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ISSN 0033-2615

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^ PSYCHE

A JOURNAL OF ENTOMOLOGY

founded in 1874 by the Cambridge Entomological Club

Vol. 89 1982 No. 1-2

i JMH .1 i 'in. , J

/ IfiDADlP^

• **■* o ft A r\ 5

CONTENTS

Dedication: Joseph C. Bequaert. Frank M. Carpenter 1

Communication, Raiding Behavior, and Prey Storage in Cerapachys

(Hymenoptera: Formicidae). Bert Hdlldobler 3

Designation of a Type-species for Cvclogaster Macquart, 1834, and the Result-
ing Synonymy (Diptera: Stratiomyidae). Norman E. Woodley 25

Orb Plus Cone-webs in Uloboridae (Araneae), with a Description of a New
Genus and Four New Species. Y. D. Lubin, B.D. Opell, W. G. Eberhard, and
H.W. Levi 29

Population Structure and Social Organization in the Primitive Ant, Amblyo-
pone pallipes (Hymenoptera: Formicidae). James F.A. Traniello 65

The Biology of Nine Termite Species (Isoptera: Termitidae) from the Cerrado
of Central Brazil. Helen R. Coles de Negret and Kent H. Redford 81

The Life History of the Japanese Carrion Beetle, Ptomascopus morio and the
Origins of Parental Care in Nicrophorus (Coleoptera, Silphidae, Nicrophini).

Stew art B. Peck 107

Tergal and Sternal Glands in Male Ants (Hymenoptera: Formicidae). Bert
Hdlldobler and Hiltrude Engel-Siegel 113

Termite-Termite Interactions: Workers as an Agonistic Caste. Barbara L.
Thorne 133

Type Designations and Synonymies for North American Silphidae (Coleoptera).
Stewart B. Peck and Scott E. Miller 151

Chemical Mimicry as an Integrating Mechanism for Three Termitophiles Asso-
ciated with Reticulitermes virginicus (Banks). Ralph W. Howard, C.A.
McDaniel, and Gary J. Blomquist 157

Parataruma, a New Genus of Neotropical Crabronini (Hymenoptera, Spheci-
dae). Lynn S. Kimsey 169

Supplementary studies on ant larvae: Formicinae (Hymenoptera: Formicidae)
George C. Wheeler and Jeanette Wheeler 175

Morphological comparisons between the obligate social parasite, Vespula aus-
traica (Panzer) and its host, Vespula acadica (Sladen). Hal C. Reed and
Roger D. Akre 183

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1981-1982

President Barbara L. Thorne

Vice-President Frances Chew

Secretary Heather Hermann

Treasurer Frank M. Carpenter

Executive Committee John Shetterly

Mary Hathaway

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
P. J. DARLINGTON, Jr., Professor of Zoology, Emeritus, Harvard
University

B. K. HOLLDOBLER, Professor of Biology, Harvard University
H. W. LEVI, Alexander Agassiz Professor of Zoology, Harvard University
R. J. McGlNLEY, Assistant Professor of Biology, Harvard University
Alfred F. NEWTON, Jr., Curatorial Associate in Entomology, Harvard
University

R. E. SlLBERGLIED, Smithsonian Tropical Research Institute, Panama
E. O. WILSON, Baird Professor of Science, 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: $11.00, domestic and foreign. Single copies, $3.50.

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 expected to bear part of the printing costs, at the rate of $27.50 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.

Psyche, vol. 88, no. 3-4, for 1981, was mailed May 28, 1982

The Lexington Press, Inc., Lexington, Massachusetts

Joseph Charles Bequaert

This issue of Psyche is dedicated to the memory of Joseph C.
Bequaert, who died in his 96th year in Amherst, Massachusetts, on
January 12, 1982.

Dr. Bequaert was born in Belgium in 1886 and was educated
there, receiving his Dr. Phil, degree in botany in 1908 from the State
University in Ghent. The next seven years he spent in the Belgian
Congo (now Zaire), at first as Entomologist on the Belgian Sleeping
Sickness Commission and later as head of botanical explorations in
the Congo for the Belgian Colonial Government. During those
years his main interest shifted from botany to entomology, in which
he subsequently did the greater part of his research and teaching. In
1917 he was appointed Research Associate in Congo Zoology at the
American Museum of Natural History. Six years later, after becom-
ing a naturalized citizen of the United States, he joined the faculties
of the Harvard School of Public Health and the Harvard Medical
School, as an assistant professor in medical entomology, and
remained there until 1945. He then accepted the position of Curator
of Recent Insects in the Museum of Comparative Zoology, succeed-
ing Nathan Banks. In 1951 he was appointed Alexander Agassiz
Professor of Zoology, a chair that he held until his retirement in
1956. Most of the remaining 26 years of his life were spent in Tuc-
son, Arizona, where he was associated with the departments of
entomology and zoology at the University of Arizona.

He was internationally known for his publications, totalling more
than 250, on medical entomology, mollusks, botany, and systemat-
ics of several families of insects.

Joe joined the Cambridge Entomological Club in 1923, as soon as
he reached the Boston area, and he was very active in the society for
the next 33 years. He was president in 1928, 1935-36, and 1942-43;
vice-president in 1937, 1941, and 1946; secretary in 1925 and 1926;
and treasurer in 1943. He also served on the editorial board of
Psyche from 1947-1956. He gave many of the scheduled talks at our
regular meetings and was chosen as the speaker for the 500th meet-
ing of the Club on December 15, 1931. In recognition of his services
and contributions to the activities of the society, he was elected an
Honorary Member in 1961.

1

Joseph Charles Bequaert

Photograph taken in Belgian Congo, 1934

I first met Joe at the September meeting of the Club in 1923, at
which he was nominated for membership. His exuberance and his
extraordinary enthusiasm for nearly every aspect of natural history
were the most obvious traits of his personality. In 1956 he wrote the
following statement of his scientific interests: ecology of flowers;
taxonomy and ecology of Bryophyta; geography and ecology of
African plants; relations of Arthropoda to disease; taxonomy and
ethology of Diptera and Hymenoptera, particularly Vespidae; mala-
cology; medical entomology. He was certainly one of the most dis-
tinguished and respected entomologists of his generation.

Frank M. Carpenter, editor

PSYCHE

Vol. 89

1982

No. 1 -2

COMMUNICATION, RAIDING BEHAVIOR
AND PREY STORAGE IN CERA PA CHYS
(HYMENOPTERA; FORMICIDAE)*

By Bert Holldobler

Department of Organismic and Evolutionary Biology,

MCZ - Laboratories

Harvard University, Cambridge, Mass. 02138 U.S.A.

Introduction

The former subfamily Cerapachyinae was recently recognized by
Brown (1975) as a tribe (Cerapachyini) within the subfamily Poneri-
nae. All of the cerapachyine ant species investigated feed entirely on
ants (see review in Wilson 1958; Brown 1975). During foraging
cerapachyine workers engage in mass expeditions during which they
raid the nests of the prey species, capturing preferably larvae and
pupae, but also occasionally adults and returning them to the raid-
ers’ nest.

Although the detailed field observations on cerapachyine forag-
ing raids reported by Wilson (1958) strongly suggest that the raiding
expeditions follow chemical trails, this has not yet been experimen-
tally investigated. In fact, almost nothing was hitherto known about
the behavioral organization of the raiding expeditions and the under-
lying communication mechanism. This paper presents the first ex-
perimental analysis of the raiding behavior of a cerapachyine ant species.

Materials and Methods

Three colonies of Cerapachys (?) turneri (turneri group) (acces-
sion #163a, b, c; voucher specimens in Australian National Insect

* Manuscript received by the editor January 22, 1982.

3

4

Psyche

[Vol. 89

Collection, ANIC, Canberra) were collected from nests in the soil in
a sclerophyl scrub pasture near Eungella, North Queensland (Aus-
tralia). One colony had a single ergatoid queen; the other colonies
had two ergatoid queens apiece. Each colony was housed in separate
glass tube nests (8cm X 0.6cm c />), with water trapped at the bottoms
behind cotton plugs. Each nest tube was placed into arenas of
varying sizes, depending on the experimental design. Histological
studies were conducted according to the procedures described in
Holldobler and Engel 1978. Additional methodological details will
be given with the description of the individual experiment, as pre-
sented below.

Results

Raiding behavior and paralysis of prey larvae

Species of the genus Cerapachvs seem to preferably prey on ant
species of the myrmicine genus Pheidole (Wilson 1958; Brown
1975). When 1 provided Cerapachvs with colonies or fragments of
colonies of a variety of species of the genera Iridomyrmex, Meranop-
lus, Monomorium, Crematogaster, Pheidole, Stigmacros, Polvrha-
chis, Camponotus (placed in a 65 X 120cm arena) they preyed freely
only on Pheidole. They also accepted Monomorium larvae as prey,
but only when these insects were directly inserted into the Cera-
pachys nest. When the Cerapachvs workers encountered workers of
the other species, or came close to their nest tubes, they usually
showed avoidance behavior. The reaction was very different, how-
ever, when individual scouts of Cerapachvs discovered the nest tube
of Pheidole (accession #209, voucher specimens in ANIC). The Cera-
pachys worker vigorously vibrated its short antennae and moved
slowly into the nest tube, which contained approximately 200 Phei-
dole workers and soldiers and about 150 larvae and pupae. It did
not venture very far into the foreign nest but left after a short while
and ran, in a somewhat meandering route, back to its own nest,
located 70cm away from the Pheidole nest. During honyng it
appeared frequently to touch the ground with its abdominal tip, as if
it were laying a chemical trail or depositing scent spots. Seconds
after it had entered the nest of its own colony, its n^stmates became
very excited. Many grouped around the scout ant, which repeatedly
raised its gaster upwards. Within one minute the scout left the nest

1982]

Holldobler — Cerapachys

5

again and moved in direction toward the Pheidole nest tube. It was
closely followed by 17 nestmates. The leading scout ant continued to
move with its abdominal tip close to the ground, but intermittently
it paused or moved much slower while raising its gaster slightly
upwards (Fig. 1). When the Cerapachys column arrived at the Phei-
dole nest tube they invaded it and attacked the Pheidole workers
and soldiers. Pheidole fought back but without any effect. The heav-
ily sclerotized and specially protected Cerapachys (Fig. 2) were not
at all affected by the mandibular grip of the Pheidole soldiers, even
when they were attacked simultaneously by 3-5 Pheidole (Fig. 3).
Although Pheidole outnumbered the Cerapachys invaders more
than 10 times, they were rapidly disabled by the obviously very

Figure 1. Recruiting Cerapachys worker, (a) Worker walking with its abdomi-
nal tip close to the ground, (b) Worker raising the gaster upwards; arrow indicates
the position of the opening of the pygidial gland.

6

Psyche

[Vol. 89

Figure 2. Longitudinal section through the head and part of the thorax (a) and
through part of the petiolus and gaster (b) of a Cerapachvs worker. Arrows indicate
cuticle projections over intersegmental membranes (IM).

1982]

Holldobler — Cerapachys

7

Figure 3. Cerapachys raiding group invading a Pheidole nest.

effective stinging attack of the Cerapachys, during which the raiders
grasped the Pheidole with their short mandibles, simultaneously
bending their gasters forward, so that in each case the tip, where the
sting extrudes, touched the opponent’s body. Each sequence usually
lasted less than 1 second. Almost immediately after such an attack
the Pheidole appeared to be immobilized. Only a few Pheidole
workers escaped from the nest tube into the arena, some of them
carrying brood. After approximately 15 minutes almost all Pheidole
adults in the nest tube were disabled or killed but not a single
Cerapachys worker was dead or visibly injured. Next the Cera-
pachys began transporting the dead and immobilized Pheidole
adults to their own nest. After the first workers of the raiding expe-
dition had returned and unloaded the booty they returned to the
Pheidole nest. Some of them raised the gaster repeatedly upwards,
upon which several additional Cerapachys workers followed them
to the Pheidole nest, where they participated in the retrieval of the
prey. Only after most of the Pheidole adults had been retrieved did
the Cerapachys begin to transport the Pheidole brood. Each larva
and pupa was briefly stung before it was picked up and carried to
the Cerapachys colony. Interestingly, after approximately half the
brood had been retrieved, Cerapachys nest workers began discard-
ing all the dead and disabled Pheidole adults, and the next day only

8

Psyche

[Vol. 89

Pheidole brood was stored in the Cerapachvs nest. Apparently the
booty of this raiding expedition was so abundant that Cerapachvs
preferred to keep only the more valuable and better preservable
brood of the prey species, and they discarded the less valuable
cadavers of the adult Pheidole. In other instances, however, where
Cerapachvs had only, adults of prey species available, I observed
Cerapachvs feeding on the gasters of dead Pheidole workers and
soldiers.

This experiment was conducted on the 25th and 26th of October

1980. At this time there was no Cerapachvs brood in the colony. On
November 10, 1980, I noticed the first large clutch of eggs in the
Cerapachvs nest tube. On December 1 1, 1980, the colony had many
large (presumably last instar) larvae, and another large cluster of
eggs (Fig. 4). The colony still contained a very good supply of
Pheidole larvae (Fig. 4), which did not grow or develop further but
which were obviously alive. Under the microscope one could see
that the prey larvae slightly moved their mouthparts. Workers,
queens and larvae of Cerapachvs all fed on the Pheidole larvae. On
December 26, 1980, there were still some prey larvae left. Many of
the large Cerapachvs larvae had pupated; in addition the nest con-
tained many medium sized larvae and another large clutch of eggs.
On January 3, 1981, a Cerapachvs worker was observed leaving the
nest tube and venturing out into the arena, for the first time since
October 27, 1981. At this time I provided another fragment of a
Pheidole colony with larval brood in the arena; and on January 5,

1981, Cerapachvs conducted another raid, very similar in details to
that just described. The fact that the captured Pheidole larvae were
kept alive inside the Cerapachvs nest chamber for a period of more
than two months (but did not pupate or visibly increase in size)
strongly suggested that they were sustained in a state of metabolic
stasis. Recently Maschwitz et al (1979) provided experimental evi-
dence that the ponerine species Harpegnathus saltator and Lepto-
genys chinensis paralize prey objects by stinging and thereby are
able to store prey a limited time. In one case the preserving paralysis
effect was observed to last for two weeks, and in no instance did the
stung prey object ever recover from the paralysis. Similar observa-
tions have been made independently by Traniello (unpublished
data) with the ponerine species Aniblvopone pal/ipes.

1982]

Holldobler — Cerapachys

9

Figure 4. Fractions of a Cerapachys colony, with paralyzed prey larvae. Q: erga-
toid queens; E: eggs; C: Cerapachys larvae; P: Pheidole prey larvae.

10

Psyche

[Vol. 89

As just noted, Cerapachys workers apparently sting each Phei-
dole larva and pupa during the raid, before they transport the vic-
tims to their nest. This appears to be a very stereotyped behavior.
For example when 1 shook a Cerapachys colony which contained
Pheidole larvae out of the nest tube into the arena, so that they had
to move back into the nest, Cerapachys workers picking up a Phei-
dole larva almost invariably went through the typical stinging
motion pattern. They did not do this, however, when they picked up
their own larvae. Although stinging behavior did not frequently
occur inside the nest, occasionally I observed a Cerapachys stinging
several larvae while reshuffling a pile.

The Pheidole larvae are small and tender and the powerful Cera-
pachys sting (Fig. 5) could easily pierce the larva and thereby kill it.
Thus the injections of a paralyzing secretion through the sting has to
be very subtle in order not to kill, but to preserve the larva. Brown
(1975) describes the differentiated pygidium (Fig. 6) with its denticu-
late margins, being present in all workers and queens of cera-
pachyine ants. Brown states that “the function of the denticle-
bordered pygidial plate is not known from direct observations, but
it is assumed to have something to do with helping the insects to
force their way through passages and cracks in soil or rotten wood,
perhaps in connection with their entry into nests of termites or ant
prey species”.

Our morphological and histological investigations have revealed
that these denticuliform and spinuliform setae on the pygidium of
Cerapachys turneri and Sphinctomyrmex steinhei/i are sensory
setae and comprise probably mechanoreceptors (Fig. 7). It is most
likely that during the stinging process these mechanoreceptors sig-
nal the gaster tip’s touch of the prey larva and the extent of the
stings’ protrusion is thereby regulated. Many of the nonsocial acu-
leate Hymenoptera, which paralyze prey by stinging, are equipped
with mechanoreceptors on the tip of the sting sheath (Oeser 1961,
Rathmayer 1962, 1978). We did not detect similar structures on the
tip of the sting sheaths of Cerapachys or Sphinctomyrmex. In addi-
tional experiments I further confirmed the suggestion that the prey
larvae, captured by Cerapachys, are preserved alive. Approximately
30 Pheidole larvae collected from a Pheidole colony were put with-
out workers in a small test tube, which was kept moist by a wet
cotton plug. A second similar test tube contained 30 Pheidole larvae
which were taken from the Cerapachys nest. In two replications the

1982]

Holldobler — Cerapachys

11

Figure 5. (a) SEM picture of the abdominal tip of a Cerapachys worker. The

picture shows the partly extruded sting, surrounded by the sensory setae at the
pygidium, and last exposed sternite. (b) Close-up of the two kinds of setae at the
pygidium.

12

Psyche

[Vol. 89

Figure 6. SEM picture of frontal view of pygidium of a Cerapachvs worker (a),
and a worker of Sphinctomvrmex steinheili (b). Note the arrangement of the two
kinds of setae on the truncated pygidial plate of both species.

1982]

Holldobler — Cerapachys

13

Figure 7. Longitudinal section through pygidial plate (a) and last exposed ster-
nite (b) of a Cerapachys worker. The structure and innervation of the setae suggest
that they function as mechano receptors.

14

Psyche

[Vol. 89

larvae taken directly from the Pheidole colony were all dead after
two weeks. On the other hand all of the larvae from the Cerapachys
colony were obviously still alive after two weeks, many of them
moving their mouthparts slightly. These findings clearly demon-
strate that Cerapachys can store living prey larvae for a considerable
period of time. This food storage system appears to enable Cera-
pachys to stay inside their nest for longer intervals. They evidently
do not conduct raids as long as a good food supply is present. The
following experiments were designed to test this hypothesis.

One day after the Cerapachys colony B had conducted a raid on
Pheidole all prey larvae were removed. As a control I manipulated
colony A in the same way, but the prey larvae were immediately
returned to colony A. A few days later I observed scouts of colony B
in the arena, where I had provided a nest tube with a fraction of a
Pheidole colony, and within a period of 4 (test 1) and 7 days (test 2)
colony B had conducted another raid. In the control colony A I
noticed a worker briefly leaving the nest tube only once and then
without venturing far into the arena. Although a tube containing
Pheidole workers and brood was also provided in the arena of
colony A, this colony did not conduct another raid until its supply
of prey had declined considerably.

Emigration behavior

Although it is still an open question whether the Cerapachyini are
nomadic, Wilson (1958, 1971) and Brown (1975) suggested that
nomadism in the ant-preying cerapachyine species could well be
adaptive to avoid depleting the food supply in a given neighbor-
hood, just as it is in the army ants. This assumption of a nomadic
life style is further supported by Brown’s observations that the nests
of many cerapachyine species appear to be impermanent, and that
the “brood show a strong tendency to be synchronized, like those of
army ants and nomadic Ponerinae”. Brown (1975) also pointed out
that the larvae of the Cerapachyini have a slender and cylindrical
shape (G. C. Wheeler and J. Wheeler 1964), which makes them easy
to transport longitudinally under the bodies of workers in the
manner of other predatory and nomadic ants, such as Eciton, Aenic-
tus, Dorylus, Leptogenys and Onychomyrmex. Although I was
unable to demonstrate periodic nomadic behavior of Cerapachys in

1982]

Holldobler — Cerapachys

15

the laboratory, I could easily initiate nest emigrations by removing
the waterplug and thereby causing the nest tube to quickly dry out.
Individual workers soon ventured into the arena and eventually
discovered a new moist nest tube located approximately 20-30 cm
away from the old nest. After exploring the new nest site the scout
moved back to the colony. When entering the nest tube it exhibited
the same behavior as when recruiting to a raid, including a repetitive
lifting of the gaster. When the scout left the nest again to return to
the newly discovered nest site, it was usually followed by several
ants. Most of these first recruits also showed the gaster raising
behavior on their return to the colony, and soon the whole colony
began to leave the old nest tube and move to the new one. The
larvae and pupae were carried in the manner Brown (1975) pre-
dicted, slung longitudinally under the bodies of the workers (Fig. 8).
Adult transport was never observed; the ergatoid queens and even
relatively freshly eclosed workers moved on their own to the nest
site. The colonies did not contain males. After the workers had
moved most of their own brood, they transported the prey larvae
( Pheidole ).

Figure 8. Cerapachys worker carrying a larva during nest emigration.

16

Psyche

[Vol. 89

From the ants’ orientation behavior it appeared that they were
following chemical trails during the nest emigration. In fact, the
recruitment behavior during nest emigrations and raiding appeared
to be identical. The following experiments were designed to analyze
further the communication mechanisms involved in both events.

Communication during emigration and raiding

Two distinct behavioral patterns were observed in Cerapachys
ants during recruitment. (1) They seem to lay a chemical trail when
returning from the target area (prey colony or new nest site) by
frequently touching the abdominal tip to the ground; and (2) when
close to or just entering the nest, they repeatedly raised their gaster
upwards into a “calling position” and continued to do so when they
moved back to the target area, usually being closely followed by a
group of recruited nestmates. Since it was easier to initiate emigra-
tions rather than raids, most of the experiments were conducted
during colony emigration. Several new exocrine glandular struc-
tures have recently been discovered in ponerine ants (Holldobler
and Haskins 1977; Holldobler and Engel 1978; Holldobler et al.
1982; Maschwitz and Schonegge 1977; Jessen et al. 1979). The
Cerapachyini were not included in these studies. We therefore con-
ducted first a histological survey for possible exocrine glands that
might be involved in the communication behavior of Cerapachys.
Besides the known glands associated with the sting, we found a
pygidial gland, which consists of a paired group of a few glandular
cells under the 6th abdominal tergite. Each cell sends a duct through
the intersegmental membrane between the 6th and 7th tergite (Fig.
9). The intersegmental membrane is laterally slightly invaginated, so
that at each side it forms a small glandular reservoir. No particular
cuticular structure on the pygidium is associated with the pygidial
gland.

In a first set of pilot experiments I dissected out of freshly killed
Cerapachys workers poison glands, Dufour’s glands, hindguts,
pygidial glands (6th and 7th tergites) and the last 3 sternites. For
each test one organ of a kind was crushed on the tip of hardwood
applicator sticks. These were then immediately inserted into the nest
tube until the tip of the applicator was 2-3cm away from the colony,

1982]

Holldobler — Cerapachys

17

Figure 9. (a) Longitudinal section through the gaster of a Cerapachys worker

showing the location of the pygidial gland (PG). (b) Longitudinal section through

the pygidial gland; GC: glandular cells; CH: glandular channels through inter-
iegmental membrane.

18

Psyche

[Vol. 89

which usually had gathered near the cotton plug. In the following 30
seconds I observed the reaction of the ants, and between each test I
waited at least 10 minutes before another sample was inserted into
the nest tube. These pilot tests (3 repetitions with each organ) clearly
indicated that only crushed poison glands and pygidial glands eli-
cited increased locomotory activity and attraction in Cerapachvs
workers. The ants did not exhibit any particular behavioral reaction
when sternites, hindgut or crushed Dufour’s glands were intro-
duced.* For the next series of experiments I first initiated colony
emigrations either by following the procedure described above, or
by shaking the colony out of the nest tube onto the arena floor.
Before each experiment the arena was provided with a new paper
floor. A new nest tube was offered 15-20cm away from the old nest
tube or the displaced colony.

Once the colony emigration to the new nest tube had commenced,
I covered the floor area between the colony and the new nest site
with a cardboard, onto which I had drawn two artificial trails, one
with a crushed glandular organ to be tested, and a second one with a
drop of water (control). The trails were made to originate either
from the entrance of the nest tube or from the periphery of the
clustered colony. Each trail (test and control) diverged through an
angle of 45° to either side from a possible natural trail (which was of
course covered by a piece of cardboard). In addition the whole
paper floor was rotated for 90°, in order to control for possible
visual orientation (Fig. 10). During the following 2 minutes I
counted the ants following the trails (10cm long) to the end. Only
trails drawn with crushed poison glands elicited a precise trail fol-
lowing behavior in Cerapachvs workers. There was some initial
following response to trails drawn with crushed pygidial glands, but
the ants followed only through the first 1-3 cm, then usually turned
or meandered off the trail. Only once was it possible to conduct a
similar test during raiding behavior of Cerapachvs. In this instance
the ants followed only an artificial trail drawn with a crushed poison
gland.

Although pygidial gland secretions did not release trail following
behavior in Cerapachvs, it clearly elicited increased locomotory

*Cerapachvs has also a very well developed sting sheath gland. It was not possible to
test whether or not secretions of the gland play a role in communication.

1982]

Holldobler — Cerapachys

19

Figure 10. Schematical illustration of the experimental arrangement during trail
tests. The colony was emigrating from nest NI to nest Nil along a natural trail a.
During the trail tests, the whole arrangement was turned 90° (arrow). The natural
trail a was covered by a cardboard, on which the test trail (T) and a control trail (C)
were offered, each deviating from a in an angle of 45°.

activity and attraction in the ants. I hypothesized therefore that the
recruiting ant might discharge pygidial gland secretions when it
exhibited the gaster raising behavior. The pygidial gland pheromone
might function as an additional recruitment signal by which the
recruiting ant keeps the raiding party stimulated when leading it to
the prey colony. In order to test this hypothesis, I tried on four
different occasions to close the opening of the pygidial gland by
applying collophonium wax between the 6th and 7th tergites. Unfor-
tunately these experiments failed; apparently the ants were too dis-
turbed by the procedure. During two raiding expeditions of

20

Psyche

[Vol. 89

Cerapachys we succeeded, however, in diverting individual ants
from the raiding column over a distance of at least several centi-
meters by presenting two applicators in front of them, one contami-
nated with pygidial gland secretions and the other with water. Both
applicators were slowly moved away from the columns in opposing
directions. Of a total of 10 ants tested, 4 responded by following for
a few centimeters behind the applicator with the pygidial gland
secretions; no ant followed the control applicator. Although these
results can be considered only preliminary, they do suggest that
pygidial gland secretions might be involved in the recruitment pro-
cess of Cerapachys. This suggestion was further supported by the
results of a series of experiments in which I offered artificial trails
drawn with crushed poison glands. I compared the trail following
response of Cerapachys (within the first two minutes) successively
either to trails drawn with poison gland secretions only or to poison
gland trails offered simultaneously with pygidial gland secretions.
For each kind a total of 6 experiments was carried out. Between
each test at least one day had elapsed. The following response
appeared to be stronger to poison gland trails when offered together
with pygidial gland secretions (5.5 ± 2.9) than to those offered with-
out pygidial gland secretions (3.0 ± 1.4) (0.1 > p > 0.05; Students
t-test). Because of lack of material this series could not be extended,
and thus the results remain only suggestive.

The two final experiments demonstrated that a trail (10cm long)
drawn with one crushed poison gland, was still effective as an
orientation cue several hours after it had been drawn. Using the
same experimental arrangement described above (Fig. 10), I was
able to show that emigrating Cerapachys would follow poison gland
trails, 2 and 6 hours old, when they were offered after the natural
trail had been covered. On the other hand, crushed poison glands
introduced into the nest tube after 2 and 6 hours, or poison gland
trails offered 2 and 6 hours after they had been drawn, did not elicit
excitement or spontaneous trail following behavior. From these
results it appears that the poison gland material might contain a
short lasting stimulating component as well as a longer lasting
orienting component.

1982]

Holldobler — Cerapachys

21

Discussion

Raiding expeditions in Cerapachys turneri are organized by indi-
vidual scout ants, that return to the colony after having discovered a
nest of the prey species. The scout lays a chemical trail with secre-
tions from the poison gland, which serve as recruitment and orienta-
tion signals for the nestmates. Circumstantial evidence suggests that
in addition the scout releases a stimulating chemical recruitment
signal from the pygidial gland. This occurs probably when the
scouts move with their gaster held slightly upwards in a calling
position.

Wilson (1958) reports the field notes made by H. Potter on the
cerapachyine species Phvracaces potteri, which contain the only
available description of the early stages of a complete raid observed
in the field. Before the raid started Potter noted a few workers
moving rapidly about, “each with its abdomen raised upwards”.
These observations match closely my findings in the laboratory and
lend further support to the hypothesis that in addition to the trails
laid with poison gland secretions, another stimulating signal is dis-
charged, presumably from the pygidial gland of the recruiting ants.

Wilson (1958) observed groups of Phvracaces moving along a
raiding trail laid down by a raiding party on the previous day. In
this case no individual leadership was involved and the foragers
seemed to emerge from the nest randomly without a special recruit-
ment stimulation by scout ants. Obviously these ants were following
an established foraging trail, leading to a previously raided Phcidole
nest which appeared to be vacated this time. Small exploratory
parties conducted brief excursions to the side, but in most cases they
turned back to the main trail. No nest suitable for raiding was found
during these explorations.

These observations strongly suggest that chemical trails laid dur-
ing raiding expeditions might still function as orientation cues one
day later and that foraging parties can follow these established trails
without the leadership of a recruiting scout ant. Indeed, my labora-
tory experiments with Cerapachys have demonstrated that artificial
trails drawn with poison gland material are effective as orientation
cues at least for several hours.

22

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[Vol. 89

Although the raiding cerapachyine ants are usually enormously
outnumbered by the worker force of the prey species, not one Cera-
pachys worker was lost during all the raiding experiments in the
laboratory. As can be seen from Fig. 2, Cerapachys and Sphinc-
tomyrmex are excellently protected by a heavily sclerotized cuticle.
The intersegmental joints, that is, the joints between head and
thorax, and between thorax, petiole and gaster, are covered by
cuticular projections so that no intersegmental membrane is ex-
posed, even if the ant is twisted and bent to an extreme degree.

In addition, Cerapachys and probably all the other cerapachyine
ants have a most powerful sting that immobilizes the opponents
within seconds. Not only the adults of the raided colony, but also
the captured larvae and pupae are stung by the raiders before they
are retrieved to the Cerapachys nest. Observations and experiments
demonstrated that the prey larvae are kept in a stage of metabolic
stasis and can thereby be stored for a period of more than two
months. This food storage system enables Cerapachys to adjust the
raiding activities to food requirement and supply. From the labor-
atory experiments we can conclude that Cerapachys does not con-
duct daily or periodic raiding expeditions. The frequency of raiding
expeditions depends on the food supply stored inside the Cera-
pachys nest.

I was unable to demonstrate periodic nomadic behavior in Cera-
pachys in the laboratory. I assume that nest emigrations might
occur relatively frequently in this species, but that they do not fol-
low a periodic pattern. Instead, environmental factors such as food
supply or physical conditions of the nest site are likely to play the
important role in inducing a Cerapachys colony to emigrate.

Acknowledgements

Many thanks to H. Engel-Siegel for technical assistance, to E.
Seling for the SEM work, and to W. L. Brown and R. W. Taylor for
identifying the ants. I am most grateful to R. W. Taylor and the
Division of Entomology, CSIRO, Canberra (Australia) for their
generous hospitality. This work was supported by a grant from the
National Science Foundation BNS 80-021613, the National Geo-
graphic Society and a fellowship from the John Simon Guggenheim
Foundation.

1982]

Holldobler — Cerapachys

23

References

Brown, W. L., Jr.

1975. Contributions toward a reclassification of the Formicidae. V. Ponerinae,
Tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichi-
ni, and Aenictogitini. Search‘5, 1-115.

Holldobler, B. and C. P. Haskins

1977. Sexual calling behavior in primitive ants. Science 195, 793-794.
Holldobler, B. and H. Engel

1978. Tergal and sternal glands in ants. Psyche (Cambridge) 85, 285-330.
Holldobler, B., H. Engel and R. W. Taylor

1982. A new sternal gland in ants and its function in chemical communication.
Naturwissenschaften in press.

Jessen, K., U. Maschwitz and M. Hahn

1979. Neue Abdominaldriisen bei Ameisen. 1. Ponerini (Formicidae: Poneri-
nae). Zoomorphologie 94, 49-66.

Maschwitz, U. and P. Schonegge

1977. Recruitment gland of Leptogenvs chinensis: a new type of pheromone
gland in ants. Naturwissenschaften 64, 589.

Maschwitz, U., M. Hahn and P. Schonegge

1979. Paralysis of prey in ponerine ants. Naturwissenschaften 66, 213.

Oeser, R.

1961. Vergleichend-morphologische Untersuchungen liber den Ovipositor der
Hymenopteren. Mitt. Zool. Mus. Berlin 37, 1-1 19.

Rathmayer, W.

1962. Das Paralysierungsproblem beim Bienenwolf, Philanthus triangulum F.
(Hym. Sphec.) Z. Vergl. Physiol. 45, 413-462.

Rathmayer, W.

1978. Venoms of Sphecidae, Pompilidae, Mutilidae, and Bethylidae. Hand-
book of Experimental Pharmacology vol. 48, Arthropod Venoms (S.
Bettini, ed.) pp. 661-690. Springer-Verlag, Heidelberg-New York, 1978.

Wheeler, G. C.

1950. Ant larvae of the subfamily Cerapachyinae. Psyche 57, 102-1 13.
Wheeler, G. C. and J. Wheeler

1964. The ant larvae of the subfamily Cerapachinae. Suppl. Proc. Entomol.
Soc. Washington 66, 65-71.

Wilson, E. O.

1958. Observations on the behavior of the cerapachyine ants. Insectes Sociaux
5, 129 140.

Wilson, E. O.

1971. The Insect Societies. Belknap Press of Harvard University Press, Cam-
bridge (Mass.).

DESIGNATION OF A TYPE-SPECIES FOR
CYCLOGASTER MACQUART, 1834, AND THE
RESULTING SYNONYMY (DIPTERA: STRATIOMYIDAE)*

By Norman E. Woodley
Museum of Comparative Zoology
Harvard University
Cambridge, Massachusetts 02138

The generic name Cyclogaster Macquart (1834) has been used in
combination with specific names for taxa of Stratiomyidae from
diverse regions of the world. It has remained more or less in synony-
my with Lasiopa Brulle (1832) since the time of Brauer (1882),
although Pleske (1901: 336) described Cyclogaster caucasica (Palae-
arctic) and Hutton (1901: 10) described C. peregrinus from New
Zealand after Brauer’s work appeared. Kertesz (1908) also consid-
ered the two names synonymous, and placed 15 species in Lasiopa.
These species are placed in at least five genera at the present time.

The purpose of this paper is to designate a type-species for Cyclo-
gaster, which to my knowledge has never been done, in order to
stabilize the generic synonymy as it is presently used by workers in
the Stratiomyidae. A brief review of the history of the name Cyclo-
gaster and generic names associated with it is necessary to under-
stand the situation fully.

Macquart (1834: 256) first proposed the name Cyclogaster in the
Diptera, and included in that taxon two species, Nemotelus villosus
Fabricius (1794: 270; Palaearctic) and Stratiomys at rata Fabricius
(1805: 83; Neotropical). No single type-species was designated.

The generic name Inermyia Bigot (1856: 82, 63) was proposed for
the South African species Stratiomys edentula Wiedemann (1824:
29). Gerstaecker (1857: 322) and Loew (1860: 7) both considered
Stratiomys edentula a member of Cyclogaster Macquart and Ker-
tesz (1908: 30) listed Inermyia as a synonym with a query. Lindner
(1972: 32) considered the species to be congeneric with the true,
Palaearctic Lasiopa, and it is listed as such by James (1980: 260).

Kirkaldy (1910: 8) noted that the name Cyclogaster was preoccu-
pied in zoology by Cyclogaster Gronovius, in the fishes (this name
will be discussed in more detail below). He proposed a replacement
name for the name in the Diptera, Neotropicalias. No reference was
made to any specific names, although one might infer he was think-

25

26

Psyche

[Vol. 89

ing of the Neotropical species that Macquart had originally included
in Cyclogaster.

Enderlein (1914: 579, 615), without any reference to Kirkaldy
(1910), but evidently realizing that the two species originally in-
cluded in Cyclogaster were not congeneric, proposed the name
Labocerina for Stratiomys at rat a Fabricius. In his paper, the new
name was spelled Labocerina twice (pp. 579, 615), and “ Labacerino ”
once (p. 615), and has subsequently been spelled “ Labocerino ” by
James (1940: 124). These latter two spellings were regarded as errors
by James (1973: 26.29). In the same paper, Enderlein considered
Cyclogaster a synonym of Lasiopa.

The name Cyclogaster Gronovius (1756: 9; 1760: 265; 1763: 55)
was in dispute, as were all of his generic names, because many
authors felt his work was not truly binomial. His Cyclogaster was
first published in 1756, but this is pre-Linnean. The 1760 work is
clearly not binomial, although this is the date of the name usually
found in zoological nomenclators, being the first post-Linnean publi-
cation of it. In 1954, the International Commission on Zoological
Nomenclature formally ruled that Gronovius’ 1763 work, as well as
an index of it subsequently published by Meuschen, be placed on
the Official Index of Rejected and Invalid Works in Zoological
Nomenclature. Thus Macquart’s Cyclogaster became the earliest
valid use of the name in zoology.

Lindner (1958: 432), while discussing “Cyclogaster” peregrinus
Hutton from New Zealand, recounted most of the above briefly,
and noted that no type-species had been designated for Cyclogaster
Macquart, but was apparently not aware of the I. C. Z. N. ruling.
He also mentioned that Nemotelus villosus Fabricius was the type-
species of Lasiopa (as had Enderlein, 1914: 613, and several other
authors), which is erroneous, as the only species name associated
with Lasiopa in Brulle’s original description was Lasiopa peleteria,
which was described concurrently and is still regarded as a valid
species.

As I interpret the situation, a type-species designation is necessary
for Cyclogaster Macquart in order to stabilize generic synonymy,
and as far as I am aware, this has never been done. In order to
preserve the presently accepted generic synonymies, I hereby desig-
nate Nemotelus villosus Fabricius, originally included in Cyclogas-
ter by Macquart, as type-species for that genus. The following

1982]

Woodley — Cyelogaster

27

synonymy for Lasiopa, the senior generic name, results:

Lasiopa Brulle, 1832: 307. Type-species: L. peleteria Brulle, 1832: 308 (by monotypy).
Cyelogaster Macquart, 1834: 256. Type-species: Nemotelus villosus Fabricius, 1794:
270 (by present designation).

Inermvia Bigot, 1856: 82. Type-species: Stratiomys edentu/a Wiedemann, 1824: 29
(by original designation, op. cit. :63).

Neotropiealias Kirkaldy, 1910: 8; replacement name for Cyelogaster Macquart, 1834,
nee Gronovius, 1763. Type-species: Nemotelus villosus Fabricius, 1794: 270 (by
autotypy).

The above type-species designation thus stabilizes the long-used
synonymy of Cyelogaster with Lasiopa, while retaining the name
Laboeerina Enderlein for the Neotropical Stratiomys atrata Fabri-
cius. The name Neotropicalias Kirkaldy became an unnecessary,
and therefore invalid, replacement name when Cyelogaster Grono-
vius was rejected by the I. C. Z. N. ruling.

Acknowledgments

I wish to thank Curtis W. Sabrosky and Margaret K. Thayer for
critically reading the manuscript.

Literature Cited

Bigot, J. M. F. 1856. Essai d’une classification generale et synoptique de l’ordre
des Insectes Dipteres. (4e Memoire.) Ann. Soc. Ent. Fr. (3)4: 51-91.

Brauer, F. 1882. Zweifltigler des Kaiserlichen Museums zu Wien. II. Denk-
schr. Akad. Wiss. Wien 44(1): 59-1 10.

Brulle, G. A. 1832. IVe Classe. Insectes. Pp. 64-395, in Bory de Saint-
Vincent (ed. ), Expedition scientifique de Moree. Section des sciences physiques
3(1) (Zool. 2), Paris. 400 pp.

Enderlein, G. 1914. Dipterologische Studien. IX. Zur Kenntnis der Stratio-
myiiden mit 3astiger Media und ihre Griippierung. A. Formen, bei denen der I.
Cubitalast mit der Discoidalzelle durch Querader verbunden ist oder sie nur in
einem Punkte beriihrt (Subfamilien: Geosarginae, Analcocerinae, Stratiomyii-
nae). Zool. Anz. 43: 577-615.

Fabricius, J. C. 1794. Entomologia systematica emendata et aucta. Secundum
classes, ordines, genera, species adjectis synonimis, locis, observationibus, de-
scriptionibus. Vol. 4. Hafniae. 472 pp.

1805. Systema antliatorum secundum ordines, genera, species adiectis

synonymis, locus, observationibus, descriptionibus. Brunsvigae. 372+ 30 pp.

Gerstaecker, A. 1857. Beitrag zur Kenntniss exotischer Stratiomyiden. Linn.
Ent. 11: 261-350.

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Gronovius, L. T. 1756. Musei Ichthyologici tomus secundus sistens Piscium
indigenorum & nonnullorum exoticorum, quorum maxima pars in Museo Lau-
rentii Theodori Gronovii, J. U. D. adservatur, nec non quorumdam in aliis
Museis observatorum descriptiones. Accedunt nonnullorum exoticorum Pis-
cium icones aeri incisae, et Amphibiorum Animalium Historia Zoologica. Lug-
duni Batavorum. [i-viii] + 1-88 pp.

1760. Animalium in Belgio Habitantium centuria prima. Acta Helvet-
ica 4: 243-270.

1763. Zoophylacii Gronoviani Fasciculus primus exhibens Animalia

Quadrupeda, Amphibia atque Pisces, quae in Museo suo adservat, rite exami-
navit, systematice, disposuit, descripsit, atque iconibus illustravit. Lugduni
Batavorum. [i-iv] + 1 136 pp.

Hutton, F. W. 1901. Synopsis of the Diptera brachycera of New Zealand. Trans.
Proc. New Zealand Inst. 33: 1 95.

International Commission on Zoological Nomenclature. 1954. Opinion
261. Rejection for nomenclatural purposes of the Index to the Zoophylacium
Gronovianum of Gronovius prepared by Meuschen (F. C.) and published in
1781. Opin. Decl. Int. Comm. Zool. Nom. 5: 281-296.

James, M. T. 1940. Studies in Neotropical Stratiomyidae (Diptera). IV. The
genera related to Cvphomyia Wiedemann. Revista Ent. 11: 119 149.

1973. Family Stratiomyidae, No. 26, in A catalog of the Diptera of the

Americas south of the United States. Sao Paulo. 95 pp.

1980. 20. Family Stratiomyidae, pp. 253-274, in Crosskey, R. W.,

ed. Catalogue of the Diptera of the Afrotropical Region. London. 1437 pp.

Kertesz, K. 1908. Catalogus Dipterorum. Volumen 111. Stratiomyiidae, Erin-
nidae, Coenomyiidae, Tabanidae, Pantophthalmidae, Rhagionidae. Buda-
pestini. 366 pp.

Kirkaldy, G. W. 1910. On some preoccupied generic names in insects. Canad.
Ent. 42: 8.

Lindner, E. 1958. Uber einige neuseelandische Stratiomyiiden Osten-Sackens im
Deutschen Entomologischen Institut in Berlin. Beitr. Ent. 8: 431-437.

1972. Uber einige Stratiomyidae des Transvaal Museums (Diptera: Bra-
chycera). Ann. Transvaal Mus. 28: 27 34.

Loew, H. 1860. Die Dipteren-Fauna Siidafrika’s. Erste Abtheilung. Abh. Naturw.
Ver. Sachsen u. Thiiringen in Halle 2: 57-402.

Macquart, J. 1834. Histoire naturelle des Insectes. Dipteres. Vol. 1. Paris.
578 pp.

Pleske, T. 1901. Studien fiber palaearktische Stratiomyiden. I. Die Gattung
Cyclogaster Macqu. Sitzungsber. Naturf. Ges. Univ. Jurjeff (Dorpat) 12:
335-340.

Wiedemann, C. R. W. 1824. Munus rectoris in Academia Christiana Albertina
aditurus Analecta entomologica ex Museo Regio Havniensi maxime congesta
profert iconibusque illustrat. Kiliae. 60 pp.

ORB PLUS CONE-WEBS IN ULOBORIDAE (ARANEAE),
WITH A DESCRIPTION OF A NEW GENUS
AND FOUR NEW SPECIES

By Y. D. Lubin,1, B. D. Opell,2, W. G. Eberhard,3
and H. W. Levi4

Introduction

Spiders of the genus Uloborus (Uloboridae) characteristically
spin horizontal orb-webs with a sticky spiral of cribellar silk. We
describe here the webs of U. conus, U. albolineatus, U. bispiralis, U.
#2072, U. trilineatus, and Conifaber parvus which are modifications
of this basic uloborid orb-web form and include cones composed of
regular arrays of threads beneath the orbs’ lower faces. The web
building and prey capture behaviors of U. conus (observations of
YDL) are also described, and descriptions of Conifaber parvus new
genus, new species and the new species U. conus, U. albolineatus,
and U. bispiralis are provided (by BDO).

Study Sites and Methods

Uloborus conus was found at three localities in Papua New
Guinea: 1) in lowland wet forest, Gogol Forest Reserve near
Madang, Madang Province, 2) in a Pandanus swamp (freshwater)
and a mangrove swamp (brackish) at Buso, Morobe Province, and
3) in the understory of klinki pine (Araucaria hunsteinii ) plantations
at 1200 m elevation in McAdam Memorial Park near Wau, Morobe
Province. Webs were built about 0.5 to 2.0 m above the ground in
gaps formed by the uppermost, generally vertical branches of small
shrubs and saplings. They were always found in humid, shaded

1. Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Panama and
Department of Zoology, University of Florida, Gainesville, Florida, 32611.

2. Department of Biology, Virginia Polytechnic Institute and State University,
Blacksburg, Virginia 24061.

3. Smithsonian Tropical Research Institute and Escuela de Biologia, Universidad de
Costa Rica, Ciudad Universitaria “Rodrigo Facio”, Costa Rica.

4. Museum of Comparative Zoology, Harvard University, Cambridge, Massachu-
setts 02138.

* Manuscript received by the editor September 25, 1981.

29

30

Psyche

[Vol. 89

locations. Several individuals were kept and observed in an insect-
ary at the Wau Ecology Institute (WEI).

Uloborus albolineatus and U. bispiralis were found on the Gazelle
Peninsula, East New Britain (ENB), Papua New Guinea. The webs
of U. bispiralis were observed on the Lowlands Agricultural Experi-
mental Station (LAES) at Kerevat, ca. 100m elevation, in cocoa
plantations and in secondary growth lowland forest and near
Malasat (ENB) at ca. 600m elevation. One web of U. albolineatus
was observed at LAES in secondary-growth forest along a river.

A single mature female of Uloborus #2072 (numbers refer to
specimen numbers placed in vials) was found (by WGE) near Dan-
deli, Karnataka, India, in the foliage of a bush growing in a teak
forest. Uloborus trilineatus is common in undergrowth of gallery
forest in eastern Colombia where WGE worked extensively. The
webs described here were found at Finca Chenevo, about 20 km SW
of El Porvenir, Meta, and Finca Mozambique, about 15 km SW of
Puerto Lopez, Meta. Conifaber parvus was also found at Finca
Mozambique (by WGE) where it occurred in periodically flooded
forest but not in surrounding savanna.

Webs were first dusted with cornstarch or talcum powder using
either the method described by Eberhard (1977a) or Carico’s (1977)
modification of this method, and then measured and photographed.
All specimens mentioned in this paper are deposited in the Museum
of Comparative Zoology.

Observations
Uloborus conus *

The Web

The web of U. conus has three parts: the inner orb, the rim, and
the cone (Fig. 1). The inner orb and rim are in nearly the same plane
and are more or less horizontal. The inner orb consists of a closed
hub, radii and a few loops of non-sticky spiral, while the rim has
several loops of sticky, cribellar spiral which end where the rim radii
join those of the inner orb. Rim radii are continuous with those of
the cone, and those of the inner orb are attached to them. Cone radii
are attached in groups of two or three to a central guy thread which

♦This is a new species decribed below.

1982]

Lubin, Opell, Eberhard, Levi — Uloboridae

31

Figures 1-2. Web of Uloborus conus. 1. Side view showing the rim sticky spiral
(RS), inner orb (I), cone (C) with jagged sticky spiral (CS) on a framework of radii
and non-sticky spiral, and cone radii (CR) converging toward a central guy thread.
Note that 2-3 cone radii are attached together to form one thread which attaches to
the cone guy thread, and that these attachments are dispersed along the guy thread so
that there is no single apical point to which all cone radii attach. 2. Top view
showing typical Uloborus- type hub and non-sticky spiral of the inner orb. The cone
with its jagged sticky spiral (CS) is seen through the plane of the orb. Note the gap
between the non-sticky spirals of the cone and inner orb on the one hand and the rim
sticky spiral on the other. The cone sticky spiral can be seen as a continuation of the
rim spiral (arrow points to beginning of cone sticky spiral).

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is in turn attached distally to a leaf or branch. The cone has a non-
sticky spiral and a few irregularly-spaced, jagged turns of cribellar
silk. This jagged sticky spiral is a continuation of the innermost
sticky spiral loop in the rim (Figs. 2, 4).

The hub of the inner orb (Fig. 2) is similar to that of other
uloborid orbs, e.g. U. diver sus (Eberhard, 1972), and its spiral
continues outward to form the non-sticky spiral of the inner orb.
There is always a large gap between the last turn of this spiral and
the innermost loop of sticky rim spiral (Figs. 1, 2).

Sticky spiral loops in the rim are more tightly spaced than are
either the non-sticky spiral loops of the inner orb and cone or the
cone’s sticky spiral. The outermost loop of rim spiral often follows a
zigzagging path, with some segments of the sticky silk found on the
radii (Figs. 2, 3). This zigzagging was more pronounced in some
webs than in others and was generally most evident on the side of
the orb which was larger (the orbs were rarely perfectly symmetrical).

Variations on this basic pattern were seen. Webs of immatures
frequently had only a narrow rim, sometimes with only a single loop
of sticky spiral. Some webs had a few loops of sticky spiral on the
inner orb, with the non-sticky spiral left intact (Fig. 4). Webs of two
adult females and several immatures had thin linear stabilimenta at
their inner hubs. Adult males were found sitting on webs similar to
those of immatures, but it was not determined if these were of their
own construction. Adult males did not build webs in captivity.

Web Building Behavior

Web building by two adult females was observed from start to
finish and various stages of web construction were seen on four
other occasions. Durations of different stages of construction were
noted for one of the adult females. Web construction began late at
night or in early pre-dawn hours. The inner orb and cone of the old
web were probably removed early in the night, but this behavior was
not observed. One WEI female was found sitting at the center of a
rudimentary web consisting of a partly collapsed rim and a few radii,
and had a ball of silk in her mouthparts which shrank visibly as it
wasj(presumably) ingested. This spider removed the rest of the rim
and added the material to the ball of silk in her chelicerae before
building the new web. Reusing frame threads from the previous
web, the spider began construction by laying new radii.

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33

Figures 3-4. Web of Uloborus conus. 3. Detail of first (outermost) loop of rim
sticky spiral showing zigzag path with sticky silk laid directly on the radii. 4. Top
view of web with 1 x/i loops of sticky spiral (IS) in the inner orb (IS). Also visible is the
cone sticky spiral (CS) continuing in from the rim spiral (RS) and the zigzag outer
loop of rim sticky spiral.

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Radii and non-sticky spiral were laid as in U. diversus (Eberhard,
1972) and their construction lasted 5 and 1.5 minutes, respectively.
Radii were laid by walking out from the hub on an existing radius
with a dragline, attaching the dragline to a frame thread, and then
doubling it by walking back to the hub with another dragline. At the
hub the dragline was attached to a succession of adjacent radii
(forming the closed hub spiral) before the next radius was laid.
When most of the radii were completed, the spider continued the
hub spiral outward to form the non-sticky spiral, laying occasional
“tertiary radii” (Le Guelte, 1966) during the process. This non-sticky
spiral did not reach the frame threads.

At the start of the sticky spiral even very faint light falling on the
spider caused her to cease spinning and bounce up and down on the
web. Consequently, observations of sticky spiral construction were
made only sporadically, using indirect lighting. The first (outer-
most) loop of non-sticky spiral was completed in 13 min. During
sticky spiral construction the spider reversed directions five times in
the larger part of the web. The sticky spiral was attached to each
radius that it crossed, and the spider broke non-sticky spiral loops as
she laid the sticky spiral. One immature female was observed laying
a zigzag outer loop of sticky spiral. The sequence of attachments of
the cribellar silk to produce the zigzag loop (Fig. 5a) was distinct
from that involved in laying the normal sticky spiral loops (Fig. 5b).

After meticulous, slow sticky spiral construction, which in one
case lasted 3 hrs. 6 min., the spider suddenly began spinning out
cribellar silk in a rapid and seemingly reckless fashion while moving
inward toward the hub at an angle of about 25° to the last turn of
the regular sticky spiral (Figs. 2, 4). After completing half a loop,
the spider reversed direction and continued spiralling toward the
hub, laying a jagged and irregularly spaced sticky spiral. The jagged
spiral was attached to only a few radii, crossing 3-7 radii and, in
some cases, several non-sticky spiral loops between attachments.
The non-sticky spiral was left intact. This entire phase was very
rapid and in one case the four jagged loops were completed in just 6
min. This jagged spiral was to become the sticky spiral of the future
cone.

After completing the cone sticky spiral, the spider moved to the
hub and slowly turned in a circle, pulling on successive radii with the
first legs. After 2 min. she went out to the end of a radius and

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35

Figure 5. Construction of U. conus web. (a) Sequence of attachments of sticky
silk to produce the outer zigzag loop of rim sticky spiral. The spider started at the
junction of the radius (Rl) and frame thread (F), attaching the cribellar thread at
point A, walked along Rl toward the hub and attached the cribellar thread at point
B, about half way between the frame thread and outer loop of non-sticky spiral (NS).
The spider then continued inward along Rl, combing out cribellar silk, reached the
non-sticky spiral and ran rapidly across it and 2/3 of the way out on R2 without
combing out additional silk. It then continued to walk out on R2, combing out
cribellar silk and attached the thread at point C, the junction between R2 and the
frame thread. The sequence was then repeated, walking in along R2, attaching
cribellar thread at point D, etc. (b) Sequence of attachments of cribellar silk to
produce the normal sticky spiral. The spider attached cribellar thread at point A on
radius Rl, walked in on Rl, combing out cribellar silk, until it reached the
temporary, non-sticky spiral loop (NS), then ran along the non-sticky spiral and out
on radius R2 without combing out cribellar silk and attached the cribellar thread to
R2 at point B.

dropped from it to a leaf below, attached her dragline to the leaf,
and went back up the dragline and across the web to its hub on a
radius, attaching the new dragline from the leaf to the hub. This
formed the central guy thread of the cone. The spider then went
down the guy thread, broke it, reattached it to a different point on
the leaf, and then returned to the hub. By this time the hub was al-
ready drawn down under tension, and the web formed a shallow
cone. The cone was then elongated by cutting radii at their
attachment to the hub, lowering their tension and then attaching
them to the central guy thread by the sequence of behaviors shown
in Fig. 6a, b.

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Figure 6. Construction of U. conus web. Sequence of thread attachments in
forming the cone (web viewed from the side). Arrows indicate direction of movement
of the spider. Dots are points where attachments were made or broken, (a) The spider
went to point X on radius R1 at the edge of the hub, cut the radius, attached its
dragline to the inner broken end and then let out additional dragline as it faced away
from the hub. This was then attached to the outer broken end which had now moved
to point Xi. Usually adjacent radii were also broken and attached to radius Ri at
point Xi (see also Fig. 1). (b) The spider then walked back toward the hub to point

Y, attached a dragline, ran to the hub and down the central guy thread (G), attaching
the dragline at point Z. Radius Rl was thus pulled down toward the apex of the cone
to form the cone radius YZ while the thread HY formed a temporary inner orb
radius, (c) To move the temporary inner orb radius up on the cone, the spider
walked out on temporary radius HY and broke it at its attachment to the cone at
point Y. The spider then attached a dragline to the broken end and walked out on
radius Rl, reattaching it at point A at the inner edge of the rim sticky spiral.

(d) The completed cone radius is indicated by line AZ and the new inner orb
radius by line HA. The section AY of the cone radius bears the cone sticky spiral. The
upper portion of the guy thread (HZ) was absent in the completed web, but it is not
known when it was removed.

After forming the cone, the spider cut most of the temporary
inner orb radii, thus collapsing the hub and leaving only a bit of silk
to which a few temporary radii were attached. The spider then
began replacing these temporary inner orb radii and at the same
time completing cone formation by incorporating into the cone the

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37

section of the original orb containing the jagged sticky spiral (Fig.
6c, d). This stage followed initial cone formation without interrup-
tion, and it was difficult to determine when cone building ended and
replacement and construction of new inner orb radii began. The
spider went out to the cone along a temporary radius, broke the
attachment to the cone and attached her dragline to the inner end of
the temporary radius, then carried the radius upward by walking
along radii and non-sticky spiral loops on the inner surface of the
cone, and finally reattached it at or just inside (below) the innermost
loop of the rim sticky spiral. She then walked back to the hub on the
new radius, thereby doubling the thread. Upon reaching the center,
she made attachments to form a new hub. The upper portion of the
guy line was absent in finished webs, but it was not determined how
it was removed.

Additional new inner orb radii were constructed in much the
same manner as “normal” orb radii. The spider went out on an
existing radius (or temporary radius) with a dragline, reached the
cone non-sticky spiral, walked across it to the next cone radius,
attached the dragline to the cone radius just below (inside) the rim
spiral, and return to the hub on the new radius (doubling it).
Consecutive radii were always laid with angles of more than 90°
between them, perhaps serving to reduce differences in tension on
all sides of the orb (Eberhard, 1981).

The last stages of web building, beginning with attachment of the
dragline and ending with completion of the inner web, lasted 23
min.

Resting Postures

The spider normally sat under the hub with legs I and II slightly
flexed and holding separate radii. When disturbed, the spider
adopted a cryptic posture with legs I and II held together and flexed
and legs III and IV pressed close to the body (Figs. 2, 4). This
posture was adopted either at the hub or under a short “dragline”
thread beneath the hub, which was attached to the hub at one end
and to a radius at the other. When disturbed repeatedly, or when
sunlight struck the web and made it visible, the spider dropped from
the hub onto the dragline thread and bounced up and down on it.
Spiders also bounced while wrapping prey and sometimes while
going out to attack an insect or upon returning to the hub. This

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bouncing may be an anti-predator behavior similar to the bouncing
flight of craneflies and the rapid vibrating of opilionids and
pholcids.

Prey Capture Behavior.

Successful captures of five fruitflies ( Drosophila-sizQ ), one 4 mm
long dolichopodid fly, one unidentified 1 mm fly, three 3^t mm
ants, and one 5 mm lepidopteran larva were observed (by YDL). Of
these, seven were trapped in the rim and three in the cone. All but
one sequence conformed to the description given below. Like other
uloborids (Marples, 1962; Eberhard, 1969; Lubin et al., 1978) U.
conus and U. bispiralis immobilize all insects by wrapping in silk.
Spiders ran out to the cone on an inner orb radius to reach insects
trapped in the rim sticky spiral, squeezed through the cone (often
turning sideways to do so) and continued out onto the undersurface
of the rim. If an insect was trapped on the cone sticky spiral, the
spider went through the cone and ran down the outer surface of the
cone. Upon reaching the insect, the spider often tapped it with legs I,
turned 180° so that it faced the hub (or upward on the cone) and
began to wrap. Initially the prey was wrapped from a distance by
throwing sheets of silk backwards with legs IV. Later the spider
moved into contact with the prey and held it with legs II and III
while wrapping. The spider interrupted wrapping to cut sticky spiral
attachments, then cut the inner radius attachment (toward the hub)
and continued to wrap while holding the end of the radius with one
leg I. Finally, the outer (distal) end of the radius was cut and the
prey was held free of the web in legs II and III while the spider hung
from the broken radius by legs I, bridging the gap with its body, and
wrapped the prey with legs IV while rotating it occasionally with the
palps or legs.

All prey were carried to the hub in the palps (with the aid of the
chelicerae), held “overhead” in characteristic uloborid fashion.
After transferring the prey package from the legs to the palps, the
spider attached a dragline to the distal end of the broken radius and
then to the proximal end, thus closing the gap. At the hub the spider
again transferred the prey from the palps to legs II and III and
wrapped it while hanging from the dragline thread beneath the hub.
In most instances the dragline thread appeared to be broken and the
spider spanned the gap with its body.

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39

Prey Capture Sequences With Different Prey Types.

The only case not conforming to this description was that of a
fruitfly caught on the inner orb; the spider wrapped it, secured it by
reattaching it to the radius and fed on the prey in situ.

U. conus rejected or ignored a number of insects offered as prey.
Five small orthopteran nymphs 3-4 mm long (probably newly
emerged) where given to adult females and all were rejected. On two
occasions, the spiders approached and tapped the insects with legs I
and then returned to the hub. In other instances the spider pulled
the radii in the direction of the orthopteran, shook the web and then
ignored it. The same individuals readily attacked fruitflies offered as
prey after the orthopterans. Fruitflies were not attacked on three
occasions when they were offered while the spider was already
wrapping a prey or feeding at the hub. Two ants ( Anopolepis
longipes, 4mm long) were rejected under the same circumstances.

Sequences With Multiple Prey.

On six occasions spiders feeding at the hub attacked second or
third prey thrown into their webs. These included two ants, two
fruitflies, a dolichopodid fly and an unidentified small fly. On all but
one occasion the spider carried the first prey in its palps as it ran out
to attack the second. In one instance a spider that had been
wrapping the first prey at the hub attached this insect to a dragline
thread below the hub before going out to attack the second insect.

The second prey was immobilized in the same manner as the first,
but rather than cut this insect out and carry it to the hub, the spider
secured the second prey at the capture site and returned to the hub
to resume feeding on the first prey. While performing immobili-
zation wrapping, the spider usually broke the radius attached to the
prey on the inner side (toward the hub), but not on the outer side.
Before leaving it at the capture site, the spider reattached the prey to
the broken end of the radius, thus securing it at both ends.

Eggsac and Eggsac Web.

The eggsac of U. conus is about 8mm long by 3mm wide, with
angular projections along the edges (Fig. 7). It is suspended in an
eggsac web on a strengthened radius of a former web, where the hub
of the inner orb had been. The web is similar to those of U. diversus
(Eberhard, 1969) and Miagrammopes sp. near unipus (Lubin et al.
1978) and consists of frame threads, a few radii and one or more

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zigzag loops of sticky silk, with some sticky silk laid directly on the
radii. The radii are attached to the main eggsac radius or to the
eggsac itself. One female had a three-dimensional eggsac web
consisting of a rudimentary cone and inner orb radii (Fig. 7) with
sticky silk in both the plane of the orb and the cone. Unlike the
eggsac webs of Miagrammopes, these webs were retained both day
and night. Insects that became entangled in the sticky threads were
attacked in the usual manner.

Females guarded their eggsacs (one per female) until the young
emerged (13 days for one eggsac). Newly emerged spiderlings
remained on the eggsac web for one or two days, then moved away
and constructed typical Uloborus-type “baby webs”, consisting of
radial threads connected by a thin sheet of very fine, non-sticky silk
(Szlep, 1961; Eberhard, 1977b) without any cone. One immature,
however, had an orb plus cone-web with a filmy “baby web” sheet
where the rim sticky spiral would normally be found and also some
“baby web” sheet on the cone. Structural spirals were present in the
rim and inner orb; there was no sticky spiral.

Uloborus bispiralis*

The cone web of U. bispiralis (Fig. 8) is similar to that of U. conus
in that the cone sticky spiral is continuous with that of the rim, and
the outer loop(s) of rim spiral follow a zigzag path, with some sticky
silk laid on the radii. Unlike webs of U. conus , the inner orb non-
sticky spiral extends right up to the innermost (last) loop of rim
sticky spiral and all webs had a few loops of sticky spiral in the inner
orb. Most webs also had a thin, linear stabilimentum of white silk
across the inner orb, with a spider-size gap at the hub.

Webs of juvenile females were similar in all respects to those of
adults. None of the webs observed showed signs of repairs. Like
those of U. conus, they are probably renewed daily. On one
occasion only, a juvenile female was seen hanging inside the cone
while an adult male fed on prey at the hub. Another adult male was
observed sitting at the edge of an adult female’s web and a third
male was found sitting in a small cone-web (no sticky spiral was
observed).

This is a new species, described below.

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41

Figures 7-9. Uloborus. 7. Eggsac and three-dimensional eggsac web of Ulo-
borus conus. The female spider can be seen sitting in a cryptic posture to the left of
the eggsac. Sticky threads (heavy white lines) occur in the plane of the former orb and
on the rudimentary cone. 8. Web of Uloborus bispiralis. 9. Tubular eggsac of
Uloborus bispiralis with female sitting in cryptic posture at one end of the eggsac
(arrow).

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The long, tubular eggsacs of U. bispiralis (34-40 mm long and 1 .5
mm wide) have no angular projections (Fig. 9) and resemble those
of Miagrammopes (Lubin et al. 1978). They are suspended along the
radius of a former web of which only a few radii and frame threads
remained intact. There was no evidence of sticky silk in the four
eggsac webs examined. Spiders sat in line with the eggsacs, with legs
I and II extended forward and legs IV grasping the eggsac, and were
reluctant to move even when prodded.

Uloborus albolineatus*

One individual of U. albolineatus was observed on a cone web
similar to that of U. bispiralis. The rim spiral had one or two zigzag
outer loops, and both the cone and inner orb had jagged loops of
sticky spiral. The inner orb non-sticky spiral extended almost to the
rim spiral. The female sat at the hub with legs I and II extended
forward and held together and legs IV extended backward.

