Luykx. Spatial distribution of castes within
colonies of the termite, Incisitermes schwarzi. 35 1

Luykx, Jeanette K. See Luykx, Peter.

Maddison, Wayne. Distinguishing the jumping spiders Eris militaris and Eris f Java in
North America (Araneae: Salticidae). 141

Michel, Jack. See Luykx, Peter.

Moffett, Mark W. Notes on the behavior of the dimorphic ant, Oligomyrmex overbecki
(Hymenoptera: Formicidae). 107

Moffett, Mark W. Evidence of workers serving as queens in the genus Diacamma
(Hymenoptera: Formicidae). 151

Opell, Brent D. The choice of web-monitoring sites by a green Miagrammopes species
(Araneae: Ulobidae). 167

Peeters, Christian, and Robin Crewe. Male biology in the queenless ponerine ant,
Ophthalmopone berthoudi (Hymenoptera: Formicidae). 277

Pollock, Gregory B. See Rissing, Steven W.

Porter, Charles C. Bicornis in Peru, with notice of an endemic species from the coastal
desert (Hymenoptera:Ichneumonidae). 51

Porter, Charles C. South American and Floridian disjuncts in the Sonoran genus
Compsocryptus (Hymenoptera: Ichneumonidae). 13

Porter, Charles C. A new arboricolous Thyredon from Costa Rica (Hymenoptera,
Ichneumonidae: Ophioninae). 1 33

Rissing, Steven W., Robert A. Johnson, and Gregory B. Pollock. Natal nest distribution
and pleometrosis in the desert leaf-cutter ant, Acromyrmex versicolor (Perhande)
(Hymenoptera: Formicidae). 177

Rasnitsyn, A. P. Review of the fossil Tiphiidae, with descriptions of a new species
(Hymenoptera). 91

Richardson, John S. See Wiggins, Glenn B.

Roth, Louis J. Blatella asahinai introduced into Florida (Blattaria: Blatellidae). 37 1

Stuart, Robin J. An early record of tandem running in leptothoracine ants:
Gottfrid Adlerz, 1890.

Tucker, Carol. W. See Herbers, Joan M.

Ward, Philip S. Functional queens in the Australian greenhead ant, Phytidoponera
metallica (Hymenoptera: Formicidae). 1

Wheeler, George C. and Jeannette Wheeler. Young larvae of Eciton (Hymenop-
tera: Formicidae). 341

380

Wheeler, Jeanette. See Wheeler, George C.

Wiggins, Glenn B. and John S. Richardson. Revision of the Onocosmoecus uni-
color group (Trichoptera: Limnephilidae, Dicosmoecinae). 187

Winsor, Donald M. See Heere, Edward Allen
Winter, Ursula. See Buschinger, Alfred.

INDEX TO SUBJECTS

All new genera, new species and new names are printed in capital type.

Acromyrmex versicolor, 167
Actizona borealis, 123
A llomer us, 321
ANELCANA, 375
Alopecosa cuneata, 127
Atoposega simulans, 153
Biconus in Peru, 51
Biconus apoecus, 53
Biconus subflavus, 57

Biosystematic revision of Epimyrma
kraussei, vandeli, and foreli, 253

Blattella asahinai, in Florida, 371
BUPON PASOHANUS, 156
Choice of web-monitoring sites, 167
Cladobethylus aquilus, 157
Cladobethylus gibus, 157
Cladobethylus japonicus, 158
Comp so cry plus, 13
Compsocryptus fasciipennis, 28
Comsopcryptus fuscofasciatus, 20
Compsocryptus melanostigma, 24
CYCLOPTERINA, 375
CYCLOPTERINIDAE, 375
Diacamma, 151
Eciton, 341
Epimyrma foreli, 253
Epimyrma kraussei, 253

Epimyrma vandeli, 253
Eris flava, 1 4 1
Eris militaris, 141
Euplectus silvicolus

Eyeless subterranean beetle, Pseudanoph-
thalmus, 47

Fossil Tiphiidae, 91

Functional queens in Rhytidoponera
metallica, 1

Geographic variation in Neaphaenops
tellkampfi, 231

Geotiphia halictina, 94
Geotiphia orientals, 97
Geotiphia sternbergi, 95
Geotiphia pachysoma, 96
gladicosa, 285
Gladicosa bellamyi, 3 1 1
Gladicosa euepigynata, 3 1 2
Gladicosa gulosa, 290
Gladicosa huberti, 305
Gladicosa pulchra, 299
Incisitermes schwarzi, 351
Isegama malaysiana, 159
Jumping spiders, 141
KRYPTOSEGA ANOMALA, 160
Kryptosega kaindeana, 162
Leptothorax longispinosus, 2 1 7
Leptothoracine ants, 103

381

Lithotiphia scudderi, 93
MAGDALIUM CUNEIFACIALIS, 164

Male biology Ophthalmopone berthoudi,
277

Miagrammopes, 167
Mischocyttarus, 321
Mycetophilid flies, 1 17

Natal nest distribution and pleometrosis
in Acromyrmex, 177

Neaphaenops tellkampfi, 23 1
Nearctic species of Gladicosa, 285

Nesting associations of wasps and ants,
321

Oligomyrme x overbecki, 107
Orthea alveusincola, 364
Onocosmoecus, 189
Onocosmoecus unicolor, 193
Onocosmoecus sequoiae, 208
Ophthalmopone berthoudi, 277
Paravespula vulgaris, 33 1
Population fluidity in Leptothorax, 217

Presumptive pheromone-emitting struc-
ture in spiders, 127

Pseudanophthalmus, 47

Pselaphidae from New Hampshire, 121

Revision of Onocosmoecus unicolor
group, 187

Rhytidoponera metallica, 1

Spatial distribution of castes of Incisi-
termes, 351

Synonymic generic checklist of Eumeni-
nae, 61

Thyreodon santarosae, 133

Web-monitoring by Miagrammopes, 167

Winter prey collection by Paravespula
vulgaris, 331

Witica, 35

Witica cayana, 44

Witica crassicauda, 4 1

Wolf spider genus Gladicosa, 285

Workers serving as queens in Diacamma,
151

382

CAMBRIDGE ENTOMOLOGICAL CLUB

BACK VOLUMES OF PSYCHE

Requests for information about back volumes of Psyche should
be sent directly to the editor.

F.M. Carpenter
Editorial Office, Psyche
16 Divinity Avenue
Cambridge, Mass. 02138

FOR SALE

Reprints of articles by W. M. Wheeler

The Cambridge Entomological Club has for sale numerous reprints
of Dr. Wheeler’s articles- that were filed in his office at Harvard
University at the time of his death in 1937. Included are about
12,700 individual reprints of 250 publications. The cost of the
reprints has been set at 5c a page, including postage; for orders
under $5 there will be an additional handling charge of 50c. A list of
the reprints is available for $1.00 from the W. M. Wheeler Reprint
Committee, Cambridge Entomological Club, 16 Divinity Avenue,
Cambridge, Mass. 02138. Checks should be made payable to the
Cambridge Entomological Club.

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