Uloborus sp. (2072)

Only a single web was seen. It consisted of a somewhat inclined
orb (43° with horizontal) with a cone underneath it which contained
loops of sticky spiral (Fig. 10a, b). This web differed from those of
U. Conus in having sticky spiral threads near the center of the
horizontal orb (Fig. 10c) as well as near its edge, as well as having
some of the “radial lines” of the cone attached directly to the frame
of the orb while others ended on radii as in U. conus webs.

At the hub the spider sat in a “crouched” position (Fig. 10a)
similar to that of Philoponella (Opell and Eberhard in prep.), and
was reluctant to move away when disturbed.

Uloborus trilineatus Keyserling

Most of the many webs of mature and immature U. trilineatus
individuals observed were typical, more or less horizontal orbs like
those spun by other Uloborus species (e.g., Szlep, 1961; Wiehle,
1927; Eberhard, 1972). Webs of mature males were similar to those
of newly emerged uloborid spiderlings (Szlep, 1961; Eberhard,

♦This is a new species, described below.

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43

Figure 10. Web of Uloborus sp. (#2072). A. Side view with spider (arrow) at hub. B. Top view. C. Enlargement of the
hub. Both cone and orb spirals are sticky. Most cone radii are attached to orb radii, but some end on frame lines. The cone
sticky spiral seems not to be continuous with the orb spiral.

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1977b). However, at both Finca Chenevo and Finca Mozambique
one immature was found at the hub of a web like that shown in
Figs. 11a, b. Each web consisted of a small, more or less horizon-
tal orb which had only a non-sticky spiral. Below this was a cone
which also had a non-sticky spiral. Only one of these spiders was
collected, the other was left on its web, and the next day the web
was deserted and an exuvium was found clinging to its hub. Identity
of the collected immature specimen is not certain, but abundance of
U. trilineatus at these sites plus the failure of extensive collecting of
orb weavers to reveal similar species in these habitats indicates that
these immatures were U. trilineatus.

Conifaber parvus *

This species was fairly common in a periodically flooded forest on
Finca Mozambique. Only mature females were found with webs.
The webs all had an “orb” similar or identical to those spun by most
newly emerged uloborid spiderlings (Szlep, 1961; Eberhard, 1977b),
having radii, hub, frames, and a non-sticky spiral as in typical orbs
but lacking a sticky spiral and having instead a dense mat of very
fine threads (so fine that in Figs. 12a, b they do not show up as
individual threads, and one only sees the grains of cornstarch).
Below this orb was a conical web consisting of radii which
converged below to a single downward-directed line, and a more or
less regularly spaced spiral, also of non-sticky silk. The hubs were
often decorated with linear stabilimenta.

The spider crouched at the hub with its legs I folded ventrally in
the typical Philoponella posture (Opell and Eberhard in prep.).
Sometimes when a spider was disturbed she let herself fall from the
hub and hung suspended inside the cone on her dragline and
bounced actively there. On other occasions spiders bounced on their
orbs.

Attack behavior was observed twice and seemed to be typical for
uloborids. The spider turned to face away from the prey and threw
silk over it with her legs IV, gradually cut it loose as she wrapped it,
then held it with the palps and/or chelicerae as she reattached the
ends of the broken radii, took it to the hub, and then resumed

These are a new genus and species, described below.

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45

Figure 11. Web of penultimate female of Uloborus trilineatus Keyserling.
A. Side view. B. Top view. Most (or all?) of the cone radii are attached to frame
lines. The central area of the cone has fewer radii than the upper portion, and gives
the impression of having been partially dismantled, perhaps during the process of
being connected to the central thread as in U. consu.

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wrapping while hanging there by her spread legs I.

The eggsacs were different from those described for any other
uloborid. They were pure white, 2-3 mm diameter spheres with
projecting spikes, and resembled the heads of maces; they were
suspended in the plane of the orb portion of the web on a radial line
(Figs. 13a, b).

Discussion

While the webs of all five species are similar in having more or less
horizontal orbs with cones below, the details are strikingly different.
The cones of U. conus, U. albolineatus, U. bispiralis, and U. #2702
have a sticky spiral while those of U. trilineatus and C. parvus do
not. In U. albolineatus, U. bispiralis, and U. sp. #2702 both the
outer (rim) and inner portions of the orb have sticky spirals, while in
U. conus the main capture surface is the rim sticky spiral and only
occasionally is a sticky spiral present in the inner orb. The “orbs” of
Conifaber parvus have no sticky spiral, but the dense mat serves as a
trapping surface, as in uloborid “baby webs”. Orb-plus-cone webs of
U. trilineatus have no sticky silk at all.

The function of the cone in webs of all four species is probably
primarily defense of the spider at the hub against predators and
parasites. The cone forms a “cage” of threads around the spider, and
a defense function is suggested both by the fact that U. conus and
Conifaber parvus drop from the orb and hang inside this cone when
disturbed or when the web becomes visible in sunlight, and by the
fact that construction of conewebs by U. trilineatus occurs only
when the spiders are about to enter the particularly vulnerable
moulting period. The sticky threads in the cones of U. conus and U.
albolineatus and U. #2702 sometimes trap prey (some U. conus webs
have almost no other sticky lines), but the fact that the cones of U.
conus, U. albolineatus, and U. bispiralis have only a few, irregularly
spaced sticky spiral loops while those of U. trilineatus and C. parvus
lack sticky threads suggests that prey capture is a secondary
consequence rather than a primary function of at least some of the
cones. Placement of sticky threads in cones could have evolved as an
additional defense of the spider against predation or parasitism.

The uloborid cones resemble the barrier meshes made by the
araneid Nephila maculata (Robinson and Robinson, 1973) at one or
both sides of their more-or-less vertical orbs; in young N. maculata

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47

Figure 12. Conifabar parvus web. A. Side view showing framework threads,
radii, mat of non-sticky spirals, and cone radii. B. Top view showing non-sticky
spiral mat, two stellate eggsacs, and the female (arrow) resting at the web’s hub.

Figure 13. Conifaber parvus web hub and eggsacs. A. Female (arrow) resting at
hub. Two stellate eggsacs and some of the horizontal web’s fine, non-sticky threads
are visible. B. Female (arrow) resting in crouched posture at the hub of a web
decorated with linear stabilimenta.

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the mesh is a cone-shaped, rudimentary orb with no sticky spiral.
The Robinsons attributed a defensive function to these structures,
and indeed the arguments developed here suggest that barrier
meshes made by a number of other araneids ( Metepeira , Leucauge,
Argiope, Arachnura, Gasteracantha, and Phonognatho) may also
function defensively.

The evolutionary origin of the orb-plus-cone web designs in
uloborids is not clear. At least two other uloborid orb-plus-cone
webs are known. Workman (1896) described the orb-plus-cone web
of Uloborus quadrituberculatus (Thorell). His apparently schematic
drawing shows a horizontal orb lacking spiral lines and a cone with
a 14 loop spiral (he did not note whether or not the spiral was
sticky). The cone is attached on all sides to surrounding vegetation
by short lines. In Sembrong Jungle near Layang-Layang, Johore,
Malaya, Frances Murphy photographed the orb-plus-cone web of a
specimen matching Workman’s (1896) description of U. quadri-
tuberculatus. This web was constructed about 1.5 m above the
ground and had a zigzag outer loop and an irregular cone spiral. An
unidentified species of Tangaroa collected in mesophyll rainforest in
the Iron Range, northeastern Queensland, Australia had an orb-plus
cone web with a zigzag outer loop of rim sticky spiral (V. Todd
Davies, personal communication). It is not known if the cone spiral
was sticky. However, a small, unidentified Tangaroa species from
Yap, Caroline Islands constructed a horizontal orb-web in both the
field and lab (Joseph Beatty and James Berry, personal communica-
tion and BDO unpublished observations, respectively), indicating
that the cone-web is not characteristic of all members of this most
primitive uloborid genus (Opell, 1979) and, therefore, does not
represent the “original” uloborid web design.

We do not know if the cones of the five species studied here are
constructed in the same manner. Certain behaviors associated with
cone construction in U. conus (and probably U. albolineatus and U.
bispiralis ) including the laying of a jagged sticky spiral with few
attachments to the radii, formation of a cone by cutting and
reattaching radii to a central line, replacement and reposition of
radii, and pulling the orb into a cone, have not been seen in other
uloborids. When one takes into account the webs of other uloborids
such as Philoponella vicina (Peters 1953, 1955), P. semiplumosa
(Lahmann and Eberhard 1979), P. oweni (Eberhard 1969), P. divisa
(Opell 1979), and P. para (Eberhard, unpub.) which are more or less

1982] Lubin, Opell, Eberhard, Levi — Uloboridae 49

reduced and modified planar or domed orbs in the midst of meshes
which include sticky as well as non-sticky threads ( P . oweni also
spins orbs without meshes— Eberhard, 1969), the “orb” of Polenecia
( =Sybota ) which lacks sticky spirals and has instead sticky radii
(Wiehle 1931), the orbs cum sheet webs spun by young spiderlings
and mature males of several species (Szlep, 1961; Eberhard, 1977b),
and the various simplified webs of Hyptiotes (Wiehle 1927, Marples
and Marples 1937) and Miagrammopes (Akermann 1932, Lubin et
al. 1978), it becomes clear that there is an extraordinary diversity of
web forms in the relatively small family Uloboridae. It is likely that,
in conjunction with morphological studies, a fuller understanding of
the webs and behavior of uloborids will shed more light on relation-
ships within the family.

Systematic Section

Conifaber new genus*

Figures 14-15, 20-29

Type. The type species of Conifaber is Conifaber parvus, new
species. The genus name is a masculine noun derived from the Latin
nouns conus and faber and means “cone craftsman”.

Diagnosis. Conifaber contains the smallest known uloborids,
females being 2.0 mm and males 1 .5 mm long. Because of their small
size members of this genus are more likely to be confused with those
of Ariston and Siratoba than with Zosis, Octonoba, and Purumitra,
to which they are more closely related. Conifaber males and females
are distinguished from those of Ariston and Siratoba (Opell, 1979;
figs. 41, 72) by having a strongly recurved anterior eye row whose
median eyes are located on a slight anterior carapace extension and
have a diameter twice that of the other eyes (Figs. 20-23). Unlike
Ariston and Siratoba females whose first femora are 1.5 and 2.0
times the carapace length, respectively, and whose thoracic grooves
are in the carapace’s posterior two-fifths, Conifaber females have
first femora equal in length to the carapace and have a centrally
located thoracic groove. Like Ariston, but unlike Siratoba, Coni-

*For nomenclatural purposes B. D. Opell is the author of the genus Conifaber and
the species C. parvus.

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[Vol. 89

faber females lack dorsal abdominal tubercles. Like Siratoba but
unlike Ariston, their clypeus height in anterior view is equal to the
AME diameter. Conifaber males lack first femoral macrosetae
present in Ariston and Siratoba males (Opell, 1979, figs. 39, 70) and,
like Ariston, lack abdominal tubercles.

Using Opell’s (1979) keys to uloborid genera, Conifaber males key
to couplet 10, which separates Octonoba and Purumitra, and
females key to couplet 10, which separates Octonoba and Uloborus.
Conifaber males are distinguished from those of Octonoba and
Purumitra by having first femora whose lengths are equal to rather
than 1.5 to 2.0 times as long as the carapace, by lacking femoral
macrosetae present in these genera (Opell, 1979; figs. 181, 183), and
by having a longer, more conspicuous tegular spur than these genera
(Fig. 14; Opell, 1979; plate 6-c, fig. 157). Conifaber females lack
dorsal abdominal tubercles present in Octonoba and Uloborus
(Opell, 1979; figs. 132, 140) and have inconspicuous, anteriorly
directed epigynal lobes (Figs. 24-26) instead of conspicuous poste-
riorly directed lateral epigynal lobes (Opell, 1979; figs. 137, 145, 178,
184).

Description. Maximum carapace width 0.84 carapace length,
attained in posterior half of female carapace and in posterior third
of male carapace (Figs. 21-22). Female carapace slopes up to a
point just behind PLE and then down to AME (Fig. 20). Male
carapace slopes more steeply up to a point slightly forward of its
center and then down to PME (Fig. 23). Shallow, transverse female
thoracic groove at carapace center; deep, U-shaped male thoracic
groove in posterior quarter of carapace. In both sexes anterior eye
row strongly recurved so that a line across AME’s posterior margins
passes in front of ALE’s by a distance equal to one ALE diameter.
Posterior eye row slightly recurved so that a line across PME’s
posterior margins passes along PLE’s anterior margins. Median
ocular area’s length and posterior width 0.8 its anterior width.
Female AME diameter 0.75 that of male AME, remaining eyes
equal to 0.66 female AME and 0.50 male AME. AME’s 1.3 as far
from one another as from ALE’s, PME’s 1 .7 as far from one another
as from PLE’s. Sternum 0.80 as wide as long, widest between first
and second coxae. Female endite 0.80 and male endite 1.00 as wide
as long. Labium 1.40 as wide as long. First femur equal in length to
carapace. Male first tibia with six or seven short and one long

1982]

Lubin, Opell, Eberhard, Levi — Uloboridae

51

Figures 14 and 15. Apical (14) and retrolateral (15) views of Conifaber parvus n.
sp. holotype male left palpus. The arrow in 15 shows the normal position of the
tegular spur (TS) embolus (E) guide as it rests in the grooved tegulum (T). MAB =
median apophysis bulb, MAS = median apophysis spur, MH = middle hemato-
docha. Scale lines are 100 yum long.

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[Vol. 89

dorsoprolateral macrosetae, two or three long proximodorsal
macrosetae, and two or three distoretrolateral macrosetae (Fig. 28).
Abdomen without tubercles or abrupt peak (Figs. 20-23). Female
abdomen 0.98 as wide and 1.38 as high as long, male abdomen 0.70
as wide and 0.93 as high as long. Distance between cribellum and
epigastric furrow 0.44 abdomen length. Abdomen and cephalo-
thorax were separated when the epigynum was removed. Examina-
tion of the severed petiole revealed no large tracheal trunks,
indicating that, as in Philoponella and Daramuliana (Opell, 1979
fig. 1), no tracheae extend into the cephalothorax or, as in Zosis,
Purumitra, and Octonoba (Opell, 1979; fig. 2), only fine tracheoles
extend into the cephalothorax.

Male Palpus. Femur without ventral tubercles. Like Zosis, Puru-
mitra and Octonoba (Opell, 1979; plates 6-c,d, 7-c,d, fig. 157),
Conifaber male palpi have a tegular spur which acts as an embolus
guide (Figs. 14-15). This tegular spur is proportionately larger than
those of other genera and rests in a tegular groove unique to
Conifaber. Members of Zosis also have a large, grooved tegular
spur, but the median apophysis bulb of Conifaber is a plate rather
than a hemisphere, and its median apophysis spur a grooved plate
rather than a hook. The tegular spur’s tip may rest in the median
apopysis spur’s distal groove.

Epigynum. Two posterior lateral epigynal lobes extend anteriorly
a short distance, concealing a pair of weakly sclerotized, anteriorly
directed oval areas (Figs. 24-25). In posterior view the epigynum’s
posterior plate is 0.6 as high as broad and has slightly curved and
rounded ventral rim about one third the height of the posterior
plate (Fig. 26). A highly coiled duct leads from each weakly
sclerotized oval to a spherical spermatheca whose short fertilization
duct appears to connect to the vagina’s ventrolateral margin (Fig.
27).

Distribution. This genus is known only from the type localities
in eastern central Colombia.

Conifaber parvus new species
Figures 14-15, 20-29

Types. Male holotype, male paratype, and female paratype
from Finca Mozambique, 15 km S.W. of Puerto Lopez in the

1982] Lubin, Opell, Eberhard, Levi — Uloboridae 53

Colombian department of Meta; collected 1978 by W. G. Eberhard,
in the Museum of Comparative Zoology. The specific epithet is a
Latin noun in apposition, referring to the small size of members of
this species.

Description. As most features of this species are presented in the
genus description, only those of color and size are given here. Total
length of female 1.92 mm, of males 1.50 mm. Female carapace 0.72
mm long, male carapace 0.66 mm long. Female sternum 0.44 mm
long, male sternum 0.38 mm long. Female AME diameter 60 jum,
male AME diameter 80 ^m, remaining eyes of both sexes 40 pm in
diameter. Female leg length (I— IV): 2.86, 1.78, 1.52, 2.42 mm. Male
leg length: 2.70, 1.56, 1.33, 1.94 mm. Female calamistrum composed
of 10 setae and 0.22 mm long, extending 0.52 the metatarsus length.
Female cribellum 180 wide, 60 ^m long. Female anterior
spinnerets 0.30 mm long, male 0.16 mm long. Female posterior
spinnerets 0.27 long, male 0.18 mm long. Female anal tubercle 0.14
mm long, male 0.10 mm long.

Except for dark circles around the eyes (Figs. 21-22) members of
neither sex have conspicuous color markings. The thoracic groove is
slightly darker than the rest of the carapace, and white guanine
deposits under the abdomen’s integument are interrupted by the
cardiac area which creates a tan median stripe (Figs. 21-22).
Lacking these deposits, the anterior third of the female’s abdomen is
also tan rather than white.

Distribution. Known only from the type locality in eastern
central Colombia.

Uloborus conus new species*

Figures 16-19, 30-35

Types: All types from Papua New Guinea. Female holotype and

paratype from Madang Prov., 40 km south of Madang, collected 21
March 1979 by H. W. Levi and Y. D. Lubin. Two male and three
female paratypes from Morobe Prov., Buso Forest Reserve, col-
lected 25 Oct. 1979 by Y. D. Lubin. Four female paratypes from

*For nomenclatural purposes B. D. Opell is the author of this species.

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Figures 16-19. Uloborus conus n. sp. 16. Retrolateral view of holotype male left
palpus (a trilobed piece of debris is lodged at the upper right). 17. Retrolateral view
of MAS. 18. Apical view of MAS. 19. Ventral view of female paratype epigy-
num. C = conductor; other abbreviations as in Figures 14 and 15. Scale lines are 100
jum long.

1982]

Lubin, Opell, Eberhard, Levi — Uloboridae

55

Central Prov., along Brown River, near Port Moresby, collected 29
April 1980 by Y. D. Lubin. One male and one female paratype
deposited in the American Museum of Natural History, the remain-
ing types are deposited in the Museum of Comparative Zoology.
The specific epithet is a Latin noun in apposition, referring to the
conical web produced by members of this species.

Diagnosis. Males and females are distinguished by a carapace

length of less than 1.00 and 1.30 mm, respectively. Males have a long,
lobed palpal femoral tubercle, a reduced, flattened median apophy-
sis, a long, broad conductor, and a blunt median apophysis spur
(Figs. 16-18). Length of female femur I less than 1.2 carapace length
rather than 1.4- 1.5 carapace length as in other uloborids. Central
region of epigynum from which lobes arise about one third rather
than half as wide as the posterior plate (Figs. 19, 34).

Description. Female. Total length 2.80-3.40 mm (X = 3.20),
carapace length 1 .00-1.30 mm (X = 1 .09), maximum carapace width
0.90-1.00 (X = 0.96), carapace width at PLE’s 0.58-0.64 mm (X =
0.60), area. All eyes except AME’s surrounded by small black circles
(Fig. 30). PLE nearer midline than in other Uloborus species.
Sternum tan. Leg I of most specimens as shown in Fig. 33, but
nearly black in two dark specimens. Dorsum of femur I of all
specimens black. Abdomen of most specimens light tan or white.
Abdomen of two dark specimens with white dorsum, black venter
and two broad, white lateral stripes extending from anterior apex to
posterior tips. Epigynum consists of two small, weakly sclerotized
posterior lobes (Fig. 19) whose combined basal width is one-third
that of the posterior plate (Fig. 34). An epigynal opening found
dorsal to each lobe leads to a large, irregular spermatheca from
whose posterior lateral margin a short fertilization duct extends
(Fig. 35).

Male. Total length 2.00-2.20 mm, carapace length 1.00 mm,
maximum carapace length 0.85 mm, carapace width at PLE’s 0.66
mm, sternum length 0.56 mm. Carapace and sternum coloration
similar to that of female except that broad gray streaks extend
anteriorly from the posterior eyes (Fig. 31). Legs light tan, tibiae
II-IV with light gray dorsal tip. Femur I with three prolateral, one
dorsal, central; and one distal, retrolateral macrosetae (Fig. 32).

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Figures 20-27. Conifaber parvus n. sp. 20. Lateral view of female. 21. Dorsal
view of female. 22. Dorsal view of male. 23. Lateral view of male. 24. Anterior
view of epigynum. 25. Ventral view of epigynum. 26. Posterior view of epigy-
num. C = conductor. D = other abbreviations as in Figures 14 and 15. Scale lines are
100 jum long.

1982] Lubin, Ope 1 1, Eberhard, Levi — Uloboridae 57

Tibia I with eight prolateral, seven dorsal, and three retrolateral
macrosetae. Sternum and abdominal venter with orange setae.
Abdomen gray with a pair of thin, white, lateral longitudinal stripes
running nearly its full length. Palpal femur with a large, lobed
retrolateral tubercle and a very small prolateral tubercle (Fig. 16).
Median apophysis bulb small and flattened (Fig. 16); median
apophysis rectangular with a blunt apex (Figs. 17-18). Conductor
long and broad, extending from median apophysis spur to area of
palp adjacent to patella.

Distribution. Known only from the type localities in Papua New
Guinea.

Uloborus albolineatus new species*

Figures 36-39.

Type. Female holotype from Lowlands Agricultural Experimental
Station, Kerevat, East New Britain, Papua New Guinea, collected 6
July 1980 by Y. D. Lubin, deposited in the Museum of Comparative
Zoology. The specific epithet is a noun in apposition, referring to
the species’ white median abdominal stripe.

Diagnosis. Males are unknown. The female is distinguished by
having reddish brown median eyes, a very convex sternum (Fig. 37),
white guanine deposits in the cardiac region (Fig. 36), and weakly
sclerotized epigynal lobes rising from the center rather than poste-
rior of a transparent epigynum (Fig. 38). Unlike many Uloborus
species, the carapace lacks a conspicuous median light stripe.

Description. Female. Total length 2.40 mm, carapace length 0.92
mm. maximum carapace width 0.74 mm, carapace width at PLE’s
0.50 mm. Carapace tan with gray, reticulate lateral markings (Fig.
36). Median eyes reddish brown. AME’s on a more conspicuous
tubercle than most Uloborus species. Sternum tan, widest at coxae I
rather than between coxae I and II as in other Uloborus species.
Legs light tan with faint gray distal rings on most segments. Tibia I
with very sparse distal setal brush. Abdomen height and width 0.9
its length, dorsum with a pair of centrolateral tubercles, posterior

For nomenclatural purposes, B. D. Opell is author of this species.

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1982]

Lubin, Opell, Eberhard, Levi — Uloboridae

59

tip projecting slightly posterior to anal tubercle’s base and separated
from anal tubercle by a distance one third the abdomen’s height.
White guanine deposits extend both in a narrow transverse band
across the abdomen’s anterior ventral surface and along the cardiac
area. A broader, more diffuse median guanine deposit extends from
the abdomen’s humps to its posterior tip. A pair of large guanine
spots is found anteriolaterally to the spinnerets. Epigynum convex
with broad posterior extension, a pair of low, weakly sclerotized
median lobes, and a transparent integument through which a single
pair of spherical spermathecae is clearly visible (Figs. 38, 39).

Distribution. Known only from the type locality in Papua New
Guinea.

Uloborus bispiralis new species*

Figures 40-48.

Types: Female holotype, three male and seven female paratypes

from Fowlands Agricultural Experimental Station at Kerevat, East
New Britian Prov., collected 2, 4, and 6 July 1980 by Y. D. Fubin.
Male and two female paratypes deposited in the American Museum
of Natural History, remaining types in the Museum of Comparative
Zoology. The specific epithet is a latin noun in apposition, referring
to the male’s doubly coiled embolus.

Diagnosis. Females are distinguished by having a single, narrow
median, posterior epigynal lobe (Figs. 42, 43) rather than a pair of
posterior epigynal lobes, and by each epigynal duct making five
rather than the usual single loop (Fig. 44). Males are distinguished
by an embolus which loops twice rather than once around the

*For nomenclatural purposes B. D. Opell is the author of this species.

Figures 28 and 29. Conifaber parvus n. sp. 28. Dorsal view of male left first
tibia. 29. Retrolateral view of expanded male left palpus (R = radix, BH = basal
hematodocha, other abbreviations as in Figs. 14 and 15).

Figures 30-35. Uloborus conus n. sp. 30. Dorsal view of female carapace.
31. Dorsal view of male carapace. 32. Prolateral view of male first femur, patella,
and tibia. 33. Retrolateral view of female leg I. 34. Posterior view of epigynum.
35. Dorsal view of cleared epigynum.

1982]

Lubin, Opell, Eberhard, Levi — Uloboridae

61

median apophysis and by a flattened, elongate median apophysis
bulb which bears a broad conductor (Figs. 45, 46). Both males and
females have a gray lateral abdominal stripe (Fig. 48).

Description. Female. Total length 3.28-3.68 mm (X = 3.47, S,
0.14, N = 8), carapace length 1.10-1.20 mm(X= 1.15, SD = 0.04),
maximum carapace width 0.94-1.04 (X = 0.98, SD 0.04), carapace
width at PME’s 0.54-0.58 mm (X = 0.56, SD 0.01). All eyes except
AME’s surrounded by small black circles (Fig. 41). Carapace with
light lateral margins, light posterior median stripe, and central gray
patch. Sternum tan. First and second legs light gray with tan
proximal ring on tibia, metatarsus, and tarsus. Tibia I without a
conspicuous setal brush. Third and fourth legs tan with gray distal
rings on tibia, metatarsus, and tarsus. Abdomen without humps,
dorsal and lateral surfaces densely covered by white guanine spots
except in cardiac region and along a faint lateral stripe similar to but
not as sharply defined as that shown in Fig. 48. Venter tan with only
sparse guanine spots. Epigynum a raised mound with single median
lobe (Figs. 42, 43), probably representing a pair of fused lateral
lobes. Under normal light microscopy a clove oil-cleared epigynum
showed only a pair of oval spermathecae with a fertilization duct
leading from the posterior lateral margin of each and a short, broad
duct extending from the median surface of each to epigynum’s
posterior margin. Examination with Nomarski optics revealed the
more extensive system of thin-walled ducts shown in Figure 44. It
was not possible to determine precisely where the ducts opened
externally, but this appears to be between the spermathecae and
near the base of the epigynal lobe.

Male. Total length 2.32-2.40 mm, carapace length 0.98-1.00 mm,
maximum carapace width 0.78-0.80 mm, carapace width at PLE’s
0.50-0.52 mm. Carapace and sternum coloration similar to that of

Figures 36-39. Uloborus albolineatus n. sp. 36. Dorsal view of female holotype.
37. Lateral view of female carapace. 38. Ventral view of epigynum. 39. Dorsal
view of cleared epigynum.

Figures 40-48. Uloborus bispiralis n. sp. 40. Male carapace. 41. Female holo-
type carapace. 42. Ventral view of holotype epigynum. 43. Posterior view of
epigynum. 44. Dorsal view of cleared epigynum. 45. Retrolateral view of male
palpus. 46. Apical view of male palpus. 47. Prolateral view of male first femur,
patella, and tibia. 48. Lateral view of male abdomen.

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females except for absence of central gray carapace spot (Fig. 40).
Legs reddish brown. Femur I with three or four prolateral macro-
setae, tibia I with nine prolateral, six or seven dorsal, and three
retrolateral macrosetae (Fig. 47). Abdomen with fewer guanine
spots than female, dorsum and lateral surface tan; gray lateral
stripe, gray venter and gray posterior tip (Fig. 48). Palpal femur
with a large proximal retrolateral tubercle and small prolateral
tubercle. Median apophysis bulb flat and elongate (0.16 mm long),
terminating in a bent median apophysis spur (Figs. 46, 47). Unlike
other members of the genus, the embolus loops twice around the
median apophysis bulb before passing into a broad, weakly scle-
rotized conductor.

Distribution. Known only from the type locality in Papua New
Guinea.

Acknowledgements

YDL and HWL thank the Wau Ecology Institute for use of the
facilities in Wau, and S. Smith for making available the facilities at
LAES (Kerevat). A portion of this study was supported by a
Smithsonian Institute Scholarly Studies Research Award (to M. H.
Robinson and YDL). A Small Projects Grant from the College of
Arts and Sciences, Virginia Polytechnic Inst, and State Univ. (to
BDO) made the S.E.M. work possible. WGE thanks Carlos Rod-
rigues, the Dixon Stroud family. Dr. Luis Arango, Dr. Madhav
Gadgil, and A. J. T. Johnsingh for help and hospitality in the field
and the Comite de Investigaciones of the Universidad del Valle,
Cali, Colombia and the Vicerectoria de Investigaciones of the
Universidad de Costa Rica for financial support. HWL thanks
National Science Foundation grant DEB 76-15568 and DEB
79-23004 for support and M. H. Robinson and B. Robinson for
being instrumental in getting him to New Guinea and flavoring his
stay with their hospitality and enthusiasm. We thank Frances
Murphy, V. Todd Davies, Joseph Beatty and James Berry for
allowing us to use their unpublished observations and M. H.
Robinson and B. Robinson for their comments on and criticism of
the manuscript.

1982]

Lubin, Opell, Eberhard, Levi — Uloboridae

63

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Carico, J. E.

1977. A simple device for coating orb webs for field photography. Bull. Br.
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Eberhard, W. G.

1969. The spider Uloborus diversus Marx and its web. PhD. thesis, Harvard
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1972. The web of Uloborus diversus (Araneae: Uloboridae). J. Zool., Lond.
166:417-465.

1977a. Photography of orb webs in the field. Bull. Br. arachnol. Soc.
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1977b. The webs of newly emerged Uloborus diversus and of a male Uloborus
sp. (Araneae: Uloboridae). J. Arachnol. 4:201-206.

1981. Construction behaviour and the distribution of tensions in orb webs.
Bull. Br. arachnol. Soc. 5(5): 189-204.

Lahmann, E. and W. G. Eberhard

1979. Factores selectivos que afectan la tendencia a agruparse en la arana
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LeGuelte, L.

1966. Structure de la toile de Zygiella x-notata C. (Araignees, Argiopidae) et
facteurs que regissent le comportement de l’araignee pendant la con-
struction de la toile. These. Pub. Univ. Nancy: 1-77.

Lubin, Y. D., W. G. Eberhard, and G. G. Montgomery

1978. Webs of Miagrammopes (Araneae: Uloboridae) in the Neotropics.
Psyche 85(1): 1-23.

Marples, M. and B. J. Marples

1937. Notes on the spiders Hyptiotes paradoxus and Cyclosa conica. Proc.
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Marples, B. J.

1962. Notes on the spiders of the family Uloboridae. Ann. Zool. Agra 4:1-11.
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1979. Revision of the genera and tropical American species of the spider family
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Peters, H.

1953. Beitrage zur vergleichenden Ethologie und Okologie tropischen Web-
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Robinson, M. H. and B. Robinson

1973. Ecology and behavior of the giant wood spider Nephila maculata
(Fabricius) in New Guinea. Smithson. Contrib. Zool. 149:1-76.

Szlep, R.

1961. Developmental changes in web-spinning instinct of Uloboridae: con-
struction of the primary-type web. Behaviour 27.60-70.

Wiehle, H.

1927. Beitrage zur Kenntnis des Radnetzbaues der Epeiriden, Tetragnathiden,
und Uloboriden. Z. Morph. Okol. Tiere 9:468 537.

1931. Neue Beitrage zur Kenntnis des Fanggewebes der Spinnen aus den
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Workman, T.

1896. Malaysian Spiders, volume F Published by the author. Belfast.

POPULATION STRUCTURE AND SOCIAL ORGANIZATION
IN THE PRIMITIVE ANT AMBLYOPONE PALLIPES
(HYMENOPTERA: FORMICIDA E)

By James F. A. Traniello1
Harvard University,

Museum of Comparative Zoology Laboratories
Cambridge, Massachusetts 02138, U.S.A.

I INTRODUCTION

The genus Amblyopone contains the most morphologically and
behaviorally primitive species in the poneroid complex of ants, and
a detailed examination of their social structure could significantly
contribute to the reconstruction of social evolution in the Formi-
cidae. But because of their cryptic habits and distribution, the biol-
ogy of the majority of species of Amblyopone and the related genera
Mystrium, Myopopone, Prionopella, and Onychomyrmex remains
almost entirely unknown. Previous investigations have provided
information on colony foundation (Haskins, 1928; Haskins and
Enzmann, 1938; Haskins and Haskins, 1951), ecology, behavior,
and taxonomy (Wheeler, 1900; Brown, 1960; Gotwald and Levieux,
1972; Baroni Urbani, 1978), and physiology (Whelden, 1958). Still,
many of the details of social organization in Amblyopone are lack-
ing. I present in this paper the results of a two-year study on the
behavior and ecology of Amblyopone pallipes.

Material and Methods
Study areas and nest collection

Thirty-one colonies of A . pallipes were collected under stones in a
damp, white pine woodland in Westford, Massachusetts. A single
colony was taken under the bark of a rotting log. Nests generally
consisted of one or two shallow (6-10 mm) depressions in the soil
immediately beneath the stone, from which a single gallery opened
to subterranean chambers. Gentle excavation usually revealed one
or two additional loosely structured chambers. Workers, queens,

•Present address: Department of Biology, Boston University, Boston,

Massachusetts 02215

Manuscript received by the editor February 18, 1982.

65

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sexuals, and brood were found at all levels of the nest and were
quickly aspirated.

Distribution and natural history

A. pallipes has been found in the eastern United States and in the
St. Lawrence Valley in Canada in cool, moist, forested areas
(Brown, 1960). General references on the natural history of this
species are given by Wheeler (1900) and Haskins (1928).

Laboratory arrangements

Colonies were housed in artificial nests composed of a thick,
moist filter paper bottom with cotton sides approximately 6 mm
high covered with a glass plate. The nests were placed in 15 X 22cm
plastic boxes in which the humidity was kept high. The total nest
area was roughly 10cm2. A second chamber, similar in structure but
somewhat larger, was connected to the nest as a foraging arena,
where live prey were offered. Colonies were fed on whole, live geo-
philomorph and lithobiid centipedes; in addition, elatyrid, bupres-
tid, and tenebrionid beetle larvae were acceptable to the ants. This
method of culture proved successful and greatly facilitated studies
of social interaction since the activity of an entire colony could be
monitored on the stage of a dissecting microscope. Ethogram data
were compiled in this manner, and were analyzed using the methods
of Fagen and Goldman (1977).

Results and Discussion

1. Life cycle and population structure.

Nest distribution and colony size. The spatial distribution of col-
onies at the principal study site in Westford is presented in Fig. 1.
An interesting feature of this population, in addition to its clumped
distribution pattern is that three colonies were collected under
stones in 1978 precisely where colonies were found the year before.
This suggests that the colonies that were collected represented sub-
units of a large, subterranean population. Each unit is small (modal
size class = 9-16 workers). Complete collection data are presented in
Fig. 2. Although distributional data are not given, a population of
seemingly comparable density was discovered by Wheeler (1900),
who uncovered 30 nests in a three hour period. Also, the colony size
data correspond closely to the data of Francoeur (1965, 1979, and
personal communication).

1982]

Traniello — Amblyopone pallipes

67

Queen number. The frequency distribution of the number of
queens in a colony is given in Fig. 3. Of 19 queenright colonies, 10
(52%) contained more than one dealate female. Observations of
multiple queened colonies in the laboratory revealed that in at least
some of these colonies each queen was functionally reproductive.
However, many queens in apparently polygynous colonies did not
lay eggs, and engaged primarily in worker tasks.

Life cycle, colony reproduction, and population structure. Be-
cause colonies were collected and censused throughout the spring
and summer of 1977 and 1978, it is possible to outline the life cycle
of A. pallipes (Fig. 4). Eggs are laid in late April or early May and
larvae hatch and develop throughout June and July. Mature larvae
pupate in mid-July and early August, and adults eclose approxi-
mately two to three weeks later. Although small numbers of eggs
and larvae are present in most colonies throughout the spring and
summer, it appears that only one brood matures per year. The large
number of eggs found in colonies collected in August hatch before
September and overwinter as larvae (Talbot, 1957). It is possible
that the winter chilling results in the determination of these larvae as
sexuals. In late August and early September workers and sexuals
simultaneously eclose unassisted from their pupal cases. The adults
which eclose at this time are predominantly workers. In four colo-

68

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[Vol. 89

15 -«

CO

LU

O

CD

CD

CD

10 -

t — i — i — i — r

1 2 9 8 16 32

NO, WORKERS PER COLONY

Fig. 2. — Frequency distribution of colony sizes for 35 nests.

nies collected in late August which reared brood in the laboratory,
the ratio of <5:$: $ ? was as follows: 2:1:36; 0:5:13; 3:0:7; and 0:4:19.
In all cases the worker population of a colony was at a maximum at
this time. If this is considered in conjunction with the available
information on nuptial flights in A. pallipes, then it is possible to
speculate on colony reproduction and population structure.

1982]

Traniello — Amblyopone pallipes

69

NO. OF QUEENS PER COLONY

Fig. 3. — Frequency distribution of the number of dealate females in 19 queenright
colonies.

Although the complete sequence of colony reproduction has not
been observed, the studies of Haskins (1928, 1979) and Haskins and
Enzmann (1938) provide some evidence of its organization. Early in
September, winged females leave the nest and disperse over short
distances, finally alighting on the ground or low vegetation. Then,
with gaster arched and sting partially extruded, they “call” males
with a chemical sexual attractant. Males quickly locate females,
copulation ensues, and soon after insemination females shed their
wings and re-enter the soil; perhaps they return to the parent nest.

70

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[Vol. 89

At this point in the life cycle, the worker/ queen ratio is highest, as
described above, yet colonies collected in the late spring are much
smaller in size (approximately 50%). Therefore, colony reproduc-
tion by budding may occur if one or more fecundated queens depart
with a portion of the worker force. This hypothesis has previously
been considered by Wheeler (1900) and Brown (1960), and is sup-
ported by my data on colony growth, nest distribution, and nest
structure. An additional feature of the nest distribution pattern sup-
ports the hypothesis of limited dispersal. The most dense population
of colonies occurred on the south side of an early stone wall (< 1 m
high), although nest sites were abundant on both the north and
south sides, and soil, vegetational, and exposure parameters ap-
peared to be identical. Also, laboratory observations indicate that
alate females may shed their wings before mating occurs. On several
occasions newly eclosed females left the nest, shed their wings, and
returned to the nest. Because mating occurs on the ground, such
behavior does not exclude the possibility that these individuals
could eventually become inseminated. These females may then
return to the parent nest or may be adopted by a nearby colony. In
several laboratory experiments queens were introduced into other
queenright nests or orphaned colonies. In all cases they were
accepted by both workers and queens. Similarly, workers could be
transferred from one colony to another without aggression. There-
fore, populations of A. pallipes appear to be unicolonial and secon-
darily polygynous. Ecologically, the patchy distribution of this ant
correlates with this type of population structure.

2. Social organization

The social ethogram. Social ethogram data were gathered from
five colonies which were observed for a total of 73 hours, during
which 6,500 individual acts were recorded. The behavioral catalog
of a single colony of A. pallipes (2 queens, 18 workers, brood) that
was studied for 25.7 hours is given in Table I. The total number of
acts observed was 42 (95% confidence interval for catalog size [27,
47]), and the sample coverage was 0.9992. Behaviors listed in the
ethogram having a frequency of 0 were observed in other colonies
and are included as part of the species repertory. The majority of
activities are common to many ant species; those that are unusual
will be discussed briefly. Antennal tipping is a behavior previously
described in Zacryptocerus varians (Wilson, 1975) which occurred

WORKERS

1982]

Traniello — Amblyopone pallipes 71

CD

aooaa do inhd N3d

72

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[Vol. 89

Table I. — The social ethogram. N = number of acts observed in each caste.

Behavioral Act

Workers (16)
N=2525

Queens (2)
N=158

1 . Self-groom

0.3303

0.4114

2. Allogroom queen

0.0044

0

3. Allogroom worker

0.0384

0

Brood care:

4. Lay egg

0

0.0127

5. Carry egg or egg pile

0.0123

0.0696

6. Lick egg

0.0305

0.1013

7. Lick larva

0.0950

0.1139

8. Carry, drag, or role larva

0.0337

0.0633

9. Bank mature larva with soil

0.0048

0

10. Carry pupa

0.1200

0

1 1. Lick pupa

0.0824

0.0127

12. Place larvae on prey

0.0032

0

13. Assist removal of meconium

0.0004

0

14. Assist larval molt

0.0012

0

15. Lick ecdysial skin

0.0063

0.0380

Aggressive display:

16. Undirected

0.0250

0

17. To worker

0.0051

0.0127

18. To queen

0.0004

0

Predatory behavior:

19. Forage

0.0158

0

20. Sting prey

0.0040

0

21. Drag prey to nest

0.0019

0

22. Drag prey within nest

0.0048

0

23. Lick prey

0.0578

0

24. Handle prey within nest

0.0051

0

Nest maintenance and defense:

25. Guard

0.0083

0

26. Handle nest material

0.0190

0

27. Repair nest wall

0.0277

0

28. Lick nest material

0.0012

0

29. Excavate nest

0.0111

0

30. Bury noxious object

0.0004

0

3 1 . Carry or drag dead worker

0

0

32. Carry or drag live worker

0.0004

0

33. Extrude sting

0.008

0.0190

34. Remove empty pupal case

0.0008

0

35. Jitter

0.0055

0.0190

36. Jolt body

0.0135

0.0633

37. Lick meconium

0.0008

0

38. Tip antennae

0.0008

0

39. Flick antennae

0.0099

0.0127

1982]

Traniello — Amblyopone pallipes

73

Behavioral Act

Workers (16)
N=2525

Queens (2)
N=158

40. Pinch larvae

0

0.0063

41. Cannibalize larva

0.0170

0.0380

42. Discharge subpharyngeal
pellet

0

0.0063

Totals:

1.0

1.0

infrequently in A. pallipes. During this behavior the body is raised,
the gaster is curved forward, and with the mandibles agape the
antennae are held forward with their terminal funicular segments
slightly inclined toward each other. The significance of antennal
tipping is unknown, but it appeared to be part of a grooming
sequence. Vibrational displays were given by workers if the nest wall
was breached or if an individual was mechanically disturbed. If the
stimulus was intense enough, other workers would show the same
vigorous jittering behavior, consisting of rapid vertical movements
of the head and thorax. This behavior had the effect of producing a
general arousal within the colony and resulted in an increase in the
number of workers appearing at the source of stimulation. In the
case of nest damage, building behavior eventually occurred but did
not immediately follow. This signal appears to be a primitive form
of mechanical communication, in which alarm is propagated di-
rectly through body contact. A similar vibratory display has been
documented in A. australis (Holldobler, 1977).

Workers and queens of A. pallipes have retained a number of
behaviors that appear to reflect their wasp ancestry. Queens were
seen grasping larvae and squeezing them in the neck region with
their mandibles, thus causing them to regurgitate a droplet of clear
liquid which they then consumed. Workers were never observed to
regurgitate with other workers, queens, or larvae, and all individuals
fed directly on prey. Aggression was observed between workers and
queens. An aggressive display typically consisted of opening the
mandibles and rising up on the extended legs. This behavior was
usually exhibited by queens in the area of the egg pile and seemed to
produce avoidance in contacted workers. These observations raise
the question of whether queens maintain their reproductive status
through behavioral dominance or inhibitory pheromones.

Polyethism. Studies on the division of labor within the worker
caste have revealed that temporal castes are absent in the species. A

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[Vol. 89

complete account of polyethism in relation to the life history of A.
pallipes is given by Traniello (1978).

Predatory behavior. Prey were found in only three of the colonies
collected. In two colonies larvae were found clustered around litho-
biid centipedes (length = 1.5-2. Ocm), and in the third colony a
carabid beetle larva was taken. In the laboratory, colonies were
offered a variety of live arthropods that workers might encounter in
leaf litter, soil, or rotting wood. Wood lice ( Oniscus ), house centi-
pedes ( Scutigera ), and various millipedes were consistently rejected
while small elatyrid, tenebrionid, and buprestid beetle larvae were
carried to the nest and fed upon. The diet of A. pallipes appears to
be restricted to live, linear-shaped arthropods that can be captured
by workers. A related species, A. pluto, is entirely specialized on
geophilomorph centipedes (Gotwald and Levieux, 1972). When
large, robust-bodied centipedes ( Lithobius sp.) were offered to col-
onies of A. pallipes, workers were unable to grasp the prey due to
its escape movements and body diameter. It is difficult to imagine a
condition under which large prey could be captured, even if they
were “cornered” in a narrow gallery. When Lithobius of similar size
were held with forceps, workers were still unable to subdue the centi-
pede. Freshly killed centipedes were not accepted. It is therefore
difficult to support the hypothesis of a nomadic life style for A.
pallipes. Although this species of Amblyoponini does not appear to
move its colonies to the location of large, previously captured prey,
other species, such as Onychomyrmex do provide evidence linking
group predation and nomadism in this primitive group of ants
(Wilson, 1958).

Prey capture and retrieval is very stereotyped, and solitary hun-
tresses stalk prey in a highly methodical manner. As prey are
approached, workers advance cautiously, apparently orienting to
odors or air microcurrents produced by the prey. When within strik-
ing range (2-3 mm) the mandibles are opened and the head is
oriented orthogonal to the long axis of the prey. Then in a single
motion the mandibles close around the prey, the legs elevate the
body, and the gaster is swung forward. The prey is then repeatedly
stung and the venom soon shows its paralytic effects. Initially, only
the area adjacent to the cuticle penetrated by the sting is immobi-
lized, and stinging continues until escape movements stop. Subse-
quently, the retrieval of the prey begins after a brief period of self-

1982]

Traniello — Amblyopone pallipes

75

grooming. The retrieval process varies in duration depending upon
prey size, but even long (4. 0-5. 0cm, 2. 0-2. 5 mm in diameter) geo-
philomorph centipedes are easily dragged to the nest. A number of
orientation trips made between the prey and the nest generally pre-
ceded retrieval. During these orientation runs, which were made
throughout the retrieval process, workers continually checked their
position relative to the nest. The prey was then dragged several
centimeters; the worker then stopped, released the prey, and con-
tinued homeward until she contacted the nest entrance. She then
returned to the prey and repeated this sequence, alternating prey
movement with orientation trips. Once the prey was in the nest,
other workers approached and began vigorously licking the areas of
the prey’s body opened during capture. Larvae were either carried to
the prey or, if close enough, moved toward it and adjusted their
position on its body on their own accord. At times workers assisted
in positioning larvae. Additional details of feeding behavior are
nearly identical to those described by Gotwald and Levieux (1972).

Communication during foraging. At times, two or three ants
attempted to jointly carry prey, but cooperative efforts were hap-
hazard and inefficient. But cooperative retrieval seems unnecessary
due to the physical capabilities of individual ants. The critical ele-
ment in prey capture is probably not retrieving, but subduing rela-
tively large arthropods. Often after a worker began stinging a prey
item, a second or third worker approached and assisted in para-
lyzing the prey. The fact that workers were attracted to the point of
prey capture suggested that additional ants may be recruited over
short distances by orienting to prey odor, air currents, or some
signal produced by the forager. To test the hypothesis that phero-
mones are involved in this process I stimulated foragers to grasp and
attempt to sting the tip of a pair of forceps and then lowered the
worker, still attacking the forceps tip, in front of the nest entrance.
The response of workers in the nest was dramatic. In five replicates,
5.8 ± 2.3 workers/0.5 min approached the nest entrance under the
experimental conditions. Only 0.2 ± 0.1 workers/0.5 min were
attracted to the nest entrance in controls (agitated forceps alone).
This difference is statistically significant (.001 < p < .01; t = 6.1,
Student’s t-test). Although the possibility that stridulatory signals
might be involved could not be ruled out, the results of these experi-
ments suggest that chemical cues are involved in the attraction

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[Vol. 89

response. Subsequently, crushes of the head, thorax, and gaster
were offered on applicator stick tips at the nest entrance. Also,
crushes of dissected poison and pygidial glands (Holldobler and
Engel, 1979) were offered. Only head crushes produced attraction.
Whelden’s (1958) studies, in addition to our own histological investi-
gations, revealed a group of large glandular cells at the base of the
mandible. The indirect evidence described above suggests that dur-
ing prey capture the contents of these cells are released, attracting
workers in the vicinity to assist in subduing prey.

3. Ecology and social evolution

The results of this study and previous investigations suggest that
populations of A. pallipes are unicolonial. Workers from different
subnests within a population show no aggression toward each other.
Such worker compatibility has been demonstrated in Rhytidopon-
era metallica (Haskins and Haskins, 1979), whose populations
appear to be structurally similar to those of A. pallipes, but occupy
larger area geographically. Workers taken from nests three miles
apart were not mutually hostile. The lack of aggression was consist-
ent within, but not between populations. Ambylopone pallipes col-
onies appear to be similarly viscous, but do not occupy as extensive
an area.

Observations in the laboratory are in accord with Brown’s (1960)
position which states that after mating, females “always or usually
return to the parent nest”. Secondary polygyny in this species, in
addition to its patchy distribution, indicates that this species is in the
terminology of Holldobler and Wilson (1977) a habitat specialist.
The characteristic A. pallipes habitat is cool, damp, heavily shaded
woodland. Nest site and prey abundance are also important fea-
tures. Populations apparently grow slowly, and through reproduc-
tion by budding, eventually saturate the habitat. Such a scheme
does not rule out the occurrence of dispersal flights, which have
been witnessed on occasion (Haskins, 1928). As colonies become
more populous within a habitat, dispersal flights should occur more
frequently in order to colonize additional areas. Once a founding
queen locates a preferred habitat, colony reproduction again is
accomplished through budding. The strategy may be similar to that
of the mound building species Formica exsectoides. However, it
must be noted that in laboratory situations, A. pallipes queens have
never been observed to successfully found colonies (Haskins, per-

1982]

Traniello — Amblyopone pallipes

11

sonal communication). But it is difficult to determine whether this is
an abnormality which occurs only in the laboratory or represents an
inability of A. pallipes queens to found a colony alone. Newly
inseminated queens of A. australis found colonies in the partially
claustral mode (Haskins and Haskins, 1951). However, A. australis
is monogynous.

Within a habitat, A. pallipes escapes competition with the more
advanced groups of ants by additional specializations on micro-
habitat and diet. This is in contrast to other unicolonial species
which are broad generalists.

Behaviorally, A. pallipes exhibits both primitive and advanced
social traits, and many of the primitive characters are more conserva-
tive than those of Myrmecia. Age polyethism is lacking, and commu-
nication between individuals is primarily mechanical, although a
rudimentary short-range recruitment system that is mediated by
pheremones exists. Among the primitive trophic characteristics is
the use of the sting to paralyze prey, which are subsequently fed
directly to the larvae without prior dismemberment. Employing the
sting to paralyze prey appears to be widespread in the Ponerinae,
and recently Maschwitz et al. (1979) have demonstrated that the
venom of the oriental ponerine species Harpegnathus saltatus and
Leptogenys chinensis indeed has paralytic, and not toxic, effects.
Prey paralyzation also occurs in Daceton armigerum and Paltothy-
reus tarsatus (Wilson, 1962; Holldobler, pers. comm.). This is con-
trasted to myrmicine species which use the sting as a defensive
weapon. The importance of paralyzing but not killing arthropod
prey in Amblyopone pallipes is obviously related to the direct provi-
sioning of larvae; prey must be kept from decomposing until they
are consumed. Also, immobilization is necessary for successful
retrieval, and energetically it is more efficient for solitary foragers to
carry paralyzed prey. The absence of regurgitation which is charac-
teristic of the Ponerinae, also is a primitive trait. Although one of
the more distinctive features of A. pallipes and other Amblyoponini,
prey specialization, appears to be a conservative formicid trait, it is
also possible that specialization was a response to competition.

Finally, based on the theories of Malyshev (1968), Wilson (1971)
has speculated that the Amblyoponini may have approached euso-
ciality in a way very different from the partially claustral colony
founding route assumed by Haskins and Haskins (1951). Because

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[Vol. 89

these ants appear to be specialized on large arthropods, they may
have passed through a phase of subsociality similar to that of the
bethylid wasp Scleroderma. Although the evolution of ants from
Scleroderma- like ancestors has been ruled out on morphological
grounds, the possibility remains that the Amblyoponini represent an
independent venture into eusociality. The present study, which sug-
gests that A. pallipes is not dependent upon large arthropod prey,
and the studies of Haskins (1928) and Haskins and Haskins (1951)
on colony foundation, do not support or refute this theory. Addi-
tional studies must be carried out on the behavior of newly insemi-
nated females, their prey preferences during colony foundation and
their reproductive physiology to test this hypothesis.

Acknowledgements

This research was carried out while the author was a doctoral
candidate at Harvard University, and was supported by the Ander-
son and Richmond Funds, NSF Grant BNS 80-02613 (B. Holl-
dobler, sponsor), and NSF predoctoral grant DEB 78-16201. Drs.
Gary Alpert, Bert Holldobler, and Edward Wilson provided useful
comments on the manuscript. I would especially like to thank Dr.
Caryl Haskins for sharing with me his great wealth of knowledge of
primitive ants. Finally, I thank Michelle and Eric Scott, who were
indispensable in the field.

Summary

1. The behavior of ecology of the primitive ponerine ant Amblyo-
pone pallipes was studied in the laboratory and the field. Thirty-
three colonies were collected over a two-year period, 94% of which
were excavated from one locality where 68% of the colonies were
strongly clumped in their spatial distribution. Workers and queens
could be transferred between these nests without hostility.

2. The inability of workers to recognize members of other colo-
nies within a population, the colony life cycle, limited dispersal, the
presence of multiple queens in nests, and circumstantial evidence on
the adoption of newly inseminated females by their parent nest
suggest that A. pallipes is secondarily polygynous and unicolonial.
Although dispersal flights do occur, colony reproduction seems to
be accomplished through budding.

1982]

Traniello — Amblyopone pallipes

79

3. Studies on the ethology of A. pallipes show that this species has
retained many conservative behavioral traits. Among these are the
absence of age polyethism and the provisioning of larvae with whole
prey (chiefly chilopods and beetle larvae). Observations of preda-
tory behavior do not support the hypothesis that colonies are
nomadic. Prey are paralyzed by stinging and are then retrieved.
Larvae feed directly on the body of the prey.

4. A primitive form of alarm communication, presumably trans-
mitted through body contact, is mediated by a vibratory display.
Workers show attraction to head crushes, and mandibular gland
pheromones appear to be involved in a weak form of recruitment.

5. Because of the lack of precise information on the behavior of
colony founding queens, the question of whether sociality in the
Amblyoponini arose in a manner different from the partially claus-
tral colony founding mode of Myrmecia remains an enigma.

References

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Fagen R. M., Goldman R. N., 1977. — Behavioral catalogue analysis methods.

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12-47.

Gotwald W. H., Levieux J., 1972. — Taxonomy and biology of a new west

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Haskins C. P., Enzmann E. V., 1938. — Studies of certain sociological and

physiological features in the Formicidae. Ann. New York Acad. Sci., 37, 97-162.
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between populations of Rhvtidoponera metallica. Psyche 86, 301-312.
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Holldobler B. K., Engel H., 1979. — Tergal and sternal glands in ants. Psyche,

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85, 285-330.

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trait in ant evolution. Naturwissenschaften, 64, 8-15.

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evolution (tr. from Russian by B. Haigh; Richards O. W., Uvarov B., eds.).
Methuen and Co., London, viii +319 pp.

Maschwitz U., Hahn M., Schoenegge P., 1979. — Paralysis of prey in ponerine

ants. Naturwissenschaften, 66, 213.

Talbot M., 1957. — Populations of ants in a Missouri woodland. Ins. Soc., 4,

375-384.

Traniello J. F. A., 1978. — Caste in a primitive ant: absence of age polyethism

in Amblvopone. Science, 202, 770-772.

Wheeler W. M., 1900. — The habits of Ponera and Stigmatomma. Biol. Bull., 2,

43-69.

Wilson E. O., 1958. — The beginnings of nomadic and group-predatory behavior

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tocerus varians (Fr. Smith). Anim. Behav., 24, 354-363.

THE BIOLOGY OF NINE TERMITE SPECIES
(ISOPTERA: TERMITIDAE)

FROM THE CERRADO OF CENTRAL BRAZIL

By Helen R. Coles de Negret1 and Kent H. Redford2
Introduction

The Neotropical region is second to the Ethiopian in numbers of
described termite species (Araujo 1970). However, little is known of
their biology. The literature on Brazilian termites is largely re-
stricted to isolated taxonomic descriptions of species from the
Amazon Basin and southern states of Brazil (Araujo 1961, 1969,
1977 and Fontes 1979). Exceptions to this include information re-
lating termite species and their distribution to vegetation types in
Mato Grosso State (Mathews 1977), the effect of deforestation on
termites in the Amazon (Bandeira 1979) and data on the ecology
and defense of termites in the cerrado vegetation of the Distrito
Federal (Coles 1980).

The present study was done in conjunction with a study on
mammalian termite predators, in particular the giant anteater,
Myymecophaga tridaetyla (Coles 1980 and Redford in prep.). Six
aspects of termite biology of importance in defense by termites
against mammalian predators were studied for nine of the most
common mound-building termite species in the Distrito Federal,
Brazil. Reported here are individual weights, morphology of soldier
castes, worker-soldier ratios, mound sizes and forms, mound hard-
nesses and nest materials, distributions and abundances of nests and
feeding habits for these nine species.

All species studied were from the family Termitidae (see Fig. 1 for
comparison of soldier heads), subfamily Apicotermitinae, Grigioter-
mes metoecus (Matthews); subfamily Nasutitermitinae, Armitermes

•Laboratoria de Zoologia e Ecologia Animal, Universidade de Brasilia, Brasilia D. F.
80910, Brazil.

2Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138; and
Department of Zoological Research, National Zoological Park, Smithsonian Institu-
tion, Washington, D.C. 20008.

Manuscript received by the editor March 3, 1982.

81

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euamignathus (Silvestri), Cornitermes cumulans (Kollar), Cortariter-
mes silvestri (Holmgren), Nasutitermes sp., Procornitermes araujoi
(Emerson), Syntermes dirus (Burmeister), Velocitermes paucipilis
(Mathews); subfamily Termitinae, Orthognathotermes gibberorum
(Mathews).

Methods and Results

This study was conducted primarily in the Distrito Federal, Brazil
(15 47'S 47 56'W) with supporting work done in Emas National
Park, Goias State (18 19'S 52 45'W). Both areas are located within
the cerrado sensu latu vegetation type. Cerrado (sensu latu ) is a
semi-deciduous xeromorphic savanna vegetation found in the inter-
mediate rainfall (750-2000 mm/ yr) area of Brazil. It is characterized
by woody plants with thick bark and coreaceous leaves and a sea-
sonal ground layer of grasses and herbs. Although geographically
and floristically the cerrado vegetation zone is very uniform, physi-
onomically it shows considerable variation (Eiten 1972). The types
of cerrado sensu latu which were examined in this study are campo
limpo (grassland), campo sujo (grassland with shrubs), cerrado
sensu strictu (woodland) and cerradao (dense, tall cerrado). Within
the cerrado zone, gallery forest vegetation is found on the wet, more
fertile soils along river courses; however this was excluded from the
present study as it supports a termite fauna which differs greatly
from that of the other vegetation types (Coles 1980).

I. The Termites
A. Comparative Morphology

Figure 1 depicts soldiers of the eight species of termites examined
in this study, with a worker head of the soldierless species Grigio-
termes provided for comparison, while Tables 1 and 2 provide
information on the fresh weights and total body lengths. Table 2
also provides measurements of mandible length, nasus length, head
length, head width and head depth for the soldiers (position of
measurements depicted in Figure 2).

As can be seen from these data, the termite species in this study
can be placed along a spectrum based on soldier and head shape.
The two ends of this spectrum are ‘well-developed nasus/ vestigial
mandibles’ (such as Nasutitermes) and ‘no nasus /verv well-devel-

1982]

Negret & Redford — Termite Species

83

Figure 1. Soldier heads of eight of the species of termites studied; Grigiotermes
metoecus worker included for comparison: a, Grigiotermes metoecus; b, Armiter-

mes euamignathus ; c, Cornitermes cumulans; d, Cortaritermes silvestri; e, Pro-
cornitermes araujoi; f, Nasutitermes sp.; g, Syntermes dir us; h, Velocitermes
paucipilis; i. Orthognat hotermes gihberorum.

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loped mandibles’ ( Orthognathotermes ). Intermediate positions are
occupied by forms with ‘slight nasus development /well-developed
mandibles’ (such as Cornitermes ) and ‘well-developed nasus/ well-
developed mandibles’ ( Armitermes ). Grigiotermes, with no soldier
caste, cannot be placed on this spectrum.

These data also show that soldiers with very well- to well-
developed mandibles and poorly developed nasi are both heavier
and longer than soldiers with vestigial mandibles and well-developed
nasi, Armitermes once again occupying an intermediate position.

Complete taxonomic descriptions for Grigiotermes metoecus,
Armitermes euamignathus, Cortaritermes silvest ri, Velocitermes
paucipilis, and Orthognathotermes gibberorum can be found in
Mathews (1977). Procornitermes araujoi is fully described in Emer-
son (1952). Samples of Cornitermes cumulans collected during the
study in Brasilia were identified following Emerson (1952). Al-
though the general head and mandible forms were consistent with
the published description, head length and width measurements
were much lower than those previously described for this species.
However, Emerson indicated that there is considerable variation in
mean measurements between colonies from different localities. The
samples from Brasilia were compared with various other species in
the Museu Zoologia de Universidade de Sao Paulo (MZSP). The
most closely related species was C. villosus which was clearly differ-
ent in that it had a greater number of setae and differently shaped
mandibles. As a result of this divergence the best classification
appears to be C. cumulans. Specimens from Brasilia have been
deposited in the MZSP and the Museum of Comparative Zoology,
Harvard University.

Samples of Nasutitermes sp. collected from the Distrito Federal
were compared extensively with material in the MZSP but differed
from all species examined. N. coxipoensis most resembled the Nasu-
titermes we studied but differed in being smaller and in having a
more oval shaped head. Further studies on these two forms are
necessary to determine whether these differences are sufficient to
warrant calling it a new species.

B. Weights

Fresh weights were measured on a Mettler balance. Fifty workers
and fifty soldiers from each of three different nests were weighed,
except for Syntermes for which only fifteen individuals of each caste

1982]

Negret & Redford — Termite Species

85

from the three nests were weighed and Nasutitermes for which five
nests were sampled. The results are presented in Table 1 and are
ordered from heaviest soldiers to lightest soldiers. Syntermes dims
has workers and soldiers much heavier than the next heaviest spe-
cies, Cornitermes. The termite species with soldiers possessing
strong or long mandibles are heavier than those termites whose
soldiers have vestigial mandibles, and well developed nasi. These
latter soldiers are also lighter than their workers, a relationship
reversed in the other termite species.

Table 1. Individual wet weights of termites (measurements expressed in micro-
grams; mean with standard deviation in parentheses).

Species

Workers

Soldiers

Syntermes dims

42.75a

117.3

(2.34)

(11.1)

Cornitermes cumulans

9.30

19.83

(0.36)

(1.07)

Orthognathotermes gibberorum

6.91

19.09b

(0.75)

(0.69)

Procornitermes araujoi

6.63

8.26

(0.76)

(0.40)

Grigiotermes metoecus

6.27

(0.95)

—

Armitermes euamignathus

3.48

4.19

(0.15)

(0.52)

Cortaritermes silvestri

3.23

2.08

(0.12)

(0.20)

Nasutitermes sp.

3.46c

1.56

(1.06)

(0.42)

Velocitermes paucipilis

2.52c

1.31b

(0.55)

(0.09)

a Equal number of all three morphs weighed,
b Only major soldiers weighed,
c Mixture of two worker types weighed.

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C. Morphology of Soldiers

The positions of measurements taken on soldier heads are indi-
cated in Figure 2 (adapted from Coles 1980). Total body length was
measured from tip of mandible or nasus, whichever extended
further, to the end of the abdomen. The figures presented in Table 2
are the averages of 15 individual soldiers and are ordered from
greatest to least mandible length. As can be seen, these five morpho-
logical measurements are, on the whole, positively correlated with
each other, with total body length and with weight (Table 1). The
major exception is Orthognathotermes, which has mandibles and a
nasus of a different shape than the other species.

D. Worker-Soldier Ratios

Worker-soldier ratios were calculated by counting all of the
workers and soldiers in a piece of termite mound. The piece was
rapidly removed from the surrounding mound so as to prevent a
change in the normal worker-soldier ratio. For all species except P.
araujoi, A. euamignathus, S. dims and C. silvestri, five pieces of
mound from at least three different mounds were counted. The
result obtained from a piece of mound was not used if the piece
contained less than 600 individuals. Because of the large variation
obtained in the first five counts for P. araujoi, an additional three
pieces were counted. The fifth count used for A. euamignathus was
an average of 45 samples and was taken from Domingos (1980).
Only four counts were taken for C. silvestri.

The large diffuse mounds inhabited by S. dims and the rapid
retreat of soldiers and workers made it impossible to obtain worker-
soldier ratios from populations within the mound for this species.
Instead, the value presented in Table 3 is an average of counts made
on eleven foraging parties. The method used (Coles 1980) was to
plug the exit at least one hour after foraging had begun. After
spraying with pyrethrin aerosol insecticide all soldiers and workers
were collected and counted. Table 3 presents the data on worker-
soldier ratios ordered from greatest to least percent soldiers.

Those termite species with soldiers having chemical-based defen-
sive systems have fewer workers per soldier than the other termite
species. In fact, for these species, Velocitermes, Nasutitermes and
Cortaritermes, there is little variation between species in this
worker-soldier ratio. Similarly, Cornitermes and Procornitermes,

Figure 2. Positions of morphological measurements of soldier heads: lh= Lat-
eral head length; ln = nasus length; lm = mandible length; Wh = maximum
head width; dh = head depth including postmentum.

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Table 2. Morphological measurements of soldiers (measurements expressed in
millimeters; mean with standard deviation in parenthesis).

Species

Mandible

Length

Nasus

Length

Lateral

Length

of

Head

Maximum

Head

Width

Head

Depth

Total

Body

Length

Orthognathotermes

2.96

0

3.06

2.09

1.83

9.25

gibberorum

(0.11)

—

(0.08)

(0.06)

(0.05)

(0.40)

Svntermes

2.45

0.16

5.20

5.15

3.17

15.57

dirus

(0.11)

(0.02)

(0.19)

(0.15)

(0.14)

(0.65)

Cornitermes

1.36

0.36

3.90

2.67

1.95

9.55

cumulans

(0.78)

(0.03)

(0.11)

(0.09)

(0.07)

(0.42)

Proeornitermes

1.14

0.45

2.46

1.98

1.57

7.47

araujoi

(0.05)

(0.03)

(0.03)

(0.04)

(0.05)

(0.21)

Armitermes

0.58

0.91

2.05

1.10

1.04

5.35

euamignathus

(0.02)

(0.05)

(0.05)

(0.04)

(0.05)

(0.20)

Nasutitermes

0.17

0.63

1.65

1.05

0.82

4.32

sp.

(0.03)

(0.02)

(0.64)

(0.35)

(0.34)

(0.14)

Velocitermes

0.15

0.80

1.65

1.05

0.82

4.32

paucipilis

(0.18)

(0.03)

(0.64)

(0.35)

(0.34)

(0.14)

Cortaritermes

0.15

0.61

1.64

1.08

0.80

3.95

silvestri

(0.02)

(0.03)

(0.06)

(0.06)

(0.06)

(0.25)

Note: Grigiotermes is excluded for it has no soldiers.

two similar species have very similar workers-soldier ratios. Armi-
termes occupies an intermediate position while Orthognathotermes
has a large number of workers per soldier.

II. The Mounds

A. Mound size and form

Table 4 presents data on mean heights, widths and lengths of ten
mounds for each of the nine species of termites. Figure 3 (a-r) con-
sists of two photographs of each species mound, one of an entire
mound and the other of a mound in transverse cross-section. As can
be seen from the data and the photographs, the shapes of these
mounds range roughly from an inverted cone ( Cornitermes ) to a low
dome ( Orthognathotermes ).

1982]

N egret & Redford — Termite Species

89

Table 3. Proportion of workers in nests (mean with standard deviation in
parentheses).

Species

Worker-

Soldier

%

Soldiers

Velocitermes paucipilis

4.00

25.80

(0.72)

(4.23)

Nasutitermes sp.

4.06

25.50

(0.83)

(5.56)

Cortaritermes silvestri

5.12

21.20

(1.64)

(6.90)

Svntermes dir us*

9.66

11.10

(2.72)

(3.02)

Armitermes euamignathus

13.82

7.68

(3.79)

(2.57)

Procornitermes araujoi

30.12

5.10

(18.30)

(3.76)

Cornitermes cumulans

30.23

3.48

(7.61)

(3.14)

Orthognathotermes gibberorum

80.75

1.30

(18.18)

(0.32)

♦Figures derived from foraging parties. See text. Grigiotermes excluded as it has
no soldiers.

The nature and form of individual mounds vary greatly and the
characteristics listed below are generalized descriptions of mounds
found in the Distrito Federal and Emas Park.

Cornitermes cumulans (Fig. 3 a,b): The mound has a very hard
outer shell of soil surrounding a soft inner core of carton (fecal
material, communited plant material add bits of soil) which often
extends below ground as much as 40 cms. The galleries are large and
unlined.

Nasutitermes sp. (Fig. 3 c,d): The mound is domed with the outer
several centimeters softer than the inner core (as in arboreal Nasuti-
termes and Constrictotermes) and often extends 25cms under-
ground. The internal structure consists of thin-walled, convoluted.

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Table 4. Dimensions of the epigeal portion of termite mounds (measurements
expressed in centimeters; mean with standard deviation in parentheses).

Species

Height

Length

Width

Cornitermes cumulans

91.6

92.8

79.5

(16.7)

(17.1)

(14.5)

Nasutitermes sp.

78.1

100.1

85.9

(14.3)

(18.2)

(16.4)

Syntermes dims

51.7

173.0

150.7

(19.4)

(26.5)

(20.5)

Velocitermes paucipilis

31.2

27.3

22.6

(4.5)

(7.0)

(5.8)

Grigiotermes metoecus

2.96

60.2

47.2

(4.5)

(7.9)

(7.2)

Procornitermes araujoi

28.8

69.5

60.0

(12.0)

(33.9)

(34.4)

Armitermes euamignathus

26.7

59.5

52.8

(5.1)

(8.8)

(8.1)

Cortartiermes silvestri

15.8

24.8

20.5

(4.7)

(3.2)

(2.6)

Orthognathotermes gibberorum

15.0

35.9

40.4

(3.0)

(11.3)

(13.6)

irregular galleries with a mottled black and soil-colored lining of
fecal origin.

Syntermes dims (Fig. 3 e,f): This species builds low-domed termi-
taria, the major parts of which are below ground level (often to
depth of 1.5 m.). The galleries are large and diffuse, often containing
grass stores and are lined with regurgitated soil in which individual
pellets are clearly visible.

Velocitermes paucipilis (Fig. 4 g,h): The mounds are pyramidal,
very soft, crumbly and are generally built around a grass tussock.
They often extend several centimeters underground in a series of
very diffuse galleries which are lined with a discontinuous layer of
black material of fecal origin. Large amounts of cut plant material
are found inside the mound.

1982] TV egret & Redford — Termite Species 91

Figure 3. Mounds of the termite species studied; external view and longitudinal
section: a and b, Cornitermes cumulans; c and d, Nasutitermes sp.; e and f,

Syntermes dims.

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[Vol. 89

Grigiotermes metoecus (Fig. 4 i,j): These medium-sized domed
mounds are often occupied by other species of termites and ants.
The galleries are distinguished by smooth, shiny soil-colored floors
and by small pieces of stone incorporated into the ‘ceilings.’ Indi-
vidual deposits of fecal material used in construction are visible on
the mound surface.

Procornitermes araujoi (Fig. 4 k,l): These medium-sized, rounded
mounds are often characterized by a thin layer of loose soil covering
the outer shell. These mounds are quite brittle and homogenous and
have galleries with a mottled lining of black soil and colored parti-
cles, probably of fecal origin. They rarely extend below ground.

Armitermes euamignathus (Fig. 5 m,n): This species builds very
characteristic slightly domed mounds. The walls are very hard but
the mound itself is only loosely held to the substratum with a cavity
frequently occurring between it and the soil. The internal structure
consists of large irregular chambers connected by very small galler-
ies. During the alate flight season mounds of this species are charac-
terized by earthen turrets several centimeters high built on the outer
surface and serving as ‘launching platforms’ for alates.

Cortaritermes silvestri (Fig. 5 o,p): This species builds soft, low
rounded mounds with large irregular galleries. The mounds are fre-
quently built around grass tussocks and extend several centimeters
underground as in Velocitermes.

Orthognathotermes gibberorum (Fig. 5 q,r): The low mounds
built by this species are covered with loose soil and bound together
by living grass stems. The galleries are regular and homogenous
throughout. The mound frequently extends several centimeters
underground but can be separated easily from surrounding soil
when pried up.

B. Mound hardness and nest material

The ‘hardness’ of a mound was measured using a soil penetrome-
ter which measures the force necessary to push a metal cone into the
soil. The resistance to penetration is obtained by dividing the load of
penetration (force applied) by the area at the base of the cone, which
was 637.939 mm3. The resistance to penetration was taken as a
measure of hardness of the mound surface.

A termite mound is not a solid structure but consists of a complex
system of galleries and chambers. The outer wall is often thick
enough for penetration of the whole cone. However, at times, the

1982]

Negret & Redford — Termite Species

93

Figure 4. Mounds of the termite species studied; external view and longitudinal
section: g and h, Velocitermes paucipilis; i and j, Grigiotemies metoecus; k and

1, Procornitermes araujoi.

cone pushed into a gallery and a low reading was obtained. In order
to obtain a representative figure for the whole mound ten measure-
ments were taken, each from different positions, e.g. base, middle,
top.

The hardness of any mound varies considerably throughout the
year with the amount of rainfall. To reduce these variations all the

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Figure 5. Mounds of the termites species studied; external view and longitudinal
section: m and n, Armitermes euamignathus; o and p, Cortaritermes silvestri; q

and r, Orthognathotermes gibberorum.

measurements were made in one month (April) at the end of the
rainy season. Some variation in hardness occurs from day-to-day
and so on any one day of recording, one mound from each of the
eight species was examined. Ten mounds from each species were
examined and ten measurements were made from each mound. Care
was taken to select approximately the same size of mound for the
ten mounds of any one species.

The mean values for the hardness of termite mounds in each
species are shown in Table 5. As the range is large (15.24-0.11
Newtons/ mm3) the data were transformed (\f~x) and the differences

1982]

Negret & Redford — Termite Species

95

Table 5. “Hardness” of outer mound and materials used in mound construction
(In column 1, any two means not followed by the same letter are significantly
different at p = 0.05. In columns 3 through 6, ++ = usually used; + = occasionally
used).

Resistance to

Penetration (Newtons mm3) Nest Construction Material

Species

Termite

Mound

Soil at
Base

Soil

Regurgitated

Soil

Fecal

Material Saliva

Velocitermes

0.11a

0.48

++

++

paucipilis

(0.05)

(0.16)

Nasutitermes

0.25b

0.42

++

++ ++

sp.

(0.05)

(0.15)

Cortaritermes

0.25b

0.44

++

++

silvestri

(0.04)

(0.18)

Procornitermes

0.36b

0.42

++

. ++

++ ++

araujoi

(0.11)

(0.14)

Orthognat hotermes

0.48

—

++

++

gibberorum *

(0.15)

—

Syntermes

0.57c

0.42

+

++

+

dims

(0.13)

(0.14)

Grigiotermes

1.25d

0.70

+

++

metoecus

(0.17)

(0.18)

Armitermes

4.66e

0.36

++

+

euamignathus

(1.08)

(0.10)

Cornitermes

15.241

0.37

+

++

++

cum u Ians

(5.36)

(0.16)

* Determined for only 4 mounds so no statistics performed.

between these means tested for significance using Hartley’s multiple
range test. The ranking obtained from this analysis is shown in
Table 5 with the mean values of the raw data. Velocitermes, Nasuti-
termes, Cortavitermes and Procornitermes had the softest nests while
Cornitermes had the hardest nest, 140 times harder than the softest,
Velocitermes.

The composition of material used to build mounds was deter-
mined by direct observation of workers. Observations were made on
at least ten mounds per species, at different times of the day and
year. The results are presented in Table 5. Four types of material

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were observed to have been used by termite workers in nest construc-
tion: soil, regurgitated soil, fecal material and saliva. In some cases,
such as Procornitermes nests, all four were used. Soil and/or re-
gurgitated soil were always the most common forms of building
material.

C. Distribution and Abundance of Nests

Information on the distribution and abundance of termite
mounds in each vegetation type was collected from a variety of
sources and the results are presented in Table 6. Different sampling
methods can produce different results, depending on the spatial
distribution of the termite mounds, the size of area sampled and the
number of areas sampled. It is often difficult to interpret figures on
termitaria densities because investigators do not report whether all
termitaria examined contained the mound-building species. Thus,
the specific methods used to obtain each of the densities reported in
Table 6 are detailed below.

Method a: (Coles 1980); method b (Domingos 1980); method
e (Coles de Negret et al. in prep.).

Blocks of 50 X 50 meters were selected randomly in each of the
four vegetation types studied in the Distrito Federal. As some of the
termite species in the present study were occasionally found in
mounds built by other species, in these methodologies, all the epi-
geal mounds in the area were completely excavated. The abundance
of each species was thus expressed in numbers of nests per hectare.
In order to exclude sites with only foraging termites, a “nest” was
defined as a structure in which termite nymphs and larvae were
present.

Method d: (Redford in prep.).

Twelve separate transects, each of 100 by 20 meters were marked
out in the campo limpo vegetaton of Emas National Park, Goias.
All the mounds built by Cornitermes cumulans in each transect were
counted. The figure in Table 6 is the mean calculated from these
twelve transects (standard deviation = 16.1).

Method e: (Brandao in prep.).

Two blocks, 100 by 100 meters were marked out in separate areas
of campo sujo and two others, of the same size, in areas of cerrado
vegetation in the Distritb Federal. All the Syntermes dims mounds
present in each area were counted. As this species frequently con-
structs small soil domes, apparently for storing food, nests were

1982]

Negret & Redford — Termite Species

97

Table 6. Distribution and densities of termite nests/mounds per hectare in four
vegetation types (Letters correspond to different sampling methods — see text for
details).

Species

Campo

Limpo

Campo

Sujo

Cerrado
Sensu Stricto

Cerradao

Grigiotermes

40a

28a

24a

48a

metoecus

4c

Armitermes

84a

1 12a

1 16a

124a

euamignathus

236b

116b

152b

120b

4 1 f

156c

Cornitermes

0a

12a

32a

0a

cumulans

58d

0c

Cortaritermes

40a

12a

4a

0a

silvestri

Nasutitermes

48a

32a

0a

0a

sp.

16c

Procornitermes

4a

12a

52a

4a

araujoi

12c

Syntermes dirus

4a

20a

0a

0a

33e

Oe

54e

8e

Velocitermes

40a

96a

32a

0a

paucipilis

lOlf

24c

27g

Orthognat hotermes

12a

0a

16a

4a

gihberorum

again defined as structures in which termite nymphs and larvae were
present.

Method f: (Curado et al. in prep.).

All the mounds built by Armitermes euamignathus and Ve/oci-
termes paucipilis in an area of campo sujo ( 100 by 100 meters) in the
Distrito Federal were sampled and counted.

Method g: (internal report. University of Brasilia).

Mounds of Velocitermes paucipilis present in a transect 230 by 10
meters extending from campo limpo to campo sujo in the Distrito
Federal were counted.

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III. Feeding Habits and Foraging Behavior

Feeding habits were deduced from field observations, examina-
tion of worker mandibles and gut contents, information in the liter-
ature and in some cases, from laboratory food preference experiments.
Results are summarized in Table 7. Details of foraging behavior,
methods of investigation and food sources are given below.

Grigiotermes metoecus

Field observations and examinations of worker mandibles and
gut contents indicate that this species is entirely geophagous. It
excavates subterranean galleries in the soil surrounding its mound
and is also frequently found in old, disused termite workings, pre-
sumably rich in organic material.

A rmitermes euamignathus

In the cerrado and cerradao vegetations foraging workers can be
found under the bark of living trees and sound, dead trees. How-
ever, this species also occurs with equal frequency in campo limpo
where few or no woody shrubs exist. Field observations on the
foraging behavior of 100 colonies of this species show that in the
absence of woody vegetation they can exploit the root systems of
grasses (Domingos 1980). Laboratory food preference experiments
carried out by the same author on five colonies of A. euamignathus
indicates that when presented with a range of food sources, all
colonies selected wood in preference to bark, litter and grass roots.
Further field observations confirmed that this species selects dead,
sound wood in preference to live and to dead, decomposed wood.
The workers forage diurnally and reach the food source via subter-
ranean galleries. On average, mounds are 0.4 and 0.3 meters from
their food sources in cerradao and cerradao respectively and 1.2 and
1.0 meters in campo sujo and campo limpo, respectively (Domingos
op. cit.).

Cornitermes cumulans

Field observations on foraging parties indicate that workers of
this species feed on living and dead grasses and herbs, which they
reach through subterranean tunnels, occasionally foraging under a
fine layer of soil-sheeting. Small pieces of grass are cut from stand-
ing grass tussocks and carried to the mound. Feeding in situ has
been observed occasionally. Preliminary food preference experi-

1982]

Negret & Redford — Termite Species

99

Table 7. Modal feed
consumed).

ing habits (+ + = commonly consumed; + =

occasionally

SPECIES

FOOD SOURCE

Grass &

Sound Decomposing

Herbaceous

Humus Wood Wood

Litter

Grigiotermes

++

metoecus

Armitermes

++

_|_

euamignathus

1

Cornitermes

++

cum u Ians

Cortaritermes
si Ivest ri

+(?)

+ (?)

Nasut iterates sp.n.

+ ' ! + '

++

Procornitermes

++

araujoi

Svntermes dirus

++

Velocitermes

paucipilis

-++

Orthognat hotermes
gihherorum

++(?)

ments carried out on laboratory colonies showed that workers col-
lect dead grass in greater amounts than live. When presented with
only dead roots or dead grass blades, they fed more on the latter.

Cortaritermes si /vest ri

Field observations made in the Distrito Federal and information
presented in Mathews ( 1977) indicate that this species feeds in grass
tussocks among the roots and stems. It is not clear, however,
whether it feeds on the organic residues in the soil or on the grass
roots themselves.

Nasutitennes sp.

These termites have not been observed foraging in the open and
rarely construct runways over the ground as do many other species
in this genus. It is probable that they excavate underground tunnels
to their food source, the exact nature of which is not known. Recent

100

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[Vol. 89

experiments on laboratory colonies have shown that this species can
feed on a range of plant material including sound wood and both
living and dead grass.

Procornitermes araujoi

Field observations have been made on above-ground foraging
parties in the open and under soil sheeting. Workers cut and collect
grass litter, generally at night, but occasionally on dull, humid days.

Svntermes dims

This species forages above ground in the open, at night, and
crepuscularly. Workers and soldiers leave the tunnels from small
exit holes which are plugged with several millimeters of soil during
inactive periods. These foraging holes may be on the mound or at
distances of up to 20 meters from it. The above-ground foraging
parties consist of major workers and soldiers. At the end of a partic-
ular trail the workers spread out over several centimeters and start
cutting grass. Some climb up stands of vegetation and cut long
pieces of grass which drop to the ground. Other workers cut these
into smaller pieces and carry them to the nest. Consumption in situ
has not been observed.

Ve/ocitermes paucipiiis

These termites feed on grass and surface litter which they collect
at night in the open. The workers form trails to the food source
where they spread out to cover a large area, cut small pieces of grass
and leaves, and return with them to the nest. The workers are
flanked at regular intervals by soldiers oriented with their raised
heads pointing outwards.

Orthognathotermes gibber ovum

Examination of worker mandibles and gut contents together with
information from Mathews (1977) suggests that this species feeds on
organic residues in the soil. Observations of foraging behavior have
not been made.

Food sources were divided into four categories: humus, sound
wood, decomposing wood, and grass and herbaceous litter. The few
termites eating sound wood and the many eating grass and herba-
ceous litter probably reflect the fact that most of the vegetation
types included in this study were open with few trees. Examination
of the termite fauna within the gallery forests would reveal many

1982]

Negret & Redford — Termite Species

101

more wood-eating species. The predominance of grass-eating ter-
mites is understandable because of the large biomass and rapid
turnover of their food source.

Of the 54 species of termites in the cerrado vegetation of the
Distrito Federal (excluding gallery forests) only nine mound-
building species were examined in this study. Many of the other
species do not build mounds and are found instead living within
mounds built by one of these nine species. It is probable that many
of these non-mound-building species will be found to be geophagous
or humivorous, feeding in or near the mounds they inhabit.

Discussion

The cerado vegetation of the Distrito Federal, Brazil has a diverse
termite fauna with at least 54 species present (excluding those found
in gallery forest vegetation) (Coles 1980). Estimates of the termite
density in savanna areas in other continents are much lower with
only 19 species in the Sahel, Senegal, 19 in northern Guinea, Nige-
ria, 23 in southern Guinea, Nigeria and 36 in savannas of the Ivory
Coast (Wood and Sands 1978).

A survey by Coles (1980) indicated that most cerrado species were
present in all the physionomic vegetation types; however, in terms of
abundance, certain species were more common in one particular
type of vegetation. This is clearly illustrated by the data in Table 6.
Nests of Nasutitennes sp., Velocitermes paucipilis, Cortaritermes
si/vestri, Syntermes dirus and Cornitermes cumulans were all more
abundant in the open vegetation types (campo limpo and campo
sujo). Grigiotermes metoecus and Armitermes euamignathus were
equally common in all types while Procornitermes araujoi was more
common in woodland areas. Orthognathotermes gibberorum had
an irregular distribution being less common in the cerrado sensu strictu
of the Distrito Federal but more common in the campo limpo of
Emas Park. These preferences for particular vegetation types can, to
some extent, be related to the feeding habits of each species (Table
7); however, abundance of a species is also influenced by other
species present. In some areas conditions were particularly favorable
for one species, an example of which was found in Emas National
Park where populations of Cornitermes cumulans were exception-
ally high, with other species much less common.

The variation in abundance of a species in different regions can be

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accompanied by variations in mound form and size. Howse (1979)
gives several different examples of termite species which build very
different mounds in different regions. Macrotennes subhvalinus in
western Uganda builds mounds with very thick walls and no open-
ings but on the Serengeti Plains, where the soil is volcanic ash, the
mounds are low with many pit-like openings. In the semi-arid
regions of eastern Africa they are different again, being steeple-
shaped and constructed around a central chimney. Even though
regional differences can exist, the characteristics of mounds investi-
gated in this study showed a remarkable consistency throughout the
cerrado region reinforcing observations by Emerson (1938).

In constructing a mound, galleries are excavated within the soil
by the termites and particles are often transported from considera-
ble depth and incorporated in the epigeal portion of the mound.
This not only increases aeration of the soil but can also alter its
chemical composition (Lee and Wood 1971). Soil used in building is
reinforced with excreta and in some instances wood and other plant
material.

Studies on the chemical composition of termite mounds in the
cerrado have recently been started in Brasilia. Preliminary results
indicate that both Ve/ocitermes and Armitermes mounds have much
higher concentrations of calcium, phosphorus, potassium and alu-
minum than the soil surrounding the mound (Curado et al. in
prep.). However, an analysis of Table 5 shows that the materials
used in mound building are not directly related to the hardness of
the outer layer of the mound. Such factors as the way in which the
material is deposited by the workers at the actual site of construc-
tion as well as the size and arrangement of galleries and the thick-
ness of walls also contribute to the overall hardness of the mound.

The mounds are constructed entirely by the worker caste. This
caste takes little active role in the defense of the mound, a role
performed by the soldier caste. The proportion of these two castes
varies with the species and is apparently finely regulated by phero-
mones produced by the queen and the soldiers (Luscher 1961).
Haverty (1977), in a comprehensive work, summarized the data
available on the relative proportion of workers and soldiers in 1 12
species of termites. Unfortunately, many of these data, gathered by
different investigators, are not strictly comparable because of differ-
ences in sampling techniques and types of groups sampled. The

1982]

N egret & Red ford — Termite Species

103

homogeneity in methodology used in calculating worker-soldier
ratios in this study allows for precise comparison between species
within the limits of accuracy of this method. The worker-soldier
ratios were found to vary greatly between nests in some species (i.e.,
Procornitermes ) and remain quite constant in others (i.e., Velocitermes).

The behavior of nasute soldiers, which respond to a break in the
nest by rapidly recruiting to the break, can greatly alter the worker-
soldier ratio calculated. As an example of this, on one occasion the
number of soldiers counted from a piece of Nasutitermes mound,
which had been excised from the surrounding mound but left in
place for 30 seconds, was almost half again the number of soldiers
counted from a piece taken from the same mound but removed
immediately following excision. Although comparison can be made
between the nine species of termites it must be noted that these data
were taken during one period of the year and present a static picture
of the proportions of workers and soldiers in given nests. It seems
probable that in the species examined, as in other species (Sands
1965), the worker-soldier ratio varies seasonally and possibly also
with the age and size of the nest.

It is evident from the data that some species have proportionally
many more soldiers than other species. Even though the proportion
of soldiers in a colony varies, in all cases (when there is a soldier
caste) the soldier caste is largely responsible for the defense of the
colony and has morphological features which allow it to do this.
The type of defense used by soldier termites tends to be based on
chemicals, mechanical defense or a combination of both. The sol-
dier type using a chemical-based defense has vestigial mandibles
(Table 2), is lighter than its workers (Table 1), and produces poten-
tially toxic and repellent secretions which are ejected from the tip of
a long tube or nasus at the front of the head (Nutting et al. 1974,
Eisner et al. 1976; Howse 1975; Prestwich 1979). Of the termites
studied in this work, Velocitermes, Nasutitermes and Cortaritermes
fall into this category. The soldier type using a mechanical-based
defense rarely produces defensive secretions and has a large head,
and strong, sharp mandibles. Orthognathotermes is the only species
within those here studied that has no development of the nasus,
relying solely on its mandibles for defense. Syntermes, Cornitermes
and Procornitermes all have strong mandibles which can pierce
human skin, drawing blood, together with a greatly reduced level of

104

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[Vol. 89

chemical defense (see ‘nasus length’ Table 2 as one indicator of the
extent to which chemicals are used in defense). Armitermes stands
in an intermediate position between the principally chemical and the
principally mandibulate type soldiers, with a long nasus and mandi-
bles which can pierce human skin but not draw blood. Grigiotermes
is very interesting in that it has no soldiers; the workers however
produce a large drop of liquid on either side of the abdomen when
disturbed, which may serve a defensive purpose.

Termites are probably the dominant form of animal life in many
areas of central Brazil, both in number of species and biomass. They
play major roles in herbivory, decomposition, soil formation and
alteration, and as an important source of food for other animals.
Ants are probably the major predators of termites, but in central
Brazil mammals are common and important predators as well. The
aspects of termite biology reported in this study are all important in
defense by termites against mammalian predators. The small size of
termites, the type of soldier defense and the proportion of soldiers to
workers are all factors influencing feeding by mammals once the
termite mound has been opened. The shape, size and hardness of a
mound influence the ways in which a mammalian predator can
break into a nest while the distribution and abundance of nests are a
measure of the spatial availability of termites as a food source.
Lastly, the feeding habits of termites are important in determining
when, and if, termites are available outside of the mound. Food
preference tests with large and small mammalian predators and
observation of wild giant anteaters (Redford in prep.) have shown
that all of these aspects of termite biology interact in determining
which species of termites are preferred as food and how available
they actually are to mammalian predators.

Acknowledgements

Helen Coles de Negret would like to thank the Trustees of the
Royal Society Leverhulme Scholarships and the Science Research
Council-Shell Research CASE award for financing this research.
The data form part of a Ph.D. thesis submitted to Southampton
University in 1980 under the supervision of Dr. P. E. Howse.

Kent Redford would like to thank the National Geographic
Society, the Museum of Comparative Zoology, the Organization of
American States and Sigma XI for help in financing this research.

1982]

Negret & Redford — Termite Species

105

Special thanks to the members of the Order of Saint Benedict and
the Laboratory of Ecology, University of Brasilia. Both authors
thank Barbara L. Thorne, Alan E. Mill, James F. A. Traniello and
Bert Holldobler for reading and criticizing the manuscript.

Literature Cited

Araujo, R. L.

1961. New genus and species of Brazilian termite. Revta. Bras. Biol. 21,
105-111.

1969. Notes on Dentispicotermes with description of a new species. (Isoptera,
Termitinae). Revta. Bras. Biol. 29, 249-254.

1970. Termites of the Neotropical Region. In: Biology of Termites, Vol. II,
(Ed. by K. Krishna and F. M. Weesner) pp. 527-571, Academic Press, N. Y.

1977. Catalogo dos Isoptera do Novo Mundo. Academia Brasileira de Cien-
cias. Rio de Janeiro, RJ.

Bandira, A. G.

1979. Ecologia de cupins (Insecta: Isoptera) da Amazonia central: efeitos do
desmatamento sobre as populacoes. Acta amazonica 9, 481-499.

Brandao, D. in prep.

Ecologia de duas especies simpatricas de Svntermes (Isoptera; Nasu-
titermitinae) no Distrito Federal do Brasil.

Coles, H. R.

1980. Defensive strategies in the ecology of Neotropical termites. Ph.D. thesis
Southampton University. 243 pp.

Coles de Negret, H. R., Domingos, D. J. and Fontes, E. G. in prep.

Spatial distribution of termite mounds in the cerrado vegetation, Dis-
trito Federal, Brazil.

Curado, W., Coles de Negret, H. R., Haridasan, M. in prep.

Composition of the nest material of two termite species and the soil of
their bases.

Domingos, D. J.

1980. Biologia, densidade e distribuigao espacial de duas especies de Armi-
termes (Termitidae) em cinco formagoes vegetais do cerrado. M.Sc.
thesis Universidade de Brasilia. 22 pp.

Eisner, T., Kriston, I. and Aneshansley, D. J.

1976. Defensive behaviour of a termite Nasutitermes exitiosus. Behav. Ecol.
Sociobiol. 1, 83-125.

Eiten, G.

1972. The cerrado vegetation of Brazil. Bot. Rev. 38, 201-341.

Emerson, A. E.

1938. Termite nests. A study of the phytogeny of behaviour. Ecol. Mono-
graphs. 8, 247-284.

1952. The Neotropical genera Procornitermes and Cornitermes (Isoptera,
Termitidae). Bull. Am. Mus. Nat. Hist. 99, 429-471.

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Fontes, L. R.

1979. Atlantitermes novo genero de cupim, com duas novas especies do Brasil.
(Isoptera, Termitidae, Nasutitermitinae) Rev. Bras. Ent. 23, 219-227.

Haverty, M. I.

1977. The proportion of soldiers in termite colonies: a list and a bibliography.
Sociobiology 2, 199-216.

Howse, P. E.

1975. Chemical defenses of ants, termites and other insects: some outstanding
questions. Proc. 1USS1. (Dijon), 23 29.

1979. The uniqueness of insect societies: aspects of defense and integration. In:
Biology and Systematics of Colonial Organisms (Ed. by G. Larwood and
B. R. Rosen), pp. 345-374. Academic Press, New York.

Lee, K. E. and Wood, T. G.

1971. Termites and Soils. Academic Press, New York.

Luscher, M.

1961. Social control of polymorphism in termites. In: Insect Polymorphism
(Ed. by J. S. Kennedy), pp. 57-67. Roy. Entomol. Soc., London.

Mathews, A. G. A.

1977. Studies on termites from the Mato Grosso State, Brazil. Academia Bra-
sileira de Ciencias, Rio de Janeiro, RJ. 267 pp.

Nutting, W. L., Blum, M. A. and Fales, H. M.

1974. Behavior of the North American termite Tenuirostritermes tenuirostris
with special reference to the soldier frontal gland secretion, its chemical
composition and use in defense. Psyche, 81, 167-177.

Prestwich, G. D.

1979. Chemical defense by termite soldiers. J. Chem. Ecol. 5, 459-480.

Sands, W. A.

1965. Mound population movements and fluctuations in Trinervitermes
ebenerianus Sjostedt (Isoptera, Termitidae, Nasutermitinae). Insect.
Soc. 12, 49-58.

Wood, T. G. and Sands, W. A.

1978. The role of termites in ecosystems. In: Production biology of ants and
termites (Ed. by M. V. Brian), pp. 245-292. Cambridge University Press.

THE LIFE HISTORY OF THE JAPANESE CARRION
BEETLE PTOMASCOPUS MORIO AND THE ORIGINS OF

PARENTAL CARE IN NICROPHORUS (COLEOPTERA,
SILPHIDAE, NICROPHORINI).*

By Stewart B. Peck

Department of Biology, Carleton University,

Ottawa, Ontario, K1S 5B6, Canada

Introduction

The subject of the origin and evolution of sociality in insects has a
rapidly growing literature. Most of this pertains to the Hymen-
optera. Within the Coleoptera, presocial or subsocial parental care
and division of labor are known in at least nine families (Wilson,
1971). The most advanced form of parental care known in beetles is
that of the Nicrophorus carrion or burying beetles (tribe Nicro-
phorini). This generalization is based on the study of six European
species by Pukowski (1933, 1934) which has since been abstracted
and popularized by many (e.g., Balduf, 1935; Milne and Milne,
1944, 1976; Wilson, 1971, 1975). Briefly, a male and female form a
conspecific pair at a carcass of a mouse or other small vertebrate.
They work cooperatively to exclude competitors, to bury the
carcass, and to shape it into a ball in a crypt. The male leaves after
oviposition but the female tends the developing larvae, calling them
to the carrion by stridulation, and repeatedly feeds them by
regurgitation. Such behaviors do not exist in the other tribe of
silphid carrion beetles, the Silphini.

The only work on the life cycle of a North American Nicrophorus
is a short note by Leech (1934) on N. defodiens (under the name N.
conversator). Thus, it is not really known how general or wide-
spread is the phenomenon of parental care in the genus, nor if all
species are equally advanced behaviorally. There are about 20
species in the New World, and at least 65 species in all the world, in
several lineages within the genus.

As part of a series of studies on the comparative biology and
evolution of silphid beetles, I undertook a study of the life history of
Ptomascopus morio Kraatz of Japan, to learn something of the

* Manuscript received by the editor October 29, 1981.

107

108

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[Vol. 89

origin of parental care in Nicrophorus. Ptomascopus is the only
other genus in the tribe Nicrophorini and contains only two Asian
species, P. morio being more common and widespread than P.
plagiatus Menetries (Hlisnikowski, 1942). It is illustrated in many
general Japanese insect books such as Esaki et al. (1932, 1956),
Nakane et al. (1963), and Nakane (1980). The larvae are illustrated
by K. Kurosa in Kawada (1959).

The genus shares with Nicrophorus many derived morphological
characters relative to the Silphini: adults with stridulatory files,
reduced second antennal segment, fused gular sutures, sexually
dimorphic membranous anticlypeus; larvae with abdominal para-
notal projections and cuticular sclerotization reduced, and with only
one pair of ocelli.

The main morphological characters in which Ptomascopus is
more primitive than Nicrophorus are in its possession of a normally
clavate antennal club, rather than with a strongly capitate club
formed from the last four segments, and in its less fossorial tibiae.

Methods and Materials

Four pairs of P. morio were collected in August, 1980, at carrion
baits in a warm-temperate mixed mesophytic forest in the Omogo
Valley of Mount Ischizuchi Quasi-National Park, Shikoku, Japan.
They were brought to Ottawa, Canada, and placed in culture at
18° C, with a normal daylight regime, from September to December.
The pairs were kept in separate seven cm deep boxes of clear plastic,
floored with five cm of coarse damp sand. Two cm cubes of chicken
neck were given as carrion food at required intervals. Observations
were made daily. The data gained are variable in quantity and
quality and are usually not abundant enough for tests of signifi-
cance. Only simple means, sample sizes, and ranges are reported,
but these are sufficient for comparative purposes.

Results

Both sexes dug irregular tunnels in the sand but not in direct
association with the carrion. Most of their time was spent in these
tunnels. They fed at the carrion and sporadically dug under it, but
there was no direct indication of digging with the intention of
burying the food, or of manipulating the food into a food ball, or of
forming a crypt for it. Mating was observed occasionally but no
indication of a courtship ritual was noted.

1982]

Peck — Life History of Ptomaseopus morio

109

Eggs were laid singly in the sand several cm to the side of the
carrion. A mean of 13 eggs (N = 9, r = 9-16) were laid per female in
6 days (N = 9, r = 5-8), and a new clutch was started after a
refractory period of 6 more days (N = 8, r = 5-8). The eggs hatched
in 5 days (N = 30, r = 4-7). Frequent adult attempts to fly and leave
the culture containers after the egg clutch was laid may indicate that
post-mating (for the male) or post-oviposition dispersal is normal,
and that the adults are normally not present with their young.

The larvae fed together under and directly on the carrion. There
was no indication of parental attendance to, or feeding of, the
larvae. The adults and larvae may feed on fly larvae or other insects
associated with carrion in nature, but carrion alone is adequate
for complete development of larvae in culture. There were 3 larval
instars; the first lasted 1 day (N = 30, r = 1-2), the second 2 days
(N = 30, r = 2-3). The third instar larvae fed for 7 days (N = 30,
r = 6-9) before crawling away from the carrion and burrowing into
the sand to form pupal cells. In total, over 300 larvae were pro-
duced, of which about 50 were preserved for morphological study.

Prepupae had a high mortality due to a fungal contamination.
The prepupal phase seems to be about 30 days in duration (N = 7,
r = 28-40). The pupal phase also seems to last about another 30 days
before emergence of the adult (N = 2, r = 25-35). At culture
temperatures the parental generation adults died by early Decem-
ber, for a longevity of at least four months. This could be con-
siderably different in the field depending on their sensitivity to cool
fall temperatures and whether or not they overwinter as adults.

Discussion

There was no indication of any subsocial or other behavioral
association between the larvae and the adults as known in Nicro-
phorus. The brood size, reduced fecundity, and shorter larval
developmental times are similar to those reported in Nicrophorus ,
but otherwise the life cycle characteristics are generally similar to
those reported for the carrion-feeding Silphini (Balduff, 1935;
Brewer and Bacon, 1975; Cole, 1942; Cooley, 1917; and Ratcliffe,
1972). It should be noted that some Silphines appear to have derived
feeding characteristics, being strict predators and phytophages.
How this may have changed behavior and life cycle characteristics is
not known.

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The results were verified by Dr. Kazuyoshi Kurosa of Tokyo
(pers. comm.) who reared the beetle some 30 years ago in Oita
Prefecture, Japan, but did not publish the results. He found no
parental care, no sign of burying the food, and no parental
attendance on the larvae, which grew well on fresh beef. Still,
further observations with a natural forest soil substrate and natural
food items like mouse or shrew carcasses would be desirable. How
the beetles survive and “partition resources” in the face of what
seemed to me to be severe competition from the diverse fauna of
Japanese carrion beetles remains unknown.

Conclusions

It appears that the origin of parental care of larvae did not occur
in an ancestor common to Ptomascopus and Nicrophorus, but
seemingly in Nicrophorus itself, after the differentiation of the
genus. If the origin was sometime after that of the genus itself we
may expect a wider range of parental care and related behaviors in
Nicrophorus than is generally assumed in the recent literature on
these beetles. A greater number of Nicrophorus species should be
studied to investigate the questions of the origin and evolution of
sub-sociality within the genus, and the results should be evaluated
with reference to a cladistic (phylogenetic) analysis of the evolution
of morphological characters.

Acknowledgments

I thank Dr. Shun-Ichi Ueno of Tokyo and Dr. Kazuo Ishikawa of
Matsuyama for making my Japanese field work possible and
exceptionally informative. Field support was from operating grants
of the Canadian Natural Sciences and Engineering Research Coun-
cil. The manuscript was read and helped by comments from R.S.
Anderson, A.F. Newton, K. Kurosa, R.B. Madge, and D.S. Wilson.

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Balduf, W. V.

1935. The bionomics of entomophagous Coleoptera. J. S. Swift Co., St. Louis.
220 pp. Reprinted in 1969 by E. W. Classey, Hampton, England.
Brewer, J. W. and T. R. Bacon

1975. Biology of the carrion beetle Silpha ramosa Say. Ann Entomol Soc Am
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111

Cole, A. C., Jr.

1942. Observations of three species of Silpha (Coleoptera: Silphidae). Am
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Cooley, R. A.

1917. The spinach carrion beetle. J Econ Entomol 10: 94-102.

Esaki, T., H. Hori, S. Hozawa.

1932. Iconographia Insectorum Japonicorum. Hokuryukan, Tokyo. 4404 pp.

Esaki, T., T. Ishii, T. Kawamura.

1956. Iconographic Insectorum Japonicorum. Editio Secunda, Reformata.
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Hlisnikowski, J.

1942. Coleopterologische Notizen. Mitteil Miinchner Entomol Gesells 32:
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Kawada, A.

1959. Illustrated Insect Larvae of Japan. Hokuryukan Co., Ltd., Tokyo. 712
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Leech, H. B.

1934. The family history of Nicrophorus conversator Walker. Proc British
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Milne, L. J. and M. J. Milne

1944. Notes on the behavior of burying beetles ( Nicrophorus spp.). J New
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Milne, L. J. and M. J. Milne

1976. The social behavior of burying beetles. Scientific American, 235: 84-89.

Nakane, T., K. Ohbayshi, S. Nomura, and Y. Kurosawa

1963. Iconographic Insectorum Japonicorum, Colore naturali edita, Volumen
II (Coleoptera). Hokuryukan, Tokyo. 443 pp.

Nakane, T.

1980. Coloured illustrations of the insects of Japan, vol. I, Coleoptera.
Enlarged and revised, edited by the Japan Coleopterological Society.
Hoikusha Pub., Osaka. 275 pp.

Pukowski, E.

1933. Okologische Untersuchungen an Necrophorus F. Zeit Okol Morph Tiere
27: 518-586.

Pukowski, E.

1934. Die Brutpflege des Totengrabers. Entomol Blatter 30: 109 113.

Ratcliffe, B. C.

1972. The natural history of Necrodes surinamensis (Fabr.) (Coleoptera:
Silphidae). Trans Am Entomol Soc 98: 359-410.

Wilson, E. O.

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

Wilson, E. O.

1975. Sociobiology, the new synthesis. Belknap Press, Harvard University
Press, Cambridge, Mass. 697 pp.

TERGAL AND STERNAL GLANDS IN MALE ANTS*

By Bert HOlldobler and Hiltrud Engel-Siegel

Department of Organismic and Evolutionary Biology,
MCZ-Laboratories, Harvard University,
Cambridge, Massachusetts.

Introduction:

Several recent morphological investigations have uncovered a
variety of hitherto unknown or neglected exocrine glandular struc-
tures in ant workers (Holldobler and Haskins 1977; Holldobler and
Engel 1978; Kugler 1978; Jessen et al 1979; Holldobler et al 1982;
Holldobler 1982; Jessen and Maschwitz in press). The behavioral
functions of several of these glands have already been determined
(For review see Holldobler 1982).

These studies dealt almost exclusively with ant females and except
for the results of Janet’s (1902) classical histological investigations
of the internal anatomy of males of Myrmica rubra, nothing is
known about exocrine glandular structures in the gaster of ant
males. Since we consider this information important not only for a
further analysis of the behavior of ant males, but especially for our
understanding of the evolution of pheromone glands and chemical
communcation in ants, we have undertaken a histological study of
exocrine glandular structures in ant males. In this paper we present
a survey of the abdominal glands not directly associated with the
gonads. The purpose of this paper is not to give detailed descriptions
of each gland found, but rather to present a comparative account of
abdominal glands detected in representative species in the different
subfamilies.

Materials and Methods:

For histological investigations live specimens were fixed in alco-
holic Bouin or Carnoy (Romeis 1948), embedded in methyl methac-
rylate, and sectioned 8 /x thick with a D-profile steel knife on a Jung
Tetrander I microtome (Rathmayer 1962). The staining was Azan
(Heidenhain). Especially small objects were embedded in a water
soluble plastic (JB-4 embedding kit. Polysciences, Inc., Pennsyl-

* Manuscript received by the editor May 1, 1982

113

114

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[Vol. 89

vania) and sectioned 4-6^u thick with glass knives on a rotary micro-
tome. In this case the staining was Hematoxylin-Eosin (triple
strength). The SEM pictures were taken with an AMR 1000 A
Scanning Electron Microscope. In a few cases only specimens were
available which had been preserved in 70% ethanol.

Results:

The major results are summarized in table I. In the following we
will discuss some of the details of our findings.

Penis and subgenital plate glands:

Janet (1902) described in males of the myrmicine species Myrmica
rubra two major glandular structures directly associated with the
copulatory apparatus. ( 1 ) The first comprise the penis glands, paired
clusters of glandular cells located inside the penis valves (Fig. 1).
Each cell sends a duct through a membrane into the lumen formed
by the valves (sperm gutters). This gland was also detected in males
of Formica rufa (Clausen 1938) in Conomyrma brunnei and Fore-
lius sp. (Marcus 1953; cit. in Forbes 1954), in Camponotus pennsyl-
vanicus (Forbes 1954), in Neivamyrmex harrisi (Forbes & Do-Van-
Quy 1965) and we found it in representative species of all major
subfamilies of ants. The size of the paired penis gland clusters
(which are also called aedeagal gland, Forbes 1954) varies greatly
among different species. In some it is a major gland (Fig. 1). In
others it is represented only by a few glandular cells, and sometimes
we were unable to identify the opening of the glandular ducts. (2)
The other major gland, associated with the copulatory apparatus is
located in the 9th sternite, which together with the coxopodites
comprise the subgenital plate (Weber 1954). We therefore named
these paired clusters of glandular cells “subgenital plate gland”.
Each glandular cell sends a duct through the intersegnental mem-
brane into the ventral part of the genital chamber (Fig. 1,2). The
subgenital plate gland was found in representative species of all
subfamilies studied.

Tergal glands:

In his study of the workers and males of Myrmica rubra, Janet
(1898, 1902) discovered a pair of clusters of a few glandular cells
under the 6th abdominal tergite. Each cell is drained by a duct that
penetrates the intersegmental membrane between the 6th and 7th

19821 Holldobler & Engel-Siegel — Glands in Male Ants 115

A

Fig. 1 A. Schematic drawing of a longitudinal section through the gaster of a
Novomessor B. Longitudinal section through 6th, 7th, 8th and 9th abdominal
segments of a Novomessor albisetosus 51. A=anus; P=part of penis with penis
gland; PG=pygidial gland; PPG=post-pygidial gland; SPG=subgenital plate gland.

116

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[Vol. 89

Fig. 2 A. Longitudinal section through pygidial gland of Novomessor a/bisetosus
(5- B. Longitudinal section through subgenital plate gland of N. albisetosus
ft. CS= cuticular structure; GC=glandular cells; DO=openings of glandular ducts.

1982] Holldobler & Engel- Siegel — Glands in Male Ants 1 17

abdominal tergites. In recent investigations this gland was found in
workers of representative species belonging to all subfamilies,
except in the Formicinae. Although the structure and size of the
gland varies greatly, its wide distribution led us to conjecture that
this gland might be a primitive monophyletic trait in ants generally,
perhaps reaching back to the typhioid (or mutilloid) wasp ancestors
of ants. In fact, we have recently found first indications that this
gland is also present in some living typhiid wasps.

Since this gland is anatomically closely associated with the last
exposed tergite in female ants (7th abdominal tergite = pygidium)
Kugler (1978) suggested that it be called the pvgidial gland. Of the
several tergal glands recently discovered, the pygidial gland appears
to be the most frequent in occurrence. Moreover, in several species
its secretions have been found to serve as pheromones (Holldobler
et al 1976; Holldobler and Haskins 1977; Maschwitz and Schonegge
1977; Kugler 1979; Holldobler and Traniello 1980 a,b; Traniello and
Jayasuriya 1981). The pygidial gland seems to be homologous with
the “anal glands” of the dolichoderine ants described by Pavan and
Ronchetti (1955). As we pointed out previously (Holldobler and
Engel 1978) the term “anal gland” is misleading, because the gland
does not exit from the anal or cloacal opening of the gaster, but
between the 6th and 7th abdominal tergites. We therefore suggested
to refer the dolichoderine structure to the pygidial gland. Recently
Jessen and Maschwitz (in press) proposed to name the pygidial
gland in honor of its discoverer Charles Janet. Thus we have now
three names for this tergal gland: anal gland, pygidial gland and
Janet’s gland.

Because the anatomical designation of the organ in ant workers (a
criterion we prefer) has been used in several recent publications, we
will continue to call the tergal gland opening between the 6th and
7th abdominal tergites pygidial gland.

Table 1. (Following pages) List of species that were investigated
histologically, and of the types of tergal and sternal glands found. When
the histological series was incomplete and we could not make a definite
statement, or when we could not clearly identify glandular ducts, the
column is marked with a “?”. r=with reservoir; c=with cuticular
structure.

TABLE 1

Subfamily / species

Collector and Locality

M YRMECIINAE
Myrmecia pilosula

PONERINAE

B. Holldobler, Brindabella Ranges,
Australia

Diacamma australis
Ectatomma ruidum

B. Holldobler, Townsville, Qld.,
Australia

J. Traniello, BCI, Panama

Ectatomma tuberculatum

J. Traniello, BCI, Panama

Leptogenys diminuta

B. Holldobler, Kuranda, Qld., Australia

Pachycondyla apiacalis
Pachycondvla obscuricornis

J. Traniello, BCI, Panama
J. Traniello, BCI, Panama

Paltothvreus tarsatus

B. Holldobler, Shimba Hills, Kenya

Rhvtidoponera metallica
DORYLINAE

B. Holldobler, Brindabella Ranges,
Australia

Eciton

A. Aiello, R. Silberglied, BCI, Panama

Neivamvrmex

A. Aiello, R. Silberglied, BCI, Panama

PSEUDOMYRMECINAE
Pseudomvrmex pallidus

P. Ward, Texas, USA

MYRMICINAE
Catalacus intrudens
Leptothorax ( Macromischa)
alardvcei

B. Holldobler, Shimba Hills, Kenya
B. Cole Florida Keys, USA

Novomessor a/bisetosus

B. Holldobler, Arizona, USA

Novomessor cockereUi

B. Holldobler, Arizona, USA

Orectognathus versicolor
Pogonomyrmex barbatus

B. Holldobler, Eungella, Queensland,
Australia

B. Holldobler, Arizona, USA

NOTHOM YRMECIINAE

Nothomvrmecia macrops

R. W. Taylor, Eyre Peninsula, Australia

ANEURETINAE
Aneuretus simoni
DOLICHODERINAE

Anula Jayasuriya, Sri Lanka

Iridomyrmex purpureus

B. Holldobler, Canberra, Australia

Liometopum apiculatum

B. Holldobler, Arizona, USA

FORMICINAE
Formica perpi/osa
Mvrmecocystus mendax
Oecophvlla longinoda

B. Holldobler, Arizona, USA
B. Holldobler, Arizona, USA
B. Holldobler, Shimba Hills, Kenya

Intersegmental tergal glands Intersegmental sternal glands

IX VIII VII VI V IV IX VIII VII VI V/ IV

VIII VII VI V IV III VIII VII VI V IV III

+ +
r

9

9

r

+

r,c

+

r,c

9

r,c

+

r,c

+

+ + +

r

+ +

+

+

r

TABLE ! (continued)

Subfamily 1 species

MYRMECIINAE
Mvrmecia pilosula

Other tergal glands

PONERINAE
Diacamma austra Us

Ectalomma ruiclum

Ectafomma tuberculatum

Eeptogenys dim inula

Pachycondyla apiacal is
Pachycondyla obscuricornis

glandular cells in 7th and 8th segment
ducts open dorsally into genital chamber

Pa It othyrei is tar sat i is
Rhytidoponera metaUica

IXth tergite; ducts open into genital
chamber

DORYLINAE

Eciton

lllrd

Neivamyrmex

PSEUDOMYRMECINAE
Pseudomyrmex pallidus

MYRMICINAE
Catalacus intrudens
Leptot borax (Macromischa)
alardycei

lllrd

N o \ 'omessor alb isetosus

Novomessor cockerel! i

Orectognathus versicolor

Pogonomyrmex bar bat us

NOTHOM YRMF.CIINAE
Not homy rmecia macrops

ANEURETINAF,

A new el us simoni
DOLICHODFRINAF.
Iridomyrmex purpureus

l.iometopum apiculatum

FORMICINAF
Formica perpilosa
Myrmecocystus mendax
Oecophylla bnginoda

postpetiole gland opens between lllrd
tergite and postpetiole
postpetiole gland

Other sternal glands

Tergo-sternal

glands

Sub-

genital

plate Penis Anus

gland gland gland

VUIth

+

+

Between 4/5; 5/6; +

6/7 segments

Between 4/5; 5/6; +

6/7 segments

Between 4/ 5;5 / 6; +

6/7 segments

glandular cells in petiole; ducts +

open ventrallv through cuticle

VII Ith ? +

Vlllth ? +

Between 4/5; 5/6; ?

6/7; 7/8 segments

9 9

9

9

IMrd Vlllth IXth

+

+

Illrd Vlllth IXth

+

+

+ + +

9

9

122

Psyche

[Vol. 89

As mentioned before Janet found this gland not only in workers
of M. rubra but also in males. Ant males differ from the workers in
having one more exposed segment (8th segment); often even part of
the 9th segment is visible. Thus in ant males the pygidial gland does
not open at the last exposed tergite (Fig. 1).

As can be seen from tab. 1 we found a pygidial gland in species of
the subfamilies Myrmeciinae, Ponerinae, Dorylinae, Pseudomyr-
mecinae, Myrmicinae, Nothomyrmeciinae and Dolichoderinae. In
Aneuretus simoni (Aneuretinae) we detected a few glandular cells,
but we could not clearly see glandular ducts. In the males, as in the
workers, there exists a considerable variation in the morphology of
the pygidial glands, even within a single subfamily. In some species
large clusters of glandular cells are associated with a special cuticu-
lar structure on the 7th tergite (Fig. 1, 2, 6c). Some species possess
more or less developed reservoirs, composed of an invagination of
the intersegmental membrane (Fig. 3). In other species there are
only a few glandular cells that send dorsolaterally ducts through the
intersegmental membrane.

In ant males a post pygidial gland is almost as common as the
pygidial gland (Fig. 1). It also consists of paired clusters of glandu-
lar cells that open through the intersegmental membrane, but
between the 7th and 8th tergites (Tab. 1). This gland is especially
well developed in Nothomyrmecia macrops males, where the inter-
segmental membrane forms a large reservoir. There it closely
resembles the pygidial gland found in workers. Interestingly, the
males of this species have only a few glandular cells between the 6th
and 7th tergites. Thus in comparison with Nothomyrmecia females
(including queens) the major tergal gland in the males is shifted one
segment posteriorly. In most other ant species examined, however,
the gland between the 6th and 7th tergites (pygidial gland) is the
major tergal gland in both workers and males.

The doryline males are a remarkable exception. They, too, have
large pygidial glands, consisting of paired complex glands and a
large reservoir. But the same structure is present in the next 3 seg-
ments anteriorly (6th-5th; 5th-4th; 4th-3rd) (Fig. 4, 5). In addition,
intersegmental glandular cells were found between the 7th and 8th
tergites. In the 3rd tergite we also found paired groups of glandular
cells, the ducts of which penetrate the sclerotized cuticle of the 3rd
tergite dorsolaterally.

1982] Holldobler & Engel- Siegel — Glands in Male Ants 123

Fig. 3 Longitudinal section through pygidial gland of Pogonomyrmex barbatus
S- GC=glandular cells; R=reservoir.

Sternal glands:

More than in any other subfamily, the males of the doryline ants
are also richly endowed with sternal glands (Tab. 1; Fig. 4). We
found major complex glands with intersegmental reservoirs between
the 3rd and 4th, 4th and 5th, 5th and 6th sternites. Glandular clus-
ters are also present between the 6th and 7th sternites — but without
a pronounced reservoir. In addition, clusters of glandular cells
whose ducts penetrate the sclerotized cuticle were found in the 3rd
sternite, they are also strongly developed in the 8th and 9th sternites
(Fig. 4). Similar sternal glands were found in the 8th sternite of
males of Myrmecia pilosula, Pachycondvla apiaealis and P. obscuri-
cornis. In both Pachycondvla species the glandular ducts open in
bundles into cuticular cups located in the 8th sternite (Fig. 7a).

In the ponerine species Leptogenys diminuta the males possess a
huge intersegmental sternal gland between the 7th and 8th sternites.
This gland consists of large paired clusters of glandualr cells. Each
cell sends a duct into wider collecting channels which lead into a
paired large reservoir, consisting of ventro-lateral invaginations of
the intersegmental membrane (Fig. 6). There is a second paired
sternal gland between the 8th and 9th sternite; but this gland is

124

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[Vol. 89

A

Fig. 4 A. Schematic drawing of a longitudinal section through the gaster of a
Neivamyrmex spec. illustrating the segmental glandular structures. B. Longitudi-
nal section through a intersegmental complex-gland (between IVth and Vth tergites).
C. Longitudinal section through a intersegmental sternal gland of Eciton spec.
Q. A=anus; GC=glandular cells; P=part of penis with penis gland; R= reservoir.

1982] Holldobler & Engel- Siegel — Glands in Male Ants

125

Fig. 5 A. SEM micrograph of the tergite with the attached intersegmental mem-
brane of a Neivamyrmex B. Larger magnification showing clearly the glandular
duct openings in the cuticle (GO) and the intersegmental membrane which consists of
a mat of bristle-like structures. This mat-membrane forms the intersegmental glandu-
lar reservoir. See also Fig. 4B.

126

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[Vol. 89

much smaller, and no pronounced reservoir could be detected. Lep-
togenvs workers have two well developed sternal glands between the
5th and 6th, and the 6th and 7th sternites respectively (Holldobler
and Engel 1978; Jessen et al 1979). Leptogenvs males lack these
structures but do possess sternal glands in the 7th and 8th sternites.
In males of the ponerine ant Palt hot hy reus tarsatus we found large
paired clusters of glandular cells in the 8th sternite. The glandular
ducts open through the intersegmental membrane between the 8th
and 9th sternites. In addition Paltothvreus males possess unpaired
intersegmental sternal glands, similar to those found in Paltothvreus
workers (Holldobler and Engel 1978), but smaller, between the 5th
and 6th, and 6th and 7th sternites.

Other abdominal glands;

As indicated in table 1 we found several other abdominal glands
in males which cannot directly be assigned to the group of tergal or
sternal glands. In a few species ( Novomessor , Leptogenvs) we
detected glandular cell clusters in the petiole. In Ectatomma, Dia-
camma, Paltothvreus we found small tergo-sternal glands. The
ducts of the glandular cells composing them open laterally through
the pleural membrane. We found similar small glandular cell
bunches in males of Pachycondvla and Rhvtidoponera, but we
could not clearly identify the glandular ducts. These tergo-sternal
glands resemble closely similar structures described by Jessen et al
(1979) in workers of several ponerine species. Finally we found
small groups of glandular cells directly at the anus of males in
Pachycondvla, Ectatomma, Neivamyrmex, Eciton, Mvrmecocystus
and Liometopum (Fig. 7b). These anus glands vary considerably in
size, and it is possible that they are present in more species than we
were able to document. We first found them in workers of Dorv/us
(Holldobler and Engel 1978). The anus glands should not be mis-
taken for the rectal gland, an invagination of a glandular epithelium
of the rectum, recently discovered in Oecophvlla workers by Holl-
obler and Wilson (1978). It is interesting to note that males of
Oecophvlla also possess a small rectal gland.

Discussion:

Except for the glands associated with the ovipositor and sting
apparatus, which the males lack, ant males are as richly endowed
with exocrine glands as the females. In many species of ants the
males have well developed mandibular glands, pro- and post-

1982] Holldobler & Engel-Siegel — Glands in Male Ants 127

Fig. 6 A. Longitudinal section through gaster of a Lepidgenys ciiminuta show-
ing the reservoir ( R) of the large sternal gland between 7th and 8th sternites. A=anus.
B. Section through the large cluster of glandular cells of the sternal gland, opening
into the reservoir between 7th and 8th sternite. S=secretion in reservoir. C. Longi-
tudinal section through the pygidial gland of Leptogenys ciiminuta Note that the
glandular cells (GC) are considerably larger than the glandular cells of the sternal
gland (Fig. 6B). CS=cuticular structure

128

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[Vol. 89

pharyngeal glands, maxillary glands, salivary glands, and meta-
pleural glands, although the size of these various structures can vary
considerably between the female castes and males. In this paper we
surveyed specifically the abdominal sternal and tergal glands in ant
males.

In almost all species studied we encountered two major glandular
structures that Janet (1907) had already described in Myrmica
rubra, the penis glands and the subgenital plate glands. Also quite
generally present in males (except in the Formicinae) are the pygi-
dial glands. The males share these organs with the females, although
less well developed in some species. An interesting case is Notho-
myrmecia: here the males have a rudimentary pygidial gland but a
weil developed postpygidial gland (between the 7th and 8th tergites).

The males of the doryline ants are unusually well endowed with
abdominal glands, in which they differ markedly from the workers.
Although doryline workers have well developed pygidial- and post-
pygidial glands (Holldobler and Engel 1978), the males have mas-
sive glandular structures in each segment. In this context the
findings by Whelden (1963) are of considerable interest. Whelden
described a series of exocrine glands in the gaster of Eciton queens
as follows: “Each of the segments of the gaster, including those
telescoped together in the posterior part, contains a pair of these
glands which are smaller in the anterior segment than those in the
following segments”. We were not yet able to section a doryline
queen and therefore cannot compare the queen organs with those
we found in males. It appears, however, that the males possess a
glandular equipment very similar to that of the queens. Presumably
in doryline queens these massively developed exocrine glands play
an important role in the queen’s chemical control of the worker ants
and in her high attractiveness to workers, (Watkins and Cole 1966).
We hypothesize that the males imitate queen pheromones, which
might enable them to penetrate a foreign colony in order to get
access to the wingless virgin female reproductives (Franks and Holl-
dobler unpublished). In fact, this might also be the function of the
massively developed sternal gland in Leptogenvs males. In this genus,
as in the dorylines ergatoid reproductive females presumably mate
in the nest, so that males flying in from other nests have to penetrate
a foreign colony.

1982] Holldobler & Engel-Siegel — Glands in Male Ants 129

Fig. 7 A. Longitudinal section through sternal gland in 8th sternite of a Pachy-
conciyla apiacalis D=glandular duct; CU=cuticular cup. B. Longitudinal sec-
tion through 8th tergite of a Ectatomma ruidum A=anus; AG=anus gland.

130

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[Vol. 89

But these are speculations. We know nothing about the function
of the abdominal glands in ant males. Jessen and Maschwitz (in
press) suggested that some of the numerous intersegmental glands
they discovered in workers of the ponerine Pachyeondyla tridentata
might function as lubrication glands, reducing the friction between
the segments when the workers bend the gaster during the act of
stinging. In ant males some of the intersegmental glands could pro-
duce lubricants in order to keep the abdomen flexible during mating
behavior or to assist the extrusion of the copulatory apparatus. On
the other hand some of the well developed tergal and sternal glands
seem almost certainly to produce allomones or pheromones. The
recent morphological investigations of glandular structures in ants
have opened a new phase in the study of chemical communication in
ants.

A cknowledgments:

We would like to thank all the collectors mentioned in Table 1,
and W. L. Brown, R. Snelling, R. W. Taylor for helping us with the
identification of many species, and Ed Seling for his assistance
during the SEM work. This work was supported by NSF grant
BNS80-02613.

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1955 Studi sulla morfologia esterna e anatomia interna dell’ operaia di Irido-
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mecina. Atti Soc, Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano 94: 379-477.

132

Psyche

[Vol. 89

Traniello, J. F. A. and A, K. Jayasuriya

1981 Chemical communication in the primitive ant Aneuretus simoni: The
role of the sternal and pygidial gland. J. Chem. Ecol. 7: 1023-1033.
Watkins, J. F. and T. W. Cole

1966 The attraction of army ant workers to secretions of their queens. Texas
J. Sci. 18: 254-265.

Weber, H.

1954 Grundriss der Insektenkunde, Gustav Fischer Verlag, Stuttgart.
Whelden, R. M.

1963 The anatomy of adult queen and workers of the army ants Eciton bur-
chelli Westwood and Eciton hamatum Fabricius. New York Entomol.
Soc. 71: 90-1 15.

TERMITE-TERMITE INTERACTIONS:
WORKERS AS AN AGONISTIC CASTE*

By Barbara L. Thorne
Museum of Comparative Zoology
Harvard University
Cambridge, Massachusetts 02138

Termite soldiers are a defensive caste. Their heavily sclerotized
head capsules can be equipped with hard mandibles capable of
crushing, pinching, piercing, or slashing predators. Soldier castes of
many phylogenetically advanced species have well-developed fron-
tal glands and are capable of exuding or spraying chemical secre-
tions. Such chemical armaments are toxic, irritable, or oily fluids
which can impair physiological, sensory and/or mechanical facul-
ties of the recipient (Prestwich, 1979). Termite soldiers are thus
formidable opponents for ants and vertebrate predators. Soldiers are
fed by workers and their behavior within the colony is generally
limited to signaling alarm, participating in defense, and organizing
foraging expeditions (Stuart, 1969; Traniello, 1981).

Despite their specialization, however, soldiers are not the only
defensive caste in a termite colony: worker termites of some species
(from four families) are known to be able fighters in termite-termite
aggressive interactions [Kalotermitidae (Grassi and Sandias, 1 896—
1897; Dropkin, 1946); Hodotermitidae (Nel, 1968); Rhinotermitidae
(Pickens, 1934; Clement, 1978); Termitidae (Dudley and Beaumont,
1889a, b; Andrews 1911)]. This paper explores intra- and interspe-
cific agonistic encounters among termites, and focuses on the roles
of workers and soldiers in such conflicts.

The report is presented in two sections, corresponding to two sets
of experiments on this topic. Section A describes a field manipula-
tion inducing intraspecific encounters among colonies of Nasuti-
tennes corniger in Costa Rica. Section B presents data on
laboratory experiments examining intra- and interspecific interac-
tions among four species of Panamanian termites.

* Manuscript received by the editor February /, 1982

133

134

Psyche

[Vol. 89

A. Intraspecific Field Experiment: Nasutitermes corniger

Nasutitermes corniger (Motschulsky) is a common arboreal ter-
mite ranging through much of Central and northern South America
(Thorne, 1980). Large carton nests contain up to 800,000 termites
(Thorne and Noirot, 1982) and, with the addition of foragers, total
colony size may exceed a million individuals. Distinct foraging
trails, covered by carton galleries, are visible issuing from a nest and
proceeding along tree branches, trunks, and the ground surface. The
termites also travel underground and in galleries located within trees^
or fallen logs. N. corniger foraging trails can radiate many meters
from the parent colony.

Given the density of N. corniger colonies in primary forest (7.0 ±
1.8 per hectare in the Hubbell Plot of Barro Colorado Island,
Panama, N = 4 hectares) and in areas of young second growth (27 in
one hectare in Frijoles, Panama) (Thorne, unpub. data), it is likely
that, at least occasionally, foraging parties from different colonies
encounter one another in the midst of exploring or exploiting a local
food source. Observation of a natural inter-colony encounter would
be difficult. It would require tracking single foraging trails, which
would undoubtedly result in disturbance as one cleared away the
forest litter to locate foragers. Even if trails could be accurately
followed without disruption, it would be rare to view simultaneous
interception with an active trail known to be from a second colony.
Because the odds of witnessing such a natural event are low, I forced
an encounter through a transplant experiment.

METHODS

On the morning of 18 August, 1978 three Nasutitermes corniger
nests were collected from separate areas of second growth near
Sirena headquarters of Corcovado National Park, Osa Peninsula,
Costa Rica. The nests measured 29.8, 52.7, and 41.9cm in height
and 26.0, 29.8, and 26.0cm in diameter, respectively. Nests were
sawed from their host trees and hand-carried to the experimental
site. Each colony was suspended on a wire from one of two branches
of a large tree (Fig. 1). The nests were hung in an equilateral triangle
such that the distance from their base to the ground was 60cm, the
distance from the edge of each colony to each neighbor was 50cm,
and the original compass orientation of each colony was main-

19821 Thome — Termite-Termite Interactions 135

Figure 1. Field set-up for intraspecific encounter experiment involving Costa
Rican Nasutitermes corniger.

tained. A coat of tanglefoot was placed at the base of each wire
strand (near the point of attachment to nest support branches) to
prevent termites from crawling up the wires. A 3.5 X 3.5 m2 plot
beneath the nest triangle was completely cleared of leaf litter, forest
debris and herbaceous plants so that movement patterns of the
termite trails could be monitored. At 7:30 p.m. that evening stick
“ramps” (90cm in length, 1 cm in diameter) were installed to connect
the nests to the ground. The tips of the sticks were shallowly inserted
into both the nest carton and ground surface for support. Bases of
the ramps also ended in vertices of an equilateral triangle on the
ground, 20cm from tip to tip.

RESULTS

Hanging above the ground from a single strand of wire, each nest
was an island in mid-air: no escape routes were open for the ter-
mites. By dusk of the day of collection, soldiers and workers from
all colonies were crawling over the peripheries of their nests. This
activity gained participants and momentum: at 7:30 p.m. each nest
was a seething mass of termites. The stick ramps were embedded to

136

Psyche

[Vol. 89

connect the nests to the ground. Immediately following implanta-
tion termites swarmed onto the ramps, soldiers in the lead followed
by a mixture of soldiers and workers. Only four white immatures
were seen leaving the nests throughout the experimental period.

As they reached the ground groups fanned out: termites from a
single colony divided into several ribbons heading in different direc-
tions. Because of the close proximity of the ramp exits, it was inevi-
table that encounters occur between trails from different colonies.
The meetings were not passive. Soldiers oriented towards (and
apparently squirted) termites from other colonies, but this did not
seem much of a deterrent to recipients. The major defense stemmed
not from the soldiers, but from the Nasutitermes workers.

Workers from different colonies grabbed each other with their
mandibles and locked in one-on-one conflict. Pairs of workers
squirmed and bent with vigor, often until the death of both. Occa-
sionally a third or fourth worker would join the engagement, but
usually only temporarily. Workers avidly attacked soldiers as well,
grabbing at the legs and occasionally biting the abdomen.

The next morning worker carcasses littered the arena over an area
of approximately 1,500cm2, with some battle “patches” as far as
1.5 m from the center of the ramp triangle. The density of bodies was
often quite high (25-50 dead in a 4.0cm2 area). Surviving termites
did not appear to cannibalize the dead. Ants, flies, staphylinid bee-
tles, and wasps began scavenging the termite carcasses.

How were foraging trail routes influenced by the intersection bat-
tles? Agonistic confrontations were instigated when at least one
colony was in the process of establishing or changing a foraging
route, i.e. in a scouting phase. Preliminary observations (Fig. 2)
suggest that both colonies’ foraging pathways were displaced by
encounters — trails were repulsed from the meeting site following
battles lasting 10-30 minutes. One colony may maintain a trail
tangential to the “battle field”, but I did not see one continuing
through an area of dead termites.

It is difficult to determine the effect of agonistic encounters on
final foraging path location. In isolation a colony establishes forag-
ing routes by scouting in a broad network, but several hours later
this highly branched fan collapses into a single actively travelled
ribbon with few side trails. Thus the fact that termites have trav-
ersed a given area in no way assures that route as the path of a final

1982]

Thorne — Termite-Termite Interactions

137

.m

10-20 am
8/20/78

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I

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•rti

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8/20/78

5

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6

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Figure 2. Periodic maps of ground foraging trails departing from the tips of exit
ramps attached to suspended Nasutitermes corniger colonies (Section A). Central
dots indicate positions of the ramps. Maps are 3.5 X 3.5 m2.

138

Psyche

[Vol. 89

foraging trail. Battles may be a deterrent to formation of a given
path, but their influence is difficult to assess independently.

Foraging path trajectories from the three nests were monitored
for two and one half days following initiation of the experiment (see
Fig. 2 for the final 24 hour period). Once a scouting fan condensed
into a single pathway the positions were relatively stable. Minor
adjustments in path locations did occur periodically, and activity on
specific trails varied from day to day and even hour to hour. Occa-
sionally (usually in the evening) new scouting parties would emanate
from the ramp tip or as a tributary of the main trail network.

Construction of trail covers varied from colony to colony. Colony
III began covering both its ramp and ground trails quickly (a total
of 64cm of trail covered by 7 am, 20 August). In contrast. Colonies I
and II had only 19cm and 3.2cm of covering, respectively, at 7 a.m.
on 20 August. These same relative speeds were repeated when the
initial experiment was replicated from 21 to 22 August. Building
behavior also showed distinct inter-colony variation, mainly in the
amount of advance siding deposited before the trails were roofed.

To replicate the first nights’ encounters, I removed the three
ramps and scraped clean the entire 3.5 X 3.5 m2 grid at noon on 21
August. This caused attrition of those individuals on the ground and
out foraging, but the established trails had to be destroyed to induce
active scouting. New ramps (90cm long) were installed at 6:30 p.m.,
with ends touching the ground in an equilateral triangle of side
length 25 cm.

Members of Colony I came down their ramp fairly rapidly and
began three major paths from its tip, one to the southeast, one to the
north, and one due west. At 7 p.m. termites from Colony II began
coming down their ramp and immediately began to fight with Col-
ony I’s southeast-bound foragers. When the first workers from Col-
ony III came down their ramp and encountered foreigners, they
rapidly reversed direction and returned en masse to the mouth of
their nest, after which a large group of Colony III termites stormed
down the ramp. The possibility of worker recruitment in these cir-
cumstances should be investigated. The battle between Colonies I
and III was vehement for 20 minutes; after 30 minutes Colony I’s
southeast trail was abandoned. Colony III established a new trail
180° away from the direction of original interference with Colony I.

1982]

Thome — Termite-Termite Interactions

139

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Psyche

[Vol. 89

In these and several other encounters that night, aggressive interac-
tions among colonies were similar to those described earlier.

B. Intra- and Interspecific Laboratory Experiments:
Termite-Termite Interactions Among Four
Panamanian Species

To examine conflicts among colonies and species of sympatric
termites I staged laboratory encounters in pairwise tests: soldiers vs
soldiers, soldiers vs workers, and workers vs workers. Characteris-
tics of the four species used in these experiments are summarized in
Table I. Amitermes beaumonti soldiers have curved mandibles;
Armitermes chagresi soldiers have long, curved mandibles and a
prolonged nasus; and soldiers of Nasutitermes corniger and N.
ephratae are nasutoid with vestigal mandibles (Fig. 3).

METHODS

Experimental trials were conducted in March and April, 1981 in
the Smithsonian Tropical Research Institute laboratory on Barro
Colorado Island (BCI), Panama (9° 09' N, 79° 51' W). All Ami-
termes beaumonti and Armitermes chagresi were collected from
nests on BCI; samples of Nasutitermes corniger and N. ephratae
were from colonies in Frijoles, Panama (4km east of BCI). Pairwise
encounters were staged in petri dish arenas (4.6cm diameter) lined
with moist filter paper. Equal numbers of termites, soldiers or
workers, from each colony were introduced to an arena simultane-
ously. Dishes were then covered and left undisturbed in darkness for
12 hours. After the interaction period survivors were counted: con-
spicuously injured individuals were considered as dead. Most
worker-worker trials involved 50 individuals from each colony (only
large workers ($) were used from Nasutitermes colonies; Amitermes
and Armitermes have monomorphic worker castes). When soldiers
of Amitermes or Armitermes were involved in a trial, and in occa-
sional trials involving Amitermes or Armitermes workers, fewer
individuals were available so experiments proceeded with less than
50 termites from each colony. In Table II the number of individuals
from each colony used in each trial is indicated in parentheses fol-
lowing the survival percentages. A minimum of three trials were
conducted for each intra- and interspecific interaction. Each such

1982]

Thorne — Termite-Termite Interactions

141

Figure 3. Termite-termite interactions (Section B). a. Nasutitermes corniger
workers (dark heads) vs Armitermes chagresi soldiers (light heads), b. N. corniger
soldiers vs Amitermes foreti workers, c. N. corniger intraspecific worker-worker
encounter, d. Amitermes fore/i soldier vs N. ephratae worker, e. Armitermes cha-
gresi soldier vs N. ephratae worker.

142

Psyche

[Vol. 89

trial pitted termites from different colonies. After the 12 hour
encounter all dead termites from selected trials were collected and
examined for injuries under a dissecting microscope.

RESULTS

Survival percentages of termites involved in each trial are pre-
sented in Table II. For conspecific interactions among members of a
single caste (soldier vs soldier or worker vs worker), it was not
possible to differentiate colony affiliation so a single survival per-
centage is indicated. These figures indicate whether or not a fight
ensued, although it is impossible to determine if one colony suffered
more or less mortality than the other.

Variability within and between blocks of Table II is high. Among
some replicates mortality is low for both groups of interacting ter-
mites (signified by a at the base of the block). Some encounters
suggest consistent “victors”, represented by an arrow pointing in the
direction of that party. Other groups of interactions indicate agonis-
tic behavior on both sides (‘+’), without clear assignment of a
“winner” or “loser”.

All interspecific worker-worker encounters resulted in a fight, often
with a trend suggesting a “dominant” species but with sufficient
variation among trials to prevent assigning a “winner”. Such varia-
tion may result from relative differences in individual colony nutri-
tion, age, health, and history. For example, an interaction between
two strong colonies may be quite different from a similar encounter
between members of a weak and a strong colony. It should be noted
that soldiers are absent during worker-worker trials, which may
affect the excitability and response of workers.

Intraspecific worker-worker engagements demonstrated variable
aggression within Nasutitermes corniger and N. ephratae, and no
lethal attacks in any of the Amitermes or Armitermes trials. Fight-
ing among conspecific Nasutitermes colonies is variable and appar-
ently influenced by as yet uninvestigated factors. In these experi-
mental trials, aggressive interactions generally occurred, although in
all but one 1 V. corniger trial well over half of the workers survived
the 12 hour meeting. I have previously observed both extremes in
conspecific Nasutitermes corniger encounters: 100% mortality and
100% survival, even among colonies from distant locations. Dudley
& Beaumont (1889a,b) report that mixing two N. corniger colonies

1982]

Thorne — Termite-Termite Interactions

143

resulted in lethal fights. Variance in response may be due to experi-
mental protocol, particularly isolation of a colony’s soldiers and
workers. Under natural conditions a colony’s soldiers and workers
may interact with one another in recognition of and response to
foreign termites. The soldier secretion has been demonstrated an
alarm pheromone in N. exitiosus, although workers showed little
reaction to fresh secretion presented on an applicator (Eisner et al.
1976).

In these experiments soldiers and workers from different N. cor-
niger or Armitermes chagresi colonies did not fight, while soldier-
worker conflict did occur in N. ephratae and Amitermes beaumonti
conspecific encounters. In the field manipulaton involving Costa
Rican N. corniger (Section A), soldier-worker battles were observed.

Summarizing other general trends, N. ephratae workers scored
relatively well in worker-worker inter-specific encounters, although
they were not consistent victors over Armitermes workers. Both
Amitermes and Armitermes soldiers faired relatively well in most
encounters while Nasutitermes soldiers were less successful. In intra-
specific Armitermes chagresi interactions, only soldier-soldier con-
flict was observed; among Amitermes beaumonti, only meetings between
soldiers and workers stirred fighting. Such patterns imply species
differences in communication, meaning, and recognition of any
colony-specific odors.

Injuries suffered by the dead during the interaction experiments
were scored for several trials, and are summarized in Table III.
Presence or absence of damage to the abdomen was scored, though
no analysis of extent of abdominal injury was recorded because the
exact number of wounds or punctures was difficult to assess. The
percentage of dead with abdominal wounds is generally high.

Table II (Following pages): Survival Percentages of Panamanian Termites in

Paired Laboratory Encounters

Paired encounters were staged matching equal numbers of termites (number of individu-
als from each colony given in parentheses following trial results). The total percentage of
individuals surviving the 12 hour meeting is given for intraspecific soldier-soldier and
worker-worker interactions: inability to identify colony affiliation prevented comparative
percentages. All other trials report the survival percentage of the termites listed to the left
over that of termites listed on top. Arrows at the base of a block point in the direction of a
consistent “winner”: arrows in parentheses note a less pronounced tendency,
indicates few deaths on either side (no fight): '+' signifies lethal interactions among
the termites but with no consistent trend toward a victor.

Table 11: Survival Percentages of Panamanian Termites in Paired
Laboratory Encounters

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146

Psyche

[Vol. 89

Table 111: Analysis of Injuries for 7 Paired Worker-Worker Interactions

Percentage

of

Individuals

with

Abdominal

Wounds

Antennal

Legs Heads

Mean Number
of

Non-Abdominal
Wounds Per
Individual

A rmitermes chagresi ( 10 of 12)

100%

5

6

1

1.20

vs

Nasutitermes ephratae ( 16)

87.5%

7

6

2

0.94

Armitermes chagresi (20 of 21)

3

13

3

1.00

vs

Nasutitermes ephratae (43)

—

41

61

0

2.37

Nasutitermes ephratae (8)

87.5%

2

6

1

1.12

vs

Nasutitermes corniger (41)

78.0%

30

125

1

3.83

Nasutitermes ephratae (12)

100%

3

13

0

1.25

vs

Nasutitermes corniger (43)

90.7%

27

169

0

4.56

Amitermes beaumonti (32 of

38) 65.6%

36

32

20

2.75

vs

Nasutitermes ephratae (46)

82.6%

39

46

0

1.85

Nasutitermes corniger (19)

73.7%

12

25

1

2.05

vs

Amitermes beaumonti (38)

50.0%

24

35

19

2.05

Nasutitermes corniger (37)

67.6%

47

113

0

4.32

vs

Amiterme (38)

50.0%

24

35

19

2.05

Nasutitermes corniger (37)

67.6%

47

113

0

4.32

vs

Amitermes beaumonti (39)

35.9%

28

53

16

2.51

Numbers in parentheses following species names are the number of individuals exam-
ined (killed). The colony with the fewest deaths is listed first for each interaction. Descrip-
tions of the injury categories and criteria are described in text Section B under Results.

1982]

Thorne — Termite-Termite Interactions

147

Damage to each antennae and leg was scored separately, and a
pierced or decapitated head was scored as one head injury, even if
multiple punctures were present. Thus for 10 dead termites, a max-
imum of 20 antennal, 60 leg, and 10 head injuries were possible.
That number of cumulative injuries divided by the total number of
termites examined yielded the mean number of non-abdominal
injuries per individual. This index gives some indication of the
intensity of attack.

The frequency of antennal, leg, and head injuries shows that Ami-
termes beaumonti heads appear relatively vulnerable (at least when faced
by Nasutitermes ), and that N. corniger was prone to numerous leg
injuries.

DISCUSSION

Worker termites of some species join in aggressive encounters
with members of other colonies and are often adept fighters, partic-
ularly against other workers. Intensity and outcome of agonistic
encounters is variable depending on the species, colonies, and
castes involved. Soldiers also participate in termite-termite conflicts.
This study suggests that mandibulate soldiers are more effective
one-on-one inter-specific antagonists than are Nasutitermes sol-
diers, although nasute soldiers are generally present in higher
numbers per colony and may be more effective in groups.

Research on termite-termite encounters has not been extensive,
but is of interest because the recognized defensive caste, the soldiers,
are not the sole participants, and may not join in such interactions
at all. After viewing the attack of an introduced Termes worker by
nymphs and larvae in a Calotermes colony, Grassi and Sandias
write in their 1896-1897 account (p. 283), . .similar observations

have been made several times, and show. . .that the soldiers purpose-
fully reserve themselves for more important foes.” Pickens (1934)
noted that workers of established Reticulitermes hesperus colonies
will attack and kill founding pairs and incipient colonies which
settle nearby.

Andrews (1911) did an extensive series of intra-specific interac-
tion experiments with Nasutitermes ripper tii. Nearly all staged
encounters resulted in immediate conflict involving both soldiers
and workers, although occasionally two colonies were completely
docile towards each other. Andrews also came to the conclusion

148

Psyche

[Vol. 89

that “different communities (colonies) have different grades of hos-
tility” (p. 218). Studying Coptotermes acinaciformis, Howick and
Creffield (1980) similarly report inter-colony variance in degree of
aggression.

The mechanism by which termites recognize non-colony mem-
bers, and the associated stimuli for aggressive response, are not
understood. Termites of some species quickly discriminate between
colony-mates and foreigners; other species seem oblivious to “ali-
ens”. Certain treatments have been shown to interfere with recogni-
tion or aggressive behaviors: water washes (Andrews 191 1), chilling
termites to immobilization (Dropkin 1946; Howick & Creffield
1980), and laboratory rearing (Nel 1968). Unpigmented immatures
of some species appear immune from attack (Andrews 1911, Sands
& Lamb 1975). Dudley and Beaumont (1889a) postulated that col-
ony members bear like “odors”, and that they can thereby differen-
tiate colony-mates from foreigners. This idea of what is now termed
colony-specific recognition pheromones is still viable (reviewed by
Stuart 1970), perhaps involving cuticular hydrocarbons as recogni-
tion cues (Howard et al. 1978, Blomquist et al. 1979). The degree of
intraspecific pheromone variation, the environmental components
of response, and the mode of aggressive stimulus remain unknown.

Workers may be particularly useful as a capable, defensive unit in
termite-termite encounters because they are the numerically domi-
nant caste and although they are accompanied by soldiers on forag-
ing forays, they are vulnerable to predation and competition while
foraging. Termite mounds and arboreal nests have few exposed
openings, and any which exist are guarded, usually by soldiers. In
contrast, worker foragers cannot rely on nest protection, and their
armada of soldier escorts may be insufficient to stay competitors.
Mandibulate soldiers are normally in low proportion relative to
workers (Haverty 1977). The ratio of termite soldiers to workers in
Nasutitermes is relatively high (Haverty 1977), but the soldiers have
vestigal mandibles and their chemical sprays are apparently not a
complete defense against other termites. Soldiers may be especially
proficient at repelling ant and vertebrate attacks (although workers
may assist, eg. Eisner et al. 1976), while worker castes are adept at
joining with soldiers to defend resources against other termites.

1982]

Thorne — Termite-Termite Interactions

149

ACKNOWLEDGEMENTS

I thank C. Justine Allen and Kent H. Redford for special assist-
ance, and the Organization for Tropical Studies and the Smithso-
nian Tropical Research Institute for logistical support. Kenneth P.
Sebens, James F. A. Traniello, and Edward O. Wilson read earlier
versions of this paper. This research was funded by NSF disserta-
tion improvement grant DEB-80-16415 and a predoctoral fellow-
ship from the American Association of University Women.

References

Andrews, E. A.

191 1. Observations on termites in Jamaica. J. Anim. Behav. 1: 193-228.
Blomquist, G. J.; R. W. Howard and C. A. McDaniel.

1979. Structures of the cuticular hydrocarbons of the termite Zootermospis
angusticollis (Hagen). Insect Biochem. 9: 365-370.

Clement, J.

1 978. L’agression interspecifique et interspecific des especes frangaise du genre
Reticulitermes (Isoptere). C. R. Acad. Sc. Paris. 286: 351 354.

Dropkin, V. H.

1946. The use of mixed colonies of termites in the study of host-symbiont
relations. J. Parasit. 32: 247-251.

Dudley, P. H. and J. Beaumont.

1889a. Observations on the termites, or white-ants of the Isthmus of Panama.
Trans. N. Y. Acad. Sci. 8: 85 1 14.

1889b. The termites or so-called “white-ants” of the Isthmus of Panama. J. N. Y.
Microscop. Soc. 5: 59-70 and 1 1 1-1 12.

Eisner, T.; I. Kriston and D. J. Aneshansley.

1976. Defensive behavior of a termite ( Nasutitermes exitiosus). Behav. Ecol.
and Sociobiol. 1: 83-125.

Grassi, B. and A. Sandias.

1896-1897. The constitution and development of the society of termites:
Observations on their habits: with appendices on the parasitic protozoa
of Termitidae, and on the Embiidae. Quart. J. Microsc. Sci. 39: 245-322;
40: 1-82.

Haverty, M. I.

1977. The proportion of soldiers in termite colonies: a list and a bibliography
(Isoptera). Sociobiology 2(3): 1 19-216.

Howard, R. W.; C. A. McDaniel and G. L. Blomquist.

1978. Cuticular hydrocarbons of the eastern subterranean termite, Reticuli-
termes flavipes ((Collar) (Isoptera: Rhinotermitidae). J. Chem. EcoL
4(2): 233-245.

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Howick, C. D. and J. W. Creffield.

1 980. Intraspecific antagonism in Coptotermes acinaciformis (Froggatt) (Isop-
tera: Rhinotermitidae). Bull, of Ent. Res. 70: 17-23.

Nel, J. J. C.

1968. Aggressive behaviour of the harvester termites Hodotermes mossambi-
cus (Hagen) and Trinervitermes trinervoides (Sjostedt). Ins. Soc. 15(2):
145-156.

Pickens, A. L.

1934. The biology and economic significance of the western subterranean ter-
mite Reticulitermes hesperus. In: Termites and Termite Control (C. A.
Kofoid, ed.) Ch. 14 pp. 157-183. Univ. of Calif. Press, Berkeley.

Prestwich, G. D.

1979. Chemical defense by termite soldiers. J. of Chem. Ecol. 5(3): 459-480.

Sands, W. A. and R. W. Lamb.

1975. The systematic position of Kaudernitermes. gen. n. (Isoptera: Termitidae,
Nasutitermitinae) and its relevance to host relationships of termitophi-
lous staphylinid beetles. J. of Ent. Series B Vol. 44 Pt. 2, pp. 189-200.

Stuart, A. M.

1969. Social behavior and communciation. In: Biology of Termites
(K. Krishna and F. M. Weesner, eds.). Vol. I pp. 193-232. Academic
Press, N. Y.

1970. The role of chemicals in termite communication. In: Advances
in Chemoreception Vol. I: Communication by Chemical Signals (J. W.
Johnson, Jr.; D. G. Moulton & A. Turk, eds.). pp. 79-106. Appleton-
Century-Crofts, N. Y.

Thorne, B. L.

1980. Differences in nest architecture between the Neotropical arboreal ter-
mites Nasutitermes corniger and Nasutitermes ephratae (Isoptera: Ter-
mitidae). Psyche 87: 235-243.

Thorne, B. L. and C. Noirot.

1982. Ergatoid reproductives in Nasutitermes corniger (Motschulsky):
Isoptera, Termitidae. International J. of Insect Morph, and Embryol., in
press.

Traniello, J. F. A.

1981. Enemy deterrence in the recruitment strategy of a termite: Soldier-
organized foraging in Nasutitermes costalis. Proc. Nat. Acad. Sci. 78(3):
1976-1979.

TYPE DESIGNATIONS AND SYNONYMIES FOR
NORTH AMERICAN SILPH IDAE (COLEOPTERA)

By Stewart B. Peck1 and Scott E. Miller2

The purpose of this paper is to provide type data and lectotype
and neotype designations for North American Silphidae described
by J. L. LeConte, Thomas Say, M. H. Hatch, and J. W. Angell, and
new synonymies of other species. We are engaged in ongoing revi-
sionary work on North American silphids (e.g. Miller and Peck,
1979) and have found considerable nomenclatural confusion be-
cause of varying interpretations of poor descriptions and names not
fixed to types. Publication of these data is also necessary for their
inclusion in the forthcoming fascicle on this family in “A Catalog
of the Coleoptera of America North of Mexico”, U.S. Dept. Agric.
Handbook 529 (J. M. Kingsolver, editor-in-chief). Improved identi-
fication keys and characterizations of all United States and Cana-
dian silphids are given in Peck (1982a).

In the LeConte and Horn collections of the Museum of Compara-
tive Zoology (MCZ), Harvard University, the specimens that bear
“type” labels have not been formally validated, and these were
placed on the assumed types (supposedly the first in each series)
during routine curation early in this century. The Say neotypes were
selected from the LeConte collection because it is generally agreed
that the original Say material is lost, and that LeConte had the
opportunity to compare his specimens with those in Say’s collection
(see Lindroth and Freitag, 1969; Miller and Peck, 1979). The speci-
mens designated do not differ in characters from the original pub-
lished descriptions.

Primary types from the Hatch collection have been deposited by
Oregon State University in the United States National Museum of
Natural History (USNM).

Types have recently been designated for Silpha aenescens Casey,
Silpha raniosa Say, Agvrtes longulus LeConte, and Necrophilus
pettitii Horn (Miller and Peck, 1979; Peck, 1974 and 1982b).

'Department of Biology, Carleton University, Ottawa, Ontario, Canada K1S 5B6.

2Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts

02138.

151

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[Vol. 89

Herman (1964) has shown that the correct spelling of the genus of
the sexton or burying beetles is Nicrophorus Fabricius, 1775, and
not Necrophorus Thunberg, 1789 (see Madge, 1980).

Type Specimens of Nominal Species

Necrophorus lunatus LeConte 1853: 277. lectotype (here desig-

nated), a male with a yellow disc and a red square label with white
dorsal margin “Type 3144” and white label “N. sayi Lap/ lunatus
Lee.,” and our designation label, in MCZ. The yellow circle in
LeConte’s code is supposed to mean “Central Valley or Western
States (Ohio, Illinois, Indiana, Missouri, Western Tennessee,
Kentucky, Iowa, and southern Great Lakes)”. However, the spe-
cies localities were published by LeConte as New York and Geor-
gia. The name is a junior synonym of Nicrophorus sayi Laporte,
1840; and a junior homonym of Nicrophorus lunatus Fischer,
1842, of Eurasia. Harold ( 1868) supplied Necrophorus luniger as a
replacement name for Necrophorus lunatus LeConte.
Necrophorus confossor LeConte 1854: 20. Described from a

single specimen, the holotype: a male with a dark blue disc (indi-
cating Oregon and Washington), and a red square label with
white dorsal margin “Type 3146” and white label “N. confossor/
Cooper Lee.”, and white label “maratimus 4”, in MCZ. The pub-
lished type locality is Prairie Paso, which is in Washington. The
species is a synonym of Nicrophorus investigator Zetterstedt,
1824.

Necrophorus pollinctor LeConte 1854: 19. lectotype (here desig-

nated) a male with dark blue disc (meaning Oregon and Washing-
ton), red label with white dorsal margin “Type 3145”, white label
“N. pollinctor/Cooper Lee.”, white label “vespilloides 9”, and our
designation label, in MCZ. Accompanied by male paralectotype
with blue disc, red label with white dorsal margin “Type/2/ 3145”
and white label “vespilloides 10” and male paralectotype with blue
disc and red label with white dorsal margin “Type/2/3145” and
white label “vespilloides 1 1”, both in MCZ. The published distri-
bution is from Fort Vancouver to the Yokolt Plain. The species is
a synonym of Nicrophorus clefodiens Mannerheim, 1846, and a
junior homonym of Necrophorus pollinctor Mannerheim, 1853
(which is a synonym of Necrophorus investigator Zetterstedt,
1824).

1982]

Peck & Miller — North American Silphidae

153

Necrophorus orbicollis Say 1825: 177. neotype (here designated),

a male in LeConte collection with white label “N. orbicollis/ Say/
Hallii Kirby”, and our designation label, in MCZ. Published
localities are from “the N. W. Territory” (which at the time meant
Ohio, Indiana, Illinois, Wisconsin, and Michigan), and “very rare
in the Middle States”, and “one specimen from Dr. T. W. Harris
of Milton, Massachusetts”. The Harris collection is preserved in
the MCZ and was examined but no specimens of the species were
found that would help to validate Say’s species concept.

Nicrophorus hecate immaculosis Hatch 1957: 15. The name was

validated by this use as a subspecies, not by its earlier use as an
aberration (Hatch 1927: 362) according to Article 10b of the
ICZN. The holotype, which we have not seen, is a specimen from
“California” in the I nstitut fur Pflanzenschutzforschung Zweig-
stelle Eberswalde, East Berlin (formerly Deutsches Entomolo-
gisches Institut). Our examination of much west coast material
(and unpublished data of R.S. Anderson) shows that immaculosis
represents part of a range of variation in elytral patterning within
N. hecate Bland 1865, and, furthermore, that hecate represents a
portion of variation within N. guttu/a Motschoulsky 1845. We
therefore propose immaculosis as a new synonym of hecate,
and hecate as a new synonym of guttula.

Necrophorus maritimus Eschscholtz, in Guerin-Meneville 1835,
Iconographie, plate 17, fig. 8. This name was last used by Hatch
(1957: 14) as Necrophorus investigator maritimus, for material
mostly from coastal islands of British Columbia with reduced
elytral fascia. It was originally described from Sitcha Island,
Alaska, but we have not seen authentic type material. Our exami-
nation (and unpublished data of R. S. Anderson) of much west
coast material shows this to be one part of variation, which is not
geographically coherent, of N. investigator Zetterstedt 1824, and
we therefore propose maritimus as a new synonym.

Necrophorus grandior Angell 1912: 307. lectotype (here

designated), a male with labels “California”, “Janson”, “Original/
type”, “Necrophorus / grandior / Angell”, “Necrophorus / grandior /
2917 Ang./Det. M. H. Hatch 1925”, “Necrophorus/germanicus/
ab. bipunctatus/<3 Kr./M. H. Hatch-1926” and our designation
label, in USNM. Hatch (1927) cited this as the “type” (holotype),
but it was published as a syntype. The species is a synonym of Nicro-

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[Vol. 89

phorus germanicus (Linnaeus), 1758. The location of the second
syntype, a specimen of N. humator (Gleditsch) 1 767 (according to
Hatch 1927) is unknown. It is not in the Hatch collection at
Oregon State University (G. L. Peters, pers. comm.).

Si/pha truncata Say 1823: 193. neotype (here designated), a

male with dark green disc (meaning New Mexico) and white label
“S. truncata Say.”, and our designation label, in MCZ. The pub-
lished locality is “Arkansa” and “near the Rocky Mountains”,
which we take to mean the upper reaches of the Arkansas River in
eastern Colorado. Say collected the single specimen while he was
a naturalist on Major Long’s 1819 party exploring the upper
reaches of the Platte, Arkansas, Canadian, and Red Rivers, of the
southern Great Plains and foothills of the Rockies (Weiss and
Ziegler, 1931). The species is now Thanatophi/us truncatus (Say).

Si/pha bituberosa LeConte 1859b: 6. Described from a single

specimen, the holotype: a female with pale green disc (meaning
Nebraska, Kansas, North Dakota, South Dakota, Oklahoma,
Colorado, Wyoming, and Montana), and red label with white
dorsal margin “Type 8952” and white label “S. bituberosa/
Drexler Lee.”, in MCZ. The published type locality is “near Fort
Bridger”, now in SW Wyoming. The species is now in the genus
Aclypea (in some literature as Blitophaga ), but the former name is
given priority, following Seidlitz (1888: 31 1) as the first reviser in
accordance with article 24 (a) (i) of the International Code of
Zoological Nomenclature.

Si/pha caudata Say 1823: 192. The species was described from

material collected by Thomas Nuttall “on the upper Missouri”
River and by Say from “near the Rocky Mountains.” LeConte
(1859c) recognized this species as a synonym of Si/pha lapponica
Herbst, now Thanatophi/us lapponicus (Herbst), and this has been
accepted ever since. We do not designate a neotype for this name,
for it is not necessary in the interests of stability of nomenclature.

Necrophi/us tenuicornis LeConte 1859a: 84. Described from a

single specimen, the holotype: a female with dark blue disc
(meaning Oregon and Washington), and red label with white dor-
sal margin “Type 3 147” and white label “N. tenuicornis/ P. Sound
Lee. ’’and white label “Pt./tenuicorne/(Lec)’\ in MCZ. The pub-
lished type locality is Puget Sound, Washington. The species is
now Aptero/oma tenuicorne (LeConte).

1982]

Peek & Miller — North American Silphidae

155

New Subgeneric Synonymy

Our studies, as well as those of R. S. Anderson, R. B. Madge, and
A. F. Newton (all unpublished), have not provided data to support
retention of the monotypic subgenus Neeroeharis Portevin 1923 for
Nicrophorus carolinus (Linnaeus 1771). Therefore we consider
Neeroeharis a new synonym of Nicrophorus Fabricius 1775.

Acknowledgements

We thank A. F. Newton (MCZ), T. J. Spilman (USDA c/o
USNM), M. D. Schwartz and G. L. Peters (both Oregon State
University) for their courtesy in allowing study of specimens under
their care. A. F. Newton and R. B. Madge reviewed the manuscript
and have provided much helpful advice throughout our silphid stud-
ies. R. S. Anderson (and his unpublished Masters thesis at Carleton
University on distribution and biology of Silphidae in Canada and
Alaska) helped clarify questions of species identities. Miller’s work
was done while at the Santa Barbara Museum of Natural History
and the Smithsonian Institution.

Literature Cited

Angell, J. W.

1912. Two new North American species of Necrophorus (Coleop.). Ent. News, 23:
307.

Guerin-Meneville, M. F. E.

1835. Iconographie du Regne Animal de G. Cuvier. Insects, vol. 7. Bailliere,
Paris. 576 pp, 104 plates, (plate 17 dated January, 1835 by Cowan, 1971,
J. Soc. Bibliog. Natur. Hist., 6: 18-29: text dates to 1844).

Hatch, M. H.

1927. Studies on the Silphinae. J. New York Ent. Soc., 35: 331-371.

1957. The beetles of the Pacific Northwest. Part II: Staphyliniformia. Univ.
Washington Publ. Biol., 16. 384 pp. Univ. Washington Press, Seattle.
Herman, L. H., Jr.

1964. Nomenclatural consideration of Nicrophorus (Coleoptera: Silphidae).
Coleop. Bull., 18: 5-6.

LeConte, J. L.

1853. Synopsis of the Silphales of America, north of Mexico. Proc. Acad. Nat.
Sci., Philadelphia, 6: 274-267.

1854. Descriptions of some new Coleoptera from Oregon, collected by Dr. J. G.
Cooper of the North Pacific R.R. Expedition, under Gov. J.J. Stevens.
Proc. Acad. Nat. Sci., Philadelphia, 7: 16-20.

1 859a. Catalogue of the Coleoptera of Fort Tejon, California. Proc. Acad. Nat.
Sci., Philadelphia, 1859: 69-90.

156

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[Vol. 89

1859b. The Coleoptera of Kansas and eastern New Mexico. Smithsonian
Cont. to Knowledge, 11(6): 1-66, 3 plates.

1859c. The complete writings of Thomas Say on the entomology of North
America. S.E. Cassino and Co., Boston. Two volumes, XXIV + 412 pp.
+ 54 plates and IV + 814 pp.

Lindroth, C. H. and R. Freitag

1969. North American ground-beetles (Coleoptera, Carabidae, excluding Cic-
indelinae) described by Thomas Say: designation of Lectotypes and Neo-
types. Psyche, 76: 326-359.

Madge, R. B.

1980. A catalogue of type-species in the family Silphidae (Coleoptera). Ent.
Scand., 11: 353-362.

Miller, S. E. and S. B. Peck

1979. Fossil carrion beetles of Pleistocene California asphalt deposits, with a
synopsis of Holocene California Silphidae (Insecta: Coleoptera: Silphi-
dae). Trans. San Diego Soc. Nat. Hist., 19: 85-106.

Peck, S. B.

1974. A review of the Agyrtes (Silphidae) of North America. Psyche, 81:
501-506.

1982a. Silphidae and the associated families Agyrtidae and Leiodidae, in D.
Dindal, ed., Soil Biology Guide. Wiley and Sons, in press.

1982b. Distribution and biology of flightless carrion beetle Necrophilus pettitii
in eastern North America (Coleoptera; Silphidae). Ent. News, 92:
181-185.

Say, T.

1823. Descriptions of Coleopterous insects collected in the late expedition to
the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of
War, under the command of Major Long. Jour. Acad. Nat. Sci., Phila-
delphia, 3: 139-216.

1825. Descriptions of new species of Coleopterous insects inhabiting the Uni-
ted States. Jour. Acad. Nat. Sci., Philadelphia, 5: 160-202.

Seidlitz, G.

1887-1891. Fauna Baltica. Die Kafer (Coleoptera) der Deutschen Ostseepro-
vinzen Russlands. Zweite neu bearbeitete Auflage. Konigsberg: Har-
tungsch Verlagsdruckerei. (Silphidae issued in 1888)

Weiss, H. B. and G. M. Zeigler

1931. Thomas Say, Early American Naturalist. Charles C. Thomas Pub.,
Springfield, 111. 260 pp.

CHEMICAL MIMICRY AS AN INTEGRATING MECHANISM
FOR THREE TERMITOPHILES ASSOCIATED WITH
RETIC ULITERMES VIRGINICUS (BANKS)12

By Ralph W. Howard,34 C. A. McDaniel,* 2 3 4 5
and Gary J. Blomquist6

Introduction

The mechanisms by which termitophiles integrate themselves into
the social life of termite colonies have long intrigued entomologists
(Kistner, 1979). Various authors have suggested that plausible inte-
gration mechanisms might include the using of “appeasement chem-
icals” (Wilson, 1971), passing as morphological mimics (Kistner,
1968), or engaging in behavioral mimicry (Kistner, 1979). We
recently reported (Howard et al., 1980a) that the host-specific,
highly integrated termitophile Trichopsenius frosti Seevers asso-
ciated with Retieulitermes flavipes (Kollar) possesses the same com-
plex mixture of cuticular hydrocarbons as its termite host. We
suggested that this was an example of chemical mimicry which func-
tioned to integrate this beetle into the termite society.

Retieulitermes virginicus (Banks) is sympatric with R. flavipes
throughout much of its range and, as predicted (Howard et al.,
1978; Blomquist et ah, 1979), the two species possess distinctly dif-
ferent cuticular hydrocarbons which function as species recognition
cues (Howard et ah, 1982). They also have different termitophilous
cohorts. Thus, T. frosti is associated only with R. flavipes whereas
T. depressus Le Conte, Xenistusa hexagonalis Seevers (both Sta-
phylinidae: Trichopseniinae), and Philotermes howardi Kistner and
Gut (Staphylinidae; Aleocharinae) are associated only with R. vir-
ginicus. We now report that the three R. virginicus staphylinids also
appear to use chemical mimicry as an integrating mechanism; i.e..

'Manuscript received by the editor June 3, 1982.

2Isoptera: Rhinotermitidae.

3 Forestry Sciences Laboratory. Southern Forest Experiment Station, P. O. Bo x 2008
GMF, Gulfport, MS 39503.

4 Author to whom correspondence should be addressed.

5 National Monitoring and Residue Analysis Laboratory, VS DA Animal and Plant
Health Inspection Service, P. O. Box 3209, Gulfport, MS 39503.

6 Department of Biochemistry, University of Nevada- Reno, Reno, NV 89557

157

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[Vol. 89

they have the same complex mixture of cuticular hydrocarbons as
their host termite. In addition, we report that at least one of these
beetles (X. hexagonalis ) biosynthesizes its hydrocarbons.

Methods and Materials

Portions of several colonies of R. virginicus were collected in 1979
from pine logs in Harrison, Jackson, and Stone Counties, Missis^
sippi. The beetles were separated from the termites, counted by
species, and stored at — 20° C until used. A total of 230 beetles was
collected: 10 P. howardi, 140 T. depressus, and 80 X. hexagonalis.
Cuticular hydrocarbons from pooled samples (by species) were iso-
lated and separated as previously described (Howard et al., 1978).
Hydrocarbons were characterized by gas-liquid chromatography
(GC) retention times and by electron impact (El) and chemical ioni-
zation (Cl) mass spectrometry (Howard et al., 1980b; Jackson and
Blomquist, 1976). Double bond stereochemistries were determined
by comparison with standards using argentation thin-layer chroma-
tography (AgNCE-TLC) (Kates, 1972).

In vitro biosynthesis experiments were conducted as previously
described {Howard et al., 1980a) using 60 X. hexagonalis collected
from a single colony of R. virginicus in September 1979.

Radioactivity was assayed by liquid scintillation counting for 10
minutes at about 85 percent counting efficiency. All counting was
done with a standard deviation of less than 5 percent. A portion of
the isolated hydrocarbons was assayed for total radioactivity. The
remainder of the material was separated by AgN03-TLC into satu-
rated, monounsaturated, and diunsaturated components, which
then were assayed for radioactivity.

Results

The retention times of all peaks present in the GC profile of
cuticular hydrocarbons from R. virginicus (Fig. 1) match those from
the GC profile of the cuticular hydrocarbons of P. howardi (Fig. 2),
T. depressus (Fig. 3), and X. hexagonalis (Fig. 4). Confirmation of
the chemical identity for each of the hydrocarbon components in
most of the GC peaks was obtained by El and Cl mass spectrometry
(MS). In every instance, the GC-MS retention times and mass spec-
tra of the beetle hydrocarbon components were identical to those

1982] Howard, McDaniel, Blomquist — Three Termitophiles 159

previously obtained from R. virginicus cuticular hydrocarbons
(Howard et al., 1982). Likewise, concurrently obtained
AgN03 TLC retention values (Rf) were identical for all beetle
derived alkenes and R. virginicus alkenes. Components which were
identified include n-alkanes, 2-, 3-, 1 1-, 13-, and 15-methylalkanes,
1 1, 15-dimethylalkanes, Z-9-alkenes, Z,Z-7,9-dienes, and E/Z-6,9-
dienes ranging in carbon number from C2] to C40 (Table 1). Double
bond location and stereochemistries of the beetle derived alkenes
were inferred solely from GC and GC-MS retention time data,
and AgN03-TLC Rf data, since insufficient sample was available
for infrared analysis and methoxymercuration-demercuration (Blom-
quist et al., 1980). Early eluting components not identified by a
number in Figures 1 to 4 are unidentified, but have retention times
consistent with a homologous series of /7-alkanes.

The relative abundance of individual hydrocarbon components
varied from species-to-species, but no more so than that of their
termite host, whose percent composition varies considerably by
caste (Howard et al., 1982).

The in vitro radioisotope incorporation experiment was con-
ducted with X. hexagonalis to determine if this species can biosyn-
thesize its cuticular hydrocarbons de novo. Howard (1978) reported
that this species engages in frequent allogrooming with its termite
host, with the resulting possibility of acquiring host hydrocarbons
by mechanical transfer rather than by de novo biosynthesis. A com-
bination of these two alternatives is also possible. After 2 hours of
incubating beetle cuticular tissues with 10 ^uCi of [1— 14C]-acetate,
19.6 ± 8.8 pmole (mean ± SD) of [1— 14C]-acetate was incorporated
into hydrocarbon. About 87.8 ± 5.3 percent of the radioactivity was
in the alkane fraction, 10.2 ± 4.0 percent was in the alkene fraction,
and 1.9 ± 1.3 percent was in the alkadiene fraction. This closely
approximates the distribution of alkanes and olefins in X. hexago-
nalis, suggesting that this species can de novo biosynthesize its
cuticular hydrocarbons. In vitro biosynthesis experiments were not
conducted with T. depressus and P. howardi because we were
unable to collect enough beetles simultaneously.

Discussion

The striking mimicry of hydrocarbon components observed
among these three beetles (representing two subfamilies) and their

160

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[Vol. 89

Fig. 1 . GC trace of total cuticular hydrocarbons of Reticulitermes virginicus. GC
conditions: 1.83 m X 3 mm i.d. Stainless steel column packed with 3 percent (w/w)
SP-2100 on 100/120 mesh Supelcoport; temperature programmed from 150° to
325°C at 5° C/ min.

Fig. 2. GC trace of total cuticular hydrocarbons of Philotermes howardi. GC
conditions same as for Fig. I.

1982] Howard, McDaniel, Blomquist — Three Termitophiles 161

Fig. 3. GC trace of total cuticular hydrocarbons of Trichopsenius ciepressus. GC
conditions same as for Fig. 1.

Fig. 4 GC trace of total cuticular hydrocarbons of Xenistusa hexagonalis. GC
conditions same as for Fig. 1.

162

Psyche

[Vol. 89

termite host is strongly suggestive for their role as integrating fac-
tors. It also supports our earlier hypothesis that cuticular hydrocar-
bons may serve as species recognition cues (Howard et al., 1978;
Blomquist et al., 1979; Howard et al., 1980a; Howard et al., 1982).
Behavioral evidence for this interpretation comes from the finding
(Howard, unpublished observations) that live T. depressus placed
into laboratory colonies of R. flavipes were killed by the termites
within a 24-hour period (five observations). Similarly, the placing of
live T.frosti into laboratory colonies of R. virginicus results in their
being killed (five observations). Beetles can be freely exchanged
among different colonies of their hosts however. These two Tri-
chop senius spp. are nearly identical morphologically and behavior-
ally, but differ markedly with respect to cuticular hydrocarbons.
Similar transplants of workers or soldiers of R. flavipes or R. virgi-
nicus into colonies of the other species also resulted in the death of
the alien individual (five observations). Transplants of conspecific
termites into different colonies did not produce agonistic interac-
tions (five observations). As with the beetles, the two termite species
are morphologically and behaviorally quite similar. We have shown
that R. virginicus workers are antagonistic towards neutral, critical-
point dried (CPD) conspecific workers treated with R. flavipes
cuticular hydrocarbons (Howard et al., 1982), but are not aggressive
toward CPD workers treated with R. virginicus cuticular hydro-
carbons. While we cannot exclude the possibility of other biochemi-
cal differences among either the beetles or their host termites, GC
comparisons of total body extracts revealed none.

The termitophiles associated with R. virginicus (in common with
other termitophiles) possess many epidermal glands (Kistner, 1979)
which have often been postulated to be a source of chemicals which in
some manner aids in the integration of the beetles into the termite
society. While we cannot rule out this interpretation, we would like
to suggest an alternative hypothesis for the function of these glandu-
lar products. Termitophiles are never found in great abundance
(Wilson, 1971; Kistner, 1979), and the nature of termite nest-galley
systems is such as to present substantial problems in the location
and recognition of conspecifics. Perhaps these glands are producing
pheromones directed at conspecifics rather than kairomones directed
at their host. Since pheromones are usually produced in extremely

1982] Howard, McDaniel Blomquist — Three Termitophiles 163

small amounts, such an interpretation would explain the lack of GC
evidence to date for beetle derived biochemicals different from those
of their termite host. An experimental test of this hypothesis must
await the development of suitable bioassays.

Reticu/itermes virginicus and its termitophiles have been co-
evolving for a long period of time (Kistner, 1968, 1979). The beetles
are totally integrated into the social life of the colony and appear to
be chemically indistinguishable from the termites (chemical mim-
icry) vis-a-vis their cuticular hydrocarbons. Most known termite-
termitophile associations, however, occur within the family Termiti-
dae (Kistner, 1979). These associations are characterized by termito-
philes ranging in status from nonintegrated to totally integrated. If
our hypothesis is correct regarding the integrating role of cuticular
hydrocarbons then a corresponding spectrum of congruences of
hydrocarbon profiles would be predicted among the termitophiles
of these communities. We are presently testing this hypothesis.

Many species of ants are known to have inquilines associated with
them, but unlike termitophiles, these myrmecophiles are seldom
host specific (Wilson, 1971). In addition, myrmecophiles seem to
show a wider range of integration (or lack thereof) than do termito-
philes. A correspondingly greater range of integrating mechanisms
might therefore be expected, and have been found. These include
body color, appeasement substances, trichomes, unicellular epi-
dermal glands, physogastry, exudatoria and grandular antennae.
All have been superbly reviewed by Wilson (1971) and Kistner
(1979). The most recent addition to this plethora of mechanisms is
the finding that the scarab beetle Myrmecaphodius excavaticollis
(Blanchard) associated with various Solenopsis spp. (“fire ants”) has
a cuticular hydrocarbon composition which closely mimics that of
its current ant host (Van der Meer, personal communication in
Howard and Blomquist, 1982). The mechanism by which the beetles
achieve this is unknown. Each of the four ant hosts that the scarab
beetles is found with, however, has a unique hydrocarbon profile.
Perhaps ants, like subterranean termites, also use cuticular hydro-
carbons as species-recognition cues. Clearly a great deal remains to
be learned before we achieve an adequate understanding of the
diversity of relationships between social insects and their guests.

Table 1. Cuticular hydrocarbons of Reticulitermes virginicus, Philotermes howardi, Trichopsenius
depressus and Xenitusa hexagonalis.

164

Psyche

[Vol. 89

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1982] Howard , McDaniel, Blomquist — Three Termitophiles 165

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166

Psyche

[Vol. 89

Summary

The three highly integrated staphylinid termitophiles ( Philo -
termes howardi Kistner and Gut, Trichopensius depressus Le
Conte, and Xenistusa hexagonalis Seevers) associated with Reticuli-
termes virginicus (Banks), possess the same cuticular hydrocarbons
as their host. This congruence is hypothesized to be a form of chem-
ical mimicry and is postulated to function as a major way these
beetles achieve integration into the termite society.

Acknowledgement

G. J. Blomquist acknowledges the support of the Science and
Education Administration of the U.S. Department of Agriculture
under grant 7801064 from the Competitive Research Grant Office.

Literature Cited

Blomquist, G. J., R. W. Howard, and C. A. McDaniel.

1979. Structures of the cuticular hydrocarbons of the termite Zootermopsis
angusticollis (Hagen). Insect. Biochem. 9: 365-370.

Blomquist, G. J., R. W. Howard, C. A. McDaniel, S. Remaley, L. A. Dwyer,
and D. R. Nelson.

1980. Application of methoxymercuration-demercuration followed by mass
spectrometry as a convenient microanalytical technique for double-bond
location in insect-derived alkenes. J. Chem. Ecol. 6(1): 257-269.

Howard, R. W.

1978. Proctodeal feeding by termitophilous Staphylinidae associated with Re-
ticulitermes virginicus (Banks). Science 201: 541-543.

Howard, R. W., and G. J. Blomquist.

1982. Chemical ecology and biochemistry of insect hydrocarbons. Annu. Rev.
Entomol. 27: 149-172.

Howard, R. W., C. A. McDaniel, and G. J. Blomquist.

1978. Cuticular hydrocarbons of the eastern subterranean termite, Reticuli-
termes fiavipes (Kollar) (Isoptera: Rhinotermitidae). J. Chem. Ecol.
4(2): 233-245.

Howard, R. W., C. A. McDaniel, and G. J. Blomquist.

1980a. Chemical mimicry as an integrating mechanism: cuticular hydrocar-
bons of a termitophile and its host. Science 210: 431-433.

Howard, R. W., C. A. McDaniel, D. R. Nelson, and G. J. Blomquist.

1980b. Chemical ionization mass spectrometry: application to insect-derived
cuticular alkanes. J. Chem. Ecol. 6(3): 609-623.

1982] Howard, McDaniel Blomquist — Three Termitophiles 167

Howard, R. W., C. A. McDaniel, D. R. Nelson, G. J. Blomquist, L. T. Gelbaum

and L. H. Zalkow.

1982. Cuticular hydrocarbons of Reticulitermes virginicus (Banks)1 and their
role as potential species- and caste-recognition cues. J. Chem. Ecol. 8:
1227-1239.

Jackson, L. L., and G. J. Blomquist.

1976. Insect waxes. P. 201-233. In Chemistry and Biochemistry of Natural
Waxes. P. E. Kolattukudy (ed.). Elsevier, Amsterdam, Oxford, and New
York. 459 p.

Kates, M.

1972. Techniques of Lipidology: Isolation, Analysis and Identification of Lip-
ids. North-Holland Publishing Company, Amsterdam, and American
Elsevier Publishing Company, New York. 610 p.

Kistner, D. H.

1968. Revision of the African species of the termitophilous tribe Corotocini
(Coleoptera: Staphylinidae). I. A new genus and species from Ovambo-
land and its zoogeographic significance. J. N. Y. Entomol. Soc. 76:
213-221.

Kistner, D. H.

1979. Social and evolutionary significance of social insect symbionts. P.
339-413. In Social Insects. Vol. 1. H. R. Hermann (ed.). Academic
Press, New York, San Francisco, and London. 437 p.

Wilson, E. O.

1971. The Insect Societies. The Belknap Press of Harvard University Press,
Cambridge, Mass. 548 p.

PA RATA RUM A, A NEW GENUS OF NEOTROPICAL
CRABRONINI (HYMENOPTERA, SPHECIDAE)*

By Lynn S. Kimsey
Department of Entomology,

University of California, Davis, CA, 95616, USA

Crabronini are a diverse group of wasps that are found world
wide. Typical members of this tribe can be recognized by the single
forewing submarginal cell, large cuboidal head and ventrally con-
verging eyes.

The new genus, Parataruma, is found in lowland neotropical
forest in widely separated localities. This distribution can probably
be explained in several ways. Most of the neotropical lowland forest
has been poorly collected, and much of it has been destroyed. In
addition, these wasps are small and darkly colored, making them
difficult to observe.

Specimens were obtained from the following institutions: British
Museum of Natural History, London (BMNH); Museum of Com-
parative Zoology, Harvard University, Cambridge, Massachusetts
(MCZ); Entomology Museum, University of California, Davis
(UCD), and the U.S. National Museum, Washington, D.C. (USNM).

Parataruma Kimsey, new genus

Generic diagnosis

Head (figs. 3, 4): Eyes asetose, inner orbits converging strongly
below; scapal basin smooth or finely sculptured, laterally margined
by carinae; genal carina well-developed, following ocular margin to
vertex; orbital foveae absent; occipital carina well-developed,
flanged and foveate; antennal sockets touching each other and ocu-
lar margin; male flagellomeres II— III modified; palpal formula 6:4;
mandibles with a tooth on inner margin and single apical notch;
ocelli large, 1.5 times as wide as antennal sockets, forming an iso-
lateral triangle.

Thorax (fig. 1): Pronotal collar with transverse anterior and
posterior carinae, sharply angulate laterally; scutum longitudinally

*Manuscript received by the editor March 22, 1982

169

170

Psyche

[Vol. 89

ridged without anterior transverse carina; notauli indicated by cari-
nae; admedian lines absent; scutellum with deep prescutellar sulcus;
metanotum simple; postspiracular carina well developed; omalus
well-developed, continuous with acetabular carina; verticaulus
short, ending in a ventral depression; hypersternaulus and meso-
sternaulus absent; forewing recurrent vein joining submarginal cell
almost medially; jugal lobe subequal in length to submedial cell; legs
simple; propodeum finely sculptured, dorsal enclosure limited by
foveate sulcus, lateral propodeal carina present.

Abdomen: Sessile; female pygidium forming a sharp, straight
medial ridge, with deep submedial notch, terminating in a sharp
process, surrounded by stout setae (figs. 7-9).

Generotype: Parataruma leclercqi Kimsey, original designation.

Discussion.

Parataruma most closely resembles Foxita and Taruma, based on
the carinate scapal basin, wing venation, narrow female pygidium,
apically notched mandible and absence of the sternaulus. In fact,
this genus will key out to Taruma in Bohart and Menke (1976:374).
However, several characteristics of Parataruma are unusual and
immediately distinguish members of this genus from all other crabro-
nines, including Foxita and Taruma. These characteristics are the
well-developed genal and scapal carinae, the lack of any transverse
carinae on the face and the peculiar female pygidium, which has
been reduced to a narrow convex ridge and pointed apical projec-
tion surrounded by papillae-like setae.

Parataruma leclercqi Kimsey, new species
Figures 1-3, 5-8

Holotype female: Length 5 mm; head finely and densely punctate,
except along occipital and genal carinae; face with bulging brow,
scapal basin punctation obscured by pubescence; clypeal margin
medially produced into a rounded lobe subtended beneath by a tuft
of setae on either side; flagellomeres I— II 1.3 times as long as wide;
flagellomere III as long as wide; flagellomeres IV-IX wider than
long; flagellomere X 1.5 times as long as wide; mandible with tooth
on inner margin longer than diameter of antennal socket (fig. 3);
pronotum foveate along anterior and posterior carinae; scutum
finely punctate-striate with medial and lateral carinae and raised

1982]

Kimsey — Genus Parataruma

171

Figs. 1-3, 5-8 Parataruma leclercqi. Figs. 4, 9. Parataruma tropicauda. Fig. 1.
Lateral view of female. Fig. 2. Male antenna. Figs. 3-4. Complete and partial
front view of female face. Figs. 5-6. Lateral and dorsal views of male genital
capsule. Figs. 7-9. Female pygidium, dorsal (7) and lateral (8, 9) views.

172

Psyche

[Vol. 89

notauli; scutellum punctate-striate with anterior margin foveate, pos-
terior margin with 13 evenly spaced ridges; mesopleuron with fine
punctures, 1-2 puncture diameters apart, upper half with 9 lon-
gitudinal ridges; hypoepimeron ridged; propodeum finely striate
laterally, enclosure with deep medial groove, foveate above,
punctate-striate medially, transversely ridged and foveate below;
terga I V densely and finely punctate, punctures 0.5 puncture
diameter apart or less; tergum VI with large, contiguous, almost
foveate punctures; pygidial ridge sharp, abruptly notched sub-
medially, terminating in a sharp apical projection, nearly obscured
by dense papillae-like setae (figs. 7, 8); sternum 1 integument rough,
irregular; sternum II punctures about 1 puncture diameter apart
laterally, almost impunctuate medially, sterna III-V impunctate,
except transverse subapical punctate strip; sternum VI triangular,
basal half impunctate, apical half densely punctate. Body black,
except yellow scape, flagellum beneath, medial mandibular spot,
pronotal lobe and sublateral dorsal spots, scutellum laterally,
metanotum medially, T II lateral spot, apices of fore and mid-
femora, most of tibiae and tarsi. Pubescence sparse and pale, except
dense silvery appressed pubescence on clypeus, scapal basin and
gena on both sides of carina.

Male: Length 4.5 to 5.5 mm; same as female, except F-I slightly
wider than long; flagellomere II slightly longer than wide, deeply
indented beneath; flagellomere III about as long as wide (fig. 2);
scutum with longitudinal ridges, densely punctate; sterna closely
punctate, punctures 1 puncture diameter apart or less. Male geni-
talia as in figs. 5-6.

Holotype female: Barro Colorado Island, Zona del Canal, Pan-
ama, August 30, 1978, R.B. and L.S. Kimsey (USNM). Paratypes,
13 females: same data as type, except July 17, 1976 (UCD), Sep-
tember 12, 1978 (UCD) and C. and M. Rettenmeyer, April 20, 1963
(UCD); Costa Rica, Turrialba (MCZ, USNM); Colombia, Magda-
lena, 10-15 km e Santa Marta, November 26, 1974, M. Cooper
(BMNH); Venezuela, Zulia, Rosario, June 14, 1976, A.S. Menke
and D. Vincent (USNM); Trinidad: St. George, St. Augustine, June
and August 1976, F.D. Bennett and J.S. Noyes (BMNH); St.
Andrew, Oropuche, June 28, 1976, J.S. Noyes (BMNH). Two
males, which I am not designating as paratypes, were from: Brazil,
Sao Paulo, Ribeirao Preto, January 7, 1968. G.E. Bohart (UCD);

1982]

Kimsey — Genus Parataruma

173

and Mexico, Oaxaca, Oaxaca, April 22, 1959, H.E. Evans (UCD).

The diagnostic features of this species are the yellow female
antennae; yellow spots on the mandibles, metanotum and tergum II;
extensive silvery appressed setae on the scapal basin, gena and meso-
pleuron; large tooth on the inner margin of the mandibles and the
dense setae obscuring the pygidial ridge. In addition leclercqi tends
to be slightly larger than tropicauda, 5-7 mm versus 4. 5-5. Omm for
tropicauda.

I have named this species after Jean Leclercq for two reasons:
first, to acknowledge the tremendous amount of work he has done
on the Crabronini. Second, and most important, because he sent me
10 specimens of this genus to describe even though he recognized
them as new.

Parataruma tropicauda Kimsey, new species
Figures 4 and 9

Holotype female: Only diagnostic characteristics are listed below.
Length 5 mm; scapal basin coarsely and irregularly punctate; meso-
pleuron with 5 or more longitudinal ridges, polished with sparse
punctures; mandible with tooth on inner margin shorter than
diameter of antennal socket. Body black, with yellow on: underside
of scape; pronotal lobes, two pronotal dorsal spots; scutellum lat-
eral spots; fore and midfemora apically; tibiae apically, basally and
inner surfaces; tarsi. Pubescence sparse and pale, except silvery
appressed setae on clypeus and ocular side of genal carina.

Holotype female: 10-15 km e Santa Marta, Magdalena, Colom-
bia, November 26, 1974, M. Cooper (BMNH). Paratype female:
same data as type (BMNH).

This species can be distinguished from leclercqi by the lack of
yellow markings on the mandibles and metanotum, half black scape
and dark flagellum; the sparse or absent silvery pubescence on the
scapal basin and mesopleuron and sparse setae surrounding the
pygidial ridge.

The species name tropicauda, “ridge-tail”, refers to the peculiar
pygidial ridge of the female.

Reference Cited

Bohart, R.M. and A.S. Menke. 1976. Sphecid wasps of the world. 695 pp. Univ.

Calif. Press, Berkeley.

SUPPLEMENTARY STUDIES ON ANT LARVAE:
FORMICINAE (HYMENOPTERA: FORMICIDAE)1

By George C. Wheeler2 and Jeanette Wheeler2
Introduction

This article describes formicine larvae received since the prepara-
tion of our most recent supplement (1980). The larva of Proformica
has not been previously described. Also included are references to
formicine larvae in the literature and a discussion of the status of
Colobopsis.

The terms describing body profile and mandible shape are
explained in our 1976 monograph. Our own contributions are cited
by year and page only.

Tribe 4. Formicini

Genus ACANTHOMYOPS Mayr

The larvae are very active and can quickly change their posture
from circular to linear or reverse.

Genus FORMICA Linnaeus

Alpert and Ritcher 1975:289. Adults of the scarabaeid beetle
Cremastochilus armatus feed on larvae of Formica fusca and
Formica obscuripes.

Genus LASIUS Mayr
Lasius sitkaensis Pergande

Akre and Hill 1973. The pselaphid beetle Adranes taylori Wick-
ham possesses trichomes (tufts of golden hairs) on the abdomen,
tips of elytra and venter. These trichomes are highly attractive to
half-grown or smaller ant larvae, less so to larger larvae and
workers. The beetles are fed by the larvae through trophallaxis and
obtain other nutrients by feeding on dead larvae and workers.
Beetles are often seen walking about with larvae actively holding on
to the trichomes with their mouthparts; Fig. 4 (p. 531) shows a larva
so attached.

'Manuscript received by editor June 10, 1982.

2Adjunct Research Associates, Desert Research Institute, Reno, NV; present address:
326 Laurel Ridge Road, San Antonio, TX 78253.

175

176

Psyche

[Vol. 89

Genus MYRMECOCYSTUS Wesmael

Snelling (1976:22) quoted our characterization (1968:211) of the
genus and compared the larvae of this genus with those of Lasius.
Page 23: “I provided some erroneous identifications to the Wheel-
ers. These may be corrected: dugubris— cr eight oni; * mojave’=testa -
eeus; ‘ semirufus'—kennedyi (Calif.) and depilis (Ariz.)”

Page 7: “These data, albeit fragmentary, seem to indicate that
protein, such as that derived from the tissues of other insects is
essential for larval development.” Page 6: “The insect fragments are
placed among the larvae and these must fend for themselves. I have
seen no indication that larvae of these species are fed by tro-
phallaxis.”

Page 8: Larvae are subject to desiccation; hence they are to be
found in the upper chambers of the nest only in the evening and
early morning. When the surface begins to warm up the brood is
removed to deeper chambers.

Genus PROFORMICA Ruzsky

Profile pogonomyrmecoid. Integument of venter of anterior body
somites and of portions of labium, maxilla and labrum papillose.
Body hairs mostly with bifid tip. Antenna large. Head hairs few,
with 2- or 3-branched tip. Labrum large and subrectangular.
Mandible ectatommoid, with one medial tooth.

The specialization index is 18.

Proformica ferreri Bondroit

Length (through spiracles) about 3.7 mm. Profile pogonomyrme-
coid (i.e., diameter greatest near middle of abdomen, decreasing
gradually toward anterior end and more rapidly toward posterior
end, which is rounded; thorax more slender than abdomen and
forming a neck, which is curved ventrally). Anus posteroventral and
with a small posterior lip. Leg, wing and gonopod vestiges present.
Spiracles small and decreasing in diameter posteriorly. Integument
of venter of anterior somites papillose; dorsal surface of posterior
somites sparsely spinulose, the spinules minute and in short to long
transverse rows. Body hairs sparse, moderately long (0.024-0.07
mm), with simple, bifid or multifid tip. Cranium suboctagonal,
slightly broader than long. Antenna large, with 3 (or 2) sensilla, each
bearing a spinule. Head hairs few, short (0.013-0.04 mm long),
unbranched or with bifid tip. Labrum large, subrectangular, slightly

1982]

Wheeler & Wheeler — Ant larvae

177

Figure 1. Proformica ferrari. a. Head in anterior view, XI 00; b, larva in side view,
X33; c, two body hairs, X320; d, left mandible in anterior view, X320.

broader than long, with ventral border erose; anterior surface with
12 sensilla; with minute papillae near ventral border; ventral surface
papillose and with 6 sensilla; posterior surface densely spinulose, the
spinules minute and arranged in rows which radiate from the
dorsolateral angles, the rows continuous near the base but broken
distally; posterior surface with about 6 sensilla. Mandible large;
ectatommoid (i.e., subtriangular; with a medial blade arising from
the anterior surface and bearing a small medial tooth; apex curved
medially to form a tooth); anterior and posterior surfaces with
longitudinal rugae. Maxilla rather large; with paraboloidal apex;
integument papillose, the papillae bearing minute spinules; palp a
short rounded knob bearing 5 (1 encapsulated and 4 bearing a
spinule each) sensilla; galea digitiform with 2 apical sensilla. Labium
prominent; integument papillose; with a dorsal transverse welt
bearing minute spinules in transverse rows; palp a low knob with 5

178

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[Vol. 89

(1 encapsulated and 4 bearing a spinule each) sensilla; an isolated
sensillum medial to each palp; opening of sericteries wide and with a
sclerotized projection at each side. Hypopharynx with minute
spinules in transverse rows. (Masterial studied: 16 larvae from
Huesca, Spain, courtesy of X. Espadaler.)

Tribe 7. Oecophyllini

Genus OECOPHYLLA F. Smith
Hinton 1951: 169. The larvae of Wurthia aurivillii Kemner and W.
myrmecophila Roepke (Pyralididae) feed on the brood of ants of
this genus.

Tribe 9. Plagiolepidini

Genus ACANTHOLEPIS Mayr
Acantholepis frauenfeldi Mayr

Tohme and Tohme 1975: 136-138. “Les 5 stades larvaires sont
identifies grace a leur forme, leur dimension et surtout leurs poils.”
Fig. 3 (p. 136).

Tribe 10. Brachymyrmecini

Genus BRACHYMYRMEX Mayr
Brachymyrmex admotus Mayr

Length (through spiracles) about 1.6 mm. Very similar to Bra-
chymyrmex depilis (1953: 139) except in the following details. Type 2
body hairs twice as long (0.15 mm). Head hairs 2- or 3-branched:
0.038-0.075 mm long. Mandible with apical tooth slightly more
curved medially. Palp and galea subequal in height; galea more
slender. Labial palp taller. (Material studied: 6 larvae from Costa
Rica, courtesy of Jack Longino.)

Tribe 12. Camponotini

When we defined “praesaepium” (1953:180) we had overlooked
the first description (without a name) of the structure by W. M.
Wheeler and Bailey (1920:270-271): — “In a study undertaken by
the senior author and Mr. George C. Wheeler of the larvae of a large
number of other ant genera, no structure .comparable to the Pseu-
domyrmine trophothylax has been found, except in certain species

1982]

Wheeler & Wheeler — Ant larvae

179

of Camponotus of the subgenus Colobopsis. In all the species of the
latter subgenus examined the larva is very hypocephalic and the
ventral portion of the first abdominal segment projects considerably
beyond the thoracic segments and presents a pronounced concavity
or basin in the mid-ventral region precisely in the position of the
trophothylax of the Pseudomyrminae. A feeble vestige occurs in
many Camponotus larvae belonging to other subgenera. No solid
pellet is deposited in the basin of Colobopsis, but it may, perhaps,
be used to hold a supply of the liquid food regurgitated by the
workers or of the saliva secreted by the larva itself for the benefit of
its attendants.” We later found pellets in the praesaepium of Colo-
bopsis (1970:650).

Genus CAMPONOTUS Mayr
Camponotus rasilis W. M. Wheeler
Petralia and Vinson 1979. Venter — description and SEM.

Genus COLOBOPSIS Mayr

Colobopsis was established by Mayr in 1861 as a genus. In 1889
Emery “reduced it to a subgenus under Camponotus, owing to the
existence of forms intermediate between these two groups and the
relatively unimportant distinguishing characters of Colobopsis ” (W.
M. Wheeler 1904:139). And there it has remained through W. M.
Wheeler’s “Key to the Genera and Subgenera of Ants” (1922),
Emery’s “Genera Insectorum” (1925), Creighton’s “The Ants of
North America” (1950). Brown (1973:179) did not employ subgen-
era; so he had to synonymize it with Camponotus or raise it to
generic rank; he chose the former.

However, we have noticed of late a tendency among myrmecolo-
gists to elevate Colobopsis to generic rank (e.g., Snelling 1981:404).
Although we have some doubts about adult characters, we can cer-
tainly support the elevation by larval characters. In 1953:181 we
wrote: “The genera of this tribe are so similar that we cannot distin-
guish them; hence we have not attempted to key them. Colobopsis
is, however, exceptional; differences of generic magnitude separate
it not only from the other subgenera of Camponotus but also from
the other genera of Camponotini.”

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Tribe Camponotini

1. Posterior x/i of venter of All
raised to form transverse welt,
and, on either side ridges from
the welt extend forward

2. Body hairs numerous

3. Body hairs of 5 types: ( 1) 2- to
6-branched; branches all in
same plane, the most numer-
ous type; (2) simple, short,
slightly curved; (3) few, sim-
ple, long and whip-like; (4)
few, denticulate (5) few, un-
cinate

4. Antenna small

5. Head hairs numerous, long

Colobopsis

1. Praesaepium formed from
ventral surface of Till and
AI: anterior border of All
forms ventral wall; no side
walls

2. Body hairs sparse

3. Body hairs of 3 types; mostly
(1) simple or (2) bifid; few (3)
very long and whip-like; none
uncinate.

4. Antenna minute, peg-like

5. Head hairs moderately numer-
ous, short

Colobopsis pylartes W. M. Wheeler
Petralia and Vinson 1979. Venter — description and SEM.

Genus POLYRACHIS F. Smith
Hinton 1951 : 169. The larvae of Wurthia aurivillii Kemner and W.
myrmecophila Roepke (Pyralididae) feed on the brood of ants in
this genus.

Polyrhachis dives F. Smith

Hinton 1951: 167. The larvae of Batrachedra myrmecophila Snell.
(Cosmopterygidae) feed on the brood of this ant.

Literature Cited

Akre, R. D., and W. B. Hill. 1973. Behavior of Adranes tavlori, a myrmecophilous
beetle associated with Lasius sitkaensis in the Pacific Northwest. J. Kansas
Entomol. Soc. 46:526-536.

Alpert, G. D., and P.O. Ritcher. 1975. Notes on the life cycle and myrmecophilous
adaptations of Cremastocheilus armatus. Psyche 83:283-291.

1982]

Wheeler & Wheeler — Ant larvae

181

Brown, W. L. 1973. A comparison of the Hylean and Congo-West African Rain
Forest ant faunas. Pages 161 185 in “Tropical Forest Ecosystems in Africa
and South America: a comparative review.” Eds. B. J. Meggers et al.,
Smithsonian Press, Washington.

Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool.
Harvard Coll. 104:1 585, 57 pi.

Emery, C. 1889. Intorno ad alcune formiche della fauna palearctica. Ann. Mus. Civ.
Stor. Nat. Genova 27:485 520.

Emery, C. 1925, Hymenoptera, fam. Formicidae, subfam. Formicinae. Genera
Insectorum, Fasc. 183, 302 p., 4 pi.

Hinton, H. E. 1951. Myrmecophilous Lycaenidae and other Lepdoptera: a sum-
mary. Proc. South London Entomol. and Nat. Hist. Soc. 1949-1950, 1 1 1-175.
Mayr, G. 1861. Die europaeischen Formiciden. Wien I Vol.

Petralia, R. S., and S. B. Vinson. 1979. Comparative anatomy of the ventral region
of ant larvae and its relation to feeding behavior. Psyche 86:375 394
Snelling, R. R. 1976. A revision of the honey ants, genus Myrmecocystus. Natur.

Hist Mus. Los Angeles Co. Sci. Bull. 24, 163 p.

Snelling, R. R. 1981. Systematics of social Hymenoptera. Pages 370 453 in “Social
Insects” Vol. II. Ed. H. R. Hermann. Academic Press, New York.

Tohme Henriette and G. Tohme. 1975. Description des castes d'Acantholepis
frauenfelc/i Mayr et des differents stades larvaires. Bull. Soc. Entomol. Egypte
59:131-141.

Wheeler, G. C., and Jeanette Wheeler. 1953. The ant larvae of the subfamily
Formicinae. Ann. Entomol. Soc. Amer. 46:126-171, 175 217.

Wheeler, G. C., and Jeanette Wheeler, 1968. The ant larvae of the subfamily
Formicinae: supplement. Ann. Entomol. Soc. Amer. 61:205-222.

Wheeler, G. C., and Jeanette Wheeler. 1970. Ant larvae of the subfamily
Formicinae: second supplement. Ann. Entomol. Soc. Amer. 63:648-656.
Wheeler, G. C., and Jeanette Wheeler. 1976. Ant larvae: review and systhesis.

Entomol. Soc. Washington Mem. No. 7: 108 p.

Wheeler, G. C., and Jeanette Wheeler. 1980. Supplementary studies on ant
larvae: Ponerinae, Myrmicinae and Formicinae. Trans. Amer. Entomol. Soc.
106:527-545.

Wheeler, W. M. 1904. The American ants of the subgenus Colobopsis. Bull. Amer.
Mus. Nat. Hist. 20:139 185.

Wheeler, W. M. 1922. Key to the genera and subgenera of ants. Bull. Amer. Mus.
Nat. Hist. 45:631 710.

Wheeler, W. M., and I. W. Bailey. 1920. The feeding habits of pseudomyrmine and
other ants. Trans. Amer. Phil. Soc. (Philadelphia) (Art.4):235 279.

MORPHOLOGICAL COMPARISONS BETWEEN THE

OBLIGATE SOCIAL PARASITE, VESPULA AUSTRIACA
(PANZER), AND ITS HOST, VESPULA ACADICA (SLADEN)
(HYMENOPTERA: VESPIDAE)'

By

Hal C. Reed and Roger D. Akre* 2

Department of Entomology
Washington State University
Pullman, WA 99164

Introduction

Obligate social parasites (inquilines) show a vast array of be-
havioral and morphological adaptations to their unique mode of life
(Wilson 1971). The hazards of colony invasion, usurpation, and
subsequent subjugation of members of the host colony (queen
and/or workers) require special features in order to overcome col-
ony defenses and to become integrated within the host’s society. Not
only do these species have adaptations for colony takeover, but they
also lack certain social characteristics, the most notable being the
absence of a worker caste.

Such traits are exemplified in the workerless ant inquiline, Tel-
eutomyrmex sehneideri Kutter, that has enlarged tarsal claws and a
gaster with a concave venter which enables this parasite to ride on
the dorsum of its host (Wilson 1971). Among the parasitic bumble
bees ( Psithvrus spp.) a number of characteristics, such as strong
development of the sting and exoskeleton, are adapted for success-
ful colony takeover, while other social traits, such as a pollen-
collecting apparatus on the hind leg, are lacking (Alford 1975).
Similarly, vespine inquilines are distinct from their hosts in possess-
ing stronger exoskeletons, a closer fitting of their abdominal seg-
ments, stouter and more recurved stings, broader heads, more
powerfully built mandibles, and sharp bidentate clypei (Weyrauch

'Scientific Paper Number 6233, Washington State University, College of Agriculture
Research Center, Pullman. Work done under Project 0037.

2Research Assistant and Entomologist, respectively. Department of Entomology,
Washington State University, Pullman 99164.

Manuscript received by the editor May 10, 1982.

183

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1937, Beaumont 1958). These traits are presumed to function in
preventing sting penetration by host defenders and in facilitating
stinging or injuring the hosts.

Investigators have noted these unique traits primarily among the
European vespine inquilines and their hosts (Beaumont 1958,
Bischoff 1931, Carpenter and Pack-Beresford 1903, Eck 1979, Rob-
son 1898, and Weyrauch 1937). One of these inquilines, Vespula
austriaca (Panzer), has only recently been found in colonies of a
Nearctic species [ V. acadica (Sladen) Reed et al. 1979] and behav-
ioral interactions with members of the host colony have been docu-
mented (Reed 1982). Although researchers have discussed the
external morphology of V. austriaca in relation to the European
host, V. rufa (L.), no comparisons have been made between the
Nearctic host and V. austriaca. Consequently, the objectives of this
paper are: (1) a morphometric comparison between the V. acadica
queen and the parasite; (2) descriptions of certain external features
such as the stings, mandibles, femora, and abdominal sclerites; and
(3) a survey of exocrine glands of the two species.

Materials and Methods

Seven external body parts were measured in pinned specimens of
the host queen and parasite using a dissecting microscope equipped
with an ocular micrometer. These characters have been commonly
used in other biometrical studies of vespid wasps (Blackith 1958,
Eck 1979, Eickwort 1969). Specimens were obtained from local col-
lecting sites (Reed 1982) and from several North American entomo-
logical museums (Acknowledgements). Measurements of the inter-
ocular distance, mesonotal length, hind tibial length, and forewing
length followed the description and diagrams of Eck (1979). Also,
the length of the front femur was measured from the base to the
apex in the same manner as the hind tibial length, while the width
was taken at its widest point. Head width was measured in dorsal
aspect, behind the eyes along the vertex, and between the upper
edges of the genae. The mesonotal length was measured along the
midline from the anterior prescutal suture to the posterior trans-
scutal suture. The length and midline width of the first gastral ter-
gum were also determined in dorsal view.

Scanning electron micrographs (SEM) of the sting apparatus of the
host and three parasite species were prepared. Dissected stings were

1982] Reed & Akre — Vespula austraica and V. aeadiea 185

dehydrated in 100% ethanol, critical point dried, and then coated
with gold. Photographs of femora and Dufour’s glands, which had
been preserved in ethanol, were taken using conventional macro-
photographic techniques.

Exocrine glands and certain other internal features (e.g., ovaries,
ganglia) were examined in specimens preserved for dissection by
injecting Kahle’s solution under an anterior abdominal tergum until
the gaster swelled. Specimens were subsequently stored in 70%
ethanol. Seven V. aeadiea queens and 19 V austriaca females were
examined to establish the occurrence and size of the 14 known
vespine glands (Landolt and Akre 1979). Dissections were conducted
using a binocular dissecting microscope equipped with an ocular
micrometer. Gland size and conditon were compared with previous
measurements (Landolt and Akre 1979). In a few cases, exocrine
glands were inspected in freshly killed specimens. Abdominal plates,
mandibular features, and front femora were also studied in the pre-
served specimens.

Results

The morphometric analysis of selected characters revealed that
although both species are very similar in terms of overall body size
(i.e., as indicated by the width of the mesonotum and gastral tergum
I), certain body parts of V austriaca are significantly larger than
those of the host (Table 1). The head and interocular distance of the
parasite is slightly wider than that of V aeadiea. The mesonotum of
the two species are comparable with only the mesonotal length
being significantly larger in the parasite. Also, the length of the hind
tibia and the forewing are longer than the corresponding parts in the
host, although forewing length is extremely variable in both species.
The first gastral tergum, like the mesonotum, only differs signifi-
cantly in its length.

One of the more unique morphological differences is the larger
front femora of V austriaca (Fig. 1). This femur is consistently
wider and longer in the parasite females than in host queens. Fur-
thermore, the femur is quite robust in the parasite, while it is slender
and more concave on the innner side in V. aeadiea.

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Table 1. Measurements of seven external body parts of V. acadica queens (N = 57)
and V. austriaca females (N = 44).

Mean and standard deviation (mm)

Body part

V austriaca

V. acadica

Head width**

4.31 ±0.14

4.09 ± 0.09

Interocular distance**

1.90 ±0.08

1.78 ±0.06

Mesonotum

width

4.91 ±0.19

4.86 ±0.19

length**

3.80 ±0.17

3.67 ±0.19

Hind tibia

length**

3.91 ±0.16

3.45 ±0.16

Forewing

length**

13.6 ±0.50

13.0 ±0.56

Gastral tergum I

width

4.48 ± 0.20

4.39 ±0.19

length**

1.70 ±0.11

1.41 ±0.1 1

Front femur

width**

0.89 ±0.07

0.81 ±0.06

length**

3.16 ± 0.10

2.78 ±0.10

**Means are significantly different using the t test at 0.01 level of significance.

The mandibles of both species are roughly triangular when view-
ing the mesal (inner) face. The dimensions of the three sides are
approximately the same in each species; however, the mandible of
the parasite is more robust, especially at its base. This stoutness is
apparent when viewing the ventral edge of the mandible. The
ventro-basal area is distinctly wider in V austriaca, and the mesal
face lacks the concavity that is typical of the mandibles of the host
queen. The mandibles of both species have three primary or margi-
nal teeth along the truncated, cutting margin with two alternating,
secondary teeth and a molar shelf behind the margin as is character-
istic of vespines (Duncan 1979). Most yellowjackets (including V
acadica) also have a rounded projection immediately beyond the
notch on the cutting margin (see Fig. 6 in Landolt and Akre 1979,
Duncan 1939); however, this projection in V austriaca is pointed
and more tooth-like. Thus, the parasite actually has a fourth margi-
nal tooth near the dorsal edge.

1982] Reed & Akre — Vespula austraiea and V. aeadiea 187

Figure 1. Front legs of a V. austriaca parasite (left) and V. acadica queen (right).
The femur of the parasite is thicker and longer than that of the host queen. The black
line indicates 1 mm.

The terga and sterna of the gaster of the parasite are more diffi-
cult to dissect apart than those of the host. This “tough armature” is
often mentioned in regard to vespine parasites. The gastral sclerites
appear to overlap very tightly which undoubtedly prevents sting
penetration during usurpation attempts. However, the close fitting
of the abdominal segments does not appear to be due to a reduction
of intersegmental membranes or because of more sclerotization.
Instead, V. austriaea has better developed muscles (i.e., larger bun-
dles) in the abdominal sterna and terga than are present in the same
segments of a V. acadica queen. For example, the three pairs of
intersternal retractors (Duncan 1939) of a fat-laden, fall parasite are
about 1.5 times as wide as these same muscles in a fall V acadica
queen. Consequently, this parasite should be able to retract the
gastral sclerites more tightly than a host queen.

The stout, recurved sting of vespine parasites is one of the most
important morphological adaptation to their mode of life. The two
North American vespine inquilines, V austriaca and Dolichoves-

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pula arctica (Rohwer), have large and highly curved sting shafts in
comparison with those of nonparasitic queens such as V. acadica
(Fig. 2). However, a facultative social parasite, V squamosa
(Drury), does not have a recurved sting shaft, but has a large abrupt
curve at the distal end (Fig. 2E). V. austriaca and D. arctica stylets
have a similar configuration, except only the extreme tip of the
stylet is abruptly hooked (Fig. 2C). However, this condition is
entirely lacking in the V. acadica queen (Fig. 2D). Both parasite and
host have barbs on the sting lancets.

In conjunction with the curved sting, the distal tip of the seventh
abdominal sternum of V. austriaca turns down more sharply than
does the same sternum of the V. acadica queen. In addition, this
sternum has prominent lateral carinae on the ectal surface, whereas,
this ridge is less developed in the host queen. The seventh sternum in
V austriaca is a ca 0.40 mm longer along the midline than that of V.
acadica.

The exocrine glands of four late summer and two spring foun-
dresses, and one fall V acadica queen were examined and measured.
Thirteen of the 14 known glands in Vespula were present, and their
size and development fell within the ranges reported by Landolt and
Akre (1979). The endostipal gland was lacking. In contrast, only 12
glands were located in two aged and 17 preusurpation, summer
parasites since the sixth sternal and endostipal glands were absent.
The head glands, except for the hypopharyngeal, were comparable
in size to those of V acadica and other vespines (Landolt and Akre
1979). The two distinct clusters of cells of the hypopharyngeal gland
were nearly in contact with each other in the center of the suboral
plate of the labrum-epipharynx (see Fig. 5, Landolt and Akre 1979).
The clusters were on the average larger than those in V acadica, but
were usually within the size range found in V pensylvanica (Saus-
sure) (0.03 0.8 mm3). However, two parasites had clusters about

0.12 mm3. The thoracic or salivary glands were also present in V
austriaca and were similar in size to those in nonparasitic queens.

Although the seventh sternal gland, eighth tergal gland, and poi-
son gland reservoir are similar in size and development to those of
other vespines, evident differences exist between the two species in
the other gastral glands. The sixth sternal gland and the associated
sternal brush (i.e., tuft of hairs) are absent in V. austriaca. This
gland (but not the brush) is present in V. acadica, other members of

1982] Reed & Akre — Vespula austraica and V. aeadiea 189

Figure 2. The sting shaft of three social parasites and one host species ( V. acadica).
The sting shaft of V. austriaca is larger and curved (A) as compared to the smaller,
straight shaft of the host queen (B). In V. austriaca the distal tip of the stylet is
distinctly curved (C, upper right) unlike the distal end of the stylet in V. acadica (D).
The facultative, social parasite, V. squamosa, does not have a curved sting shaft, but
it is sharply bent at the distal end (E). The other Nearctic inquiline, D. arctica, also
has a recurved sting (F). The measurements are given in microns.

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the V rufa species group, and V. squamosa, but is absent in queens
of Dolichovespula and the V. vulgaris group (Landolt and Akre
1979). The seventh sternal gland and associated brush are present
and well developed in both species. The Dufour’s gland of V austri-
aca is considerably larger than in most other yellowjacket queens
(Fig. 3) and averaged 5.8 mm long (range = 4.8 - 7.5 mm, N = 15)
and 0.5 to 0.8 mm wide. In contrast, this gland was less developed in
V acadica (x = 2. 1 mm long, 0.3 - 0.5 mm wide) and other nonpara-
sitic queens (Landolt and Akre 1979, 1.5 - 2.5 mm long). The same
gland was found to be enlarged in the facultative social parasite V
squamosa, as it was ca. 0.2 mm wide and 6 mm long (Landolt and
Akre 1979). However, the most well developed Dufour’s gland is
found in the other Nearctic vespine inquiline, D. arctica. One D.
arctica female had a Dufour’s gland 14 mm long and 0.3 mm wide
(Landolt and Akre 1979) and in two parasites dissected by Jeanne
(1977) this gland was 12.8 mm and 27.2 mm long. In this study
three, early summer, D. arctica parasites were found to have very
long glands (16.5, 20.0, 20.5 mm) greatly folded around themselves
and the alimentary canal. In these three parasites and 16 other
preusurpation individuals the gland was flattened and did not con-
tain any material in the lumen. In contrast, the gland was fully
distended and filled with an oily substance in summer, preusurpa-
tion V austriaca. It was empty and flattened in new fall parasites,
while in the aged parasites the gland was only partially full and
appeared collapsed. The gland contained a yellow oily substance in
preserved specimens, but instead had a clear, oil-like material in
three V austriaca specimens killed and immediately dissected.

The ovaries consist of 12 ovarioles as do most Vespu/a and
Dolichovespula (Kugler et al. 1976). The ovaries did not fill the
entire gaster in the two aged parasites as they did in later summer
foundresses of V. acadica. Preusurpation parasites and early
summer host queens had a slight ovarian development with 1 to 6
eggs greater than 1.0 mm in length and thus probably ready to be
laid. Both species have six gastral ganglia.

Discussion

This study confirms the results of a previous morphometric analysis
of V. austriaca (Eck 1979). Eck (1979) compared the inquiline with
the European host, V rufa, and found that although both were

1982] Reed & Akre — Vespula austraica and V. acadica 191

Figure 3. Reproductive organs of a preusurpation V austriaca. The Dufour’s
gland (Dg) is filled with a clear oil material and was 6.5 mm long when fully extended.
Ov= ovaries, Ps = poison sac or poison gland reservoir.

nearly equal in overall body size, V austriaca had a wider head and
interocular distance, longer hind tibia, and longer forewing.

Some researchers (Beaumont 1958, Bischoff 1931, Weyrauch
1937) stressed the robust mandible of V austriaca, while others
(Bequaert 1916, Carpenter and Pack-Beresford 1903, Robson 1898)
found only minor differences in size and did not consider the man-
dible of V austriaca to be significantly larger. The mandible of V
austriaca is definitely robust as it is wider at the base than that of the
host. Weyrauch (1937) discussed and diagrammed the robust nature
of the mandible of the parasite Pseudovespula ingrica (Birula [= D.
ingrica (Birula)]. He stated that the mandible of the inquiline was
less triangular in shape than that of the host queen; a comparison
that was not evident in this study. Weyrauch (1937) also illustrated
the relatively pointed fourth marginal tooth in P. ingrica. The wider
head and genae of V austriaca and other vespine inquilines
(Bischoff 1931, Weyrauch 1937) undoubtedly house larger mandibu-
lar muscles. These muscles in conjunction with the stout mandibles.

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make these appendages a formidable, offensive weapon, probably as
effective as the sting during colony invasions (Reed 1982). Indeed,
Weyrauch (1937) concluded that the powerful mandibles of vespine
inquilines were adapted for fighting with the host.

Another adaptation for combat with host queens and defending
workers is the enlarged femora of the front legs. The only reference
to this feature is found in the original description of Vespa arborea
Smith (= V austriaca) (cited in Robson 1898) in which he stated
that the legs of this species were “stouter and longer” than in V rufa.
The robust front legs are not only an advantage during colony
invasion, but also are likely an adaptation for the frequent mauling
and grabbing of host workers which occurs during early occupation
of the colony (Reed 1982).

The sting is greatly curved in vespine inquilines presumably to
facilitate penetration between the vulnerable intersegmental mem-
branes of defending colony members. The sharp downward bend of
the seventh sternum, likely an accommodation for the recurved
sting, was also noted by Bischoff (1931). The abrupt curve at the
distal tip of the stylet in the inquilines, as well as in V. squamosa,
would appear to impede the thrusting of the two lancets. However,
the distal end may be curved to hook a sclerite and thus enlarge the
intersegmental membrane for further penetration by both the stylet
and lancets.

There is no obvious glandular degeneration in V austriaca, but a
hypertrophy of one exocrine gland exists. Evidently this enlarge-
ment of the Dufour’s gland has some role in vespine social parasit-
ism, but unfortunately the function in any vespine is still unknown
(Landolt and Akre 1979). Several different functions, such as sting
lubrication, have been ascribed to the gland (Spradbery 1973,
Maschwitz and Kloft 1971). The secretion is not considered toxic,
although Barr-Nea et al. (1976) found some lethality to honey bees.
Jeanne (1977) suggested that in D. arctica this gland may produce
an allomone that has some pacifying effect upon the host queen
and or workers. However, the mode of usurpation in D. arctica
differs from that in V austriaca, suggesting a different function for
the gland in the latter. D. arctica usually passively invades queen
nests and coexists with the queen prior to the emergence of the host
workers (Evans 1975, Greene et al. 1978, Jeanne 1977), while

1982] Reed & Akre — Vespula austraiea and V. aeadiea 193

V austriaea forcibly invades a host colony after worker emergence
and does not coexist with the host queen (Reed 1982). Thus, the
secretion of the Dufour’s gland does not appear to act as a pacifying
agent in V austriaea parasitism, and may function as an alarm or
dispersing chemical (Reed 1982). The possibility of differing func-
tions of this gland is indicated by the condition of the gland in the
two species prior to usurpation. In freshly dissected, preusurpation
D. arctica parasites the gland was clearly empty, whereas in preus-
urpation V austriaea females the gland was filled with a clear oil
substance. A similar relationship between an enlarged Dufour’s
gland and social parasitism is found among the slave-making ants of
the subfamily Formicinae (Parry and Morgan 1979, Regnier and
Wilson 1971) and the dulotic ant Harpagoxenus canadensis M. R.
Smith (Buschinger and Alloway 1978). In some of these slave-
makers the gland discharges a chemical that disperses the defending
host workers and attracts other slave-making workers (Regnier and
Wilson 1971).

In conclusion, V austriaea possesses morphological features
significantly different from the host species. Some, such as the pow-
erful mandibles and front legs, and large curved sting, function as
important offensive weapons during colony invasion. Other charac-
teristics, such as the large gastral retractor muscles that enable the
parasite to tightly hold the sclerites together, serve as an important
defense against stinging host workers. The function of the large
Dufour’s gland in vespine inquilines remains obscure; however, it
probably plays a key role in usurpation and control of the host
colony.

Acknowledgements

Appreciation is extended to the following institutions and
researchers for generously supplying specimens for the morpho-
metric study: American Museum of Natural History (M. Favreau);
California Academy of Sciences (P. H. Arnaud); Florida State Col-
lection of Arthropods (W. V. Weems); Museum of Comparative
Zoology, Harvard (S. M. Foster); Oregon State University (G. Fer-
guson); University of Alberta (D. Shpeley); University of British
Columbians. G. Cannings); University of California-Davis (L. S.
Kimsey); and University of Minnesota (P. Clausen).

194

Psyche

[Vol. 89

We gratefully acknowledge Larry Wright (Irrigated Agricultural
Research and Extension Center, Prosser, WA) for his help and time
in preparing the scanning electron micrographs of the sting
apparatus.

Justin Schmidt, Richard Zack, A1 Greene, and Howard Evans are
thanked for their reviews of the manuscript.

Financial support for the research was also provided by a Sigma
Xi Research Grant and a Washington State University Graduate
School Travel Grant.

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Alford, D. V.

1975. Bumble bees. Davis-Poynter Press, London. 352 p.

Barr-Nea, L., P. Rosenberg and J. Ishay.

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Beaumont, J. de.

1958. Le parasitisme social chez les Guepes et les Bourdons. Bull. Soc.
Entomol. Suisse 31: 168-176.

Bequaert, J.

1916. On the occurrence of Vespa austriaca Panzer in the northeastern United
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Bischoff, H.

1931. Zur Kenntnis der Gattung Pseudovespa. Sitz-Berlin Ges. Naturf.
Freunde Berline (1930), p. 329-346.

Blackith, R. E.

1958. An analysis of polymorphism in social wasps. Insectes Soc. 5: 263-272.
Buschinger, A. andT. M. Alloway.

1978. Caste polymorphism in Harpagoxenus canadensis H. R. Smith (Hy-
menoptera: Formicidae). Insectes Soc. 25: 339-350.

Carpenter, G. H. and D. R. Pack-Beresford.

1903. The relationship of Vespa austriaca to Vespa rufa. Entomol. Mon. Mag.
39: 230-242.

Duncan, C.

1939. A contribution to the biology of North American vespine wasps. Stan-
ford Univ. Publ. Biol. Sci. 8: 1-272.

Eck, R.

1979. Biometrische Untersuchung zur Klarung der Artunterschiede bei sozialen
Faltenwespen (Hymenoptera: Vespinae). Entomol. Abh. Mus. Tierk.
Dresden 42: 315-344.

Eickwort, K. R.

1969. Differential variation of males and females in Polistes exclamans. Evolu-
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Evans, H. E.

1975. Social parasitism of a common yellowjacket. Insect World Dig. 2: 6-13.

Greene, A., R. D. Akre and P. J. Landolt.

1978. Behavior of the yellowjacket social parasite, Dolichovespula arctica
(Rohwer) (Hymenoptera: Vespidae). Melanderia 29: 1-28.

Jeanne, R. L.

1977. Behavior of the obligate social parasite, Vespula arctica (Hymenoptera:
Vespidae). J. Kansas Entomol. Soc. 50: 541-577.

Kugler, J., T. Orion and J. Ishay.

1976. The number of ovarioles in the Vespinae (Hymenoptera). Insectes Soc.
23: 525-533.

Landolt, P. J. and R. D. Akre.

1979. Occurrence and location of exocrine glands in some social Vespidae
(Hymenoptera). Ann. Entomol. Soc. Amer. 72: 141-148.

Maschwitz, U. W. J. and W. Kloft.

1971 . Morphology and function of the venom apparatus of insects — bees, wasps,
ants, and caterpillars. Chap. 44, In Bucherl, W. and E. Buckley (eds.).
Venomous Animals and Their Venoms, Vol. III. Venomous Inverte-
brates. Academic Press, N. Y. 537 p.

Parry, K. and E. D. Morgan.

1979. Pheromones of ants: a review. Physiol. Entomol. 4: 161 189.

Reed, H. C.

1982. Biology and behavior of the forest yellowjacket, Vespula acadica
(Sladen) and the obligate social parasite, Vespula austriaca (Panzer)
(Hymenoptera: Vespidae). Ph.D. dissertation, Washington State Uni-
versity. 206 p.

Reed, H. C., R. D. Akre and W. B. Garnett.

1979. A North American host of the yellowjacket social parasite, Vespula
austriaca (Panzer) (Hymenoptera: Vespidae). Entomol. News 90: 1 10 1 13.

Regnier, F. E. and E. O. Wilson.

1971. Chemical communication and “propaganda” in slavemaker ants. Science
172: 267-269.

Robson, C.

1898. Vespa austriaca, a cuckoo wasp. Sci. Gossip (n.s.) 5: 69-73.

Spradbery, J. P.

1973. Wasps. An Account of the Biology and Natural History of Solitary and
Social Wasps. Univ. Wash. Press: Seattle. 408 p.

Weyrauch, W.

1937. Zur Systematik und Biologie der Kuckuckswespen Pseuclovespa, Pseudo -
vespula und Pseudopolistes. Zool. Jahr. (Syst.) 70: 243 290.

Wilson, E. O.

1972. The Insect Societies. Belknap Press of Harvard Univ., Cambridge, MA.
548 p.

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ISSN 0033-2615

PSYCHE

A JOURNAL OF ENTOMOLOGY

:/j£^

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Vol. 89

founded in 1874 by the Cambridge Entomological Club

1982 No. 3-4

CONTENTS

Leptothorax faberi n.sp., an Apparently Parasitic Ant from Jasper National

Park, Canada (Hymenoptera: Formicidae). Alfred Buschinger .. 197

Redescription of the Type Species of Myopsocus, M. unduosus (Hagen), and
Resulting Nomenclatural Changes in Genera and Species of Myopsocidae

(Psocoptera). Edward L. Mockford 211

Parsivoltinism in Three Species of Osmia Bees. P. F. Torchio and V. J.
Tepedino 221

A Review of the Genus Mallada in the United States and Canada, with a New
Species (Neuroptera: Chrysopidae). Phillip A. Adams and J. Allan Garland
239

Polygyny and Polydomy in Three North American Species of the Ant Genus
Leptothorax Mayr (Hymenoptera: Formicidae). Thomas M. Alloway,

Alfred Buschinger, Mary Talbot, Robin Stuart, and Cynthia Thomas . . . 249

A New Colonial Anelosimus Spider from Suriname (Araneae: Theridiidae)

Herbert W. Levi and Deborah R. R. Smith 275

Biology and Systematics of the Bee Genus Crawfordapis (Colletidae,
Diphaglossinae). Card W. Otis, Ronald J. McGinley, Lyn Garling, and Luis

Malaret 279

The Life Cycle of Heteropoda venatoria (Linnaeus) (Araneae: Heteropodidae).

John Ross, Davis B. Richman, Fadel Mans our, Anne Trambarulo, and

W. H. Whitcomb 297

Description of a New Species of Krombeinius (Hymenoptera: Perilampidae)
from the Philippines, and the Phylogenetic Relationships of the Genus.

D. Christopher Darling 307

A Description of the Ectal Mandibular Gland in the Paper Wasp, Polistes
fuscatus (Hymenoptera: Vespidae). H. A. Downing and R. L. Jeanne ... 317

Spiders Living at Wasp Nesting Sites: What Constrains Predation by Mud-

Daubers? Martin S. Obin 321

Agathidiodes Portevin, New Synonym of Stetholiodes Fall (Coleoptera:

Leiodidae: Anistomini). Alfred F. Newton, Jr 337

Fossil Tiger Beetles (Coleoptera: Cicindelidae): Review and New Quaternary
Records. Christopher D. Nagano, Scott E. Miller, and Alan V. Morgan

339

Predation on the Western Honey Bee, Apis mellifera L., by the Hornet, Vespa
tropica (L.). Michael Burgett and Pongthep Akratanakul 347

The Guild of Sawgrass-Inhabiting Ants in the Florida Keys. Blaine J. Cole

351

Defensive Spray Mechanism of a Silphid Beetle (Necrodes Surinamensis).

Thomas Eisner and Jerrold Meinwald 357

Index 371

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The Lexington Press, Inc., Lexington, Massachusetts

PSYCHE

Vol. 89 1982 No. 3-4

LEPTOTHORAX FABERI N. SP., AN APPARENTLY
PARASITIC ANT FROM JASPER NATIONAL PARK,
CANADA (HYMENOPTERA: FORMICIDAE)*

By Alfred Buschinger

Fachbereich Biologie, Institut fur Zoologie, der Technischen
Hochschule, D 6100 Darmstadt, Schnittspahnstr. 3 (FRG)

1. Introduction

The myrmicine tribe Leptothoracini comprises an astoundingly
rich variety of socially parasitic genera and species. Guest ants
(Eormicoxenus, Leptothorax provancheri), as well as slave-making
genera (Harpagoxenus, Chalepoxenus, Epimyrma) and inquilines
(Doronomyrmex) have been described (Buschinger, 1981); however,
we may suspect that only a minor fraction of the existing species is
already known to science. New species can be found nearly every-
where when populations of independent species are closely
examined.

In August, 1979, I collected leptothoracine ants in several locali-
ties of Jasper National Park, Alberta, Canada. The main object was
to find additional material of Doronomyrmex pocahontas, origi-
nally described from this locality (Buschinger, 1979). On August 19,
when inspecting rotten sticks in the coniferous forest along Mt.
Edith Cavill Road near Jasper, I found a colony of a Leptothorax
species belonging to the “L. muscorum ” group sensu lato. Among
the nearly black ants I saw a dealate female which was considerably
smaller than the ordinary queens, and more brownish in color. Its
general appearance was that of a Leptothorax kutteri queen, an
inquiline of L. acervorum in Europe (Buschinger, 1965).

* Manuscript received by the editor June 30, 1982

197

198

Psyche

[Vol. 89

The colony was kept alive for four subsequent brood periods in
artificially shortened annual cycles (Buschinger et al., 1975), and
produced (besides alates and workers of the black “ muscorum ”) a
total of 56 males, 5 females and 1 worker of the small species. From
a second colony that was established with one of the young females,
I got an additional 4 males, 2 females, and one worker. This new,
apparently parasitic species will now be described.

2. Description of Leptothorax faberi n. sp.

Figs. 1-5

Holotype female: total length 3.45 mm, head length 0.67 (exclud-
ing mandibles), head width 0.59 (behind eyes), scape length 0.49,
greatest diameter of eye 0. 16, thorax length 0.98, thorax width 0.58,
length of petiole in lateral view 0.27, width of petiole 0.22, length of
postpetiole 0.20, width of postpetiole 0.32, length of forewing 2.89,
hind wing 1.90, length of hind femur 0.58, hind tibia 0.46. The end
of the gaster is somewhat curved down so that its length (1.3mm)
cannot be determined with precision.

Paratype females (selected measures of two females): total length
3.2/3.47 mm, head length 0.68/0.79 mm, thorax length 0.97/1.18
mm, thorax width 0.50/0.66 mm.

Habitus in general similar to the queens of the genus Leptothorax,
subgenus Leptothorax sensu Smith (1950) ( —Mychothorax Ruzsky).
Mandibles with 5 or 6 teeth of normal size; one or two tiny teeth
may be present between the normal ones in the middle of the masti-
catory border. Maxillary palps 5-segmented, labial palps 3-seg-
mented. Antennae 1 1-jointed with a 3-jointed club. Anterior border
of clypeus with a feeble notch (fig. la). Three ocelli present. Thorax
(fig. lb) as in Leptothorax muscorum. Epinotal spines of moderate
size, acute; epinotal spine index (Buschinger, 1966) between 1.5 and
1.8. Wings as in L. muscorum (fig. 2). Petiole (fig. lb) not peduncu-
lated; viewed from above, the outline is nearly quadrate, with a
slight convexity of the sides. In lateral view the anterior face is
slightly concave, the posterior face distinctly so. The summit is flat,
descending backward and forming a right angle with the anterior
face, and an obtuse angle with the posterior one. A conspicuous
ventral spine forms the anterior end of a ventral, concave, triangular
field, the sharp, ventrolateral edges of which diverge towards the
postpetiole.

J982]

Buschinger — Leptothorax faberi

199

Fig. 1. Head in front view, and head, thorax, and petioles in lateral view of females
of Leptothorax. a,b, L. faberi n. sp.; c,d, its host species, “L. muscorum ”, large black
form; e, f, “L. muscorum", small brown form, for comparison.

Postpetiole (fig. lb) from above about 1.4 times broader than the
petiole, kidney-shaped with a slight anterior concavity. In lateral
view, the anterior face is slightly convex, nearly perpendicular.
Summit rounded, posterior face slightly concave, descending to-
wards the gaster.

The seemingly distinct ventral spine is formed by a sickle-shaped,
transverse protuberance with a sharp anterior edge.

Head, thorax, petiole, and postpetiole mostly coarse and rugu-
lose, gaster smooth and shining. Body sparsely covered with erect,
short and stiff hairs; legs and antennal scapes with abundant,
appressed hairs; funiculus with dense, suberect hairs. Maximal
length of hairs in head, thorax and gaster 0.06-0.08 mm.

Coloration: yellowish-brown with head, dorsal parts of thorax,
petiole, postpetiole and gaster somewhat darker brown. Legs uni-
formly brown, antennae brown with a blackish-brown club.

200

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[Vol. 89

One alate female was dissected. She had 6 ovarioles, a compara-
tively large poison gland, a Dufour’s gland of ordinary size for most
independent Leptothorax species, and an empty receptable also of
ordinary size and shape.

Allotype male: total length 3.44 mm, head length 0.63 (excluding
mandibles), head width 0.65 (behind eyes), scape length 0.26, great-
est diameter of eye 0.28, thorax length 1.21, width 0.70, length of
petiole 0.32, width 0.25, length of postpetiole 0.24, width 0.29,
length of forewing 3.09, hind wing 2.04, length of hind femur 0.79,
hind tibia 0.55, length of gaster ca. 1.05 mm. Paratype males
(selected measures of two males): total length 3.46/3.58 mm, head
length 0.60/0.65 mm, thorax length 1.22/ 1.28 mm, thorax width
0.67/0.70 mm. Habitus in general like that of other males of the
subgenus Leptothorax. Mandibles without teeth, masticatory bor-
der rounded or straight. Maxillary palps 5-segmented, labial palps
3-segmented. Antennae 12-jointed, without club. Clypeus promi-
nent, vaulted, its anterior border straight. Eyes and the ocelli as
large as usual for the subgenus Leptothorax.

Thorax with Mayrian furrows. Epinotum without distinct spines,
but their place marked by two low ridges (fig. 3).

Petiole not pedunculated, with nearly straight anterior and poste-
rior faces of the rounded node. A small ventral spine is present, with
two diverging ventrolateral edges, as in the female. Postpetiole sim-
ilar to that of the female, except that the anterior face is less steep
and more convex, and the ventral spine or transverse edge is smaller
(fig- 3).

Male genitalia: see fig. 3.

Head, sides of pronotum and of petiole coarse, dorsal and
extended lateral parts of thorax, node of petiole, postpetiole and
gaster smooth and shining. Body moderately covered with tapering,
curved hairs of variable length, in the thorax reaching 0.10 mm, on
the petiolar node 0.14 mm. Head and particularly the mandibles
with abundant, long, tapering hairs. Antennae and legs with abun-
dant, appressed or suberect hairs.

Coloration: whole body black or blackish-brown with the scutel-
lum, the metanotum, sometimes the pronotum, the mandibles and
legs somewhat lighter brown. In most males the scutellum differs so
markedly in coloration from the surrounding parts of the thorax
that this was the most valuable character for identifying the new
species’ males when they were still alive in the nest.

1982]

Buschinger — Leptothorax faberi

201

Fig. 2. Wings of females of Leptothorax. a, L. faberi n.sp.; b, "L. muscorum’\
large black form; c, “L. muscorum’’, small brown form. The dotted lines in the L.
faberi fore wing (a) indicate veins that are present in the left, and absent in the right
wing of the same specimen. Wing venation is variable in all species of this group.

202

Psyche

[Vol. 89

b

a

d

1mm

0,5 mm

Fig. 3. Epinotum and petioles of males, and male genitalia (subgenital plate,
volsella with lacinia, and sagitta) of Leptothorax. a,b, L. faberi n.sp.; c,d, its host
species, “L. muscorum" , large black form; e,f, “L. muscorum”, small brown form.
The shapes of the sagitta and volsella with lacinia vary considerably in all 3 species.

Allotype worker: total length 3.02 mm, head length 0.66 (exclud-
ing mandibles), head width 0.58, scape length 0.46, greatest diame-
ter of eye 0.16, thorax length 0.91, width 0.43, length of petiole 0.24,
width 0.20, length of postpetiole 0.17, width 0.30, length of hind
femur 0.51, hind tibia 0.42, length of gaster approximately 1.05 mm.

Habitus similar to an ordinary worker of L. muscorum, but
somewhat more stout and clumsy (fig. 4). Mandibles with 6-7 teeth,
palps with 5 and 3 joints, as in the female. Eyes of moderate size,
ocelli absent, anterior border of clypeus with a notch as in the
female. Thorax with a deep meso-epinotal suture, and the pro-
mesonotal suture clearly visible. Pronotum comparatively wide.
Epinotal spines as in the female, epinotal spine index 1.6. Petiole
and postpetiole as in the female, as well as the appendages. Head
and whole body coarse and rugulose except for the gaster, which is

1982]

Buschinger — Leptothorax faberi

203

Fig. 4. Head, thorax, and petioles of workers in lateral view, a, L. faberi n.sp.; b, its
host species, “L. muscorum”, large black form; c, “L. muscorum ”, small brown form,
for comparison.

204

Psyche

[Vol. 89

smooth and shining. Pilosity as in the female. Coloration yellowish-
brown with the head, the antennal club and the gastral tergites
darker brown.

The karyotype (fig. 5) was determined from 8 male pupae, follow-
ing the method of Imai et al. (1977). In 68 metaphase plates a
haploid number of 15 chromosomes was found, 13 of which are
metacentric or submetacentric and two are subtelocentric. The
second-largest chromosome exhibits a very characteristic banding.
In 7 metaphase plates one additional, subtelocentric chromosome
was found; this, however, may be an artefact. The host species, on
the contrary, has a haploid chromosome number of 17 as does the
second, smaller “L. muscorum ” from Jasper Park, and as occurs in
European L. muscorum.

Type locality: Jasper National Park, Alberta, Canada, a few
meters above the road from 93A to Mt. Edith Cavill parking lot, in
about 1500 m elevation. Numerous nests of the host species and also
of a smaller kind of “L. muscorum” were found inhabiting the rot-
ten sticks and logs lying on the ground of a rather open coniferous
forest.

Derivatio nominis: The ant is dedicated to my late friend, Dr.
Walther Faber, from Vienna, Austria, whom I admired for his
excellent studies in social parasitic ants.

Differential diagnosis: The new species closely resembles the
European inquiline ant Leptothorax kutteri, particularly with re-
spect to size, coloration, and the ventral spines in petiole and post-
petiole. It differs from that species through the lack of erect hairs in
the antennal scapes and the tibiae. Also, the characteristic sculpture
of the head of L. kutteri females is absent in L.faberi. The remarka-
ble light coloration of the male’s scutellum and metanotum is, as far
as I know, unique among leptothoracines belonging to the subgenus
Leptothorax and their social parasites.

The host species (fig. 1) and L.faberi are easily distinguished by
the latter’s smaller size and lighter coloration (female). They also
differ with respect to the karyotypes. L. faberi could only be con-
fused with the second, smaller Leptothorax " muscorum ” form in
Jasper Park (fig. 1), which is the host species of Doronomyrmex
pocahontas. However, this species differs in the shape of petiole and
postpetiole from L.faberi, and it has a karyotype which is identical
to that of the large, black L. “ muscorum ”, host species of L.faberi.

1982] Buschinger — Leptothorax faberi 205

< i t »**««• «*

(JtKU JtUn i « »

*

IfCltlM***** * *

Fig. 5. Karyotype of Leptothorax faberi n.sp. The normal karyotype has n = 15
chromosomes, but 2 of 18 individuals had n = 16 in 2 out of 25 and 5 out of 18
metaphase plates respectively (center line).

3. Biological observations

All observations were made under laboratory conditions, and,
due to the restricted material, they must be fragmentary. However, a
few interesting facts could be recorded, particularly with respect to
reproductive behavior. Sexuals of L. faberi were observed to leave
the nest and to become sexually active in the morning, about 3 to 4
hours after the morning rise in temperature in our artificial 15/25°C
temperature rhythm. Copulation was seen twice, the behavior being
identical to that of Leptothorax kutteri, Doronomyrmex pads and
other social parasites of this group (Buschinger, 1971, 1974, 1975).
A distinct sexual-calling behavior, resembling that of L. kutteri, was
not seen, but poison gland secretion seems to serve as sexual
pheromone as in the species mentioned above. The poison gland of
one female was squeezed onto a small piece of filter paper, and the
paper then put into a nest with L. faberi males. The males suddenly
became excited, and a few began to mount the host species workers.

206

Psyche

[Vol. 89

Mounting attempts of males on L. faberi females could also be
released by gently squeezing a female within a swarming cage with
males flying and crawling around.

One mated and dealate female was placed into a nest of the host
species; however, it had to be removed quickly because it was
seriously attacked by the workers. This same female then was put
together with two workers from the mother colony, and with a few
larvae from another host species colony, into an artificial nest. The
faberi queen became fertile, and after a hibernation I increased her
host worker stock using 25 worker pupae of Leptothorax acervo-
rum. I used L. acervorum pupae because the normal host species
colonies did not produce sufficient worker pupae: L. acervorum is
an ideal “replacement host species” for several parasitic species.
Thus, we succeeded in breeding Formicoxenus nitidulus, guest ant
of Formica, with L. acervorum (Buschinger, 1976). After a second
hibernation the original host species workers were dead, and the
colony produced 1 L. faberi male. Further acervorum worker pupae
were added, and in the third artificial brood period a total of 4
faberi males, 2 females, and 1 worker was produced. During this
period the faberi queen died.

The second laboratory-mated faberi queen was placed into the
mother colony, where it was accepted, apparently became fertile,
and survived for two artificial annual cycles alongside its mother
queen. Both died at the end of the third laboratory brood period of
this colony.

The host species of L. faberi is a comparatively large, nearly black
form which is related to L. muscorum Nyl., but it differs markedly
from this European species. I cannot identify this form yet. A
second, smaller species with more brownish coloration occurs sym-
patrically with the black form in Jasper Park. This smaller “L.
muscorum” is the host species of Doronomyrmex pocahontas
(Buschinger, 1979). It looks more similar to L. muscorum from
Europe than to North American specimens, but it seems also to
represent a distinct species. Besides the morphological differences of
size and coloration, the two Canadian “muscorum” also differ
markedly with respect to their sexual behavior. The “small brown”
species’ females exhibit a characteristic sexual calling behavior
(Locksterzeln) similar to European L. muscorum Nyl. and most
social parasites of this group. I was able to breed this species over
several generations in the laboratory. The “large black” species on

1982]

Buschinger — Leptothorax faberi

207

the contrary, seems to make a mating flight. As with European L.
acervorum, I could never induce mating in any kind of flight cage in
the laboratory.

4 Discussion

The biology of Leptothorax faberi deserves to be discussed with
respect to several features.

First of all, I am fairly convinced that this species represents an
obligatorily parasitic ant. It seems unreasonable to assume that the
one queen originally found should have run into the “muscorum”
nest by chance during collecting, that it would have been accepted
there, and that it could reproduce within the foreign nest. The very
low number of only two workers produced in two colonies is
another datum in favor of the opinion that L. faberi is a parasitic
ant.

Finally, the presence of a postpetiolar ventral spine also supports
this hypothesis, since most parasitic species among the Feptothora-
cini have it.

The production of host species sexuals within the parasitized col-
ony even in its third laboratory brood period indicates that a host
species queen must have been present. This was not checked by
dissection, but several dealate host species females were living in the
nest when it was collected. Thus, L. faberi seems to live as an inqui-
line ant alongside the fertile host colony queen(s), as do Dorono-
myrmex pads, Leptothorax goesswaldi, L. kutteri, and others.
Inquilines, however, are usually workerless. In Doronomyrmex
pads and Leptothorax kutteri, the worker caste is completely lack-
ing in the vast majority of all the colonies we ever collected or kept
in the laboratory, this being several dozen of D. pads and about 100
of L. kutteri. However, a total of 2 or 3 workers of both species have
been produced in laboratory culture, and one L. kutteri worker was
found in a field colony (Bruckner, in litt.). At present, it is impossi-
ble to decide whether the two L. faberi workers represent such rare
exceptions, or whether the species usually will produce some more
workers.

I also doubt that the exceptionally high ratio of males/ females in
the offspring of L. faberi represents the natural condition. Addi-
tional material must be collected in the field to clarify these
problems.

208

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[Vol. 89

With respect to systematic relationships, the new species clearly
supports “Emery’s rule,” under which socially parasitic ants are
always closely related to their respective host species group. No
characters linking L. faberi with European Doronomyrmex pads
or Leptothorax kutteri could be found. The new species shares a
characteristic structure in the petiole with the host species and with
Doronomyrmex pocahontas. The Canadian species have two dis-
tinct small teeth at the base of the anterior face of the petiole just
above its articulation with the epinotum (fig. 1). These teeth are
lacking in their European relatives.

The holotype female, 2 allotype males and 1 worker, and voucher
specimens of the host species are deposited in the Museum of Com-
parative Zoology, Harvard University, Cambridge, Mass. (no.
32758).

5. Summary

Male, female and worker of an apparently parasitic ant, Lepto-
thorax faberi n. sp., are described. The new ant species was found in
a queenright colony of Leptothorax muscorum (sensu lato) in
Jasper National Park, Canada. It differs from the host species in its
smaller size, in the shape of the petioles (figs. 1, 3, 4), in sculpture
and coloration. The karyotype with a haploid number of 15 chromo-
somes (fig. 5) is also different from that of the host species, which
has n = 17 chromosomes. Very few workers have been raised in two
laboratory colonies. Thus, L. faberi seems to represent an inquiline
species.

6. Acknowledgments

I am grateful to my student, Karl Fischer, who assisted me during
the collecting trip, and who also carried out the karyotype studies. I
also thank the Jasper Park authorities for having tolerated our
collecting activities. I am indebted to R. H. Crozier for critically
reading the English text.

References Cited

Buschinger, A.

1965. Leptothorax (Mychothorax) kutteri n. sp., eine sozialparasitische
Ameise (Hymenoptera, Formicidae). Ins. soc. 12: 327-334.

1982]

Buschinger — Leptothorax faberi

209

1971. “Locksterzeln” und Kopula der sozialparasitischen Ameise Leptothorax
kutteri Buschinger (Hym., Form.). Zool. Anz. 186: 242-248.

1974. Zur Biologie der sozialparasitischen Ameise Leptothorax goesswaldi
Kutter(Hym., Formicidae). Ins. soc. 21: 133-144.

1975. Sexual pheromones in ants. In Pheromones and Defensive Secretions in
Social Insects. Proc. Symp. IUSSI Dijon 1975, pp. 225-233.

1976. Eine Methode zur Zucht der Gastameise Formicoxenus nitidulus (Nyl.)
mit Leptothorax acervorum (Fabr.) als “Wirtsameise” (Hym., Form.).
Ins. soc. 23: 205-214.

1979. Doronomyrmex pocahontas n. sp., a parasitic ant from Alberta, Canada
(Hym., Formicidae). Ins. soc. 26: 216-222.

1981. Biological and systematic relationships of social parasitic Leptothoracini
from Europe and North America. In Biosystematics of Social Insects,
ed. P. E. Howse and J.-L. Clement (Academic Press, London, 1981), pp.
211-222.

Buschinger, A., Frenz, G. und M. Wunderlich

1975. Untersuchungen zur Geschlechtstierproduktion der dulotischen Ameise
Harpagoxenus sublaevis (Nyl.) (Hym., Formicidae). Ins. soc. 22:
169-182.

Imai, H. T., Crozier R. H., and R. W. Taylor

1977. Karyotype evolution in Australian ants. Chromosoma 59: 341-393.

Smith, M. R.

1950. On the status of Leptothorax Mayr and some of its subgenera. Psyche
57: 29-30.

REDESCRIPTION OF THE TYPE SPECIES OF
MYOPSOCUS, M. UNDUOSUS (HAGEN), AND
RESULTING NOMENCLATURAL CHANGES IN GENERA
AND SPECIES OF MYOPSOCIDAE (PSOCOPTERA)*

By Edward L. Mockford,

Department of Biological Sciences
Illinois State University,

Normal, Illinois 61761

The assignment of species to the major genera in the Family
Myopsocidae has been hampered by lack of detailed morphological
information about the types of these genera. The genera involved
are Myopsocus Hagen, Lichenomima Enderlein, Phlotodes Ender-
lein, and Rhaptoneura Enderlein.

Enderlein’s (1910) genera were based entirely on wing venational
characters. Some of these have later proven to be variable and of
questionable value (Badonnel 1967). Roesler (1944) synonymized
Phlotodes and Rhaptoneura at the generic level but maintained
them as subgenera. Badonnel (1955) stated that genitalic characters
justify the maintenance of Rhaptoneura and Phlotodes as genera
but did not show what characters were involved. Smithers (1964)
assigned all species which might fall in the genera Myopsocus,
Lichenomima, Phlotodes, and Rhaptoneura to Myopsocus until the
types could be studied. Badonnel (1967) following Enderlein (1910)
and Roesler (1944) assigned to Myopsocus all species with Rs and
M joined by a crossvein in the hindwing, thus synonymizing
Lichenomima with Myopsocus, and assigned all species in which Rs
and M in the hindwing are fused for a distance to Phlotodes, thus
synonymizing Rhaptoneura with Phlotodes.

The present paper reports diagnostic features of the type of
Myopsocus unduosus (Hagen), the type species of Myopsocus
(Enderlein 1910). Genus Myopsocus is re-diagnosed on the basis of
this examination, and an augmented diagnosis of Lichenomima is
included. Generic synonymies are revised, and the species now
assigned to Myopsocus and Lichenomima are listed.

* Manuscript received by the editor August 15, 1982.

211

212

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[Vol. 89

Myopsocus unduo sus (Hagen)

Psocus unduosus Hagen 1859:201.

Myopsocus unduosus (Hagen) Hagen 1866:210.

Type material and its examination. — Types consist of two males,
originally pointed, in the Museum of Comparative Zoology, Cam-
bridge, Massachusetts. Each bears a type label with MCZ number
10118 and the label “Ceylon, coll. Nietner.” I first examined these
types in January 1970, ascertained that both are males of the same
species, and selected one as lectotype. I then soaked the lectotype off
the point, placed it in 80% ethanol, mounted the right wings on a
slide in euparal, and cleared and figured the external genitalia. Early
in 1982, I borrowed the wing slide and made figures from it.

Measurements (mm). — Forewing length = 3.94; hindwing length =
3.22; posterior tibial length = 1.72; least distance between com-
pound eyes = 0.27; transverse diameter of compound eye = 0.42.

Color characters. — Forewing (Fig. 1) with fairly distinct, mottled
crossband in basal half of wing; a distinct stigmasaum darkly
marked in middle; entire margin and most of veins with alternating
dark and light marking. Hindwing (Fig. 2) unmarked except for
brown clouding at base and along anterior margin and alternating
dark and light marking along margin from distal end of Rj to distal
end of R4+5. All femora dark brown with a narrow yellowish-white
preapical ring.

Structural characters. — Forewing (Fig. 1) with relatively long
Rs-M fusion, short M-Cuj fusion. Hindwing with Rs-M fusion
slightly longer than segment of Rs before it. Hypandrium (Fig. 3)
elongate, tapering distally, with slightly bulging, shagreened area on
each side at about distal two-thirds of length; distal end on each side
with field of heavy setae, each seta tapering toward end and base.
Phallosome (Fig. 4) elongate, slender; median style separate from
lateral arms at about two-thirds distance from base to tips of arms
and extending beyond tips of arms. Epiproct (Fig. 5) semicircular
except truncated distally, the distal end beset with minute tubercles.
Paraproct (Fig. 6) bearing bluntly rounded distal process; sense
cushion with 28 trichobothria, all with basal florets.

Diagnostic Features and Synonymy of Myopsocus Hagen

Various authors have noted the constancy within and among spe-
cies of the two character states Rs and M joined by a crossvein

1982]

Mockford — Myopsocus

213

Figs. 1-6. Myopsocus unduosus (Hagen) male lectotype. Fig. 1. Forewing; scale
= 1.0 mm. Fig. 2. Hindwing; scale of Fig. 1. Fig. 3. Hypandrium; scale = 0.2
mm. Fig. 4. Phallosome (dorsal view); scale of Fig. 3. Fig. 5. Epiproct; scale =
0.2 mm. Fig. 6. Right paraproct; scale of Fig. 5.

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versus fused for a distance in the hindwing of the Myopsocid genera
under consideration. Enderlein (1910:68) stated about M. unduosus :
“im Hinterfliigel ist der Radialramus und die Media durch eine
Querader mit einander verbunden.” Obviously, the statement is not
correct. Rs and M are fused for a distance in the hindwing; however,
following Enderlein’s erroneous statement, Roesler (1944) and
Badonnel (1967) mis-assigned these two character states. Thus
Lichenomima (Rs and M joined by a crossvein) was synonymized
under Myopsocus and Rhaptoneura (Rs and M fused for a distance)
was synonymized under Phlotodes. On the basis of examination of
the type, it is evident that Myopsocus has as synonyms Phlotodes
and Rhaptoneura. Lichenomima is probably tenable as a distinct
genus.

Characters correlating with the fusion of Rs and M for a distance
in the hindwing are the following: 1) phallosome generally with a
median style (known exceptions: M. aldabrensis (New), M. minor
(New and Thornton), M. pallidus (Smithers), M. speciosus (Smith-
ers), M. splendidus (Badonnel)); 2) female subgenital plate terminat-
ing in a process tapered distally and with two large setae at the tip
plus smaller setae in some species.

Assignment of Species to Myopsocus

Given the above definition and synonymies, Myopsocus includes
the following species, grouped according to their nomenclatural
history:

1) Species originally placed in Psocus and subsequently trans-
ferred to Myopsocus:

australis Brauer 1865, Australia, Melanesia
unduosus Hagen 1859, Sri Lanka

2) Species originally assigned to Myopsocus, all subsequently
transferred, in effect, to Phlotodes, or Rhaptoneura, or both in
sequence:

clunius Thornton, Lee, & Chui 1972, Micronesia
eatoni McLachlan 1880, Europe, North Africa
furcatus Smithers 1964, Australia
griseipennis McLachlan 1866, Australia
hickmani Smithers 1964, Tasmania
incomptus Smithers 1964, Australia
*kolbei Enderlein 1903 (type of Phlotodes ), New Guinea

1982]

Mockford — Myopsocus

215

novaezealandiae Kolbe 1883, New Zealand
palauensis Thornton, Lee, & Chui 1972, Micronesia
punctatus Thornton, Lee, & Chui 1972, Micronesia
3) Species originally assigned to Phlotodes:
aenulus Badonnel 1967, Madagascar
aldabrensis New 1977, Aldabra
alticola Thornton 1981, Fiji
ambiguus Badonnel 1967, Madagascar
amicus Thornton 1981a, Tonga
angolensis Badonnel 1955, Angola, Madagascar
anomalus Smithers & Thornton 1979, Melanesia
antillanus Mockford 1974, Cuba, Hispaniola, Florida
ascoides Thornton 1981, Fiji
bellus Smithers & Thornton 1974, New Caledonia
bipunctatus Thronton 1981, Fiji
bomasus Smithers & Thornton 1974, New Guinea
brunneigenus Smithers & Thornton 1979, Melanesia
clarki Turner 1975, Jamaica
congolensis Badonnel 1949, Zaire
corticosus Smithers 1964a, Madagascar
cubanus Mockford 1974, Cuba
dentatus Smithers & Thornton 1974, New Guinea
fenestratus Smithers & Thornton 1974, New Guinea
graptus Thornton 1981, Fiji, Tonga
gregarius Smithers & Thornton 1979, Melanesia
gressitti Smithers & Thornton 1974, New Guinea
hoskinsi Smithers & Thornton 1979, Melanesia
inocellatus Smithers & Thornton 1974, New Guinea
lichenosus Enderlein 1931, Seychelles, Madagascar
lineatus Smithers & Thornton 1979, Melanesia
lyriferus Smithers 1964a, Madagascar

maculatus Smithers & Thornton 1974, New Guinea, Melanesia

marginatus Smithers & Thornton 1974, New Guinea

megops Smithers & Thornton 1979, Melanesia

minor New & Thornton 1975, Brazil

minutus Mockford 1974, Cuba, Mexico

mjobergi Karny 1925, Sarawak, Borneo

napuka Thornton 1981, Fiji

obscurus Badonnel 1967, Madagascar

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[Vol. 89

peltatus Smithers & Thornton 1974, New Guinea

pennyi New 1979, Brazil

personatus Badonnel 1967, Madagascar

pilipes Smithers & Thornton 1974, New Guinea

placidulus Smithers 1975, Australia

platyvalvulus Smithers & Thornton 1979, Melanesia

preclarus Smithers & Thornton 1974, New Guinea

punctatoides Thornton 1981, Fiji, Tonga

quadrisetosus Smithers & Thornton 1974, New Caledonia

rastafari Turner 1975, Jamaica

reptus Thornton 1981, Fiji

rimosus Smithers & Thornton 1974, New Guinea
samoanus Karny 1932, Samoa
scabiosus Smithers & Thornton 1974, New Guinea
splendidus Badonnel 1967, Madagascar
thecatus New & Thornton 1975a, Malay Peninsula
toxeres Smithers & Thornton 1974, New Guinea
venustus Smithers & Thornton 1974, New Guinea
vilazi Smithers & Thornton 1974, New Caledonia
zimmermani Thornton 1981, Fiji

4) Species originally assigned to Rhaptoneura :
africanus Badonnel 1955, Angola
ciliiferus Smithers 1964a, Madagascar
cryptus Smithers 1957, Natal

* dispar Enderlein 1910 (type of Rhaptoneura ), Paraguay
magnificus Smithers 1957, South & East Africa
muscosus Enderlein 1931, Seychelles
pallidus Smithers 1964a, Madagascar
setosus Smithers 1964a, Madagascar
speciosus Smithers 1957a, Madagascar

5) Species incertae sedis, originally assigned to Myopsocus and
best left there until they are re-examined:

bakeri Banks 1916, Philippines, Guam
cinereus Navas 1932, Argentina
enderleini Banks 1913, Philippines

fraternus McLachlan 1866, Assam (originally assigned to
Psocus )

pluviosus Navas 1934, India
taurus Banks 1941, Santo Domingo

1982]

Mockford — Myopsocus

217

Relationships of Myopsocus unduosus (Hagen)

Badonnel (1967) constructed a classification of the species from
Madagascar, and Smithers and Thornton (1974) augmented it to
include many of the Old World species. M. unduosus, being known
only from the male, and presenting such unique male characters as
the phallosome with its basal half a simple rod, and the hypandrium
with two distal fields of heavy setae, does not seem to fit into any of
the groups that have been proposed. Smithers and Thornton (1974)
noted that numerous other species could not be placed in their
classification due to paucity of information.

Augmented Diagnosis of Lichenomima Enderlein

Species assigned to Lichenomima (assigned to Myopsocus by
most authors since Badonnel 1967) have veins Rs and M joined by a
crossvein in the hindwing. Correlated with this character are
absence of a median style of the phallosome (possible exception: L.
ariasi New) and female subgenital plate distally with a transverse
sclerite, more or less separate from the main plate, and never termi-
nating in a single process tapering posteriorly.

Species assignable to Lichenomima appear to be those listed by
Smithers (1967) plus the following:

ampla Smithers & Thornton 1974 (from Myopsocus ), New
Guinea

ariasi New 1979 (from Myopsocus ), Brazil
capeneri Smithers 1973 (from Myopsocus ), South Africa
chelata Thornton & Woo 1973 (from Myopsocus ), Galapagos
Islands

clypeofasciata Mockford 1974 (from Myopsocus ), Cuba
coloradensis Banks 1907 (from Myopsocus ), Colorado
elongata Thornton 1960 (from Myopsocus ), Hong Kong
machadoi Badonnel 1977 (from Myopsocus ), Angola
medialis Thornton 1981 (from Myopsocus ), Fiji
posterior Navas 1927 (from Psocus), Costa Rica
pulchella New & Thornton 1975 (from Myopsocus ), Brazil
sanguensis New 1973 (from Myopsocus ), Nepal
varia Navas 1927 (from Amphigerontia ), Costa Rica

Note. — Myopsocus medialis Thornton (1981), assignable to Lich-
enomima on the basis of hindwing venation, appears to be so differ-
ent in several other features as to merit a distinct genus.

218

Psyche

[Vol. 89

Summary

Examination of the type of Myopsocus unduosus (Hagen), the
type species of Myopsocus, allows the genera Phlotodes Enderlein
and Rhaptoneura Enderlein to be synonymized with Myopsocus.
The species now assigned to Myopsocus are listed according to their
nomenclatural history. Species assigned to Myopsocus by most
recent authors are re-assigned to Lichenomima Enderlein.

Acknowledgments

I wish to thank the officers of the Museum of Comparative Zool-
ogy, Cambridge, Massachusetts for the privilege of examining the
type material of M. unduosus.

Literature Cited

Badonnel, A.

1949. Psocopte!4res de la Cote d’Ivoire. Rev. Fr. Entomol. 16:20-46.

1955. Psocopte!4res de l’Angola. Diamang Pub. Cult. 26:1-267.

1967. Faune de Madagascar XXIII. Insectes Psocopte!4res. Office de la Re-
cherche Scientifique et Technique Outre-Mer, Centre National de la
Recherche Scientifique, Paris, pp. 1-237.

1977. Psocopte'/ires de l’Angola V. Diamang Pub. Cult. 89: 103-152.

Banks, N.

1907. New Trichoptera and Psocidae. J. N.Y. Entomol. Soc. 15: 162-166.
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1941. New Neuropteroid insects from the Antilles. Mem. Soc. Cubana Hist.
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Brauer, F.

1865. Neuropteren. Novara-Expedition, Zoologischer Theil. 1: 1-104, pis. I,
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Enderlein, G.

1903. Die Copeognathen des Indo-Australischen Faunengebietes. Ann. Hist.

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1931. Die Copeognathen-Fauna der Seychelles Trans. Linn. Soc. Lond.
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Hagen, H. A.

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1866. Psocinorum et Embidinorum Synopsis Synonymica. Verh. Zool. Bot.
Vereins Wien 16: 201-222.

Karny, H. H.

1925. On the Copeognatha from Mt. Murud and Mt. Dulit, Sarawak.
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1932. Psocoptera. Insects of Samoa. Part VII, Fasc. 4: 1 17-129.

Kolbe, H. J.

1883. Ueber das Genus Myopsocus und dessen Species. Entomol. Nachr. 9:
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McLachlan, R.

1866. New genera and species of Psocidae. Trans. R. Entomol. Soc. Lond. Ser.
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1880. Notes on the entomology of Portugal II. Pseudo-Neuroptera (in part) &
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1974. Records and description of Cuban Psocoptera. Entomol. Am. 48:
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Navas, L.

1927. Communicaciones entomologicas 8. Socopteros del Museo de Ham-

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New, T. R.

1973. Some Psocoptera from Nepal. Orient. Insects 7: 1-10.

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New, T. R. and I. W. B. Thornton

1975. Psocomorpha (Psocoptera) collected on recent expeditions to South
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Roesler, R.

1944. Die Gattungen der Copeognathen. Stett. Entomol. Zeit. 105: 117-166.
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1957. Three new species of Myopsocidae (Psocoptera) from Natal. Proc. R.

Entomol. Soc. Lond. Ser. B, 26: 1 1-16.

1957a. Notes et descriptions sur les Psocopteres de Madagascar. Naturaliste
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1964. The Myopsocidae (Psocoptera) of Australia. Proc. R. Entomol. Soc.
Lond. Ser. B, 33: 133-138.

1964a. On the Psocoptera of Madagascar. Rev. Zool. Bot. Afr. 70: 209-294.
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1975. Additions to Australian Myopsocidae (Psocoptera). Aust. Entomol.
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Smithers, C. N. and I. W. B. Thornton

1974. The Myopsocidae (Psocoptera) of New Guinea and New Caledonia.
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Thornton, 1. W. B.

1960 New Psocidae and an aberrant new Myopsocid (Psocoptera) from Hong
Kong. Trans. R. Entomol. Soc. Lond. 112: 239-261.

1981. Psocoptera of the Fiji Islands. Pac. Insects Monogr. 37: 1-105.

1981a. Psocoptera of the Tongan Archipelago. Pac. Insects Monogr. 37:
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Thornton, I. W. B., S. S. Lee, and W. D. Chui

1972. Insects of Micronesia: Psocoptera. Insects of Micronesia 8(4): 45-144.
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1975. The Psocoptera of Jamaica. Trans. R. Entomol. Soc. Lond. 126:
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PARSIVOLTINISM IN THREE SPECIES OF OSMIA BEES*

By P. F. Torchio and V. J. Tepedino
Bee Biology & Systematics Laboratory,

Agricultural Research Service, USDA,

Utah State University, UMC 53,

Logan, Utah 84322

Individuals of most insect species follow a relatively inflexible
tempo of immature development and adult emergence that includes
a single period of diapause in one generation per year at a specific
stage in the life cycle. A few species depart from this pattern in that a
small proportion of individuals of an age cohort require an addi-
tional year or more to complete development to the adult stage
(Waldbauer 1978, Beck 1980). Among bees, for example, there are
brief reports of delayed emergence for several species (Davidson
1896, MacSwain 1958, Krombein 1967, Torchio 1975, Parker 1980,
Rust 1980) but none of these studies provides quantitative evidence
to demonstrate that delayed emergence is an integral part of the life
cycle.

In this study we supply quantitative evidence to document pat-
terns of delayed emergence in three species of megachilid bees
( Osmia montana Cresson, O. californica Cresson, O. iridis Cocke-
rell and Titus). Individuals of these species complete development in
either one or two years, i.e., the emergence pattern of each age
cohort is bimodal. Waldbauer (1978) used the term “type c” to
describe bimodal ahd polymodal emergence patterns in which the
peaks of emergence of an age cohort occur in different years. Here
we introduce the more descriptive term, “parsivoltine”, to refer to
this phenomenon. “Parsi” is adapted from the Latin pars for part or
partial; -voltine, from the Italian volta for time or cycle is used in its
usual entomological sense, as generations (cycles) per year.

Our study addresses the following questions: 1) Does the propor-
tion of one- and two-year individuals in a cohort differ between the
two years of study and/or between the two sampling sites? 2) Is
there an association between sex and time required to complete
development? 3) How are one- and two-year forms distributed

* Manuscript received by the editor September 9, 1982

221

222

Psyche

[Vol. 89

between and within individual nests? 4) Are inter-individual differ-
ences in the time required to complete development due to environ-
mental factors or to a genetic polymorphism, or both?

The three Osmia species are restricted to the western U.S. where
they are sympatric and at least partially synchronic (late spring-
early summer). Each nests gregariously in pre-existing holes, usually
in wood. The biologies of O. montana and O. californica are sum-
marized by Rust (1974); the biology of O. iridis is currently under
study (Torchio, unpub.). Briefly, nests of each species are composed
of a linear series of cells. Each cell is provided with pollen, nectar
and an egg; cells are separated by partitions constructed of macer-
ated leaf material (O. montana, O. iridis ) or mud mixed with macer-
ated leaf material ( O . californica ), and nests are plugged with one or
more partitions. Osmia montana and O. californica are oligoleges of
the Compositae; O. iridis is restricted to a non-composite host plant.

Methods

Nests of these Osmia species were obtained from trap blocks
placed at two field locations (Torchio 1976). Trap blocks of sugar
pine contained 49 drilled holes to accommodate paper soda straws
measuring 14.5 cm long and 7 mm inside diameter. One hundred
nest blocks were placed at each of two study sites during both study
years (1979-1980).

The Faust trapping site was located 42 km south of Logan, Cache
Co., Utah at 1800 m elevation. This location is on a hillside with a
SW exposure and is covered by large stands of mature aspen ( Popu -
lus tremuloides Michx.) trees surrounded by open, grassland mead-
ows. The Mendon site is located 24 km west of Logan at 1500 m
elevation on a hillside having a SE exposure. Solid stands of maple
(Acer glabrum Torr.) or aspen trees surrounded by open meadows
were characteristic of the area.

During both study years, nest blocks were attached individually
to standing trees during mid-May prior to Osmia flight and returned
to the laboratory in early July where they remained at room illumi-
nation and temperature. All nests were dissected in early September;
individual cocoons were opened to determine sex ratios of adults
(one-year forms) and position of larvae (two-year forms) in nests.
Larvae of two-year forms were weighed on an electrobalance (0.1

1982]

Torchio & Tepedino — Osmia Bees

223

Figure 1. Percent of total offspring of O. montana (a) and O. californica (b) that
were one-year forms (solid lines), and males (dashed line) at two northern Utah
sites over two years. Dotted and dashed line represents the expected sex ratio as
percent males.

mg) and all individuals were then inserted into clear, #000 gelatin
capsules. Capsules were then placed in a constant 4 degrees C
temperature cabinet on September 30 of each study year and trans-
ferred to a 26 degrees C temperature cabinet on June 1 of the
subsequent year. A photoperiod of OL:24D was maintained
throughout these treatments. Capsules containing two-year forms
(now adults) were removed from the temperature cabinet on August
30 and reweighed.

Results

Osmia montana

Almost 1 100 nests were available for examination from the four
site-years of sampling (Table 1). Nest utilization was higher in 1979
than in 1980 at both sites, and higher at Faust than at Mendon in
both years.

There were differences between sites and years in the proportion
of offspring that were one-year forms (Fig. la). At Faust a signifi-
cantly higher proportion of one-year forms was produced in 1979

224

Psyche

[Vol. 89

than in 1980 (X2 = 19.1, P < 0.001) whereas at Mendon a signifi-
cantly higher proportion of one-year cells were produced in 1980
than in 1979 (X2 = 6.3, P < 0.025). However, the Faust site yielded a
significantly greater proportion of one-year cells than did Mendon
during both years (1979, X2 = 214.1, P< 0.001; 1980, X2 = 47.4, P<
0.001).

An association between sex and the number of years required to
complete development was found (Table 1). For all site-years, there
were significantly more males than females among one-year forms,
and fewer than expected males among two-year forms (Faust 1979,
X2 = 9.8, P < 0.005; 1980, X2 = 22.6, P < 0.001; Mendon 1979, X2 =
13.9, P < 0.001; 1980 X2 = 8.8, P < 0.005). Thus the sex ratio (5/9)
of one-year forms was always higher than that of two-year forms.
However, the sex ratio of one- or two-year forms (taken separately)
was not always the same from year to year or from site to site. At
Faust the proportion of both one- and two-year males decreased in
1980 (Table 1; one-year forms, X2 = 7.4, P < 0.01; two-year forms,
X2 = 1 1.9, P < 0.01) and, as a consequence, the combined sex ratio
of offspring declined significantly from 1 .3 (1979) to 0.96 (1980) (X2
= 21.4, P < 0.001). Conversely, no such changes occurred at Men-
don (P > 0.75 all comparisons).

The incidence of one- and two-year forms appears to be con-
trolled by a genetic polymorphism rather than by the action of
environmental variables upon individual offspring. If environmen-
tal cues such as photoperiod, thermoperiod, oxygen levels, etc. act
either indirectly on the mother or directly on the progeny to deter-
mine the developmental fate of offspring, then a consistent pattern
of distribution of one- and two-year forms in mixed nests (those
containing both one- and two-year forms) should be evident. To
examine this possibility we classified mixed nests as follows: 1) one-
year forms in inner cells; two-year forms in outer cells; 2) a reversal
of 1; 3) a double switch, i.e., nests having one-year forms positioned
as bottom and top cells with a two-year form between; or, two-year
forms sandwiching a one-year form. Only mixed nests that could be
categorized with surety were counted; thus, nest totals in Table 2 are
fewer than totals listed in Table 1 because some nests were not
counted. Such a categorization of nests assumes that all nestmates
are siblings. In general this is a valid assumption; supercedure of the
nest of one female by another female is an infrequent occurrence.

Table 1 . Number of live one- and two-year male and female offspring of Osmia montana reared from two sites in two years. Offspring
are grouped by nest type, i.e., 1-yr nests contained only 1-yr offspring etc. SR = sex ratio. Number of dead cells shown in parenthesis next
to total live cells.

No. No. 1-yr. No. 2-yr. No.

Site, year and nest type nests $ 9 SR $ 9 SR cells

1982]

Torchio & Tepedino — Osmia Bees

225

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mixed 38 86 30 2.9 64 74 0.9 254

totals 62 116 65 1.8 119 121 1.0 421(163)

totals both forms #235, 9 9186, SR 1.26

226

Psyche

[Vol. 89

Table 2. Transitions within mixed nests between one- and two-year forms for
Osmia montana and californica. Transition from inner one-year forms to outer two
year forms = 1-2 yr and similarly for 2-1 yr. Double switch signifies the transitions
1-2-1 yr or 2-1-2 yr.

Osmia montana

Osmia californica

1-2 yr

2-1 yr

Double

switch

1-2 yr

2-1 yr

Double

switch

Faust

1979

22

16

47

0

1

1

1980

6

31

63

5

4

52

Totals

28

47

no

5

5

53

Mendon

1979

11

17

32

5

21

12

1980

3

12

17

3

9

22

Totals

14

29

49

8

30

34

For example, in a study of marked O. lignaria only 5 of 1 1 1 nests
(same type as used here) made in a greenhouse contained offspring
produced by more than one female (Tepedino and Torchio 1982b).

Distribution of one- and two-year forms in mixed nests is sum-
marized in Table 2. Nests with double switches were more numerous
than those in the other two categories combined in all site years;
transitions from two-year forms in inner cells to one-year forms in
outer cells were about twice as common as the reverse situation.
Thus, factors such as photo- and thermoperiods, which act on the
maternal genotype to induce diapause in the offspring of other spe-
cies of Hymenoptera (Parker and Tepedino 1982), do not seem to
influence the determination of one- or two-year forms in O.
montana.

The interspersion of one- and two-year forms in mixed nests
creates the potential for fratricide. Observations of trap-nests both
in field and laboratory demonstrated that one-year forms destroy
any two-year larval siblings positioned above them in the nest when
they emerge (Torchio, unpub.). We therefore examined the data for
mixed nests to determine the number of surviving and “doomed”
two-year offspring by sex. The category doomed was assigned to
any two-year form with a one-year form between it and the inner
limit of the nest. All two-year forms without one-year forms posi-
tioned below them were classified as surviving. Our estimates of the
percent doomed two-year offspring should be regarded with cau-

1982]

Torchio & Tepedino — Osmia Bees

227

tion. Although all three species nest in pre-existing holes in dead
wood, in natural situations it may sometimes be possible for emer-
gent one-year adults to gain egress without destroying their two-
year siblings. For example, if the nest is in a rotting log emergent
forms may be able to chew around nestmates. Thus, the estimates
given here should be regarded as maximums.

The results of these comparisons (Table 3) demonstrate that a
large portion of two-year forms was doomed in each site-year (range
40.5-61.3%). This mortality would be in addition to any losses due
to enemies or developmental arrest. It is also interesting to note that
in each site-year a significantly lower proportion of two-year
females than males would be destroyed by their siblings (X2 tests, P
< 0.001 all cases). This is because O. montana, like most bees that
construct nests in pre-existing holes (including O. californica, O.
iridis ), deposit female eggs in cells in the lower reaches of the nest
and males in outer cells (Krombein 1967). Thus, the probability that
a two-year male larva will be destroyed by an emerging sibling adult
is greater than for a two-year female.

Table 3. “Doomed” offspring by sex from mixed nests of Osmia montana and

O. californica.

Males

Females

Totals

Species,

Site, Year

N

%

Doomed

N

%

Doomed

N

%

Doomed

O. montana

Faust 1979

179

84.9

165

35.8

344

61.3

1980

158

69.6

269

23.4

427

40.5

Totals

337

77.7

434

28.1

771

49.8

Mendon 1979

81

81.5

95

35.8

176

56.8

1980

64

64.1

74

24.3

138

42.8

Totals

145

73.8

169

30.8

314

50.6

O. californica

Faust 1979

5

40.0

2

50.0

7

42.9

1980

207

69.1

69

18.8

276

56.5

Totals

212

68.4

71

19.7

283

56.2

Mendon 1979

79

46.8

40

17.5

119

37.0

1980

49

75.5

47

19.1

96

47.9

Totals

128

57.8

87

17.8

215

41.9

228

Psyche

[Vol. 89

Osmia californica

Almost 500 nests were recovered in the four-site years of sampling
(Table 4). Nest utilization was unchanged at Mendon during both
years, but a substantial increase was recorded at Faust from 1979 to
1980.

The proportion of one-year cells declined significantly at both
sites from 1979 to 1980 (Fig. lb. Table 4; Faust X2 = 221.5, P <
0.001; Mendon X2 = 30.0, P < 0.001). In agreement with results for
O. montana, the proportion of one-year forms produced at Faust
was significantly higher than that produced at Mendon during both
years (1979, X2 = 92.5, ?< 0.001; 1980, X2 = 5.7, P < 0.025).

As with O. montana, there was an association between sex and
number of years to complete development (Table 4). For all site-
years (except Faust 1979 for which insufficient numbers of two-year
forms were available for statistical tests) there was a higher propor-
tion of males among one-year forms than among two-year forms
when cells from mixed nests only were considered (Faust 1980, X2 =
4.3, P < 0.05; Mendon 1979, X2 = 20.2, P< 0.001; 1980, X2 = 10.2,
P < 0.005). When all cells were considered, the sex ratio of one-year
forms was always higher than that of two-year forms; but only one
of three comparisons was significant (Faust 1980, X2 = 2.4, P >
0.10; Mendon 1979, X2 = 1 1 .8, P > 0.001 ; 1980, X2 = 2.0, P > 0. 10).

Between-year differences in the proportion of males and females
among one- and two-year forms at each site were less evident than
for O. montana (Table 4). At Mendon the combined sex ratio of
offspring declined significantly from 4.2 (1979) to 1.8 (1980) (X2 =
30.0, P< 0.001) but no such change was evident at Faust (X2 = 0.0,
P > 0.90). The decline in the sex ratio at Mendon was due to a
significantly greater proportion of female progeny produced in 1980
for both one-year (X2 = 15.5, P < 0.001) and two-year forms (X2 =
7.5, P > 0.01). These results are the reverse of those found for O.
montana.

The distribution of one- and two-year forms within mixed nests
were similar to results obtained for O. montana (Tables 2, 4). Thus,
nests having double switches were most numerous and transitions
from one- (inner cells) to two-year forms (outer cells) were inter-
mediate.

As with O. montana, a substantial proportion of two-year forms
were ‘’doomed” (range 37.0-56.5%, Table 3) because cells contain-
ing one-year forms were often constructed lower in the nest. The

Table 4. Number of live one- and two-year male and female offspring of Osmia californica reared from two sites in two years.
Offspring grouped as in Table 1. SR = sex ratio. Number of dead cells shown in parenthesis next to total live cells.

No. No. 1-yr. No. 2-yr. No.

Site, year and nest type nests $ $ SR $ 9 SR cells

Faust 1979

1982]

Torchio & Tepedino — Osmia Bees

229

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totals 97 92 40 2.3 198 118 1.7 448(344)

totals both forms SS 290, 99 1 58, SR 1 .84

230

Psyche

[Vol. 89

proportion of “doomed” two-year females was also significantly
lower than for that of two-year males (Faust 1980, X2 = 53.2, P <
0.001; Mendon 1979, X2 = 9.8, P < 0.005; 1980 X2 = 30.5, P <
0.001).

Osmia iridis

Trap-nests were utilized by O. iridis only in 1979 (Table 5). Of the
83 nests recovered, 54 contained one-year forms exclusively; four
nests contained two-year forms; and the remaining 25 nests were
mixed. Although relatively few two-year individuals were produced
(13.1%), the proportion of two-year females recovered was greater
than that of one-year females (mixed nests, X2 = 16.1, P < 0.001,
total nests, X2 = 9.0, P < 0.005, both sites combined).

Unlike other Osmia species studied, the predominant transition
category of O. iridis in mixed nests was from the two-year form
(inner cells) to the one-year form (outer cells) 16 of 25 nests). There
were relatively few nests with either double switches (5) or with
transitions from one-year (inner) to two-year (outer) forms (4).

The tabulation of “doomed” individuals in mixed nests demon-
strated that two-year males were at greater risk than two-year
females (X2 = 6.0, P < 0.025).

Expected and observed sex ratio:

We calculated the expected equilibrium sex ratio (5/9) for each
species on the basis of male and female live weights (Table 6) as
described previously for O. lignaria propinqua (Torchio and Tepe-
dino 1980). Two interesting points emerged from this analysis. First,
for each species, the expected sex ratio was the same regardless of
whether larval or adult weights were used. Second, the expected sex
ratios of these three species were very similar to each other and to
0.1 propinqua (Torchio and Tepedino 1980). Apparently the opti-
mal size ratio between females and males is the same for many
Osmia species that nest in similar substrates.

When the expected and observed sex ratios were compared, con-
sistent biases emerged: For O. montana the observed sex ratio was
significantly biased towards females for all site-years (P < 0.005
or less, all tests). In contrast, observed sex ratios for both O. cali-
fornica and O. iridis were generally biased towards males (P <0.001
or less, all but O. calif ornica Mendon 1980). In addition, there was

1982]

Torchio & Tepedino — Osmia Bees 231

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232

Psyche

[Vol. 89

Table 6. Mean fresh weights (mg) of adults and larvae of three species of Osmia.
Weights for O. montana and californica are for two-year forms, those for O. iridis for
one-year forms. N = sample size, ESR = expected sex ratio (SI 9)-

Adults

larvae

% wgt.

loss

(5

$

<5

9

(5

9

montana

live wgt

71.9

125.3

93.1

161.7

23.0

22.6

± SD

16.1

16.9

19.2

20.3

3.0

2.1

N

21

34

21

34

21

34

ESR

1.74

1.74

californica

live wgt

73.3

121.2

94.1

156.6

22.0

22.8

± SD

14.5

19.7

18.6

21.8

4.3

3.5

N

23

16

23

16

23

16

ESR

1.65

1.66

iridis

live wgt

38.3

66.7

± SD

4.9

9.5

N

58

15

ESR

1.74

no consistent tendency for observed sex ratios to move towards
the equilibrium sex ratio in 1980 for either O. montana or O. eali-
fornica (Fig. 1).

Discussion

The data presented above are noteworthy for several reasons.
First, these three species provide the best documented examples of
parsivoltine emergence patterns in bees. Indeed, among Hymenop-
tera, detailed examples of parsivoltinism are available only for
diprionid sawflies (Prebble 1941, Griffiths 1959, Sullivan and Wal-
lace 1967, Wallace and Sullivan 1974). Previous reports of such
emergence patterns in bees have been based on small sample sizes
( Dianthidium pudicum consimile (Ashmead) (Davidson 1896), Me-
lissodes robust ior Cockerell (MacSwain 1958), Proche/ostoma phil-
adelphi (Robertson) (Krombein 1967), Perdita nuda Cockerell,
Sphecodes sp. (Torchio 1975), Osmia marginipennis Cresson (Park-
er 1980), and Hopiitis biscute/lae (Cockerell) (Rust 1980) ).

1982]

Torchio & Tepedino — Osmia Bees

233

A second point of interest is that two-year forms of all three
species undergo two periods of diapause (once as post-defacating
larvae during the first winter, and again as adults over the second
winter) whereas one-year forms diapause only as adults. For most
other insect species, diapause is stage specific and occurs only once
in the life cycle (Beck 1980); there are a few reports of non-
hymenopterous insect species that enter diapause in more than one
stage (e.g., Harvey 1967, Lounibos and Bradshaw 1975). The physi-
ological mechanisms which enable species to undergo two discrete
periods of diapause are unknown (Chippendale 1977, Waldbauer
1978).

A third unusual result of this study is the relatively large propor-
tion of individuals in each age cohort which were two-year forms
(Fig. 1). When data for live offspring were combined for all site-
years by species, 41% of all O. montana, 57% of all O. calif ornica
and 13% of all O. iridis required two years to complete develop-
ment. In contrast, the percentage of individuals requiring prolonged
periods to complete emergence in most other species with parsivol-
tine emergence patterns is low (Powell 1974, Waldbauer 1978, Sha-
piro 1979, Tauber and Tauber 1981).

A potential explanation for the high proportion of two-year
forms among these species has been provided by Cohen (1966,
1968). In his treatment of optimal reproductive strategies, Cohen
noted that when weather and/ or resources exhibit large year to year
fluctuations and, as a result, the year to year variance in reproduc-
tive success is also large, it would be adaptive for organisms to
produce offspring types that differed in the time required to reach
maturity. By this means, the effects of years unfavorable to repro-
duction would not fall upon all members of an age cohort (See also
Powell 1974, Hedrick et a/. 1976, Waldbauer 1978, Shapiro 1979,
Real 1980, Tauber and Tauber 1981). Cohen (1966, 1968) also
hypothesized that variance in reproductive success should be posi-
tively associated with the proportion of offspring that require an
extra year (or more) to complete development; and that, as the
viability of two-year forms decreased relative to one-year forms, the
proportion of two-year forms in the population should also
decrease. Thus, Cohen’s theoretical results suggest that the high
proportion of two-year forms in these Osmia species may be due to

234

Psyche

[Vol. 89

substantial temporal heterogeneity in the environment and that via-
bility of two-year forms is about the same as that of one-year forms.
In addition, the data suggest that there may be differences among
these species in the way a heterogeneous environment is expe-
rienced. The percentage of two-year forms appears to be higher for
O. montana and O. californica than for O. iridis (Fig. 1), and this
suggests that variance in reproductive success is lower for O. iridis
than for the other species.

Although there are no data available to directly address these
predictions, trap-nesting returns from northern Utah over the past
10 years (Torchio, unpub.) suggest that O. montana and O. califor-
nica populations are much more stable than are those of O. iridis.
The latter species is only occasionally abundant and, more fre-
quently, is totally absent from trap-nests. Conversely, trap-nest
returns for O. montana and O. californica fluctuate within much
narrower limits. Thus, the data available to us do not support
Cohen’s ( 1966, 1968) predictions.

Another characteristic expressed by these and other species that
does not seem to conform to Cohen’s (1966, 1968) predictions is the
relative viability of one- and two-year forms. For example, Sullivan
and Wallace (1967) reported that mortality increased and fecundity
decreased with prolonged diapause in the sawfly. Neodiprion ser-
tifer (Geoff.). Although we were unable to compare the mortality
rate of one- and two-year forms because it was impossible to assign
immature deaths in the first year to either category, it seems clear
that mortality of two-year forms must be higher than that for one-
year forms because some of the former will be destroyed when the
latter exit the nests in the first year (Table 3). Thus Cohen’s
requirement that viability of one- and two-year forms be equal
seems not to be satisfied. In this regard, the advantage of producing
a greater proportion of two-year females than two-year males may
simply be a mechanism to reduce mortality levels of two-year forms
because females almost always occur in the inner cells of the nest
where mortality due to emergence of one-year forms is minimal.
Thus some degree of linkage between sex and developmental time in
such a system would be selected for. Without linkage, sibling-
effected mortality on two-year forms would be even higher.

Despite the apparent lack of agreement between the data and the
predictions of Cohen (1966, 1968), the between year variation in

1982]

Torchio & Tepedino — Osmia Bees

235

proportion of one- and two-year forms within sites for each species
(Fig. 1) suggests that the relative fitness of these forms is determined
by environmental conditions. What these conditions are and how
they interact with the genotypes to maintain a balanced polymor-
phism (if indeed it is balanced) remains to be studied.

An interesting ramification of varying selective pressures upon
one- and two-year forms is the indirect effect upon the sex ratio of
the population. Elsewhere, we (Tepedino and Torchio 1982a) have
suggested that data from a long-term field study of O. lignaria pro-
pinqua Cresson (a univoltine species) supports Fisher’s (1958) the-
ory of an equilibrium sex ratio. In the three species studied here,
however, it appears that any approach toward equilibrium sex ratio
values is dependent upon constraints imposed by selection for parsi-
voltinism. For example, since there is an association between the
two-year form and the female sex, an increase in the relative fitness
of two-year forms in any year could divert the population away
from equilibrium and towards a female bias in subsequent years.
The potential for such diversion should depend on the genetic sys-
tem responsible for the polymorphism. However, the absence of any
consistent tendency for population sex ratios of these species to
move towards equilibrium (Fig. 1) suggests that this may be a real
phenomenon.

Summary

Offspring from nests constructed in wooden domiciles by three
non-social species of Osmia bees at two sites in northern Utah dis-
played differences in the time required to complete development to
the adult stage. Some members of each age cohort emerged in the
following year, but a substantial proportion required two years to
complete development. We propose the term “parsivoltine” to de-
scribe such emergence patterns.

There were differences in the proportion of one-year forms, both
between years, within sites and between sites, within years for each
species. The factors influencing these changes are unclear at present.
The distribution of one-year and two-year individuals within nests
suggests that environmental factors alone do not act on either the
female parent or on her offspring to determine the developmental
fate of the offspring. Many nests contained both one- and two-year

236

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[Vol. 89

forms in unpatterned linear arrangements. All three species appear
to be genetically polymorphic for the time necessary to complete
development.

There was an association between sex and time required to com-
plete development. Two-year forms were more frequently female
than male and one-year forms were more frequently male than
female. Female offspring are also typically placed in the innermost
cells of the nest. In these linear nests, if a two-year form occurs
between the nest exit and a one-year form, then the latter will fre-
quently destroy the former to gain egress from the nest. Thus, this
association between sex and developmental time may act to lower
sib caused mortality.

In addition, the association between sex and developmental time
may constrain an approach to the equilibrium sex ratio because of
selection for a particular developmental form.

Although spatiotemporal heterogeneity of weather and/or re-
sources has frequently been offered as an explanation for such
developmental polymorphisms, it is by no means clear that this is
the case for these species of Osmia. The high proportion of two-year
forms, and the differences between developmental forms in mortal-
ity, and perhaps fecundity as well, do not fit the profile which is
typically offered for parsivoltine species.

Acknowledgments

We thank Glen Trostle, Mary Klomps, Pauline Anderson, Char-
lene Roth, and Barbara Becker for their unselfish efforts in prepar-
ing nest blocks, dissections of nests, and weighing bees; and Drs. S.
D. Beck (Univ. of Wisconsin), Jerome Rozen, Jr. (Amer. Mus. Nat.
Hist.), and C. A. and M. J. Tauber (Cornell Univ.) for their helpful
comments on the manuscript.

Literature Cited

Beck, S. D.

1980. Insect Photoperiodism. Academic Press, Second Ed., New York. 387 pp.
Chippendale, G. M.

1977. Hormonal regulation of larval diapause. Annu. Rev. Entomol.

22:121-138.

Cohen, D.

1966. Optimizing reproduction in a randomly varying environment. J. Theor.

Biol. 12:119-129.

1982]

Torchio & Tepedino — Osmia Bees

237

1968. A general model of optimal reproduction in a randomly varying envir-
onment. J. Ecol. 56:219-228.

Davidson, A.

1896. Nesting habits on Anthidium consimile. Entomol. News 7:22-26.
Fisher, R. A.

1958. The Genetical Theory of Natural Selection. Dover Publ., Second Rev.
Ed. New York. 291 pp.

Griffiths, K. J.

1959. Observations of the European pine sawfly, Neodiprion sertifer (Geoff.),
and its parasites in southern Ontario. Can. Entomol. 91:501-512.

Harvey, G. T.

1967. On coniferous species of Choristoneura in North America. 5. Second
diapause as a species character Can. Entomol. 99:486-503.

Hedrick, P. W., M. E. Ginevan, and E. P. Ewing

1976. Genetic polymorphism in heterogeneous environments. Annu. Rev.
Ecol. Syst. 7:1-32.

Krombein, K. V.

1967. Trap-nesting Wasps and Bees: Life Histories, Nests and Associates.
Smithsonian Press, Washington, D. C. 570 pp.

Lounibos, L. P., and W. E. Bradshaw

1975. A second diapause in Wyeomyia smithii: seasonal incidence and mainte-
nance by photoperiod. Can. J. Zool. 53:215-221.

MacSwain, J. W.

1958. Longevity of some anthophorid bee larvae (Hymenoptera: Apoidea).
Pan-Pac. Entomol. 34:40.

Parker, F. D.

1980. Nests of Osmia marginipennis Cresson with a description of the female
(Hymenoptera: Megachilidae). Pan-Pac. Entomol. 56:38-42.

Parker, F. D. and V. J. Tepedino

1982. Maternal influence in diapause in the alfalfa leafcutting bee. Ann.
Entomol. Soc. Amer. 75:407-410.

Powell, J. A.

1974. Occurrence of prolonged diapause in ethmiid moths. Pan-Pac. Entomol.
50:220-225.

Prebble, M. L.

1941. The diapause and related phenomena in Gilpinia polytama (Hartig). V.
Diapause in relation to epidemiology. Can. J. Res. 19:437-454.

Real, L. A.

1980. Fitness, uncertainty, and the role of diversification in evolution and
behavior. Am. Nat. 115:623-638.

Rust, R. W..

1974. The systematics and biology of the genus Osmia, subgenera Osmia,
Chalcosmia, and Cephalosmia (Hymenoptera: Megachilidae). Wasmann
J. Biol. 32:1-93.

1980. Nesting biology of Hoplitis biscutellae (Cockerell) (Hymenoptera: Meg-
achilidae). Entomol. News 91:105-109.

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Shapiro, A. M.

1979. The phenology of Pieris napi microstriata (Lepidoptera: Pieridae) dur-
ing and after the 1975-1977 California drought, and its evolutionary
significance. Psyche 86:1-10.

Sullivan, C. R., and D. R. Wallace.

1967. Interaction of temperature and photoperiod in the induction of pro-
longed diapause in Neodiprion sertifer. Can. Entomol. 99:834-850.

Tauber, C. A., and M. J. Tauber.

1981. Insect seasonal cycles: Genetics and evolution. Annu. Rev. Ecol. Syst.
12:281-308.

Tepedino, V. J., AND P. F. Torchio.

1982a. Temporal variability in the sex ratio of a non-social bee, Osmia lignaria
propinqua Cresson. Extrinsic determination or the tracking of an opti-
mum? Oikos 38: 1 77-182.

1982b. Phenotypic variability in nesting success among Osmia lignaria propin-
qua females in a glasshouse environment (Hymenoptera: Megachilidae).
Ecol. Entomol. 7:453^462.

Torchio, P. F.

1975. The biology of Perdita nuda and descriptions of its immature forms and
those of its Sphecodes parasite (Hymenoptera: Apoidea). J. Kansas
Entomol. Soc. 48:257-279.

1976. Use of Osmia lignaria Say (Hymenoptera: Apoidea, Megachilidae) as a
pollinator in an apple and prune orchard. J. Kans. Entomol. Soc.
49:475-482.

Torchio, P. F., and V. J. Tepedino.

1980. Sex ratio, body size and seasonality in a solitary bee, Osmia lignaria
propinqua Cresson (Hymenoptera: Megachilidae). Evolution 34:
993-1003.

Waldbauer, G. P.

1978. Phenological adaptation and the polymodal emergence patterns of
insects, pp. 127-144. In Evolution of Insect Migration and Diapause. H.
Dingle (ed.) Springer-Verlag, New York.

Wallace, D. R., and C. R. Sullivan.

1974. Photoperiodism in the early balsam strain of the Neodiprion abietis
complex (Hymenoptera: Diprionidae). Can. J. Zool. 52:507-513.

A REVIEW OF THE GENUS MALLADA
IN THE UNITED STATES AND CANADA,

WITH A NEW SPECIES (NEUROPTERA: CHRYSOPIDAE)

By Phillip A. Adams1 and J. Allan Garland2

Analysis of the Canadian chrysopid fauna (Garland, 1981) re-
vealed an undescribed species of Mallada ranging into southern
Ontario. Accordingly, a draft description and illustrations were
transmitted to the senior author. As few of our species have been
given modern redescriptions, it is appropriate to review the status of
all four known members of this genus from the U.S. and Canada.

The taxonomic status of Mallada was discussed by Adams 1975,
and a detailed treatment of genitalic morphology given by Principi
1977. The genus is characterized by: left mandible toothed, inner
gradate crossvein of forewing ending in a branch of radial sector,
not on pseudomedia (Fig. 20); pseudomedia not comprising any
crossveins; micropoculae or cuticular glands present on male prono-
tum, microtholi absent, tignum and gonapsis present, arcessus nor-
mal; ectoprocts and hypovalva (eighth and ninth sternites) without
unusual projections, larva trash-carrying and overwintering (Seme-
ria, 1977).

Mallada is primarily an Old World genus, constituting a major
part of the chrysopid fauna of Europe, Africa, India, Southeast
Asia, and Australia. Although New 1980 does not subdivide the
Australian “Chrysopa” into genera, or species groups, it is possible
tentatively to assign species on data given; 15 of the 47 species of
Chrysopinae fall into Mallada. Tjeder 1966 points out that 19 of the
39 African “chrysopas” (Saurius + Glenochrysa + Chrysoperla +
Brinckochrysa + Apertochrysa + Anisochrysa) are assignable to
Anisochrysa (i.e., Mallada ), and places 22 additional Old World
species in that taxon. Aspock et al. 1980 list 15 European Mallada
species (as Anisochrysa). In the New World, there are only 5 known
species, M. (Triadochrysa) triangularis Adams 1978 from Mexico,
and the other North American species discussed below.

•Department of Biology, California State University, Fullerton, California 92634.
department of Entomology, Macdonald College of McGill University, Ste-Anne-
de-Bellevue, Quebec H9X ICO
Manuscript received by the editor August 16. 1982.

239

240

Psyche

[Vol. 89

Mallada macleodi sp. nov.

Description. Head narrow, pale green, eyes large, antennae
unmarked; genae with a shiny narrow black band from margin of
eye, extending to anterior lateral edges of clypeus; labrum black
posterolaterally, green medially (Fig. 3); palpi blackish throughout;
frons raised anteriorly; face with scattered setae, some longer anteri-
orly on clypeus.

Thorax pale green, pronotum with two anterolateral brown
patches; setae whitish. Legs green, pretarsal claws deeply excised.

Abdomen green, setation normal, microtholi absent.

Male terminalia (Fig. 2). Sternites VIII+IX fused, elongate; dor-
sal apodeme long, prominent; ventral apodeme absent. Genitalia
(Fig. 1) with broad transverse tignum; gonarcus expanded laterally,
rectangular dorsally; entoprocessus prominent, expanded ventro-
medially; arcessus broadly continuous with dorsum of gonarcus,
sclerotised proximally, produced caudad and slightly down-curved,
apex bluntly pointed; gonosaccus rudimentary, with only a few
small straight gonosetae positioned between the ventromedial ex-
pansions of the entoprocessus; gonapsis (Figs. 9, 10) three-pronged,
with lateral wings narrow, the caudal process dorsoventrally expan-
sive proximally and tapering as an acuminate downcurved hook,
broad internal saccus terminates in a vertical lobe; gonocristae
sparse and only minutely developed on hypovalva.

Female terminalia. Subgenitale membranous proximally, with
many microthecae; apical lobe notched; transverse callus prom-
inent, with an ental excavation but not a cavity. Spermatheca (Fig.
4) pillbox-shaped; vela tubular, conspicuously bent toward sperma-
thecal bulb.

Wings. Pterostigmata prominently marked, brownish. Venation
narrowly margined with brownish amber, especially in forewing,
gradates of forewing dark, of hind wing amber. Many crossveins of
forewing dark; costals all dark, male with 19 (22.7) 26, female with

Fig. 1-4, Mallada macleodi: 1, dorsal aspect of male genitalia (Ontario); 2, same,

terminalia with structures everted; 3, labrum and mandibles, female, Ontario; 4, same
as 3, spermatheca. Fig. 5-8. Mallada perfectus: 5, dorsal aspect of genitalia;

British Columbia; 6, same as 5, male terminalia with structures everted; 7, labrum
and mandibles, female, British Columbia; 8, same as 7, spermatheca. Scale is for
genitalia and spermathecae.

1982]

Adams & Garland — Genus Mallada

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[Vol. 89

21 (24.2) 26 (mean in parentheses). Inner gradate series of forewing
normal for the genus, terminating on a branch of the radial sector
(Fig. 20). Forewing length: male 10.0 (11.5) 12.5, n=10; female 12.0
(13.0) 13.7, n=10 (mm, mean in parentheses). Type Material. Holo-
type: Texas, Erath Co.: Stephenville, 20. iv. 1981, 3, C. W. Agnew
(MCZ No. 32576).

Paratypes. Arizona. Santa Cruz Co.: 2 mi SW Patagonia,
30. VII. 1948, F. Werner & W. Nutting, rich willow-cottonwood bot-
tom, 4050 ft. (MCZ). Yavapai Co.: Granite Dells 4 mi. N. of Pres-
cott, 28.vii. 1970, 1 <$, L. Martin (LACMNH). Kansas. Manhattan:
VI, 1 <5, R. C. Smith (CNC, det. Smith [as Chrysopa cockerelli\)\ VI
1 <5; 17.VI.1920, 1 9; 19.VIII.1920, 1 9; 9. VII. 1921, 13; 8.IX.1921,
19; 23.VI.1922, 13; 1. VIII. 1922, 13; 6.VIII. 1931, 2 3, 19; 11. VIII.
1931, 1 3, (R. C. Smith, KSU). Ontario. Durham Co.: Kendal,
17.VII.1967, 1 9; 24.VII.1967, 19, J.C.D. Riotte & L. Kohalmi,
ultra-violet light (ROM). Lambton Co.: Pinery Prov. Pk., Ausable
River near riverside campground, 3. VII. 1977, sweeping understory
shrubs, 1 3, E. Oleksuik (ROM #770108). Renfrew Co: White Lake,
4. VIII. 1966, 1 9, P- D. Hebert, ultra-violet light (ROM, in fluid).

Texas. Chisos Mts., 9-10. VII, 1 ? [abdomen missing], 9-12. VII, 1
3, W. Nutting & F. Werner (MCZ Paratype No. 32576). Erath Co.,
Stephenville, 8.V.28.V.1981, 52 specimens, C. W. Agnew (MCZ,
PAAC, CWA). Burnett Co.: Inks Lake St. Park, 4.iv.l981, 1 9, C.
W. Agnew (CWA). Randall Co.: Palo Duro Canyon St. Park,
ll.v.61, 50 specimens, L. Martin, R. H. Reid, W. A. Rees, R. J.
Ford (LACMNH). Maryland. Howard Co.: 1 2. vii. 1 967, 1 3, 1 2,
at white light, E. MacLeod [labelled as “ Chrysopa sp. indet., det. E.
G. MacLeod] (PAAC).

Remarks. The specific epithet recognized Ellis G. MacLeod, who
many years ago collected material of this species, pointed out its
existence to the senior author, and generously presented material
for study. This species is easily separable from luctuosus and sierra
by its lack of black pronotal markings, but is easily confused with
perfectus, which differs in having an entirely black labrum, usually
fewer and paler costal crossveins, and more prominently brown-
bordered venation. It is safest to verify identifications by genitalic
dissection.

1982]

Adams & Garland — Genus Mallada

243

Fig. 9-10. Mallada macleodi: gonapsis, lateral and dorsal views, Texas. Fig.
11-12, M. perfectus: gonapsis, lateral and posteroventral views, Shasta Co.,
Calif. 13-14, M. sierra: gonapsis dorsal, head and thorax, dorsal; Fig. 15-19,
M. luctuosus: 15-16„gonapsis, ventral and lateral views; 17, spermatheca; 18, head
and thorax, dorsal; 19, gonapsis and arcessus, ventral. Fig. 20, M. macleodi,
venation of male forewing, Ontario, showing inner gradate vein ending on a branch
of radial sector (and an extra crossvein, in the last gradate cell, a not-uncommon
condition).

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[Vol. 89

The new species clearly was the insect which the late Dr. R. C.
Smith studied from Manhattan, Kansas (Smith 1922, as Chrysopa
cockerelli). His immatures were debris carriers and overwintered as
larvae, confirming the generic assignment. Smith found adults in
June (CNC), others in August in association with willows; he
deserves credit for observing that the “black lines to mouth not
connecting, though the labrum is light brown” (Smith op. cit.: 1367,
including Fig. 163). Consequently, we have a fairly complete descrip-
tion of the immature stages of the new species, and it is the only
Nearctic representative of Mallada to have been studied in such
detail.

On present evidence, the new species occupies the central part of
the continent, ranging into Canada in southern Ontario and coming
into contact with the more western M . perfectus in Arizona and
New Mexico.

Mallada perfectus (Banks 1895)

Chrysopa perfecta Banks 1895: 516-517. Holotype 9 MCZ No. 1 1914, El Taste, Baja
Calif.

Chrysopa cockerelli Banks 1903:154-155, new synonymy. Holotype 9 MCZ No.
1 1375, East Las Vegas, N. M.

Chrysopa marginalis Banks 1906a:5 (not C. marginalis Navas 1905).

Chrysopa injusta Banks 1906b:98-99, new synonymy. Holotype 9 MCZ No. 1 1374,
Mts. nr Claremont, Calif. (Baker).

Mallada perfectus (Banks), Adams 1975:172.

Description. Genal stripe black, labrum (Fig. 7) wholly black,
palpi black, antennae pale. Body light green with no middorsal
stripe, pronotum with two cinnamon-brown patches. Wings with
costal veinlets black at ends, pale in 'middle, $ costal veinlets:
16-(18.7)-21, N=10; $: 1 8— (20.3)— 22, N=10 (mean in parentheses);
transverse veins conspicuously brown-bordered.

Male terminalia. Apodeme of ninth tergite articulates on short
apodeme of sternites 8+9 (Fig. 6), arcessus (Fig. 5) short, broad,
with lateral subapical projections; gonapsis (Fig. 1 1-12) with spatu-
late emergent process, well-developed arms and simple internal
bulb. Gonocristae small, but larger than in M. macleodi.

Female. Spermatheca with broad-based usually short erect vela.

Distribution. Calif., Ore., Wash., British Columbia, Wyoming,
Utah, Colo., Ariz., N.M., Baja Calif.

1982]

Adams & Garland — Genus Mallada

245

Remarks. This species occurs throughout the Western United
States, but is commonest in the Southwest. The short mediuncus
and spatulate process of the gonapsis readily distinguish the males
from those of M. macleodi, and the females are identifiable by the
broad-based erect vela. Some Arizona specimens have the vela
nearly as elongate as that of macleodi, but never curved.

Mallada sierra (Banks) new combination
Chry sop a sierra Banks 1924:431.

Description. This species is structurally and colorationally like
M. perfectus, except for the following: pronotum with 2 black spots
each surrounded by a patch of cinnamon brown (Fig. 14). Meso-
prescutum with 2 black spots. Wings with black spot at base of
costal area; forewing with black spot at intersection of 2A2 and 3A.
Crossveins darker than in perfectus, and brown-bordering of veins
less pronounced. Gonapsis (Fig. 13) with chisel-shaped reduced
medial process and reduced bulb. Gonocristae less developed than
in perfectus.

Material Examined. Holotype 9, Calif., [Los Angeles Co.], San
Gabriel Mts., Sister Elsie Peak, 10-vi [F. Grinnell] MCZ No. 14858.
Additional: CALIF., Shasta Co., 10 mi. N. Redding, Mountaingate,
1000 ft., 4-8. vi. 1981, 7 9, R. B. Miller (PAAC), 8 mi. N. Redding,
800 ft., 30.v-6.vi. 1981, 4 3, 3 9, R. B. Miller (PAAC). ARIZ. Santa
Rita Mts., 24.vii. 1927, R. H. Beamer (PAAC ex R. C. Smith). ORE.
Jackson Co.: Green Springs, 27.viii.1962, J. S. Buckett, 1 9-
WASH. Yakima Co.: Ft. Simcoe, l.viii.1962, J.F.G. Clarke, 1 9»
(USNM).

Remarks. This species is readily distinguished by the pronotal and
mesonotal markings, and in the male by the simplified structure of
the gonapsis. There has been some question as to whether sierra is a
distinct species or merely a colorational variety of perfectus. In
Shasta County, where these two species are sympatric, sierra
appears early in the season, and perfectus somewhat later, pointing
to the possibility of some seasonal isolation; in 1981, sierra was
relatively common while perfectus was scarce (R. B. Miller, pers.
comm.) thus providing some indication of the independence of popu-
lation fluctuations in these two taxa. The interaction of these two

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[Vol. 89

species is at present under study by C. A. Tauber (pers. comm.),
who reports that they interbreed readily in the laboratory. Despite
this, because of the consistence of the colorational and male geni-
talic differences, plus slightly divergent seasonality, it seems prefer-
able to regard sierra as distinct.

Mallada luctuosus (Banks)

Chrysopa luctuosa Banks 191 1:343.

Mallada luctuosus (Banks) Adams 1975:172.

Description. Green, antennae pale, head and thorax marked with
black and brown as in Fig. 18; brown stripes continue over meta-
thorax and abdomen. Thorax with longitudinal pleural stripe.
Forewings with bases of longitudinal veins, except costa and radius,
dark, transverse veins dark; hind wings less prominently dark-
veined. Abdominal sternites heavily dark-marked.

Genitalia. Arcessus (Fig. 19) elongate. Gonapsis (Fig. 15, 16)
with emergent process thin, ribbonlike apically with seta-like projec-
tions; anterior pocket wide-based. Spermatheca with tubular arcu-
ate vela inserted in doughnut shaped body, ventral impression
small.

Material Examined. Holotype <3, N.M., Ft. Wingate, 26. vi,
MCZ No. 11383. Additional: Arizona. Madera Can., Santa Rita
Mts., 16.viii. 1949, P. Adams (PAAC); Cochise Co.: Huachuca Mts.,
Sunnyside, 14.vii.58, L. Martin (PAAC); Chiricahua Mts., S.W.
Research Station, 5 mi. W. Portal, 5400 ft., 1 . viii. 1966, R. E. Dietz
(PAAC), 28. vi. 1960, J. M. Linsley (U. Calif. Davis); Globe, Pinals,
18.vii.1948, W. Nutting, F. Werner (MCZ). Colorado. Mesa Verde
Nat. Park, Campground, 12.vii.1959, J. & C. Northern (LACMNH).
Nebraska. Meadville, 10.vi.31, B. Patterson (FMNH, Chicago).

Remarks. This species is immediately recognizeable among Mal-
lada species by the dark longitudinal veins and conspicuous black
and brown body markings. It is interesting to note that the forms of
the arcessus, gonapsis, and spermatheca are more similar to those of
macleodi than are those of perfectus, despite the extreme colora-
tional differences.

1982]

Adams & Garland — Genus Mallada

247

Acknowledgements

D. K. McE. Kevan encouraged one of us (J.A.G.) to study the
Canadian chrysopid fauna, which led to discovery of the Ontario
specimens and preparation of the draft manuscript and species de-
scription, including privately financed travel to Ottawa and Boston.
Material for study was loaned by H. D. Blocker, Kansas State
University (KSU); Mary Hathaway, and K. Jepson, Museum of
Comparative Zoology, Harvard University (MCZ); J. E. H. Martin,
Biosystematics Research Institute, Agriculture Canada, Ottawa
(CNC); C. W. Agnew, Texas Agricultural Experiment Station; G.

B. Wiggins and B. D. Marshall, Royal Ontario Museum (ROM),

C. L. Hogue, Los Angeles County Museum of Natural History
(LACMNH). R. B. Miller collected critical material of M. sierra
and perfectus.

References

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Aspock, H., U. Aspock, and H. Holzel

1980. Die Neuropteren Europas. Goeke and Evers, Krefeld, 2 vol.

Banks, N.

1895. Some Mexican Neuroptera. Proc. Calif. Acad. Sci. (Ser. 2) 5:515-522.

1896. A new species of Meleoma. Ent. News 7:95-96.

1903. A revision of the Nearctic Chrysopidae. Trans. Am. ent. Soc. 29:
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1906a. Descriptions of new Nearctic Neuropteroid insects. Trans. Am. ent. Soc.
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1096b. Three new species of Neuroptera. Psyche Camb. 13:98-100.

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1924. Descriptions of new Neuropteroid insects. Bull. Mus. Comp. Zool.
65:419-455, p. I-IV.

Bickley, W. E., and E. G. MacLeod.

1956. A synopsis of the Nearctic Chrysopidae with a key to the genera (Neur-
optera). Proc. ent. Soc. Wash. 58:177-202.

Garland, J. A.

1981. The taxonomy of the Chrysopidae of Canada and Alaska (Insecta: Neu-
roptera). Ph.D. Thesis, McGill University.

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New, T. R.

1980. A revision of the Australian Chrysopidae (Insecta: Neuroptera). Aust. J.
Zool. Suppl. 77:1-143.

Principi, M.

1977. La morfologia addominale ed il sue valore por la discriminazione gene-
rica nell’ambito delle Chrysopinae. Boll. 1st. Entomol. Univ. Bologna
31:325-360.

SfeMfeRIA, Y.

1977. Discussion de la validite taxonomique du sousgenre Chrysoperla Stein-
mann (Planipennia, Chrysopidae). Nouv. Rev. Ent. 7:235-238.

Smith, R. C.

1922. The biology of the Chrysopidae. Cornell University Agric. Exp. St a.
Mem. 58:1287-1372, pi. LV-LVI11.

Tjeder, B.

1966. Neuroptera — Planipennia. The lace-wings of southern Africa. V. Family
Chrysopidae. S. Afr. anim. Life 12:228-534.

POLYGYNY AND POLYDOMY IN THREE NORTH
AMERICAN SPECIES OF THE ANT GENUS
LEPTOTHORAX MAYR (HYMENOPTERA: FORMICIDAE)1

By

Thomas M. Alloway,2 Alfred Buschinger,3 Mary Talbot,4
Robin Stuart,2 and Cynthia Thomas2

General Introduction

This paper deals with certain behavioral and ecological factors
which may be relevant to the evolution and maintenance of social
parasitism in ants. We will argue that some of the same factors
which might predispose one species to evolve into a social parasite
might make resistance to parasitism difficult for a closely related
species.

After their mating flight, the queens of most nonparasitic ant
species found new colonies alone. A queen of such a species finds a
suitable nesting place, excavates a small cavity, and seals herself
inside. She then lays a clutch of eggs and feeds her first larvae a
special “baby food” derived metabolically from the degeneration of
her wing muscles and fat body. These larvae mature to become
female workers which forage for food, enlarge the nest, feed the
queen, and rear subsequent broods of workers and reproductives.
Mature ant colonies usually occupy only one nest (monodomy).
However, the number of queens in typical mature colonies varies.
Colonies of some species never contain more than one functional
queen (monogyny), while colonies of other species often have multi-
ple queens (polygyny) (Buschinger 1974).

However, the queens of all known obligatory slave-making, in-
quiline, and temporary-parasite species found colonies non-inde-

1. This research was supported by grants to Thomas Alloway from the Natural
Sciences and Engineering Research Council of Canada and to Alfred Buschinger
from the Deutsche Forschungsgemeinschaft.

2. Erindale College, University of Toronto, Mississauga, Ontario, CANADA
L5L 1C6.

3. Fachbereich Biologie, Institut fur Zoologie, Technische Hochschule, 61 Darm-
stadt, Schnittspahnstr. 3, Federal Republic of Germany.

4. The Lindenwood Colleges, Saint Charles Missouri, U.S.A. 63301.

Manuscript received by the editor August 5, 1982.

249

250

Psyche

[Vol. 89

pendently. The parasite queen finds a colony of her host species,
enters it, and somehow usurps the role of a host-species queen. The
host-species workers then raise the parasite queen’s brood.

Species of temporary parasites possess a completely functional
worker caste. At first, the temporary-parasite workers and the host-
species workers exist alongside one another. However, when the
host-species workers die, they are not replaced; and a pure colony of
the temporary-parasite species develops. The workers of slave-
making parasites are highly specialized for fighting and raiding the
nests of host-species colonies; and as a consequence of their raids
during which they capture host-species worker pupae and larvae, a
force of host-species workers (or “slaves”) is maintained. Inquiline
parasites either have no worker caste at all; or, if one is present, the
workers seem to play no role in maintaining the colony. In some
cases, a continuing supply of host-species workers is maintained by
the host-species queen’s coexisting with the inquiline queen (Busch-
inger, 1970; Wilson, 1971).

This paper presents data concerning several aspects of the behav-
ioral biology of three North American species of the ant genus
Leptothorax Mayr: L. ambiguus Emery, L. curvispinosus Mayr,
and L. longispinosus Roger. These species interested us because
they are hosts to three closely related parasite species. All three
species are enslaved by the obligatory slave-makers L. duloticus
Wesson and Harpagoxenus americanus (Emery); and L. curvispino-
sus is the host of the workerless inquiline species L. minutissimus M.
R. Smith (Alloway, 1979; Creighton, 1950). Thus, studies of the
behavior and ecology of these three nonparasitic species may eluci-
date the ethological and ecological circumstances under which
social parasitism evolves and is maintained.

Number of Queens and the Sex of Broods in Nests

Headley (1943) and Talbot (1957) reported that the number of
queens in nests of L. curvispinosus and L. longispinosus is quite
variable. Some nests contain several dealate queens, some contain
one, and some contain none at all. Observations indicated that the
number of queens in nests of L. ambiguus is also variable (Alloway,
unpublished data). In addition, we found that many queenless nests
of all three species contained broods which either included worker
and queen pupae at the time of collection or matured into worker
and queen (as well as male) pupae.

1982] Alloway, Buschinger, Talbot, Stuart & Thomas

251

These observations raised a number of hypotheses. Nests contain-
ing more than one dealate queen suggested that some colonies of L.
ambiguus, L. curvispinosus, and L. longispinosus are polygynous.
The production of female pupae in queenless nests raised at least
three possibilities which are not mutually exclusive. First, a queen-
less nest might be part of a polydomous colony with the female
pupae being the progeny of one or more queens located in another
nest at the time of collection. Second, these species might possess
numerous ergatomorphic reproductives, individuals which resemble
workers morphologically but which have a spermatheca, can be
inseminated, and are capable of laying fertilized female eggs (Busch-
inger 1975, 1978). Third, a queenless nest might be the remnant of a
colony whose queen had died.

Materials and Methods

Over a two-year period, nests of L. ambiguus, L. curvispinosus,
and L. longispinosus were collected during late March, April, May
and early June; and weekly collection of L. ambiguus and L. longi-
spinosus were obtained throughout June, July, and August of one
summer. We recorded the number of queens present in every nest.
In nests containing pupae at the time of collection, the kind of
pupae present (queen, worker, and/or male) was also noted.
Finally, nests of all three species were collected during the early
spring of one year and cultured in the laboratory to determine the
sex and caste of the pupae which matured from larvae present in the
nests at the time of collection.

Results

Table 1 contains data regarding the proportions of nests collected
during the springs of two years which contained 0, 1, or more than 1
queen. About 1/5 of the nests contained more than one dealate
queen; about 1/3 contained no queen; and the remainder contained
1 queen. Tables 2, 3, and 4 reveal that the proportion of queenless
nests was similar across years and throughout the season.

Table 2 presents the numbers and proportions of nests of all three
species collected in the spring and containing pupae of various
kinds. Table 3 presents similar data for nests of L. ambiguus and L.
longispinosus collected throughout the summer. These tables reveal
that many freshly collected queenless nests contained female (worker
and queen) pupae. Table 4 presents data concerning the broods

252

Psyche

[Vol. 89

Table 1. Number and Percent of Nests of L. ambiguus, L. curvispinosus, and
L. longispinosus Containing 0, 1, or More Than 1 Queen

Number of
Queens

L.

ambiguus

L.

curvispinosus

L.

longispinosus

Total

0

453 (29.7%)

177 (36.3%)

237 (37.0%)

867 (32.7%)

1

765 (50.3%)

228 (46.7%)

311 (48.6%)

1304 (49.2%)

More than 1

304(20.1%)

83(17.0%)

92(14.4%)

479 (18.1%)

Total

1522 (100.0%)

488 (100.0%)

640(100.0%)

2650(100.0%)

which matured from queenless and queenright nests of the three
species collected in the early spring and then cultured in the labora-
tory. Once again, many queenless nests produced female pupae.

Discussion

First, we want to stress that variability in the number of queens in
nests of L. curvispinosus and L. longispinosus, first noted by Head-
ley (1943) and Talbot (1957), is not a local or transitory pheno-
menon and note that the number of queens in nests of L. ambiguus
is also quite variable. However, of far greater importance is the large
proportion of queenless nests of all three species which produce
female (as well as male) pupae. This fact raised questions about the
possible existence of ergatomorphic reproductives and polydomy.

Polygyny and Worker Fertility

To demonstrate that a species of ant is facultatively polygynous,
one must show that two or more fertile inseminated females can
coexist in nests. Headley (1943) and Talbot (1957) reported the
occurrence of multiple queens in some nests of L. curvispinosus and
L. longispinosus. However, these authors did not determine whether
more than one queen was inseminated and egg-laying. Wilson
(1974a, b) observed several multiple-queen nests of L. curvispinosus
and reported that all the queens laid eggs. However, as we shall
show, uninseminated queens and workers sometimes lay eggs. Thus,
the question of the occurrence of polygyny involving fertile in-
seminated queens remained open. In addition, the production of
female pupae in many queenless nests of L. ambiguus, L. curvispino-
sus, and L, longispinosus suggested, as one possibility, the hypothe-
sis that these species might possess frequent ergatomorphic female
reproductives.

1982] Alloway, Buschinger, Talbot, Stuart & Thomas

253

Table 2. Number and Percent of Queenright and Queenless Nests of L.
ambiguus, L. curvispinosus, and L. longispinousus Containing Pupae and/or
Alate Reproductives of Various Types at the Time of Collection (1977-78)

Species

9 and/or
$ Only

Queenright Nests

9 and/or
$ and $

3 Only

Total

L. ambiguus

180 (83.3%)

31 (14.4%)

5 (2.3%)

216(100.0%)

L. curvispinosus

23 (53.5%)

19(44.2%)

1 (2.3%)

43 (100.0%)

L. longispinosus

76(66.1%)

33 (28.7%)

6 (5.2%)

115(100.0%)

Total

279 (74.6%)

83 (22.2%)

12(3.2%)

374(100.0%)

Species

9 and / or
§ Only

Queenless Nests

9 and/or
$ and $

3 Only

Total

L. ambiguus

90 (80.4%)

16(14.3%)

6 (5.4%)

112(100.0%)

L. curvispinosus

19(59.4%)

9(28.1%)

4(12.5%)

32(100.0%)

L. longispinosus

35 (50.0%)

24 (34.3%)

11 (15.7%)

70(100.0%)

Total

144 (67.3%)

49 (22.9%)

21 (9.8%)

214(100.0%)

Materials and Methods

To determine whether polygyny involving inseminated queens
occurs in these species, we dissected all the queens present in sam-
ples of nests containing more than one dealate queen. To determine
whether ergatomorphic female reproductives occur frequently, we
dissected all the “workers” from five queenless nests of each species
which had produced female broods when cultured in the laboratory.

For each queen or worker dissected, we noted the following
characteristics:

a. the number of ovarioles.

b. the length of the ovaries. In young virgin queens, the ovaries are
thin and about 3/4 the length of the queen’s gaster. When a
queen becomes fertile, her ovaries grow until they eventually
become as long as her entire body. In old fertile queens, the
folded and coiled ovarioles enlarge until they almost completely
fill the gaster.

c. the presence or absence of any growing oocytes in the ovarioles.
The ovarioles of sterile individuals contain no oocytes; and in
hibernating fertile queens, the oocytes are transparent. As yolk is

254

Psyche

[Vol. 89

deposited in growing oocytes, they become opaque; and ripe
eggs are white.

d. the presence or absence of corpora lutea in the bases of the
ovarioles. These yellowish residues of nutritional cells remain in
the ovaries when eggs have been laid.

e. the presence or absence of a full or empty spermatheca. Individ-
uals with no spermatheca or an empty spermatheca are incap-
able of laying fertilized eggs which develop into workers or
queens. An empty spermatheca appears as a small, transparent
bladder on the common oviduct. When full of sperm, the sperma-
theca is white and superficially resembles a ripe egg in size and
coloration.

Results:

Our dissections enabled us to distinguish several physiologically

different kinds of queens. To simplify the presentation of data, we

Table 3. Number of Queenright and Queenless Nests of L. ambiguus and
L. longispinosus Collected during June, July, and August and the Composition of
their Broods

L. ambiguus
Queenless nests

Queenright nests

Type of Brood

Type of Brood

9,

9, S,

$ only 9 and $ and $

$ only 9 and $

and S

June

0

5 0

0 28

0

July

0

11 17

1 37

71

August

2

9 8

2 32

77

L. longispinosus

Queenless nests

Queenright nests

Type of Brood

Type of Brood

9,

9,

S only 9 and $ and S

$ only 9 and $

and $

June

0

0 1

0 2

1

July

0

0 4

0 1

12

August

0

0 1

0 0

3

1982] Alloway, Buschinger, Talbot, Stuart & Thomas

255

will employ a set of terms developed by Buschinger (1968) to de-
scribe these individuals. These terms are defined as follows:
A-queen: An inseminated, fully fertile queen. The ovaries are as
long, or nearly as long, as the whole body. The ovarioles con-
tain many developing oocytes; and conspicuous corpora lutea
are present. The spermatheca is full of sperm. Such queens are
normally more than a year old.

b-queen: An inseminated young queen. At the time of our study (in
mid-summer), the ovaries were about half their eventual length
and contained developing oocytes. Sometimes a small corpus
luteum was visible in the base of one or two ovarioles. The
spermatheca was full. We believe that these females had mated
the previous summer and were in the process of becoming fully
fertile. After mating, newly inseminated queens have very short
ovarioles with no developing oocytes. If a nest, before the mat-
ing season, contains one or more A-queens and one or more
b-queens with growing oocytes, we conclude that that nest
represents all or part of a colony which adopted one or more
newly mated queens the previous summer.

Table 4. Number and Percent of Queenright and Queenless Nests of L. ambiguus,
L. curvispinosus and L. longispinosus Collected in the Spring of 1979 which
Produced Broods of Various Compositions when Cultured in the Laboratory

Queenright Nests

Species

9 and/or
5 only

9 and/or
5 and $

S only

Total

L. ambiguus
L. curvispinosus
L. longispinosus
Total

68 (47.5%)
95 (65.5%)
42 (56.0%)
205 (56.5%)

60 (42.0%)
49 (33.8%)
22 (29.3%)
131 (36.1%)

15 (10.5%)
1 ( 0.7%)
11 (14.7%)
27 ( 7.4%)

143 (100.0%)
145 (100.0%)
75 (100.0%)
363 (100.0%)

Queenless Nests

Species

9 and/or
$ only

9 and/or
5 and $

S only

Total

L. ambiguus
L. curvispinosus
L. longispinosus
Total

37 (52.1%)
35 (43.2%)
12(38.7%)
84 (45.9%)

24 (33.8%)
38 (46.9%)
12(38.7%)
74 (40.4%)

10 (14.1%)
8 ( 9.9%)
7 (22.6%)
25 (13.7%)

71 (100.0%)
81 (100.0%)
31 (100.0%)
183 (100.0%)

256

Psyche

[Vol. 89

c-queen: An uninseminated, old, sterile female. The ovaries are
short and contain no oocytes. The spermatheca, if present, is
empty; but it may not be present. The wing muscles are degen-
erate and have been replaced by fat body. (The term d-queen
would denote a young dealate female which had not been
inseminated. The reproductive organs resemble those of c-
queens, but the wing muscles have not yet degenerated. We
found no d-queens, probably because we performed our dissec-
tions before the sexual brood had eclosed.)

C-queen: An uninseminated, egg-laying female with ovarioles like
those of an A-queen. Sometimes there is no spermatheca. In
this paper, we report the occurrence of significant numbers of
individuals of this type for the first time in Leptothoracine ants.
However, they occur rather frequently in colonies of Formica
polyctena Foerster (Ehrhardt 1970) and Monomorium pha-
raonis (L.) (Petersen & Buschinger 1971). The origin of these
females in nests of L. ambiguus, L. curvispinosus, and L. longi-
spinosus is unclear. They may be old individuals which were
once inseminated but whose supply of sperm has been ex-
hausted. However, the existence of egg-layers with no sperma-
theca indicates that insemination is not a necessary prerequisite
for fertility. Recently U. Winter (personal communication)
found that Harpagoxenus sublaevis males often transmit very
little or no sperm during their first copulation. Thus, a queen
which had mated only once with such a male might become
fertile after receiving only the secretions of the males’s acces-
sory glands. Perhaps a similar mechanism accounts for the
existence of C-queens in these species of Leptothorax.

The results of the dissections of queens of each species and of
workers will be presented separately.

1 . Leptothorax ambiguus

A total of 88 dealate females from 30 multiple-queen colonies was
dissected. Only about 1 /2 the multiple-queen nests contained more
than one A-queen and were thus “truly polygynous” (see Table 5).
Three of these truly polygynous nests also contained one or two
b-queens and were thus in the process of developing polygyny to a
higher degree.

1982]

Alloway, Buschinger, Talbot, Stuart & Thomas

257

Table 5. Number and Type of Dealate Females in Multiple-Queen Colonies of

Leplothorax ambiguus

Colony

No.

n Dealate

99

A-99

b-99

c-99

c-99

Remarks

,

2

2

_

_

_

2

2

2

-

-

-

3

2

2

-

-

-

4

2

2

-

-

-

5

2

2

-

-

-

6

2

2

-

-

-

Colonies No.

7

2

2

-

-

-

1-15 are truly

8

2

2

-

-

-

polygynous

9

4

2

2

-

-

10

3

2

-

-

i

1 1

3

3

-

-

-

12

3

2

1

-

-

13

3

2

1

-

-

14

5

4

-

-

i

15

13

2

-

3

8

16

2

1

1

-

-

17

2

1

1

-

-

18

2

1

1

-

-

Colonies No.

19

2

1

1

-

-

1 6-23 are

20

3

1

2

-

-

becoming

21

3

1

1

-

1

polygynous

22

3

1

I

-

1

23

2

-

2

-

-

24

2

1

-

-

1 +

Colony fragment?

25

2

-

1

-

1 +

+C-9 without

26

2

1

-

-

1

spermatheca

27

2

1

-

-

1

28

3

-

1

-

2

29

2

-

-

2

-

Colony fragment?

30

6

-

-

2

4

Total

88

43

16

7

22

Another 7 nests (No. 16-23 in Table 5) were in the process of
becoming polygynous. They contained 1 A-queen and 1 or 2 b-
queens. One nest (No. 23) contained 2 b-queens only and was thus
also becoming polygynous, although it lacked an A-queen. A
number of nests contained one or more C-queens. Most of these

258

Psyche

[Vol. 89

individuals were living with A-queens. Two C-queens without a
spermatheca were found in this sample (in nests No. 24 and 25).

2. Leptothorax curvispinosus

A total of 64 dealate queens from a sample of 23 multiple-queen
nests was dissected. As was the case for L. ambiguus, we found all
four categories of dealate females in L. curvispinosus (see Table 6).
However, approximately 3/4 of the curvispinosus nests (74%) con-
tained multiple A-queens, as compared to only about 1/2 of the
ambiguus nests. In addition, all 7 of the multiple-queen curvispino-
sus nests which had only 1 A-queen contained one or more b-queens
and were thus becoming polygynous. The total number of C-queens
was much lower in curvispinosus than in ambiguus. However, we
found 3 C-queens with no spermatheca; and 2 of these were fully
fertile.

3. Leptothorax longispinosus

A total of 79 queens from a sample of 26 multiple-queen nests
was dissected. The proportion of nests containing more than one
A-queen was 65%; and all but one of the other nests contained either
one or more b-queens living with an A-queen or more than one
b-queen without an A-queen (see Table 7). The only exception was
nest No. 23 which contained 7 C-queens living with a single A-
queen. One of these C-queens had no spermatheca.

4. The number of ovarioles in queens

Table 8 shows that queens of L. ambiguus usually have 6 ovari-
oles (both ovaries combined). Six is the usual number of ovarioles
for most European species of the subgenus Leptothorax sensu
stricto (=Myrafant M. R. Smith 1950) and for species of the subge-
nus Mychothorax ( ^Leptothorax sensu M. R. Smith) (Buschinger,
unpublished data). However, L. curvispinosus queens most
commonly have 8 ovarioles; and L. longispinosus queens most com-
monly have 7. Moreover, the number of ovarioles in L. longispino-
sus queens is very variable; and the distribution of ovarioles in single
specimens of this species can be quite asymmetrical. One queen with
10 ovarioles had 6 on the left side and 4 on the right; another with 1 1
ovarioles had 4 on the left and 7 on the right. There was no evidence
that the number of ovarioles is correlated with a queen’s function in
a nest. The number of ovarioles often varied considerable among

1982] Alloway, Buschinger, Talbot, Stuart & Thomas

259

Table 6. Numbers and Type of Dealate Females in Multiple-Queen Colonies of

Leptothorax curvispinosus

Colony

n Dealate

No.

9$

A-99

b-99

c-99

c-99

Remarks

1

2

2

.

.

.

2

2

2

-

-

-

3

2

2

-

-

-

4

2

2

-

-

-

5

2

2

-

-

-

6

2

2

-

-

-

Colony No.

7

2

2

-

-

-

1-16 truly

8

2

2

-

-

-

polygynous

9

2

2

-

-

-

10

3

3

-

-

-

11

3

3

-

-

-

12

3

3

-

-

-

13

4

4

-

-

-

14

4

2

2

-

-

15

4

3

1

-

-

16

4

3

-

i +

+C-9without

17

2

1

1

-

-

spermatheca

18

2

1

1

-

-

Colony No.

19

4

1

3

-

-

17-23

20

3

1

1

l+

-

becoming polygynous

21

3

1

1

-

l

+c-$ without

22

3

1

1

-

l

spermatheca

23

4

1

3

+C-9 without
spermatheca

Total

64

46

14

i

3

queens in single nests, especially in L. longispinosus. Moreover, b-
and C-queens on average had no fewer ovarioles than A-queens.

5. Workers

All the queenless nests whose workers were dissected contained
one or more egg-laying individuals (see Table 9). However, none of
the fertile workers possessed a spermatheca. Thus, we presume that
all their offspring are males. Workers invariably had only two ovari-
oles (one per ovary); and these were never as long and never con-
tained as many corpora lutea as the ovarioles of egg-laying A- and
C-queens. Thus, the number of eggs produced by a fertile worker is
probably much less than that produced by a queen.

260

Psyche

[Vol. 89

Table 7. Numbers and Type of Dealate Females in Multiple-Queen Colonies of
Leptothorax longispinosus

Colony

No.

n dealate

$9

A-9$

b-9$

c-99

Remarks

1

2

2

_

_

2

2

2

-

-

3

2

2

-

-

4

2

2

-

-

5

2

2

-

-

6

2

2

-

-

7

2

2

-

-

8

2

2

-

-

Colonies No. 1-17 truly

9

2

2

-

-

polygynous

10

2

2

-

-

11

3

3

-

-

12

3

3

-

-

13

4

4

-

-

14

4

4

-

-

15

6

6

-

-

16

3

2

1

-

17

3

2

1

-

18

2

1

1

-

19

2

1

1

-

Colonies No. 18-22 and

20

3

1

2

-

No. 24-26 becoming

21

3

1

2

-

polygynous

22

4

I

3

-

23

8

1

-

7+

+ lC-9 without spermatheca

24

2

-

2

-

25

4

-

4

-

26

5

-

5

-

Total

79

50

22

7

Table 8. Ovariole Numbers
spinosus, and L. longispinosus

in

Queens of Leptothorax ambiguus,

L. eurvi-

n. ovarioles 4 5

6

1

8

9

10

11

n$$

L. ambiguus 1 1

82

4

-

-

-

-

88

x = 5.99
s = 0.39

L. curvispinosus

-

11

44

6

-

-

61

x = 7.92
s = 0.53

L. longispinosus 1 4

19

32

18

1

2

2

79

x = 7.05
s = 1.22

1982]

Alloway, Buschinger, Talbot, Stuart & Thomas

261

Discussion:

These data establish two important points. First, polygyny involv-
ing multiple inseminated queens occurs in some nests of L. am-
biguus, L. curvispinosus, and L. longispinosus; and polygynous
nests imply the existence of polygynous colonies. Polygyny in these
three members of the subgenus Leptothorax sensu stricto as well as
in L. schaumi and L. flavicornis (Buschinger, unpublished observa-
tions) is somewhat surprising in that the majority of European
members of the subgenus are strictly monogynous (Buschinger
1967). The form of polygyny exhibited by L. ambiguus, L. curvispi-
nosus, and L. longispinosus is also interesting in that the frequent
joint presence of A- and b-queens indicates that colonies of these
species can adopt young conspecific queens. We will argue below
that this tendency to adopt queens is important for understanding
the evolutionary origins of parasitic colony foundation.

Second, although our dissections of workers in queenless colonies
which produced female pupae revealed that some workers lay eggs,
our failure to find any workers with a spermatheca indicates that
ergatomorphic reproductive females of the kind seen in the slave-
maker Harpagoxenus sublaevis are at least not common in L. am-
biguus, L. curvispinosus, and L. longispinosus.

Polydomy

This latter finding suggested two possibilities which are not mutu-
ally exclusive:

a. Some queenless nests of these species which produce broods
containing female pupae may be parts of polydomous colonies.
In such cases, the female pupae would be the progeny of queens
located in other nests at the time of collection.

b. Some queenless nests may represent declining colonies with no
queen. The female pupae are the offspring of a dead queen.

Materials and Methods

We collected groups of acorn nests which were very close together
in nature and brought the nests back to the laboratory where the
ants were established in artificial nests. We then arranged the artifi-
cial nests in arenas to duplicate the spatial arrangement of the natur-
al nests and observed the ensuing behavioral interactions. As
controls, we tested the effect of placing nests from different parts of

262

Psyche

[Vol. 89

Table 9. Numbers and Percent of Sterile and Fertile Workers in Queenless Nests
of L. ambiguus, L. curvispinosus, and L. longispinosus

L. ambiguus

Nest No.

Sterile Workers

Fertile Workers

Total

1

7 (64%)

4 (36%)

1 1

2

9 (60%)

6 (40%)

15

3

12(71%)

5 (29%)

17

4

12(86%)

2 (14%)

14

5

24 (80%)

6 (20%)

30

L.

curvispinosus

Nest No.

Sterile Workers

Fertile Workers

Total

1

13 (93%)

1 ( 7%)

14

2

18 (72%)

7(18%)

25

3

17 (81%)

4(19%)

21

4

16 (70%)

7 (30%)

23

5

15 (71%)

6 (29%)

21

L. longispinosus

Nest No.

Sterile Workers

Fertile Workers

Total

1

12 (67%)

6 (33%)

18

2

5 (56%)

4 (44%)

9

3

5(21%)

19 (79%)

24

4

17 (74%)

6 (26%)

23

5

7 (50%)

7 (50%)

14

the same collection site much closer together than they had been
found and of placing nests from different sites together.

Two kinds of arenas were employed. One type consisted of a 1 m2
area on a table top. The other was a square plexiglass enclosure
having an area of 2025 cm2 surrounded by plexiglass walls 6 cm
high. The ants were confined to the arenas by a thick barrier of
petroleum jelly. Colonies were fed an artificial ant diet (Bhatkar &
Whitcomb 1970) three times a week; water was continuously availa-
ble. The experimental room was kept on a 15-h light and 9-h dark
photoperiod at a temperature of 22° C ±1°C.

Results

A total of 28 experiments involving 96 nests of L. ambiguus and 5
experiments involving 1 1 nests of L. longispinosus were performed.

1982] Alloway, Buschinger, Talbot, Stuart & Thomas

263

Certain pertinent facts about each experiment are contained in
Table 10.

The most frequent result for nests which had been close together
in nature was so-called “fusion”. After a day or two, the ants from
the different nests peacefully moved into one of the artificial nests
and remained there indefinitely. We are not sure why fusion
occurred so frequently in the laboratory. One factor may have been
that our artificial nests are somewhat larger than the average acorn.
In any case, these peaceful mergers suggest that the ants from adja-
cent nests were members of the same colony and are thus compati-
ble with the polydomy hypothesis.

Other experiments (e.g. L. ambiguus experiments 9, 10, and 23
and L. longispinosus experiment 5) supported the polydomy hy-
pothesis more dramatically. The ants continued to occupy more
than one nest among which they maintained a more or less contin-
ual exchange of workers, brood, and queens. Thus, over a period of
several days, a nest was sometimes polygynous, sometimes monog-
ynous, and sometimes queenless. In other experiments, (e.g. L.
ambiguus experiments 12, 13, 14, 15, 16, 22, 24), it appeared that we
observed interactions between two polydomous colonies or between
a polydomous and a monodomous colony. For example, in experi-
ments 15 and 16, we had examples of four nests which had been
found in two close pairs separated by a somewhat greater distance.
The ants from each pair of nests quickly fused, but there was pro-
longed fighting among the ants from the different pairs of nests.

The results of the control experiments also supported the poly-
domy hypothesis. Ants from nests not found close together in
nature did not usually coexist peacefully. When nests from different
parts of the same collection site or from different sites were placed
near one another, the result was usually widespread and protracted
fighting. However, we observed two exceptions to this rule. In L.
ambiguus experiment 19, 3 nests which had been an average of
96 cm apart in nature were placed together in a 2025-cm2 arena.
There was no fighting; and after 12 days, the ants from two queen-
right nests which had been 1 18 cm apart in nature peacefully moved
into one nest. Even more surprising was the fusion of ants in two
queenright nests from different collection sites which we observed in
L. ambiguus experiment 18. We cannot explain these anomalous
results, although we speculate that these species have a limited

Table 10. Results of Polydomy Study
Experiments with L. ambiguus

264

Psyche

[Vol. 89

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1982] Alloway, Buschinger, Talbot, Stuart & Thomas

265

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Experiments with L. longispinosus

266

Psyche

[Vol. 89

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1982] Alio way, Buschinger, Talbot, Stuart & Thomas

267

number of possible “colony odors”. Ants from colonies with differ-
ent odors fight, but ants from colonies with sufficiently similar
odors do not fight and may merge (for a discussion, see Holldobler
and Wilson, 1977).

Finally, although some data indicate that polydomy occurs in L.
ambiguus and L. longispinosus, other data indicate that monodomy
also occurs. As already noted, many apparently polydomous colo-
nies became monodomous in the laboratory. Similar fusions may
occur in nature. In addition, several experiments suggested interac-
tions either between a polydomous and a monodomous colony or
between two or more monodomous colonies. There were several
instances in which the ants from two or more nests merged and then
attacked the ants from another single nest. These results suggest that
the ants which merged had comprised a polydomous colony and
that the ants which were attacked belonged to another colony.
Finally, the results of L. ambiguus experiments 25 and 26 suggested
interactions among three monodomous colonies; and those of
experiments 27 and 28 suggested interactions between two monod-
omous colonies.

Discussion

The evolutionary significance of polydomy and the question of
what proportion of the queenless nests producing broods containing
female pupae can be accounted for by polydomy will be discussed
below. Here we simply note that some of the queenless nests of L.
ambiguus and L. longispinosus which produce broods containing
female pupae are almost surely parts of larger polydomous colonies
in which there happened to be no queen at the time of collection. In
the absence of data, it would be premature to conclude that polyd-
omy occurs in L. curvispinosus. However, queenless nests are com-
mon in L. curvispinosus; and this fact and the many other similarities
between L. curvispinosus on the one hand and L. ambiguus and L.
longispinosus on the other suggest that L. curvispinosus is also
facultatively polydomous.

Colony Foundation

One sign of an incipient ant colony is a nest containing one or
more queens, an immature brood, and no workers. Such apparently
incipient colonies of L. ambiguus, L. curvispinosus, and L. longi-

268

Psyche

[Vol. 89

spinosus are not easy to find. Under oak and hickory trees where
there have been abundant nut falls, most inhabited nuts are occu-
pied by more mature colonies. However, over several years, we
discovered several apparently incipient colonies of L. ambiguus and
L. longispinosus.

Materials and Methods

We searched for incipient colonies of L. ambiguus and L. longi-
spinosus in late summer and early autumn. An incipient colony was
defined as a nest containing one or more dealate queens with a
brood, but no workers.

Results

A total of 15 apparently incipient nests was found, 8 of L. am-
biguus and 7 of L. longispinosus. Table 1 1 lists the number of
queens and the type of brood present when the nests were collected.

We tried to culture incipient colonies in the laboratory. However,
perhaps because the artificial nests lacked a source of moisture, we
had little success. Although the queens (perhaps unnaturally) for-
aged for food and water, their broods gradually languished and
died. Only L. ambiguus nest 7 and L. longispinosus nest 5 produced
workers in the laboratory.

Table 1 1 shows that the number of queens in apparently incipient
nests of L. ambiguus ranged from 2 to 10; and the number of queens
in apparently incipient nests of L. longispinosus ranged from 1 to
15. These data indicate that queens of L. longispinosus found new
colonies on a facultatively polygynous basis. So far we have failed to
find an instance of apparently monogynous colony foundation in L .
ambiguus. However, it would be premature to conclude that polygy-
nous colony foundation in L. ambiguus is obligatory.

Since we were mainly interested in the behavior of colony-
founding queens, we did not dissect the foundresses to determine
their reproductive status. However, the presence of male pupae in L.
ambiguus nests 4 and 8 suggests that one or more of the queens may
have become fertile without insemination.

Multiple colony foundresses showed no hostility toward one
another. To the contrary, apparently “cooperative” acts were com-
mon. All brood was kept in a single pile and seemed to be tended
jointly. Mutual grooming was frequent; and queens often regurgi-
tated to one another upon returning from foraging trips. Some
groups of queens “took turns” foraging.

1982]

Alloway, Buschinger, Talbot, Stuart & Thomas

269

Table 1 1. Apparently Incipient Colonies of L. ambiguus and L. longispinosus

Incipient Colonies of L. ambiguus

Colony No. Number of 9$ Brood When Collected

1

2

3

4

5

6

7

8

5

3
2

10

2

2

4
4

eggs and larvae
eggs and larvae
eggs, larvae, pupae
eggs, larvae, pupae
larvae

eggs and larvae
eggs and larvae
eggs, larvae and pupae

Incipient Colonies of L. longispinosus
Colony No. Number of $9 Brood When Collected

1

2

3

4

5

6
7

1
1
1

5

2

1

1

Nil

eggs, larvae, pupae
Nil

eggs and larvae

larvae

larvae

eggs, larvae, pupae

Discussion

These data indicate that colonies of L. longispinosus can be
founded either by a single queen (haplometrosis) or by more than
one queen (pleometrosis) and that colonies of L. ambiguus can be
founded pleometrotically. These preliminary findings indicate that
the colony-foundation behavior of L. ambiguus, L. curvispinosus,
and L. longispinosus deserves more thorough investigation. Among
the questions remaining to be answered are the following:

a. Can pleometrosis in these species lead directly and smoothly to
polygynous mature colonies; or is there an obligatory period of
monogyny between a colony’s pleometrotic beginnings and the
later adoption of supernumerary queens (Holldobler & Wilson
1977)?

b. How closely related are multiple colony foundresses? Are they
always sisters? If so, how do they get together to found a new
colony?

c. Is foraging for food and water a laboratory artifact; or do
colony-founding queens of these species normally forage?

270

Psyche

[Vol. 89

Finally, although we have no direct evidence, we suppose that
many colonies of these species must originate when a queenright
portion of a polydomous colony becomes permanently separated
from the other parts, a process known as “budding”. Incipient nests
containing only queens and an immature brood seem too rare to
account for all colony foundation in these species.

General Discussion

We can now reconstruct the colony life histories of these species
in some detail. New colonies of L. longispinosus can be established
either by a single newly mated young queen (haplometrosis) or by
two or more such individuals (pleometrosis). New colonies of L.
ambiguus are established pleometrotically; and it seems likely that
further research will establish that colonies of this species and of L.
curvispinosus can be founded either pleometrotically or haplome-
trotically. Young colonies of these species probably occupy only one
nest (monodomy). However, as they grow, some colonies of L.
ambiguus and L. longispinosus come to occupy two or more nests
(polydomy) among which there can be an exchange of workers,
brood, and queens. Mature colonies of all three species containing
one or more fully fertile inseminated queens also sometimes adopt
additional conspecific queens. Finally, we hypothesize that new col-
onies can be formed as a result of the break-up of polydomous
colonies into two or more autonomous units (budding).

When considering these facts, one immediately notes a large
amount of behavioral variability. Although we do not yet know
whether any individual queen is potentially capable of doing more
than one thing, young queens as a class can either join an estab-
lished colony, found a new colony alone, or found a new colony in
the company of one or more other queens. Colony life cycles and
demographies are also variable. A colony can apparently have one
or more queens at almost any stage of its development and can
occupy one or more than one nest when mature enough to produce
reproductives. Such behavioral variability is unusual, and its adap-
tive significance is obscure. Thus, the behavioral ecology of these
three species offers many opportunities for empirical and theoretical
analysis.

Two problems are particularly salient. First, we have demon-
strated that some queenless nests are parts of polydomous colonies;

1982] Alloway, Buschinger, Talbot, Stuart & Thomas

271

and we presume that others are remnants of declining colonies.
However, we can neither distinguish the two kinds of nests nor
determine their relative frequencies. Reference to the proportion of
nests producing all-male broods is not helpful because, in some
species of Leptothoracine ants, female larvae can hibernate twice
before pupating (Buschinger et al. 1975). Thus, a queenless colony
might continue to produce female pupae for one or two years.
Further work is needed to devise a simple means of distinguishing
declining colonies from the queenless nests of polydomous colonies.

Second, we would like to know how frequently these species
employ the various modes of colony foundation which we have
observed and postulated. Altogether, we report observations of 872
nonincipient nests of L. ambiguus, 342 nonincipient nests of L.
longispinosus, and of 8 and 7 apparently incipient nests of these two
species. If one assumed that the frequency of apparently incipient
nests represented the frequency of incipient colonies in the popula-
tion, one would have to conclude that the average lifespan of a
colony is unreasonably long. Thus, we were led to propose budding
as a frequent means of colony foundation. This proposal needs
verification.

However, it was the degree to which L. ambiguus, L. curvispino-
sus, and L. longispinosus are subject to social parasitism which
initially aroused our interest; and several of the behavioral processes
which we have described suggest means by which social parasitism
might either evolve or be maintained. Colonies of all three species
sometimes adopt newly mated conspecific queens, and colonies of
L. ambiguus and L. longispinosus are sometimes founded pleo-
metrotically. Since both these forms of polygyny require the peace-
ful coexistence of queens and of workers which are the offspring of
different queens, both forms of polygyny are factors which might
render these species susceptible to social parasitism. To be accepted
by a host-species colony, a parasite queen must somehow convince
the host workers and perhaps the host queen or queens that she is a
legitimate potential colony member. Since the queens and workers
of these species naturally accept supernumerary queens, the parasite
female’s task is probably simplified.

Moreover, the tendency to seek adoption by existing colonies and
the tendency to join pleometrotic foundress associations may repre-
sent preadaptive traits from which parasitic modes of colony founda-

272

Psyche

[Vol. 89

tion might have evolved in such a group of closely related species.
The queens of slave-making, temporary, and inquiline parasites
found new colonies by securing adoption in a host-species colony
(Buschinger 1970; Wilson 1971). Although the colony-foundation
behavior of such social parasites often involves an element of vio-
lence which is probably lacking from the processes by which colo-
nies of L. ambiguus, L. curvispinosus, and L. longispinosus adopt
additional conspecific queens or additional foundresses join associa-
tions (Wesson 1939; Alloway, personal observations), the tendencies
to join conspecific colonies or foundress associations could form a
basis from which more elaborate parasitic colony-foundation might
evolve.

Finally, the kind of polydomy seen in L. ambiguus, and L. longi-
spinosus also embodies factors which may be both preadaptive for
the evolution of socially parasitic behavior and significant in render-
ing a species subject to social parasitism. Polydomy in these species
can involve a more or less continuous exchange of workers, brood,
and queens among a colony’s multiple nests. Such commerce
requires a worker caste which is adept in carrying brood and adults
in a fashion which might be preadaptive for slave-raiding (Busch-
inger 1970). In this context it is noteworthy that Wilson (1975) and
Alloway (1980) have shown that L. ambiguus, L. curvispinosus, and
L. longispinosus sometimes behave like facultative slave-makers.

Polydomy also requires workers in one nest to accept and tend a
brood from another nest even though it may carry a somewhat
unfamiliar “nest odor”. Yet, any tendency to care for unfamiliar
brood might render a species vulnerable to social parasitism. The
more ready host-species workers are to accept unfamiliar brood, the
less exactly a parasite’s brood need mimic that of the host.

Summary

New colonies of L. longispinosus can be founded by a single
young queen; and colonies of L. ambiguus and L. longispinosus can
be founded by groups of two or more young queens. Mature colo-
nies of these two species and of L. curvispinosus can become polyg-
ynous or enhance the degree of their pre-existing polygyny by
adopting young conspecific queens. Some colonies of L. ambiguus
and L. longispinosus occupy more than one nest and exchange

1982] Alio way, Buschinger, Talbot, Stuart & Thomas

273

workers, queens, and brood among nests (polydomy). Other colo-
nies have only one nest (monodomy). The significance of these find-
ings for understanding the evolutionary origin and maintenance of
social parasitism is discussed.

References

Alloway, T. M. 1979. Raiding behaviour of two species of slave-making ants,
Harpagoxenus americanus (Emery) and Leptothorax duloticus Wesson (Hymen-
optera: Formicidae). Animal Behaviour, 27: 202-210.

Alloway, T. M. 1980. The origins of slavery in Leptothoracine ants (Hymenop-
tera: Formicidae). American Naturalist, 115: 247-261.

Buschinger, A. 1967. Verbreitung und Auswirkungen von Mono- und Polygynie
bei Arten der Gattung Leptothorax Mayr (Hymenoptera: Formicidae). Inaugu-
ral Dissertation, Wurzburg. 1 14 pp.

Buschinger, A. 1968. Mono- und Polygynie bei Arten der Gattung Leptothorax
Mayr (Hymenoptera: Formicidae). Insectes Sociaux, 15: 217-226.

Buschinger, A. 1970. Neue Vorstellungen zur Evolution des Sozialparasitismus
und der Dulosis bei Ameisen (Hymenoptera: Formicidae). Biologisches Zen-
tralblatt, 88: 273-299.

Buschinger, A. 1974. Monogynie und Polygynie in Insektensozietaten. In G. H.
Schmidt (Ed.), Sozialpolymorphismus bei Insekten. Wissenschaftliche Verlags-
gesellschaft m.b.h., Stuttgart, 974 p.

Buschinger, A. 1975. Eine genetische Koponente im Polymorphism der duloti-
schen Ameise Harpagoxenus sublaevis. Naturwissenschaften, 62: 239.

Buschinger, A. 1978. Genetisch bedingte Entstehung geflungelter Weibchen bei
der sklavenhaltenden Ameise Harpagoxenus sublaevis (Nyl.) (Hymenoptera:
Formicidae). Insectes Sociaus, 25: 163-172.

Buschinger, A., and T. M. Alloway. 1977. Population structure and polymor-
phism in the slave-making ant Harpagoxenus americanus (Emery) (Hymenop-
tera: Formicidae). Psyche, 83: 233-242.

Buschinger, A., and T. M. Alloway. 1978. Caste polymorphism in Harpa-
goxenus canadensis M. R. Smith (Hymenoptera: Formicidae). Insectes Sociaux,
25: 339-350

Buschinger, A., and T. M. Alloway. 1979. Sexual behavior 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: Formicidae). Zeitschrift fur Tierpsychologie, 49:
113-119.

Buschinger, A., G. Frenz, & M. Wunderlich. 1975. Untersuchgen zur Gesch-
lechtstierproduktion der dulotischen Ameise Harpagoxenus sublaevis (Nyl.)
(Hym., Formicidae). Insectes Sociaux, 22, 169-182.

Creighton, W. S. 1950. The ants of North America. Bulletin of the Museum of
Comparative Zoology (Harvard), 104: 1-585.

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Ehrhardt, H. H. 1970. Die Bedeutung von Koniginnen mit steter arrhenotoker
Parthenogenese fur die Mannchenerzeugung in den Staaten von Formica polyc-
tena Foerster (Hymenoptera: Formicidae). Inaugural Dissertation, Wurzburg,

106 pp.

Headley, A. E. 1943. Population studies of two species of ants, Leptothorax lon-
gispinosus Roger and Leptothorax curvispinosus Mayr. Annals of the Entomo-
logical Society of America, 36: 743-753.

Holldobler, B., and E. O. Wilson. 1977. The number of queens: An important
trait in ant evolution. Naturwissenschaften, 64: 8-15.

Petersen, M., and A. Buschinger. 1971. Untersuchungen zur Koloniegrundung
der Pharaoameise Monomorium pharaonis (L.). Anzeiger fur Schadlingskunde
und Pflanzenschutz, 44: 121-127.

Smith, M. R. 1950. On the status of Leptothorax Mayr and some of its sub-
genera. Psyche, 57, 29-30.

Talbot, M. 1957. Population studies of the slave-making ant Leptothorax dulo-
ticus and its slave, Leptothorax cuvispinosus. Ecology, 38: 449-456.

Wesson, L. G. 1939. Contributions to the natural history of Harpagoxenus
americanus (Hymenoptera: Formicidae). Transactions of the American Ento-
mological Society, 35: 97-122.

Wesson, L. G. 1940. Observations on Leptothorax duloticus. Bulletin of the
Brooklyn Entomological Society, 35: 73-83.

Wilson, E. O. 1971. The Insect Societies. Belknap Press of Harvard University
Press, Cambridge. X + 548 pp.

Wilson, E. O. 1974a. Aversive behavior and competition within colonies of the
ant Leptothorax curvispinosus. Annals of the Entomological Society of Amer-
ica, 67: 777-780.

Wilson, E. O. 1974b. The population consequences of polygyny in the ant Lepto-
thorax curvispinosus. Annals of the Entomological Society of America, 67:
781-786.

Wilson, E. O. 1975. Leptothorax duloticus and the beginnings of slavery in ants.
Evolution, 29: 108-1 19.

A NEW COLONIAL ANELOSIMUS SPIDER FROM
SURINAME (ARANEAE: THERIDIIDAE)

By Herbert W. Levi1 and Deborah R. R. Smith2

Until recently, only a few colonial spiders were known. The recent
increase in field work in the tropics revealed a number of new colon-
ial species (Buskirk, 1981). Some of these colonial spiders belong to
the theridiid genera Anelosimus and Achaearanea. The genus Ane-
losimus in America was revised by Levi, 1956, and the knowledge
updated in 1963 and 1972, with new species described in 1967 and
1979. Those Anelosimus species known to be colonial are: A. studi-
osus, A. eximius, A. rupununi and A. lorenzo. Another species has
now been found in Suriname. While reexamining some of the
related species in preparation for this description, it was found that
the synonymy of A. jabaquara Levi 1956 with A. dubiosus (Keyse-
rling, 1891) in Levi (1963) was in error. While A. jabaquara was
illustrated in 1956, A. dubiosus is here illustrated for the first time
since its description in 1891 (Fig. 4).

Anelosimus saramacca new species
Figures 1-3

Type. Male holotype from Voltzberg-Raleighvallen Nature Re-
serve, Saramacca Province, Suriname [lat. 04°40'N, long. 56° 10'W],
Feb. 1982 (D. Smith Trail), with 1<J, 5? paratypes in the Museum of
Comparative Zoology; 1(5, 2$ paratypes in the Cornell University
collection kept at the American Museum of Natural History; 2$
paratypes in the British Museum, Natural History.

Description. Female. Carapace orange, lighter on sides. Sternum
orange with some black pigment. Legs yellow-white with distal part
of articles darker. Dorsum of abdomen with some black and white
pigment, sides orange-white. Venter of abdomen with some black
and white pigment, black patch anteriorly and behind genital
groove, and black patch in front of spinnerets. Eyes subequal in size.

'Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02138.

2Field of Neurobiology and Behavior, Department of Entomology, Cornell Univer-
sity, Ithaca, NY 14853

Manuscript received by the editor September 14, 1982.

275

276

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[Vol. 89

Anterior median eyes their diameter apart, 0.3 diameters from later-
als. Posterior median eyes slightly more than their diameter apart,
their diameter from laterals. Total length, 3.2 mm. Carapace, 1.3
mm long, 0.9 mm wide. First femur, 1.7 mm; patella and tibia, 1.7
mm; metatarsus, 1.3 mm; tarsus, 0.8 mm. Second patella and tibia,
1.4 mm; third, 1.1 mm; fourth, 1.5 mm.

Male. Carapace, sternum orange. Legs yellow-white. Abdomen
orange to black. Eyes subequal in size, spacing as in female. Total
length, 2.3 mm. Carapace, 0.9 mm long, 0.6 mm wide. First femur,
1.2 mm; patella and tibia, 1.3 mm; metatarsus, 0.8 mm; tarsus, 0.6
mm. Second patella and tibia, 1.0 mm; third, 0.8 mm; fourth, 1.0
mm.

Diagnosis. Unlike A. jabaquara and A. dubiosus, Anelosimus
saramacca has a short terminal embolus (Fig. 3). The female has an
epigynum with a subtriangular depression enclosing a transverse
mark; the openings appear posteriorly at the ends of the mark (Figs.
L2).

Natural History. A single colony of A. saramacca was found in an
area of swampy lowland rainforest, approximately midway between
Voltz Berg and Van Stockum Berg. The web was similar to that of
Anelosimus eximius, but much smaller. It was located in a small
sapling, about 30 cm above the ground. The web consisted of a
nearly circular hammock or sheet of silk about 80 cm in diameter,
and a pyramid shaped barrier web about 1 m tall. In the center of
the hammock were retreats consisting of green leaves, some of
which were curled.

The colony contained at least 1000 individuals, including males,
females and immatures. There were many more adult females than
adult males. Large numbers of females with egg cases were found in
the leaf retreats. A quick inspection revealed at least 140 females
with egg cases.

The egg cases closely resemble those of A. eximius and A. studiosus
— they are pale brown, nearly spherical, and 1.5 mm in diameter.
Several egg cases were collected, but many later proved to be empty
or hatched out. Six egg cases containing eggs or embryos had a
clutch size of 15.2 ± 1.8 eggs.

Like A. eximius, A. saramacca shows cooperative behavior. Sev-
eral adults and immatures were seen feeding together on large prey
items, and the web appears to be a product of cooperative effort.

1982]

Levi & Smith — Colonial Anelosimus

277

Figs. 1-3. Anelosimus saramacca new species. 1, 2. Epigynum. 1. Dorsal, cleared.
2. Ventral. 3. Left male palpus.

Fig. 4. Anelosimus dubiosus (Keyserling). Left male palpus. Scale lines, 0. 1 mm.

Anelosimus dubiosus (Keyserling)

Figure 4

Theridium dubiosum Keyserling, 1891, 3: 187, pi. 6, fig. 133, Male holotype
from N. Freiburg (Nova Friburgo, Est. Rio de Janeiro), Brazil in the British
Museum, Natural History, reexamined.

Description. Carapace, legs orange. Abdomen white with a dorsal
gray band. Total length, 3.4 mm. Carapace, 1.7 mm long, 1.2 mm
wide. First femur, 2.2 mm; patella and tibia, 2.5 mm; metatarsus, 1.5
mm; tarsus, 0.8 mm. Second patella and tibia, 1.9 mm; third, 1.3
mm; fourth, 1.8 mm.

Note. Anelosimus jabaquara Levi, 1956 is not a synonym of this
species as thought in 1963. Anelosimus dubiosus differs by having a
much longer filamentous embolus (Fig. 4) than A. jabaquara (Levi,
1956, fig. 18) and A. saramacca (Fig. 3).

Acknowledgments

The specimens of A. saramacca were collected on the Second
Cornell Entomology Expedition to Suriname. The cost of this

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[Vol. 89

expedition was defrayed in part by the Explorers Club, Sigma Xi,
the Grace Griswold Memorial Fund, the Cornell Insect Collection,
and members of the Cornell Department of Entomology (Dr.
William L. Brown, Penelope Kukuk and Maurice Tauber). We also
wish to thank the staff of ST1NASU, the Suriname Nature Conser-
vancy, for their help and cooperation. The senior author thanks the
National Science Foundation for grant no. 81-20492 for research
and publication support, and Paul Hillyard for the loan of a speci-
men from the British Museum (Natural History).

References Cited

Buskirk, R. 1978. Sociability in the Arachnida. in H. R. Hermann, ed., social
insects. Academic Press, 2: 282-367.

Keyserling, E. 1891. die spinnen amerikas, Niirnberg, vol. 3.

Levi, H. W. 1956. The spider genera Neottiura and Anelosimus in America (Ara-
neae, Theridiidae). Trans. Amer. Microscop. Soc. 75:333-422.

1963. The American spiders of the genus Anelosimus (Araneae, Theri-
diidae). Trans. Amer. Microscop. Soc. 82: 30-48.

1967. The theridiid spider fauna of Chile. Bull. Mus. Comp. Zool. 136:

1-20.

1972. Taxonomic-nomenclatural notes on misplaced theridiid spiders

(Araneae, Theridiidae) with observations on Anelosimus. Trans. Amer. Micro-
scop. Soc. 91: 533-538.

1 979. in Fowler, H. G. and H. W. Levi, A new quasisocial Anelosimus

spider (Araneae, Theridiidae) from Paraguay. Psyche 86: 11-18.

BIOLOGY AND SYSTEMATICS OF THE
BEE GENUS CRAWFORDAPIS
(COLLETIDAE, DIPHAGLOSSINAE)

By Gard W. Otis1, Ronald J. McGinley2, Lyn Garling3,
and Luis Malaret3

Crawfordapis luctuosa (Smith) is a robust, dusky-haired bee, pres-
ently known from only a few localities in Mexico and Central Amer-
ica. Individuals can be as long as 24 mm and superficially resemble
the more familiar diphaglossine bees of the genera Ptiloglossa and
Caupolicana to which they are related. All three genera are placed in
the Caupolicanini which is characterized by the complete pre-
episternal groove and very elongate first flagellar segment. While
Crawfordapis is currently considered to be monotypic,
Michener (1966) raised the possibility that the material from the
more northern localities (Mexico and Guatemala) may represent a
distinct species. Much more material from different localities is
needed before that problem can be considered.

The purpose of this paper is to present biological observations
made on Crawfordapis by three of us in Costa Rica (L.G., L.M.,
G.O.). In addition, the larva of Crawfordapis is described and the
systematic interrelationships of diphaglossine genera are reviewed
(R.M.).

Biology

Description of the Site

Two nesting aggregations of Crawfordapis luctuosa were observed
approximately 5 km east of Monteverde, Province of Puntarenas,
Costa Rica (10°18'N, 84°47'W) on trails at 1540 m elevation. The
surrounding vegetation is best described as elfin forest, with some
characteristic plant species being Lycopsidium cernuum, Senecio
megaphyllus, Clibadium sp., Gunnera sp. and Myrica phanerodouta

■Environmental Biology Dept., University of Guelph, ON NIG 2W1, Canada.

2Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138.
Present address: Dept, of Entomology, N.H.B. 105, Smithsonian Inst., Wash-
ington, D.C. 20560.

3Dept. of Zoology, University of Florida, Gainesville, FL 326F1
Manuscript received by the editor July 7, 1982

279

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(details in Lawton and Dryer, 1980). Frequent rains deposit approxi-
mately 3000 mm of precipitation annually on this area. Heavy mists
augment this precipitation. Mean annual temperature is about 16°C
(max = 27° C, min = 10°C) and during prolonged rainstorms, the
temperature can remain at 15— 16°C for 3-4 days. The wind generally
blows 15 to 20 km/ hr, but ranges from nearly calm to winds in excess
of 100 km/ hr during rainstorms (R. Lawton, pers. comm.).

Nesting aggregation No. 1 was directly on the Continental Divide,
on a narrow ridge known as “La Ventana”. The aggregation was first
noticed in August 1975 and was still active but reduced in size to only
5 nests in February 1981. W. Guindon (pers. comm.) indicated the
site was active as early as 1966. Bees apparently maintained nests in
this area throughout the year (R. Lawton, pers. comm.). Nests were
built in the lee of a slope which partially protected them from mist
and rain (Fig. 1). In July 1977, there were 97 nests with tumuli in the
approximately 18 m aggregation. The majority of the nests were
within an elliptical area of about 8 m. Of the nests 29 were completely
exposed in the trail, 50 were on nearly level ground and partially
obscured by grasses and other herbs, and 18 were on the face of the
sheltering embankment.

Nesting aggregation No. 2, located at the head of the valley on the
road to Penas Bancas, was exposed to high winds and unprotected
from rain (Fig. 2). In February 1978, the aggregation consisted of
not more than 100—150 nests, but had enlarged to at least four
hundred active nests by August 1978. Nests were found both on the
edge of the road and down the adjacent steep, bare slope. By the last
visit to the site in February 1981, the number of active nests had
declined to 230. The adjacent slope had become covered with dense
vegetation and lacked nests.

Description of the Nest

Entrances to active nests had tumuli approximately 7.5 cm in
diameter and 4-7 cm in height (Fig. 3). The frequent rains obliter-
ated tumuli of all nests except those in which bees were actively
digging. Nest entrances consistently measured 1.0 cm in diameter.
In each of three nests excavated in horizontal ground, the tunnel
began nearly vertically for 7—14 cm and then continued downward
at an angle of approximately 75°. In the diagrammed nest (Fig. 4)

1982] Otis, McGinley, Gar ling & Malaret — Craw for dapis 281

Figures 1-2. Crawfordapis luctuosa nesting sites. Fig. 1. Nest aggregation No. 1.
Most nests were on level ground, either exposed or partially obscured by grasses. A
smaller number of nests were in the nearly vertical embankment which sheltered the
site from wind and mist. Fig. 2. Nest aggregation No. 2.

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[Vol. 89

Figure 3. Nest tumulus. Nests in which bees were actively digging were readily
discerned by the presence of a dirt mound around the nest entrance.

the tunnel changed directions again at at a depth of 24 cm, con-
tinued downward at a 75° angle another 8 cm, and diverged into
two tunnels. One of these angled toward the embankment at an
angle of 50° from vertical for another 12 cm, then at a depth of
41 cm continued slightly below horizontal for another 18 cm. A
single, terminal cell was found at the end of that tunnel. The other
tunnel continued downward another 16 cm from the branching
point before becoming nearly horizontal at a depth of 47 cm. This
tunnel could not be followed because the soil was too soft.

Two additional nests were excavated on the nearly vertical
embankment at aggregation No. 1. These nests differed from those
in level ground in having very short (3-5 cm) vertical portions of the
tunnel before becoming nearly horizontal. Nest B (Fig. 5) had a
single horizontal tunnel that extended 22 cm into the embankment.
Along the slight downward slope of this main tunnel were 6 nearly
horizontal lateral tunnels 4-8 cm long. The two closest to the exte-
rior contained pupae and the tunnels had been filled with soil. The
next three contained larvae, and the distal cell was empty. Nest C

1982] Otis, McGinley, Garling & Malaret — Crawfordapis 283

i i

10cm

Figures 4-6. Diagrammatic representation of three C. luctuosa nests. Fig. 4. Nest
at base of the embankment. Figs. 5-6. Nests excavated in the embankment.

(Fig. 6) had one cell (contents unrecorded) only 4 cm from the
vertical entrance tunnel. Another 8 cm further down, the main tun-
nel diverged into two. One tunnel continued to slope downward and
contained two terminal cells with pupae. The other tunnel sloped
slightly upward and had two lateral cells with larvae and an empty
terminal cell which was 34 cm from the face of the embankment.

Each completed cell was lined with a shiny, cellophane-like mem-
brane which is characteristic of Colletidae. It was not possible to lift
the cell and contents out of the soil as described for Ptiloglossa
guinnae Roberts (Roberts, 1971). Cell contents were soupy; fermen-
tation odors were not recorded. Cocoons were tough, nearly trans-
parent membranes 17 mm in diameter by 35 mm in length.

General Activity Pattern

Females of Crawfordapis luctuosa were active aboveground
primarily between the hours of 0930 and 1400 hr, with few bees
leaving the nests after 1300 hr on observation days February 19 and
20, 1978 (Fig. 7). A similar activity pattern was recorded on July 16,
1977. This sharply contrasts with the crepuscular activity pattern of
Ptiloglossa guinnae which occurs in similar habitats (Roberts,
1971). In some instances, bees were seen returning to the nests in the
morning before any bees left the nesting aggregation. It is possible

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[Vol. 89

that these bees had spent the night away from their nests as occurs
with Bombus species at high elevations (O. R. Tayor, pers. comm.).
Before leaving the nest for the first time in the morning, females
often remained just below the nest entrance for a few moments and
then upon exiting, hovered nearby for a short time before flying off.
Temperatures within the nest entrance remained at 12°C through-
out the day on February 20 when ambient temperature was between
10-1 1°C.

Males remained outside the nests at all times. During the activity
period of the females, the males flew over the nesting aggregation
and nearby at heights of 1-3 m. They rapidly approached any flying
object, including female C. luctuosa, swallows, a hummingbird, a
ctenuchid moth and a dragonfly. Males often seized females return-
ing to the nests but it was not ascertained whether copulations
occurred.

Nest Visitation Behavior

On February 19 and 20, 1978, 46 nests within a 2. 16 m subarea of
aggregation No. 2 were mapped and numbered. Sixteen female bees
were captured while leaving nests. Each bee and her corresponding
nest of origin were given an identifying color combination. The bees
were marked by paint spots on the thorax, while their nests of origin
were indicated by a wooden chip about 1 cm in length placed near
the entrance. All observed departures from and arrivals to mapped
nests were recorded by noting time, markings (or lack thereof) of
bees and nest number or color. A nest “visit” was defined as the
disappearance of the bee beneath the ground surface for any length
of time.

Of the 46 nests mapped, 40 (80%) were entered at least once by a
bee. The number of observed visits per nest made by marked or
unmarked bees to the 16 color-coded nests ranged from 0-15 over
the two days (Table 1).

Of the 16 marked bees, four were not observed again. The remain-
ing 12 marked bees visited nests a total of 78 times. Four of the
marked bees (BG, GO, YBY, OB) concentrated their visits on a
single nest, while others entered up to 12 different nests over the two
days (Table 2).

The duration of visits of both marked and unmarked bees varied
widely from less than 1 minute to a maximum of 151 minutes. The

1982] Otis, McGinley, Garling & M alar et— Craw for dapis

285

Figure 7. Number of bees exiting and entering nests in 10-minute intervals on February 19, 20, 1978.

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Table 1. Number of visits by all bees to color-coded nests

Nest Color-Code

YB

OY

YO GO BG YBY Y OBO O OB BY

YG

B GB

OG

BOY

No. of
visits

8

15

4 7 11 804282

0

7 4

3

0

Table 2. Number of visits made by marked bees to particular nests.

Marked

bee

Nest Categories

1°

2°

3° 4° 5° 6°

7° 8° 9° 10° 11°

12° Total visits

YB

3

2

2 2 2 2

1* 1 1 1 1

1 19

OY

6*

4

2 111

1111

19

YO

2*

2

2 111

1 1

11

GO

5

1

1 1 1

8

BG

5*

5

Y

1

1

1

3

YBY

3*

3

OB

3*

3

OBO

2

2

O

1

1

2

BY*,YG

Each visited one nest once.

♦Asterisked entries indicate that the visits were made to the nest from which the bee
was originally captured.

visits of 1 minute or less were likely to have been exploratory rather
than “working” visits. Of a total of 69 timed visits, 30 (43%) were in
the “exploratory” category. The average duration of the remaining
39 “working” visits was 18.1 minutes (S = 26.36) (Fig. 8).

The duration of foraging trips of 4 of the marked bees was noted
by timing of their absences from the nest area. Whether or not they
returned with pollen was not noted. Absences ranged from 19-42
minutes, the average being 30.4 minutes (n = 11, s = 13.79). It
seemed that bees returned from foraging trips in distinct pulses of
several bees at a time. The foodplants are not known.

This preliminary data on the movements of marked bees to differ-
ent nests generates several hypotheses for further testing.

1) Multiple nest entering is part of searching behavior for a female’s
own nest.

1982]

Otis, McGinley, Garling & Malaret — Crawfordapis 287

d

E

<

OC

ZD

Q

siisia do adaiAiriN

Figure 8. Number of visits by bees to nests as grouped in 4-minute intervals. The first category of visits of^ 1 minute may represent
searching behavior and is set apart from the other categories.

288

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[Vol. 89

2) Individuals are locating abandoned nests to provision rather
than starting completely new ones.

3) Individuals are “stealing” provisions from nests of other
individuals.

4) Individuals are usurping nests which other individuals are
actively excavating or provisioning.

5) Several individuals are contributing to excavation, provisioning
or oviposition within a single nest.

The latter hypothesis is particularly interesting since no species of
the family Colletidae is known to be parasocial (Michener, 1974). It
is plausible that females not only construct their own nests but also
usurp partially constructed or provisioned nests of other females as
has been reported in some other Hymenoptera that nest in aggrega-
tions (Brockmann and Dawkins, 1979; Brockmann et al., 1979;
Eickwort, 1975; Eickwort, 1981; Eickwort et al., 1977). Further stu-
dies of Crawfordapis luctuosa are needed to better understand its
social behavior and biology.

Systematics

Description of Larva

The following description follows the format used for describing
other colletid larvae (McGinley, 1981).

head (Figs. 9-1 3): (2) Labrum nonspiculate; (3) epipharynx and

(4) hypopharynx spiculate; (5) maxilla spiculate on inner surface. (7)
Head size normal in comparison to body (head not relatively large
as in Xeromelissinae); (8) head capsule somewhat elongate, slightly
produced in lateral view; (10) frontal swellings above antennae
absent; (10a) median frontal swelling above antennae absent (pres-
ent only in Ptiloglossa). (14) Anterior tentorial pit low in position
(high in all other known diphaglossines); (15) posterior tentorial pit
at junction of hypostomal ridge and posterior thickening of head
capsule; (15a) tentorial development unknown (tentorium of speci-
men examined was incomplete, probably due to nearness of speci-
men to pupation). (16) Posterior thickening of head capsule
moderately developed (17) straight medially, not curved forward;
(19) median longitudinal thickening of head capsule absent; (20)
hypostomal ridge well-developed; (25) epistomal ridge complete but
thin, (26) arching dorsally to level of antennae. (27) Parietal bands
distinct, broad and shallow. (28) Antennal prominence absent; (29)

1982] Otis, McGinley, Garling & Malaret — Crawfordapis 289

Figures 9-13. Mature larva of Crawfordapis luctuosa. Figs. 9, 10. Head capsule,
frontal and lateral view. Figs. 11-13. Right mandible, dorsal, adoral and ventral
view.

antennal papilla a moderate-sized convexity, (31) bearing three sen-
silla. (32) Clypeus moderate in length; (34) labrum not projecting in
lateral view; (35, 36, 37) labral tubercles very well-developed, nar-
rowly rounded and strongly projecting (unlike those of other
diphaglossines); (38) labral apex emarginate, (39) without sensilla-
bearing swellings as in other diphaglossines. (41) Mandibles elon-

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gate, (42) moderately slender in dorsal view, (43) broad basally in
adoral view; (44) mandibular spiculation absent; (46) outer surface
of mandible smooth, distinct tubercle and setae absent; (51) apical
portion of mandible, in adoral view, attenuate; (52) cusp moderately
well-defined; (53) cuspal projection absent; (54) cuspal region multi-
dentate; (55, 55a) dorsal apical edge with distinct, moderately large
teeth; (57) apical concavity weakly developed; (60, 60a) ventral api-
cal edge smooth, teeth absent. (61) Labiomaxillary region produced;
(62) labium and maxilla distinct, (63) subequal in length. (65) Inner
apical surface of maxilla rounded, not produced mesiad; (66) unlike
all other known bee larvae except those of Ptiloglossa, maxilla with
a longitudinal groove on adoral surface; (67) cardo and stipes sclero-
tized; (69, 70) maxillary palpus moderately elongate and slender,
(71) apically positioned on maxillary apex in lateral view; (72) galea
absent. (73) Labium divided into prementum and postmentum; (75)
palpus elongate and slightly decurved, (76) subequal to maxillary
palpus in length. (77) Salivary lips well-developed; (78, 79, 80) sali-
vary opening narrow, circular, at end of long spoutlike salivary lips,
(83) which project from a well-defined platelike structure at apex of
labium; (82) apical labial swellings absent. (84) Hypopharynx nor-
mal in size, (85) bilobed, (86) exceeded by labium and maxilla; (87)
hypopharyngeal groove distinct, sclerotized laterally.

body (Fig. 14): (88) Integument spiculate, density of spicules

greater on dorsum than on venter; (93) body moderate in length,
(94) robust, (95) widest posteriorly in lateral view; (96) interseg-
mental lines moderately incised; (97) intrasegmental lines indistinct;
(98) dorsal tubercles weakly developed, most prominent on abdom-
inal segments 5-7; (103) lateral tubercles absent (present on one
specimen from Panama); (104) ventrolateral tubercles absent; (106)
abdominal segment 10 moderate in length, (107) rounded, (108)
dorsal in attachment to segment 9; (109) venter of segment 10
slightly produced (very weakly so in specimen from Panama), (110)
without conspicuous, darkly pigmented spiculation; (111) dorsal
surface of segment 10 smooth, without lines or ridges; (113) anus
apical. (114) Spiracles large, (115) not on elevations; (117) atrium
very broad and shallow, (118) not produced above body surface;
(119) atrial wall faintly ridged, (120) with four to five broken rings
of spicules; (121) atrial rim absent; (122) peritreme wide; (123) pri-
mary tracheal collar absent; (126) subatrium apparently extremely

1982] Otis, McGinley, Gar ling & Malaret — Crawfordapis 291

Figure 14. Mature larva of Crawfordapis luctuosa.

short. (The structure of diphaglossine larval spiracles remains
poorly understood, especially with regard to the subatrium. The
atrium is connected to the trachea by a long, nonringed tube. While
this tube is characteristic of diphaglossines, its internal structure and
homologies are not known.)

material studied: Two postdefecating larvae; 5 km east Monte-
verde, Puntarenas Province, Costa Rica; July 16, 1977 (G. W. Otis);
specimens in the larval bee collection of the American Museum of
Natural History. Two postdefecating larvae; Bouquete, Chiriqui
Province, Panama; April 25, 1981 (R. W. Brooks); specimens in the
personal collection of R. W. Brooks.

Analysis of Larval Characters

McGinley (1981) described the mature larvae of 30 colletid species
including those of seven diphaglossines. Two cladograms for the
diphaglossine genera appeared to be most strongly supported by lar-
val characters. One of the cladograms, for reasons discussed in the
above mentioned paper, appeared to be the preferable working
hypothesis of diphaglossine phylogeny. This cladogram is presented
in Figure 15, with Crawfordapis now included. The polarities of the
characters listed in Table 3 were determined by out-group compari-
son, i.e., consideration of character state distributions in all other
bee larvae as well as in nonapoid larvae, especially those of specoid
wasps.

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[Vol. 89

Figure 15. Cladogram of diphaglossine genera based on larval characters. Numbers
refer to the characters listed in Table 3. Black rectangles represent presumed apo-
morphic characters; white rectangles represent plesiomorphic characters. Character
109 appears to be variable in Crawfordapis.

The larval cladogram corroborates the currently recognized
diphaglossine classification based on adult characters (Michener,
1966). Recognition of the Caupolicanini (Ptiloglossa, Crawfordapis,
Caupolicana) is supported by the presence of the unusual salivary
plate (character 83, Fig. 9) and the rounded, nonprojecting inner
maxillary surface (character 65).

1982] Otis, McGinley, Garling & Malaret — Crawfordapis 293

Table 3. Larval Characters Used in Diphaglossine Cladogram
Plesiomorphic Apomorphic

5. Maxilla spiculate

10. Median frontal swelling absent

17. Median portion of posterior
thickening of head capsule
straight

25. Epistomal ridge complete

53. Cuspal region of mandible
without distinct projection

55. Teeth on dorsal apical edge of
mandible distinct basally

65. Inner apical surface of maxilla
produced mesiad

66. Inner surface of maxilla
smooth

75. Labial palpus moderately
elongate, straight

78. Salivary lips transverse, not
spoutlike

83. Salivary plate absent

84. Hypopharynx normal in size

85. Hypopharynx bilobed

109. Venter of abdominal segment
10 not produced

Nonspiculate

Present

Curved forward
Incomplete

Cuspal projection present

Teeth fused basally, forming distinct
platelike wedge

a. Inner surface rounded

b. Inner surface strongly produced
forward

Inner surface of maxilla with
longitudinal groove

Extremely elongate, decurved
Elongate, spoutlike
Present

Conspicuously narrow

Rounded

Produced

The sister-group relationship of Ptiloglossa and Crawfordapis is
strongly supported by the presence of a longitudinal groove on the
inner maxillary surface (character 66). Weaker support for this rela-
tionship is indicated by character 109, the projection of the venter of
abdominal segment 10 (this projection is conspicuous in some spec-
imens of Crawfordapis but is only weakly developed in one speci-
men from Panama).

294

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[Vol. 89

Adult Characters

In a study of adult diphaglossines, Michener (1966) discussed the
similarities of Crawfordapis to Ptiloglossa and Caupolicana. Four
characters found in Crawfordapis were said to be more or less
Caupolicana- like: (4) outer hind tibial spur of male normal, articu-
lated at base like inner spur; (6) lateral extremities of terga of male
without areas of short, dense, erect hair; (7) sixth tergum of male
with posterior margin not thickened or sulcate; (9) eighth sternum
of male with apical process rather heavily pigmented, not down-
curved. The similarity based on character 4 is definitely symplesio-
morphic as the fusion of the hind tibial spur and the tibia is found
only in Ptiloglossa. The other three characters appear to be plesio-
morphic as well in that they represent the absence of some rather
unusual features.

Similarities between adult Crawfordapis and Ptiloglossa appear
to be apomorphic for diphaglossines: (1) clypeus strongly elevated
above level of adjacent parts of face; (2) marginal cell prolonged
basally as a narrow sinus to apex of stigma; (3) expanded second
and third hind tarsal segments of female considerably expanded
above. This evidence also supports the Crawfordapis- Ptiloglossa
sister-group relationship indicated by larval characters, but must be
considered tentative until a comprehensive cladistic analysis of adult
colletids has been performed.

Summary

Crawfordapis luctuosa, a large colletid bee, was studied at two
nest aggregations in the mountains of Costa Rica. The aggregations
were in exposed sites formed by landslides or clearing. Female bees
slowly abandoned the aggregations as they became overgrown with
vegetation. Several nests are described. In contrast to the crepuscu-
lar habits of the closely related genus Ptiloglossa, Crawfordapis was
active primarily between 0930 and 1400 hrs. Some individually
marked females showed a high degree of constancy in nest visita-
tion, while others visited several nests in succession. The exact
explanation of this behavior is not yet known. The previously
unknown larvae of Crawfordapis luctuosa are described. Informa-
tion from these larvae supports the placement of the genus in the

1982] Otis, McGinley, Garling & Malaret — Craxvfordapis 295

tribe Caupolicanini that was suggested from the systematic study of
adults, and indicates that Craxvfordapis may be the sister-group of
Ptiloglossa.

Acknowledgments

We wish to thank C. D. Michener and J. G. Rozen, Jr., for
making helpful suggestions on the manuscript and R. W. Brooks for
the loan of Craxvfordapis larvae from Panama. The Tropical
Science Center, San Jose, Costa Rica provided permission to study
Craxvfordapis within the Monteverde Cloud Forest Reserve. The
Organization for Tropical Studies provided logistical support for a
portion of the field work. Thanks are due also to the community of
Monteverde for its friendly support of biologists and appreciation
of their work.

Literature Cited

Brockmann, H. J. and R. Dawkins.

1979 Joint nesting in a digger wasp as an evolutionary stable preadaptation to
social life. Behavior 71:203-245.

Brockmann, H. J., A. Grafen, and R. Dawkins.

1979 Evolutionarily stable nesting strategy in a digger wasp. J. Theor. Biol.
77:473-496.

Eickwort, G. C.

1975 Gregarious nesting of the mason bee Hoplitis anthocopoides and the
evolution of parasitism and sociality among megachilid bees. Evolution
29:142-150.

Eickwort, G. C.

1981 Presocial Insects in Social Insects, Vol. II, ed. Henry Hermann, Aca-
demic Press, NY, pp. 199-280.

Eickwort, G. C., K. R. Eickwort, and E. G. Linsley.

1977 Observations on nest aggregations of the bees Diadasia olivacea and D.
diminuta (Hymenoptera: Anthophoridae). J. Kansas. Ent. Soc. 50:1-17.

Lawton, R. and V. Dryer.

1980 The vegetation of the Monteverde Cloud Forest Reserve. Brenesia
18:101-116.

McGinley, R. J.

1 98 1 Systematics of the Colletidae based on mature larvae with phenetic anal-
ysis of apoid larvae (Hymenoptera: Apoidea). Univ. Calif. Pub. Ent.
91:1-307.

296

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Michener, C. D.

1966 The classification of the Diphaglossinae and North American species of
the genus Caupolicana (Hymenoptera, Colletidae). Univ. Kansas Sci.
Bull. 46:717-751.

Michener, C. D.

1974 The Social Behavior of the Bees. Harvard University Press, Cambridge,
Massachusetts.

Roberts, R. B.

1971 Biology of the crepuscular bee Ptiloglossa guinnae n. sp. with notes on
associated bees, mites and yeasts. J. Kansas Ent. Soc. 44:283-294.

THE LIFE CYCLE OF
HETEROPODA VENATORIA (LINNAEUS)
(ARANEAE: HETEROPODIDAE)1’2

By

John Ross3, David B. Richman3, Fadel Mansour4,

Anne Trambarulo3, and W. H. Whitcomb3

The giant crab spider, Heteropoda venatoria (L.), is known to
occur throughout much of the tropics and subtropics of the world
where it is valued as a predator of cockroaches (Guthrie and Tindall
1968, Hughes 1977, Edwards 1979). Its feeding habits, like those of
most spiders, vary somewhat and it has also been known to eat
scorpions and bats (Bhattacharya 1941), although it is questionable
as to whether it normally attacks such prey. This spider is often
found associated with human habitation, possibly due to the abun-
dance of prey (Subrahamanyam 1944, Edwards 1979). Although
biological notes on H. venatoria have been published by several
workers (Lucas 1871, Minchin 1904, Bristowe 1924, Bonnet 1930,
Ori 1974, 1977), the only life history work to date was published by
Bonnet (1932) and Sekiguchi (1943, 1944a, b, 1945). Bonnet (1932)
based his study on only 12 spiders (of which seven matured) and
lacked data on the postembryonic stages. Sekiguchi (1943, 1944a, b,
1945) presented a more nearly complete study, but the papers are
difficult to translate and they still lack some data, especially in
regard to variation in the number of instars and carapace width. We
have raised H. venatoria in the laboratory and present here our data
on life cycle of this important beneficial arthropod.

Materials and Methods

Spiders were obtained from avocado groves in south Florida,
near Homestead, Dade County. Egg sacs taken from our laboratory

•This study was partially supported by the United States-Israel BARD Fund as
Research Project No. 1-2-79.

2Florida Agric. Exp. Sta. Journal Series No. 3798.

3Dept. of Entomology and Nematology, Univ. of Florida, Gainesville, FL 32611.

4Agricultural Research Organization, Newe Ya’ar, P.O. Haifa, Israel.

Manuscript received by the editor September 10, 1982

297

298

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[Vol. 89

population of H. venatoria were used to obtain data for eggs, first
and second postembryos and spiderling instars. Immature spiders,
through 4th-5th instars, were housed in Tygon® flexible plastic
tubing, an adaptation of methods developed by Peck and Whitcomb
(1967). Two tubing sizes were employed, 13 mm and 24 mm diame-
ter. The smaller bore tubing was cut to 10 cm lengths while the 24
mm tubing was cut into lengths of 20 cm to house spiders from 5th
to 9th instars. Plastic foam culture-vial stoppers for 14-19 mm
openings sealed the end of the tubing. Tube ends had only to be
dipped into water weekly to maintain adequate moisture and
humidity levels for the spiders. An open cell foam plug allowed for
adequate ventilation while preventing the spider’s escape from the
tube. While these cages were not as large as would perhaps be ideal,
they were easily maintained and stored in a relatively small area,
and the spiders stayed healthy in them.

Moist cotton swabs were used to clean the tubes when clear vision
into them was obscured by prey debris, spider wastes or mold. A
rolled piece of 9 cm diameter filter paper was inserted into the 24 mm
diameter tubes to further reduce cleaning frequency as the spiders
tended to retreat onto the papers and defecate. Changing the filter
paper at regular intervals maintained a high degree of sanitation.

Adult spiders were housed in 0.5 1 clear plastic cups. A heated
cork boring tool was used to fashion holes in lids in which were
inserted open cell, plastic culture tube stoppers which allowed for
ventilation. Paper can lids were inverted as bottoms to the plastic
cup spider cages and these were lined with 9 cm filter paper to
facilitate cleaning.

First instar H. venatoria were reared on adult vestigial-winged
fruit flies, Drosophila melanogaster Meigen, for which the spiders
showed a clear preference over an occasional cabbage looper larva,
Trichoplusia ni (Hiibner). Later instars were fed on adult native
fruit flies (family Drosophilidae, genus unknown), which were
larger than D. melanogaster, but the spiderlings showed greatest
weight gain on mealworm larvae (Tenebrio molitor L.). Mealworms
became the mainstay of the spiders’ diet through the 10th and 11th
instars, when the spiders were fed adult crickets, Acheta domesticus
(L.), to extend feeding intervals. Houseflies, Musca domestica L.,
were introduced in the pupal stage during the middle instars and
were fed on as the adult flies emerged.

1982] Ross, Richman, Mansour, Trambarulo & Whitcomb 299

Earlier instar spiders were maintained in a laboratory room and
transferred during penultimate or adult stages to an environmental
growth chamber. Temperatures in the room were stabilized at 27° C
in the summer and 24° C in the winter, ±2°C. The spiders were kept
under fluorescent lights. The eggs, postembryos, and first instars
used for later observations were all maintained in the environmental
chamber, which was kept at a constant 26.7° C on a 13:1 1 L:D light
period. Humidity was controlled within the chamber by a supersat-
urated NaCl solution bath in a 20 X 15 X 8 cm tray. The tray was
partially filled with small pebble-sized rocks to increase the surface
area available for moisture exchange. The humidity control method
was adapted from a technique described by Winston and Bates
(1960) and it stabilized humidity levels within the 60-70% range as
monitored by a hygrothermograph.

Mating was observed in plastic gerbil cages, which were modified
to prohibit escapes by gluing taffeta-like cloth between the upper
and lower portions of the cage.

Carapace widths were measured at the widest points with an ocu-
lar micrometer and a binocular microscope.

Results and Discussion

The courtship and mating of H. venatoria was described by
Bonnet (1932) and Sekiguchi (1944b). Our observations generally
agree with these published accounts except where noted in the fol-
lowing discussion. In the current study, males introduced to a cage
with a female were observed to construct a sperm web approxi-
mately 2 hours prior to mating. After sperm induction male spiders
groomed their pedipalps for 5-25 seconds. The males vibrated their
bodies prior to mounting, as described in detail by Rovner (1980).
After mounting, the male rubbed his first pair of legs on the female’s
abdomen before and sometimes during insertion of the pedipalps.
Copulation occurred in bouts lasting from one to six hours over a
period of 24 hours. The pedipalps were inserted alternately, for an
average of 20.4 seconds for each insertion (n = 70, SD = 6.8
seconds). Bonnet (1932) reported that insertion lasted 6-7 seconds,
not counting transfer time. Males were often cannibalized by the
female after mating, which could account for the higher proportion
of females to males found in the field.

300

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[Vol. 89

Approximately 12-14 days after mating, a circular, flattened,
creamy white egg sac was produced by the female. The size of the
sac ranged from 1.27 to 2.54 cm in diameter, and was from 3.18 to
6.35 mm thick. A network of silk was deposited on the underside of
a flat surface, such as a leaf or plastic housing container lid. The
eggs (each ca. 1 .5 mm dia.) were deposited on this base, and covered
with another layer of silk. After the egg sac was sealed around the
edges and removed from the foundation, the female carried it with
her pedipalps underneath her body during the incubation period.
The female usually did not eat during this time. Infertile egg sacs
were sometimes dropped or eaten by the female. A large number of
infertile egg sacs (54% of those produced in the laboratory) were
constructed by the reared spiders. This might be expected due to the
artificially imposed mating schedule. An average of 2.16 fertile egg
sacs were produced per female, with five the highest number. An
average of 163 eggs were laid in each fertile egg mass (n = 13 egg
masses, SD = 28.97) constructed by the experimental spiders.
Bonnet (1932) reported 207 spiderlings emerging from the one egg
mass from a female he had raised after obtaining it as an immature
spider on bananas shipped from Africa. Sekiguchi (1944a) obtained
188-436 eggs/ mass. In field observations we have found as many as
400 spiderlings in one egg sac. This may indicate that a high degree
of variability in egg mass size is common. No data were taken on the
numbers in consecutive egg sacs.

Peck and Whitcomb (1970) included a discussion of the postem-
bryonic stages and reviewed the terminology used in the literature to
describe them. The definitions used in the present study follow
theirs and are given below to avoid confusion. The first postembryo
is defined as being that stage after the chorion of the egg had been
shed from most of the embryo, but remained as a crumpled mass at
the posterior end. The second postembryo is defined as being that
stage after the vitelline membrane had been shed and the embryo
was completely free, with legs able to move. After the first molt the
spiderling was considered to be a first instar. This molt occurred
inside the egg sac. Bonnet (1932) and Sekiguchi (1944a) considered
the emerged spiderlings to be second instars.

Several egg sacs were removed from CCh-anesthetized female
spiders, opened, and placed in covered petri dishes for observation.
The egg stage lasted from 8-14 days (n = 6 egg masses). Eclosion

1982] Ross , Richman, Mansour, Trambarulo & Whitcomb 301

required approximately 4 hours and began with assistance from a
pair of brownish “egg teeth” positioned on the patellar region of
each pedipalp. The chorion was split anterioventral to the leg region
with swelling pulses (30-60 seconds between pulses and 2-3 pulses
per set) to about 2/3 of the diameter of the embryo. The membranes
were drawn towards the spinnerets by an alternating combination
of abdominal contractions and withdrawal movements of the legs,
similar to those observed in molting. The shed membrane remained
attached to the spinnerets until it was discarded at the beginning of
the second postembryonic stage along with the vitelline membrane
surrounding the legs. The first postembryonic stage lasted from 1-2
days for four egg masses, but four other egg masses required 5-6
days to become second postembryos (total n = 8).

During the second postembryonic stage the specimens were rela-
tively quiescent. Eye pigment began to appear about the fifth or
sixth day of the second postembryonic stage, with markings around
the carapace margin and on the abdomen becoming visible soon
afterwards. Dark setae appeared shortly before the first molt
occurred. The duration of the second postembryonic stage was 9-10
days (n = 3). The first instar spiderlings remained in the egg sac
approximately one week before emerging. One female which mated
on the 28th of March, 1980, produced an egg sac 12 days later.
From this egg sac 277 first instar spiderlings emerged after 32 days.
All of the longevity data came from this group of spiderlings.

The duration of each instar is summarized in Table 1 . Males were
more likely (as noted by Bonnet 1932) to have one or two fewer
molts than females, but this was not an absolute rule. Sekiguchi
(1945) recorded complete data for only one female H. venatoria and
found a total of 1 1 instars. He, however, apparently included the
second postembryo as the first instar. Of the adults in our study for
which complete data are available, the males (n = 3) had 8-10
instars (X = 8.7, SD = 1 .2) which lasted 241 .7 days (SD = 56.2) and
the females (n = 13) had 9-12 instars (X = 10.6, SD = 1.0) which
lasted 315.6 days (SD = 21.0). The survival rate from first instar to
adult in the laboratory was approximately 85%. Total length of life
from egg to death for our laboratory reared specimens was for males
(n = 4) 355-586 days (X = 464.5 days, SD = 1 12.0) and for females
(n = 16) 298-710 days (X = 580.3 days, SD = 128.6). Rovner (per.
com.) found that some females of H. venatoria can survive for three
years as adults in the laboratory.

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Table 1. Carapace width and duration of stadia in laboratory reared Heteropoda
venatoria (L.).

Instar

Carapace

width

(mm)

n

S.D.

Duration
of stadium
(days)

n

S.D.

1st

1.37

10

0.06

11.80

44

1.56

2nd

1.69

32

0.08

14.68

40

1.33

3rd

2.20

27

0.13

26.32

37

5.28

4th

2.60

22

0.16

27.97

30

4.21

5th

3.10

41

0.26

28.52

31

5.67

6th

3.44

38

0.21

36.11

37

5.41

7th

3.79

39

0.16

36.42

33

6.60

8th

4.06

39

0.29

44.46

28

10.56

9th

4.53

43

0.57

40.69

26

16.63

10th

5.36

43

0.55

35.48

23

12.57

11th

6.67

33

0.71

28.11

9

6.57

12th

—

—

—

23.0

4

7.83

The mean carapace width for each instar (not separated by sex) is
shown in Figure 1 to have a nearly linear relationship with the
stadia, as might be expected. This and the large number of stadia
seem to agree with a suggestion made by Hagstrum (1971) that large
spiders have added stadia, rather than accelerated growth between
successive molts. Sekiguchi (1945) shows similar data for the female
of H. venatoria in Japan. The carapace widths for our spiders are
summarized in Table 1.

The ratio of females to males for reared spiders was 2.4/1 (22
females/ 9 males). The sex ratio for adult specimens collected in
Homestead, Dade Co., FL, on August 14-19, 1981 was 3.4/1 (71
females/ 21 males). The sex ratio in Homestead might be due to
cannibalism of the males by females, as mentioned previously. Of
the females collected, 18.3% were carrying egg sacs, and all instars
were observed to be present in the field. Summer seemed to be the
major period of egg production both in the laboratory and in the
field.

This spider probably offers one of the best possibilities for the use
of spiders in biological control as it is well adapted for living in close
association with humans and is readily reared. As these spiders
habitually feed on cockroaches, H. venatoria behavior and ecology
may be an important key in the biological control of one of man-
kind’s oldest pests.

1982] Ross, Richman, Mansour, Trambarulo & Whitcomb

303

INSTAR

Figure 1. Relationship between logarithm of mean carapace with and stadia for
laboratory-reared Heteropoda venatoria (L.). Standard deviations are shown for
each point.

Acknowledgements

We would like to thank Dr. Stratton H. Kerr and Dr. Martin H.
Muma for suggestions regarding the manuscript, and Takuji Haya-
kawa and John Watts for help with the collection and rearing of the
spiders. Mr. Hayakawa also provided valuable help in translating
the papers published by Dr. Sekiguchi.

Summary

The giant crab spider, Heteropoda venatoria (L.) was reared in
the laboratory. These spiders reached adulthood after 8-10 instars

304

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[Vol. 89

for the males and 9-12 instars for the females and took approxi-
mately one year to mature from the egg. The postembryonic stages
were found to last approximately 2 weeks.

Literature Cited

Bhattacharya, G. C. 1941. Food and habits of the house-spider ( Heteropoda
venatoria Linn.). J. Bombay Nat. Hist. Soc. 42(4):82 1 —5.

Bonnet, P. 1930. Observation sur deux Heteropodes de la Guinee, etc. Ann. Soc.
Entomol. France 99:49-64.

Bonnet, P. 1932. Cycle vital de Heteropoda regia Fabr. Livre du Centenaire Soc.
Entomol. France: 497-503.

Bristowe, W. S. 1924. Notes on the habits of insects and spiders in Brazil. Trans.
Entomol. Soc. London, 1924:475-504.

Edwards, G. B. 1979. The giant crab spider, Heteropoda venatoria (Linnaeus)
(Araneae: Sparassidae). Florida Dept. Agric. Entomol. Circ. 205:1-2.

Guthrie, D. M. and A. R. Tindall. 1968. The biology of the cockroach.
Edward Arnald Publ. Ltd.

Hagstrum, D. W. 1971. Carapace width as a tool for evaluating the rate of
development of spiders in the laboratory and the field. Ann. Entomol. Soc.
Amer. 64:757-60.

Hughes, I. W. 1977. Cockroaches. Bermuda Dept. Agric. and Fish. Monthly
Bull. 47(9):69-72.

Lucas, H. 1871. Observations sur une ponte d 'Olios venatorius. Ann. Soc. Ento-
mol. France Bull. 5th Ser, 143:43, 58, 60 and 64.

M inchin, E. A. 1904. Exhibition of a specimen of the spider Heteropoda regia,
captured at University College, London. Proc. Zool. Soc. London, 1:229.

Ori, M. 1974. Studies on spiders as natural enemies of insect pests. 1. Observa-
tions on the spiders in houses in Nagasaki prefecture [in Japanese, English
summary]. Japan J. Sanit. Zool. 25(2): 153-60.

Ori, M. 1977. Studies on spiders as natural enemies of insect pests. 5. Species of
spiders as natural enemies of the house fly, and evaluation of their predacious
capacities [in Japanese, English summary]. Japan J. Sanit. Zool. 28(2): 175-8.

Peck, W. B., and W. H. Whitcomb. 1967. An adaptable method for rearing
spiders and cannibalistic insects. Turtox News 45(10): 242-4.

Peck, W. B., and W. H. Whitcomb. 1970. Studies on the biology of a spider,
Chiracanthium inclusum (Hentz). Arkansas Agric. Exp. Stn. Bull. 753:1-76.

Rovner, J. S. 1980. Vibration in Heteropoda venatoria (Sparassidae): A third
method of sound production in spiders. J. Arachnol. 8:193-200.

Sekiguchi, K. 1943. Life history of Heteropoda venatoria Linnaeus [in Japa-
nese], Acta Arachnol. 8(3):66-77.

Sekiguchi, K. 1944a. Life history of Heteropoda venatoria Linnaeus 2 [in Japa-
nese]. Acta Arachnol. 8(4):98— 1 17.

Sekiguchi, K. 1944b. Life history of Heteropoda venatoria Linnaeus 3 [in Japa-
nese]. Acta Arachnol. 9(1/ 2): 1-21.

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Sekiguchi, K.. 1945. Life history of Heteropoda venatoria Linnaeus 4 [in Japa-

nese]. Acta Arachnol. 9(3/4): 107-1 1 1.

Subrahmanyam, T. V. 1944. Reoccurrence of the house spider (Heteropoda
venatoria) in the field. J. Bombay Nat. Hist. Soc. 44(3):493.

Winston, P. W., and D. H. Bates. 1960. Saturated solutions for the control of
humidity in biological research. Ecology 4l(l):232-7.

DESCRIPTION OF A NEW SPECIES OF
KROM BEINIUS (H YMENOPTERA: PERILAMPIDAE)

FROM THE PHILIPPINES, AND THE PHYLOGENETIC
RELATIONSHIPS OF THE GENUS*

Bv D. Christopher Darling
Department of Entomology,

Cornell University, Ithaca, N.Y. 14853

The genus Krombeinius (Hymenoptera: Perilampidae) was re-
cently described (Boucek 1978) to include perilampids with an
amalgam of the characters of Euperi/ampus Latreille and Peri/am-
pus Walker. The habitus suggests Euperi/ampus, and there are two
synapomorphies to unite these two genera (Darling 1983): postspi-
racular sclerite reduced to a narrow triangle, less than one-half as
wide as the adjacent pronotum; and pronotum massive, at least
one-third the length of the mesoscutum. However, Krombeinius
exhibits the wing venation, presence of a marginal rim on the scutel-
lum, and large third metasomal tergite characteristic of Perilampus.
I regard these as plesiomorphic similarities. The genus is character-
ized by the absence of the defining apomorphic characteristics of
Euperi/ampus, i.e., by symplesiomorphy.

In this paper I present new information on the structure of the
male genitalia and labrum of the type species of Krombeinius. These
structures have proved to be of considerable value in defining gen-
era in the Perilampidae (Darling 1983). From this analysis 1 suggest
autapomorphies for defining Krombeinius. In addition, I describe a
new species of Krombeinius from the Philippines, and discuss the
affinities of the three included species.

Taxonomic studies of Krombeinius have been hampered by the
scarcity of material. The type species, K. eumenidarum, was de-
scribed by Boucek (1978) from a series of specimens (2 male, 2
female) reared from the larvae ol an eumenine wasp in Sri Lanka.
All specimens were prematurely killed and had to be liberated from
the pupal cuticles, producing some abnormalities in the type mate-
rial. Also included by Boucek (1978) in this genus was Perilampus
megalaspis Cameron, known only from the type material (3 females)

* Manuscript received by the editor September 22, 1982.

307

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and an additional female, all from Sarawak, Malaysia. During my
study of material in the U.S. National Museum of Natural History,
Washington [USNM], I located an additional male specimen of K.
eumenidarum. This specimen [India: Kerala Survey, 12.5 Pechipa-
rai, 25-27 August 1974] was dissected and re-mounted and is the
basis for the description of the labrum and male genitalia. In addi-
tion, the C. F. Baker Collection [USNM] contained a new species of
Krombeinius from the Philippines, which I describe herein.

Abbreviations used in text: Fl-7: funicular segments 1-7; MSC:
length of mesoscutum along midline; OOL: length of ocular-ocellar
line; PN: length of pronotum along midline; POL: length of poste-
rior ocellar line; SC: length of scutellum along midline; T2-8: meta-
somal tergites 2-8.

Krombeinius

Krombeinius Boucek, 1978: 302, Figs. 1,2.

Type species: Krombeinius eumenidarum Boucek, 1978: 302, Fig. 1. [original
designation].

Diagnosis:

Hymenoptera: Chalcidoidea: Perilampidae (sensu Graham, 1969).
Species of Krombeinius can be distinguished from Monacon Water-
ston, Burksilampus Boucek, Steffanolampus Peck and Perilampus
Latreille by the narrow postspiracular sclerite, less than one-half the
width of the adjacent pronotal panel, and from Euperilampus by
having the marginal vein longer than the postmarginal (Fig. 1).

All known species are moderately large, 3 to 5 Vi mm long, black
without metallic reflections and are restricted to the Oriental region.
There are three species currently placed in Krombeinius: K. eume-
nidarum Boucek, K. megalaspis (Cameron) and K. saunion, n.sp.

A revised key to the species of Krombeinius is not presented. The
key of Boucek (1978) separates K. eumenidarum and K. megalaspis.
An additional character to separate these two species is the inner
orbits: costate in K. eumenidarum (Fig. 8), and smooth in K. meg-
alaspis (Fig. 12). K. saunion is readily distinguished from these two
species by the prominent spine at the apex of the scutellum (Fig.
1,15). The apex of the scutellum is truncate in the other two species
(Figs. 7,1 1).

1982]

Darling — New Species of Krombeinius

309

Fig. 1. Krombeinius saunion, lateral habitus.

Description:

Head: supraclypeal area smoothly convex, without horn or ridge;
scrobal cavity deep, extending to lower ocular line or to middle of
clypeus; clypeus and supraclypeal area separated by distinct suture
or by faint line; inner orbits carinate; frontal carina separating the
median and posterior ocelli; malar sulcus absent; malar region with
strong oblique costae; posterior ocellus located high on vertex, POL
approximately equal to OOL; labrum with a single narrow stalk,
expanded distally with 7 digits, each with a terminal seta, and with
pair of sessile setae not associated with digits, strongly excised
medially [n = 1, K. eumenidarum , Fig. 3],

Mesosoma: dorsum of pronotum smoothly convex, without
transverse elevations; pronotum massive, about one-third length of
mesoscutum, not narrowed medially; mesothoracic spiracle located
between pronotum and sidelobe of mesoscutum; postspiracular
sclerite fused to the pronotum but delimited by surface sculpture;
postspiracular sclerite less than one-half width of adjacent pronotal
panel, with many or a single puncture; notauli distinct; scutellum
vaulted, jutting over propodeum and base of metasoma; apex of
scutellum acuminate, or truncate or with a distinct spine; propo-
deum with median area foveate, or with a short median ridge, sub-

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Figs. 2-5. 2,3 Krombeinius eumenidarum. 2. Male genitalia. Inset: enlargement of
paramere. 3. Labrum. 4,5. Euperilampus triangularis (Say). 4. Apex of male genita-
lia. 5. Labrum. [Scale lines 0. 1 mm.]

1982]

Darling — New Species of Krombeinius

311

median areas with weak transverse rugae or aciculate; basitarsomere
not conspicuously lengthened. Forewing with marginal vein longer
than postmarginal, postmarginal vein long, about 3 times length of
stigmal vein, stigmal vein making either a right or oblique angle
with marginal vein.

Metasoma: petiole short, transverse, the tergum forming a ridge
along anterior face of gaster, sternum shifted posteriorly; gaster
triquetrous, T2 and T3 fused, covering most of dorsum; T2 without
distinct basal fovea; T3 much longer than T2, subquadrate, slightly
wider than length along midline; ovipositor ventral, not upturned,
sheaths not distinctly exserted; male genitalia with distinct para-
meres [n = 1, K. eumenidarum, Fig. 2],

Discussion:

The male genitalia of Krombeinius eumenidarum (Fig. 2) are
similar to those of species of Perilampus : the parameres are distinct,
and strong setae are distributed on these lobes. This configuration
occurs throughout the Chalcidoidea (see Domenichini 1953) and is
regarded as plesiomorphic. In Euperilampus a derived condition is
found (Darling 1983): distinct parameres are lacking, and the basi-
paramere has a patch of strong setae distributed on transparent
areas laterad of the ventral lobe (Fig. 4).

The labrum of Krombeinius eumenidarum (Fig. 3) has a narrow
central stalk, not found in other perilampid genera (Riek 1966;
Domenichini 1969; Darling, unpublished). However, the labrum
does share synapomorphies with species of Euperilampus (Fig. 5)
including a reduced number of digits (7 or 8), a pair of smaller,
sessile setae not associated with digits, and a strong median exci-
sion. The narrow stalk distinguishes the labrum of Krombeinius
from those of Euperilampus, and is postulated as an autapomorphy
of Krombeinius. All other perilampid labra are 1 0— 1 2-digitate, and
not as strongly excised medially.

The host association of the type species of Krombeinius [larva of
Vespidae: Eumeninae] is different from that of any other described
perilampid, although solitary Sphecidae are attacked by some Peri-
lampus species (e.g., Perilampus nitidus, primary parasitoid of
Ectemnius paucimaculatus. ltrombein 1964, as P. canadensis). This
behavioral character is regarded as an additional autapomorphy for
the genus Krombeinius.

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Figs. 6 13. 6 9. Krombeinius eumenidarum. 6. Head, dorsal. 7. Mesosoma, dor-
sal. 8. Head, frontal. 9. Head, lateral. 10 13. K. megalaspis. 10. Head, dorsal. 11.
Mesosoma, dorsal. 12. Head, frontal. 13. Head, lateral. [Scale line 1 mm.]

1982]

Darling — New Species of Krombeinius

313

Krombeinius sauion n.sp.

(Figs. 1, 14 17)

Type Locality: Philippines, Mindanoa, Surigao.

Type Material: Holotype: Female [Baker Collection, USNM].
Etymology: The specific epithet is a noun in apposition, Greek for
“javelin”, and is a reference to the elongate spine on the scutellum of
this species.

Diagnosis:

This species can be immediately recognized by the prominent
spine at the apex of the scutellum (Figs. I, 15). The apex of the
scutellum is truncate in K. eumenidarum (Fig. 7) and K. megalaspis
(Fig. 11).

Description:

Female: Length, 5.4 mm. Black, except tegula and flagellum
brown, mandible reddish-brown, apex of foretibia and spur, and
tarsi yellow; wings hyaline, veins darkened.

Head: length of malar space 0.34 eye height; OOL 0.95 POL;
maximum width of scrobe 0.56 head width; head transverse, width:
height = 1.17; inner and outer orbits costate, costae convergent on
clypeus; scrobal cavity deep and wide, extending below lower ocular
line and delimiting clypeus and supraclypeal area; clypeus trans-
verse, width:height = 1.31, polished and covered with long setae;
clypeus not delimited by sutures, upper margin indicated by a faint
line, tentorial pits distinct, lower margin weakly emarginate; ocular-
ocellar region with costae radiating from posterior ocellus; vertex
with strong costae at posterior margin; supraclypeal area glabrous,
height 0.51 clypeus height; lateral wall of scrobe merging smoothly
with face; lower tooth of mandible pointed at apex; base of mandi-
ble with weak punctures; labio-maxillary complex short. Antenna;
scape narrowly linear, length 8.5 maximum width; pedicel and
funicular segments subequal in length (18 versus 14,15,17,16,15,
1 5, 1 3;F 1 — F7); pedicel 0.21 scape length; anellus 0.43 length of FI;
FI elongate, remaining flagellomeres transverse; clava 0.25 length of
funicle.

Mesosoma: pronotum massive, PN:MSC = 0.34, lateral pronotal
collar not regularly convex, suggesting bumpy shoulders; scutellum
acuminate with a long spine, SC:MSC = 1 .75; dorsum of pronotum

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punctate-reticulate, punctures coalesced to form weak irregular
transverse costulae medially; midlobe of mesoscutum and scutellum
weakly punctate, becoming punctate-reticulate along notauli; side-
lobe of mesoscutum smooth along notauli, laterally punctate-
reticulate; notauli distinct; scutellum in lateral view tapering abrupt-
ly towards apex (Fig. I; cf. Figs. 1,2 in Boucek 1978); underside of
scutellum mostly coriarious, with shallow convergent grooves;
propodeum vertical, medially about twice as long as metanotum,
with weak median ridge, submedian areas with transverse costulae,
callus reticulate-rugose; width of postspiracular sclerite 0.44 width
of adjacent pronotal collar, with about 10 foveae; axilla reticulate
above, costate below; axillula smooth. Forewing: submarginakmar-
ginakpostmarginakstigmal veins as 64:17:12:4; stigmal vein making
a right angle (90 degrees) with marginal vein.

Metasoma: T2 smoothly concave with weak coriarious sculpture;
T2 with sparse setae, without punctures, border between T2 and T3
indicated by a suture, laterotergite glabrous; T3 massive and con-
vex, about twice length of T2 along midline, length about equal to
maximum width (22 versus 25), evenly covered with long setae
except along T2 border and along margins of tergite, without
punctures.

Male: Unknown.

Discussion:

Kronibeinius saunion is more closely related to K. eumenidarum
than to K. megalaspis. Synapomorphies of these two species are: the
stigmal vein making a right angle with the marginal vein (oblique in
K. megalaspis and outgroup: Euperilampus and Perilampus ); cly-
peal-supraclypeal margin weak or indistinct (separated by distinct
suture in K. megalaspis and outgroup: Euperilampus and Perilam-
pus); lateral pronotal collar suggesting bumpy shoulders (regularly
convex in K. megalaspis and outgroup: Euperilampus and Perilam-
pus); and postspiracular sclerite with many foveae (a single fovea is
found in K. megalaspis, and in the ancestral species groups of Eupe-
rilampus, Darling 1983). Considering Euperilampus as the out-
group, the following similarities of K. eumenidarum and K. saunion
are regarded as plesiomorphic: propodeum medially about twice as
long as metanotum (equal to metanotum in K. megalaspis; autapo-
morphy); scutellum, in lateral view, not strongly convex, tapering
gradually towards the apex (highly convex in K. megalaspis, Boucek
1978: Fig. 2; autapomorphy).

1982]

Darling — New Species of Krombeinius

315

K. saunion and K. eumenidarum also have the inner orbits with
strong costae (Figs. 8,9,16,17), whereas the inner orbits of K. meg-
alaspis are smooth (Figs. 1 2, 1 3). 1 consider the costate inner orbits to
be a synapomorphy of Euperilampus + Krombeinius. As such 1
interpret the smooth orbits of K. megalaspis as an autapomorphic
reversal. A similar reversal in this character is indicated in the Eupe-
rilampus cladogram (Darling 1983).

There remain some difficulties in justifying the current composi-
tion of the genus Krombeinius. The numerous characters separating

Figs. 14 17. Krombeinius saunion. 14. Head, dorsal. 15. Mesosoma, dorsal. 16.
Head, frontal. 17. Head, lateral. [Scale line 1 mm.]

316

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[Vol. 89

K. eumenidarum + K. saunion from K. mega/aspis question the
inclusion of K. mega/aspis. A revised classification would create a
monobasic new genus for Perilampus mega/aspis, and would allow
Krombeinius to be defined by the synapomorphies of K. eumenic/a-
rum + K. saunion. Recalling that the proposed synapomorphies of
Krombeinius (structure of the labrum; host association) are not
known for K. mega/aspis, it would not be surprising if this species
were to be excluded at some later date. Clearly, more material and
associated biological information are essential to re-evaluate the
composition of Krombeinius, and any nomenclatural changes at
this time would be premature.

Acknowledgments

I would like to thank the following people for comments on the
manuscript: W. L. Brown, Jr., J. M. Carpenter, G. C. Eickwort, S.
W. Nichols, and Q. D. Wheeler.

The habitus drawing was skillfully prepared by Jim Miller.

This research was supported in part by a National Science Foun-
dation Dissertation Improvement Grant.

Lulrathrl; Citkd

Bocchk, Z. 1978. A generic key to Perilampinae ( Hymenoptera, Chalcidoidea),
with a revision of Krombeinius n. gen. and Euperilampus Walker. Entomolog-
ica scandinavica 9:299 307.

Darling, D. C. 1983. A review of the New World species of Euperilampus
(Hymenoptera: Perilampidae), with notes about host associations and phyloge-
netic relationships, hi press, Quaestiones entomologicae.

Domlnic'HIM, G. 1953. Studio sulla morphologia dell’addome delgi Hymenop-
tera Chalcidoidea. Bollettino di Zoologia Agraria e Bachicoltura, 19:183 298,
27 figs.

Domi nic him, G. 1969. Materiali per la morfologia comparata degli Hymenop-
tera Chalcidoidea. Memorie della Societa Entomologica ltaliana, 48:584 608,
53 figs.

Graham, M. W. R. dh V. 1969. The Pteromalidae of north-western Europe
(Hymenoptera: Chalcidoidea). Bulletin of the British Museum (Natural
History) Entomology Supplement 16. 908 pp.

Kromri in, K. V. 1964. Natural History of Plummers Island, Maryland XVIII.
The hibiscus wasp, an abundant rarity, and its associates (Hymenoptera: Sphe-
cidae). Proceedings of the Biological Society of Washington, 77:73 1 12.

Rii k, E. E. 1966. Australian Hymenoptera Chalcidoidea, Family Pteromalidae,
Subfamily Perilampinae. The Australian Journal of Zoology, 14:1207 1236.

A DESCRIPTION OF THE

ECTAL MANDIBULAR GLAND IN THE PAPER WASP

POLISTES FUSCATUS (HYMENOPTERA: VESPIDAE)*

By H. A. Downing and R. L. Jeanne
Department of Entomology
University of Wisconsin
Madison, Wisconsin 53706

While the ectal mandibular gland is a source of queen substance
in both honey bees and bumble bees (Butler and Simpson, 1958; van
Honk et al., 1980), little is known about this gland in the vespids.
In Vespula and Polistes spp. the mandibular gland consists of 50 to
70 ducted gland cells opening into a reservoir which in turn is said to
empty into the oral cavity at the base of the mandible (Hermann et
al., 1971; Spradbery, 1973; Landolt and Akre, 1979). Nedel (1960),
however, found that the mandibular gland of V. germanica (F.)
opens anterior to the anterior condyle and thus to the front of the
face. He describes a small brush of mechanoreceptors on the
mandible.

Because the mandibular gland is so much smaller in wasps than in
bees, Spradbery (1973) suggested that it probably has no social
function in wasps. However, the fact that this gland is the largest of
the cephalic exocrine glands in wasps and the discrepancies in the
literature concerning the locus of the gland opening caution against
such a conclusion. The purpose of the present study is to investigate
the morphology of the ectal mandibular gland in Polistes fuscatus
with reference to possible gland function.

Methods

Micrographs of the exterior opening of the ectal mandibular
gland were taken using the JELCO JSM-U3 scanning electron mi-
croscope. Quarter sections of female wasp heads containing the ectal
mandibular gland were fixed in Kahle’s solution and embedded in
Spurr Low-Viscosity embedding media (Polysciences) following the
methods of Spurr (1969). Sections 2/u thick were cut with a glass

* Manuscript received by the editor September 23, 1982.

317

318

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[Vol. 89

knife on a Porter-Blum-Microtome MT-1 (Sorvall®), stained using
Mallory’s Azure II-Methylene Blue (Richardson et al., 1960), and
examined using a light microscope.

Results and Discussion

The ectal mandibular gland in P.fuscatus females is similar to the
ectal mandibular gland Nedel (1960) described for V. germanica.
The gland reservoir, which lies appressed to the gena, opens at the
base of the mandible via a long, flattened, sclerotized duct. Gland
cells can be seen on the outer surface of the reservoir (Figure la).
The sclerotized duct of the reservoir opens on to the mandibular
surface just above a brush of bristles (Figure lb). The bristles may
be mechanoreceptors and may also serve to increase the surface area
for evaporation of the glandular secretion. A scallop of cuticle
extends ventrally from the gena, covering the brush when the man-
dible is closed. When the mandible is opened even slightly the brush
is exposed on the front of the face (Figure 2).

The position of the ectal mandibular gland opening suggests a
social rather than a physiological function. Because it does not open
into the mouth, this gland is probably not a source of digestive
enzymes or nest construction material. P. fuscatus colonies are
initiated by one or more overwintered female gynes, which work
together to raise the brood. Aggressive interactions result in the
formation of a dominance hierarchy in which the most dominant
individual is the egg-layer for the colony (Pardi, 1948). The domi-
nant female must maintain a certain level of aggression in order to
retain her dominant status, but her elevated rank is communicated
by chemical cues originating in the head (Downing, 1982). Domi-
nant wasps frequently chew on the head and thorax of their subor-
dinates, and when threatening other females will lunge toward them
with open mandibles. The ectal mandibular gland opening is
exposed at these times, suggesting that it may be the source of
chemical signals important for the communication of status during
aggressive interactions.

Acknowledgements

We would like to thank D. Post and B. J. Harrington for provid-
ing useful criticisms on early drafts of this manuscript. The research

1982] Downing & Jeanne — Mandibular Gland in Polistes 319

Figs. 1-2. Fig. 1. Ectal mandibular gland histology. Sagittal sections through a)
reservoir (arrow points to gland cells at the top of reservoir) and b) reservoir duct,
showing duct opening to the mandibular brush (arrow). Anterior is to the right.
Fig. 2. Ectal mandibular gland opening, frontal view. SEM micrograph of the
exposed mandibular brush, located just below the opening of the ectal mandibular
gland (arrow). The mandible is in an open position. Clypeus is on the left, compound
eye in the upper right.

320

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[Vol. 89

was supported by the College of Agricultural and Life Sciences,
University of Wisconsin, Madison, and by National Science Foun-
dation Grant BNS-77-0408 1 .

Literature Cited

Butler, C. G. and J. Simpson

1958. The source of the queen substance of the honey bee ( Apis mellifera L.).
Proc. Roy. Entomol. Soc. Lond. A. 33:120-122.

Downing, H. A.

1982. Glandular differences and communication of rank among females in a
dominance hierarchy of Polistes fuscatus (Hymenoptera: Vespidae).
M.Sc. Thesis, Univ. Wisconsin, Madison.

Hermann, H. R., A. N. Hunt, and W. E. Buren.

1971. Mandibular gland and mandibular groove in Polistes annularis (L.) and
Vespula maculata (L.) (Hymenoptera: Vespidae). Int. J. Ins. Morphol.
Embryol. 1:43-49.

Landolt, P. J. and R. D. Akre.

1979. Occurrence and location of exocrine glands in some social Vespidae
(Hymenoptera). Ann. Entomol. Soc. Am. 72:141-148.

Nedel, J. O.

1960. Morphologie und Physiologie der Mandibeldriise einiger Bienen Arten
(Apidae). Z. Morphol. Okol. Tiere, 49:139-183.

Pardi, L.

1948. Dominance order in Polistes wasps. Physiol. Zool. 21:1-13.
Richardson, K.. C., L. Jarett, and E. H. Finke.

1960. Embedding in epoxy resins for ultrathin sectioning in electron micros-
copy. Stain Technol., 35:313-323.

Spradbery, J. P.

1973. Wasps. Seattle: University of Washington Press. 408 pp.

Spurr, A. R.

1969. A low viscosity epoxy resin embedding medium for electron microscopy.
J. Ultrastructure Res. 26:31-43.

Van Honk, C. G. J., H. H. W. Velthuis, P. F. Roseler, and M. E. Malotaux.

1980. The mandibular glands of Bombus terrestris queens as a source of queen
pheromones. Entomol. Exp. Appl. 28:191-198.

SPIDERS LIVING AT WASP NESTING SITES:

WHAT CONSTRAINS PREDATION BY MUD-DAUBERS?

By Martin S. Obin1

The nests of mud-daubing wasps (Hymenoptera: Sphecidae) are
excellent sources of spiders (Peckham and Peckham, 1898; Rau,
1935; Muma and Jeffers, 1945; Dorris, 1970). Females of these soli-
tary wasp species construct mud nests during the late spring and
summer. They provision each brood cell with a number of spiders
which they capture and paralyze by stinging. The wasp lays an egg
on one of these spiders and, upon hatching, the larva consumes all
the spiders within the brood cell. When development is complete,
the new adult wasp chews a hole in its brood cell and emerges. A cell
in the nest of mud-daubers such as Sceliphron caementarium or
Chalybion californicum may contain in excess of 25 spiders. It
seems likely then that mud-dauber predation may be a significant
factor influencing population dynamics and evolution of those
spider genera taken as prey (see also Eberhard, 1970). But this view
of wasp and spider interactions is incomplete. The same sites at
which mud-daubers nest are also used by both wandering and web-
building spiders for capturing prey and tending eggs. Mud-dauber
nests themselves are often used by spiders for these activities. In
fact, among the group of spiders active at mud-dauber nesting sites
are species that are regularly taken as prey by those same spider-
hunting wasps. Intrigued by this fact, I initiated field studies that
addressed the following questions:

1. What groups of spiders are found living at nesting sites of
mud-daubers?

2. What is the nature of the interactions between wasps and spi-
ders at these sites?

3. If wasps do not hunt spiders at nesting sites, what factors
constrain them from doing so?

'Department of Zoology, University of Florida, Gainesville, FL 3261 1
* Manuscript received by the editor July 25, 1982.

321

322

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[Vol. 89

Materials and Methods

Procedure. Nesting sites and mud nests of three species of sphecid
wasp were observed during the summer of 1980. Three groups of
spiders were collected. These were: (a) active spiders within 2 meters
of wasp nesting sites, (b) active spiders on or next to mud-dauber
nests, and (c) paralyzed spiders from inside 110 old nests. Spiders
were identified and their total body lengths measured. No attempt
was made to census every spider at each site, as this would have
proven impossible for genera such as Tidarren and Filistata which
were numerous, mobile and often reclusive. In addition, encounters
between spiders and wasps were observed and recorded.

The Wasps. Sceliphron caementarium (Drury) (Sphecinae: Sce-
liphrini) constructs individual cells of mud collected at the edges of
ponds and streams. The nests are provisioned primarily with Ara-
neidae, Thomisidae and Salticidae (Muma and Jeffers, 1945) and
then sealed off with mud. Groups of contiguous cells are often
covered by additional layers of mud and may appear as oval or
oblong masses of up to 30 cells (see Muma and Jeffers, 1945 for
plates of relevant mud-daubers and their nests). Trypoxylon poli-
tum Say (Larrinae: Trypoxylini) builds long, tubular nests from
mud gathered at sites similar to those frequented by Sceliphron.
The “pipe organ” nests usually contain between 3 and 5 cells, each
provisioned with Araneidae of the genera Neoscona or Eustala
(Muma and Jeffers, 1945; H. J. Brockmann, pers. comm.). Rather
than constructing its own nest, the blue mud-dauber, Chalybion
californicum (Saussure) (Sphecinae: Sceliphrini) either modifies and
seals existing old cells of Sceliphron and Trypoxylon or cleans out
and reprovisions recently completed cells. Dry mud from nearby
nests is softened by mixing with water stored in the wasp’s crop. The
wet mud is then manipulated and used for sealing nests. Chalybion
specializes in hunting small Theridiidae and Araneidae (Muma and
Jeffers, 1945).

The Study Sites. Three sites in Alachua County, Florida were
selected. Two were located in the Paynes Prairie State Preserve and
were designated Boat House (BH) and Garage (G). Both sites had
females of all three species actively building and provisioning nests.
The boat house site had a 10 m X 25 m X 1 m high shaded crawl
space with an unfinished pine ceiling, dirt floor and open sides. The

1982]

Obin — Spiders Living at Wasp Nesting Sites

323

structure was within 5 m of a lake and was surrounded on three
sides by a lawn dotted with palms and turkey oak. A dense stand of
palmetto and hardwoods was located ca. 100 m distant. The Garage
site was located 150 m from the lake shoreline and was next to a
small plot of palmetto-hardwood forest. Wasp nests covered the
exterior walls of this painted wood structure and were exposed to
ambient light. The third site, Rocky Creek (RC) was two cement
bridge tunnels where State Road 121 crossed Rocky Creek. During
the study, the water level was sufficiently low such that the sand
bottom of the creek was exposed throughout most of the two tun-
nels. The tunnel entrances were fringed with tall grass, occasional
shrubs and Eupatorium sp. Light levels inside the tunnels were the
lowest among the 3 sites. The area surrounding the site was com-
posed of cleared agricultural plots interspersed with thickets and
small stands of oak and pine.

Results

Table 1 lists the spiders observed at the 3 sites. Prey species are
distinguished from non-prey species and web spiders from wander-
ing spiders.

Web-Building Spiders. Eighty-three web spiders were collected,
representing 12 genera in 4 families. Ten genera were taken as prey
by the mud-daubers nesting at the study sites. Species of three gen-
era of spiders were found living in open mud cells from which wasps
had emerged earlier in the season. Males and females of Metazygia
wittfeldae (McCook), Filistata hibernalis Hentz and Oecobius annu-
lipes (Lucas) were removed from inside old nests of Sceliphron and
Chalybion that were constructed over or close to seams and cracks
in walls. Of 1 1 cells containing M. wittfeldae, 8 also contained egg
cases. Two old cells with adult pairs and spiderlings inside were also
noted. Genera of Araneidae and Theridiidae positioned webs either
close to nesting sites (Argiope, Nephila, Micrathena and Neoscona)
or within 10-15 cm of active nests (Leucauge, Tetragnatha, Tidar-
ren, Latrodectus and Achaearanea). Webs of Argiope aurantia
Lucas were found only at Rocky Creek, where the tall grass and
bushes at the tunnel entrance afforded suitable habitat. The distribu-
tion of Micrathena sagittata (Walckenaer) appeared similarly limited
by habitat, as webs were confined to the wood’s edge behind the
Garage site.

324

Psyche

[Vol. 89

Wandering Spiders. Five families were collected, totalling 43 spi-
ders in 8 genera. Five of these genera are common prey items of
mud-daubers. A Xysticus sp. was discovered inhabiting an inactive
Sceliphron nest and a female Platy cry plus undatus (De Geer) occu-
pied a half-completed Sceliphron cell, constructed a retreat and
positioned herself at the entrance. Species of Phidippus and Thio-
dina climbed over nests, but did not remain on these structures.
However, species of Dolomedes were frequently noted on the out-
side of mud nests. They remained motionless for hours during the
day and appeared to achieve an enhanced crypsis against the nest
background.

Spider Size. Seventy-six specimens belonging to genera taken as
prey by mud-daubers were collected at the 3 sites. Of these, only 9
exceeded the upper size range of congeners found paralyzed in wasp
cells (Table 1). The 2 Argiope listed were also larger than conspecif-
ics (N=3) that could not be handled (i.e., were repeatedly dropped
after immobilization) by Sceliphron. These spiders were 15.0, 16.5,
and 16.8 mm long respectively. Two spiders dropped by Chalybion
were 15.3 mm \Pisaurina undulata (Keyserling)] and 14.6 mm long
[Peucetia viridans (Hentz)]. The largest P. undulata provisioned by
Chalybion was 1 1.8 mm (N=3), and the largest P. viridans was 13.8
mm (N=14). Extensive data for Trypoxylon, generously provided
by Dr. H. J. Brockmann, indicated that the heaviest of 289 Neo-
scona provisioned by Trypoxylon during June and July weighed
0.2400 grams. Spiders dropped by provisioning females exceed this
weight on 12 occasions, ranging in weight from 0.2537-0.4236 g.
(Spiders were weighed to the nearest 0.1 mg.)

Wasp-Spider Interactions. Surprisingly, predation by mud-daubers
on spiders living at nest sites was never observed. Brockmann (pers.
comm.), who has spent over 3,000 hours observing wasps under
bridges near Gainesville, has also never observed a single case of a
wasp preying on a spider near the nesting site. Wasps repeatedly
walked or flew within several centimeters of potential prey, display-
ing no observable taxes or predatory movements. In 2 separate
incidents, female Sceliphron that had strayed into webs of Tidarren
sisyphoides (Walckenaer) freed themselves after stinging the overly
eager spider. In neither instance did the wasp show any further
interest in the potential prey item, although in both cases the spider

1982]

Obin — Spiders Living at Wasp Nesting Sites

325

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1982]

Obin— Spiders Living at Wasp Nesting Sites

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NON-PREY SPIDERS
Web Spiders

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328

Psyche

[Vol. 89

was paralyzed. Rather, after extricating themselves, both flew to
nearby vegetation and groomed intensively before returning to their
nests.

However, a number of spiders were observed preying on mud-
daubers. Wasps carrying very large prey or those having just left the
nest sometimes blundered into webs situated directly in the flight
path of the wasp. All 3 species of wasp were found entangled in webs
of either Argiope aurantia, Neoscona domiciliorum (Hentz), Latro-
dectus mactans Fabricius, Tidarren sisyphoides, Nephila clavipes
(Linnaeus), or Filistata hibernalis Hentz. Fourteen wasps were
observed wrapped or trapped in webs. These included Trypoxylon
(7), Sceliphron (5) and Chalybion (2). One of the Sceliphron noted
was an emerging adult that was trapped in the sticky threads of F.
hibernalis that covered its cell. Micrathena webs, positioned less
than 0.5 meters off the ground, were too low to intercept wasps in
flight. Smaller spiders such as T. sisyphoides and the smaller indi-
viduals of L. mactans and N. domiciliorum did not always attack
wasps caught in webs. These spiders retreated out of the range of the
struggling wasp on 5 occasions. Biting or wrapping were delayed
until the wasp had exhausted itself and was quiescent. In contrast,
two A. aurantia observed at Rocky Creek immediately descended
from the hub, wrapped the prey, inflicted a “short bite” (Robinson,
1969) and then returned to the hub before further wrapping com-
menced. One Sceliphron and 3 Trypoxylon were dispatched in this
manner. Predation on wasps by spiders is not restricted to web-
builders. Brockmann (pers. comm.) has observed attempted preda-
tion on mud-daubers by a Dolomedes sp. which leapt off a tunnel
wall while attempting to grab a Trypoxylon hovering nearby. The
spider was unsuccessful and pulled itself up the wall by the dragline.
It is possible that large, mobile spiders of this type are preying on
male Trypoxylon that sleep in the mud nests during the night.

Discussion

Barns, old houses, bridges and the vegetation surrounding them
afford appropriate habitat for many groups of spiders. The mud-
daubing wasps are similarly attracted to such sites, for when water
and mud are available, these sites provide favorable nesting sub-
strate. It is not surprising therefore to observe spiders and spider-
hunting wasps living in close proximity. It is noteworthy, however,

1982]

Ob in — Spiders Living at Wasp Nesting Sites

329

when these spiders (76 of 107 collected in our study) are prey species
of the nesting wasps, for it suggests that these spiders escape preda-
tion. This study addressed that question in particular.

Why Mud-Daubers Do Not Hunt at Nest Sites. The inability of
mud-daubers to recognize and capture prey living amongst them
poses interesting questions. As demonstrated by Tinbergen (1935)
for the solitary wasp Philanthus triangulum (Fabricus), successful
predation may involve the hierarchical sequencing of various “ap-
petitive behaviors” (Craig, 1918), each controlled by a specific
releasing stimulus. In Sceliphron, for example, visual releasers such
as spider-sized objects on a contrasting background are known to
release a predatory pounce from a wasp in flight (Eberhard, 1970),
but it is highly probably that wasp search images vary between
habitats. Conspecifics hunting in the canopy and those hunting in
the leaf litter may respond to learned visual cues appropriate to the
particular microhabitat being searched. It is possible then that mud-
daubers were catching spiders against backgrounds different from
those presented to them at the 3 study sites. This might explain in a
proximate sense why we observed no wasps attacking spiders at
these sites. One might also propose that mud-daubers require a
minimum light level to activate particular behaviors of the preda-
tory sequence. However, nests at the Garage site were not in shade,
and no hunting by wasps was observed. Are spiders at nest sites too
large for mud daubers to immobilize and provision? The data
strongly suggest otherwise, as less than 12% of potential prey col-
lected during the study exceeded the upper range of spiders found
paralyzed in wasp nests (Table 1).

One ultimate explanation of why “leave the nest site” appears to
be a behavioral rule for foraging mud-daubers posits the importance
of spider predation on wasps. Spiders that have previously encoun-
tered a wasp may be more likely to successfully defend themselves
from subsequent wasp attack, and the probability of attacking a
spider that has previously encountered and successfully handled a
wasp may be greater close to or at nest sites than it is at a distance
from such sites. Moreover, it is possible that spiders may learn to
recognize characteristic vibrational signatures of mud-daubers. Such
pretactile prey determination has been hypothesized (Robinson and
Mirick, 1971), although Suter (1978) could not identify such a
mechanism in the araneid Cyclosa turbinata (Walckenaer).

330

Psyche

[Vol. 89

It is also possible that by not hunting at nest sites, mud-daubers
more effectivley conceal the location of their nest. Such a mecha-
nism has been proposed to explain why raptors usually do not hunt
near their own nests (Durango, 1949). Mud-dauber larvae fall prey
to a variety of parasitoids and inquilines (Rau and Rau, 1916;
Krombein, 1967). Hunting away from the nest site can reduce the
probability of parasitization if the following assumptions are met:
(1) The parasite encounters the host species at sites where the host
species hunts; (2) The parasite trails the host species back to the
nest; (3) The host species can evade the trailing parasite, the proba-
bility of so doing increasing with the distance over which the host
species is trailed.

Certain host-parasite systems involving mud daubers and sarco-
phagid flies meet the above assumptions. Flies in the tribe Milto-
grammini are larviporous parasites of many aculeates, including
sphecid wasps (Allen, 1926). The genera Amobia and Senotainia
include species of mud-dauber parasites that follow prey-laden
wasps to their nests (Chapman, 1959; Cole, 1969). The adult flies are
nectivorous, and it is likely that they encounter foraging wasps on
vegetation. Prey-laden mud daubers often fly at reduced speeds, and
their maneuverability is similarly impaired (Obin, pers. obs.). They
are presumably easier to follow at such times. Furthermore, a wasp
with prey assures a trailing fly that the wasp is nesting, that a cell is
being provisioned and is consequently open, and that there will be
food available in that cell. Sarcophagid flies have been observed
trailing C. calif ornicum females to their mud nests over distances of
3-5 m. The pursued wasps often took circuitous routes to their
nests, and in certain instances left the site altogether before reaching
their nest (Obin, unpublished data). Whether such behavior results
in successful evasion is not known, but it does suggest that wasps
may require flight distances greater than those observed in order to
evade trailing Miltogrammini. If so, selection may, on average,
favor wasps that do not hunt close to their nest.

Interactions between spiders and mud-daubers may not be exclu-
sively antagonistic, and the selective advantage accruing to wasps
that do not hunt at nest sites may be a consequence of a site-specific
mutualism between these two traditional enemies. During the study,
various parasites of mud-daubers were observed in webs at nest
sites. These included bombyliid and sarcophagid flies as well as

1982]

Obin — Spiders Living at Wasp Nesting Sites

331

chrysidid and mutillid wasps. These observations suggest that spider
predation may reduce the parasite load at mud-dauber nest sites.
Since mud-daubers are usually adept at recognizing and maneuver-
ing on webs and retreats (Eberhard, 1970; Coville, 1976), spiders at
nest sites may pose only a limited threat to wasps. We have observed
individuals of all three species of wasp successfully nidify and
provision nests positioned such that the wasp flew through or
walked behind a web on each trip to and from the nest. Wasps
became entangled in webs when their regular flight path was dis-
rupted during agonistic encounters or when they attempted to pro-
vision very large spiders. Empirical evaluation of the relative costs
(e.g., probability of predation, costs associated with increased flight
distance to foraging patches) and benefits (reduced parasitism) of
hunting away from the nest site is in progress. One predication of
the “reduced parasitism” hypothesis is that a small percentage of
mud daubers at any site may “cheat” — i.e., may occasionally prey
on spiders at nest sites. Relative to other wasps in the population,
these wasps would enjoy reductions in the time and energy costs of
hunting and transporting prey, while at the same time benefitting
from the “parasite umbrella” afforded by spiders active at nest sites.
The relative frequencies of cheating and non-cheating (hunting
away from nest sites) may perhaps be maintained by frequency-
dependent selection (Fisher, 1930) in an Evolutionarily Stable
Strategy (Maynard Smith and Price, 1973).

The Effects of Prey Size and Availability on Mud- Daubers. Vari-
ous authors have suggested that spider size constrains prey collec-
tion by Chalybion (Muma and Jeffers, 1945), Sceliphron (Muma
and Jeffers, 1945; Eberhard, 1970) and Trypoxylon (Cross et al.,
1975). Selection should favor wasps that minimize both the risks
and metabolic cost of (1) immobilizing and (2) transporting large
spiders. Do wasps refrain from attacking large prey that they can
incapacitate but not readily transport, or is the upper range of prey
size found in nests a reflection of the wasp’s inability or reluctance
to paralyze prey above a certain size? Measurements of spiders
dropped by provisioning mud daubers indicate that the wasps suc-
cessfully incapacitate spiders that exceed the upper range of prey
size noted in nests, but fail in their ability to transport or cache
them.

332

Psyche

[Vol. 89

The data differ from those of Muma and Jeffers regarding prey
selection by Chalybion and Sceliphron. In their study in Maryland,
the theridiid Latrodectus mactans constituted 25% of all prey taken
by Chalybion. Locally, L. mactans is provisioned less frequently,
comprising less than 5% of all prey taken. These differences may be
due in part to the availability of more prey species in Florida.
Spiders not found in Maryland nests but taken by Chalybion and
Sceliphron in Florida include Tetragnatha guatemalensis O. P.-
Cambridge, Tetragnatha pallescens F. P. -Cambridge, Pisaurina
undulata, Mecynogea lemnis cat a (W alckenaer), G aster acantha can-
criformis (Linnaeus), and Tidarren sisyphoides. M. lemniscata and
G. cancriformis were also found provisioned in Sceliphron cells.
Maryland constitutes the northern most distribution for M. lemnis-
cata (Kaston, 1978). G. cancriformis ranges only as far north as
North Carolina (Levi, 1978). The distribution of T. sisyphoides in
North America is restricted to the southern United States and Mex-
ico (Levi, 1955).

Nest Sites as Spider Habitat. Mud-dauber nest sites may be par-
ticularly good habitats for some spiders. Benefits to spiders at such
sites include the following:

1. Mud nests afford environmentally buffered refugia and brood
chambers and may provide cryptic backgrounds.

2. Additional prey is available, including the wasps themselves,
other spiders (Tolbert, 1975), mites and hymenopterous and
dipterous parasites of mud-daubers, “renting” Arthropods
that use empty mud-dauber cells (Dermaptera, lepidopterous
larvae and non-sphecid wasps) and nest associates such as
Psocoptera.

3. The risk of predation from wasps nesting at these sites is
reduced.

Although somewhat counter-intuitive, the probability of wasp
predation appears to be lower for spiders living at sites where wasps
nest. The greatest threat may exist for smaller, naive individuals,
and a small cost is probably incurred by those spiders whose web is
damaged after intercepting a wasp. However, the benefits of living
amidst mud-daubers may outweigh these potential costs. Associa-
tions of predator and prey at predator nests sites are not without
precedent. Nesting of passerines with raptors has been reported

1982]

Obin — Spiders Living at Wasp Nesting Sites

333

(Durango, 1949; McGillivray, 1978; Parker, 1981). Since raptors
hunt away from their nests, traditional prey species nesting close by
appear less threatened (Brown and Amadon, 1968; Parker, 1981).

Acknowledgments

The author wishes to thank Dr. H. Jane Brockmann for review of
the manuscript and Jeff Lucas for advice and suggestions. G. B.
Edwards and Jon Kochalka offered invaluable assistance identify-
ing problematic specimens. An anonymous reviewer suggested con-
sulting the avian literature. Special thanks are due the Florida
Department of Natural Resources and the rangers at the Paynes
Prairie State Preserve.

Literature Cited

Allen, H. W.

1926. North American species of two-winged flies belonging to the tribe milto-
grammini. U.S. Natl. Mus. Proc., 68(9): 1-106.

Brown, L., and D. Amadon.

1968. Eagles, hawks and falcons of the world. McGraw-Hill, New York.

Chapman, R. F.

1959. Some observations on Pachyopthahnus africa Curran, a parasite of
Eumenes macillosus De Geer. Proc. Roy. Entomol. Soc. London, 34:
1-6.

Coville, R. E.

1976. Predatory behavior of the spider wasp Chalybion calif ornicum (Hymen-
optera: Sphecidae). The Pan-pacific Entomologist, 52: 229-233.

Cole, F.

1969. The flies of western North America. University of California Press,
Berkeley.

Craig, W.

1918. Appetites and aversions as constitutents of instincts. Biol. Bull., 34:
91-107.

Cross, E. A., M. G. Smith, andT. R. Bauman.

1975. Bionomics of the organ-pipe mud-dauber, Trypoxylong politum (Hy-
menoptera: Sphecidae). Ann. Ent. Soc. Amer., 68: 901-916.

Dorris, P. R.

1970. Spiders collected from mud-dauber nests in Mississippi. J. Kans. Ent.
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Durango, S.

1949. The nesting associations of birds with social insects and with birds of
different species. Ibis, 91: 140-143.

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Psyche

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Eberhard, W.

1970. The predatory behavior of two wasps, Agenoideus humilis (Pompilidae)
and Sceliphron caementariuim (Sphecidae) on the orb-weaving spider,
Araneus cornutus (Araneidae). Psyche, 77(2): 243-251.

Fisher, R. A.

1930. The Genetical Theory of Natural Selection. Claredon Press, Oxford.

K ASTON, B. J.

1978. How To Know The Spiders. Third Edition. Wm. C. Brown and Com-
pany. Dubuque, Iowa.

Krombein, K.

1967. Trap-nesting wasps and bees. Life histories, nests and associates. Smith-
sonian Press. Washington, D.C.

Levi, H. W.

1955. The spider genera Chrysso and Tidarren in America (Araneae: Theridii-
dae). J. New York Ent. Soc., LXIII. 59-81.

1978. The American orb-weaver genera Colphepeira, Micrathena, and Gaste-
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Zool., 148(9): 417-422.

Maynard Smith, J. and G. R. Price.

1973. The logic of animal conflicts. Nature, 246: 15-18.

McGillivray, W. B.

1978. House Sparrows nesting near a Swainson’s Hawk Nest. Can. Field-Nat.,
92: 202-203.

Muma, M. H., and W. F. Jeffers.

1970. Studies of the spider prey of several mud-dauber wasps. Ann. Ent. Soc.
Amer., 38: 245-255.

Parker, J. W.

1981. Nest Associates of the Mississippi Kite. J. Field Ornithol., 52(2):
144-145.

Peckham, G. W., and E. G. Peckham.

1898. Instincts and habits of the solitary wasps. Bull. Wise. Geol. and Nat.
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Rau, P.

1935. The spider prey of the mud wasp. Sceliphron caementarium (Aranea,
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Rau, P., and N. Rau

1916. The biology of the mud-daubing wasps as revealed by the
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Robinson, M. H.

1969. Predatory behavior of Argiope argentata (Fabricius). Am. Zool., 9:
161-173.

Robinson, M. H., and H. Mirick.

1971. The predatory behavior of the golden web spider, Nephila
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Suter, R. B.

1978. Cyclosa turbinata (Aranea, Araneidae): Prey discrimination via web-
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Tinbergen, N.

1935. liber die Orientierung des Bienenwolfes ( Philanthus triangulum (Fabr.))
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Tolbert, W. W.

1975. Predator avoidance behaviors and web defensive structures in the orb
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A GA TH I DIODES PORTEVIN,

NEW SYNONYM OF STETHOLIODES FALL
(COLEOPTERA: LEIODIDAE: ANISOTOMINI)*

By Alfred F. Newton, Jr.

Museum of Comparative Zoology
Harvard University, Cambridge, Mass. 02138

Examination of type material of many obscure genera of Leiodi-
dae for a work in preparation on the suprageneric classification of
the family has revealed a new generic synonymy in the tribe Aniso-
tomini (=Agathidiini).

Stetholiodes Fall, described for a single species S. laticollis Fall
from Indiana, USA (Fall 1910), has recently been redescribed by
Wheeler ( 1981) who discussed the close relationship of the genus to
Agathidiuni Panzer.

The genus Agathodes Portevin was described for a single species
A. striatipenne Portevin from Kashmir, India (Portevin 1926). Port-
evin later (1944) proposed the new name Agathidiodes to replace
Agathodes Portevin 1926 (not Guenee 1854). He considered Aga-
thidiodes to be closely related to Agathidiuni.

Stetholiodes and Agathidiodes are each known only from the
holotype male of the type species. Direct comparison of these two
specimens (examined dry with a dissecting microscope and on tem-
porary slides in lactophenol with a compound microscope) shows
that the two species are extremely similar in all characteristics that
have been used at the generic and subgeneric level in Anisotomini. 1
therefore propose the following synonymy:

Stetholiodes Fall
= Agathidiodes Portevin, new synonymy
= Agathodes Portevin (not Guenee)

The two included species, Stetholiodes laticollis Fall and S. stria-
tipennis (Portevin) (new combination), show slight differences in
shape, sculpture, male secondary sexual characters and the shape of

* Manuscript received by the editor May 12, 1982

337

338

Psyche

[Vol. 89

the median lobe and parameres of the aedeagus. They are thus
evidently not conspecific. In S. striatipennis the basal three tarso-
meres of the protarsus and basal two tarsomeres of the mesotarsus
are dilated and bear tenent setae, while in S', laticollis the basal three
tarsomeres of both legs are similarly modified. It should be noted
that Portevin (1926) erred in describing this character for striati-
pennis as well as in attributing a 5-5-5 tarsal formula to this species
(tarsi are 5-5-4 segmented in S. striatipennis and S. laticollis).

The genus Stetholiodes has been well characterized by Wheeler
(1981), whose description is virtually unmodified by the addition of
S. striatipennis. I would add that both Stetholiodes species lack an
epistomal suture and have a supraocular carina and groove that
separate the side of the head (including the eyes) from the dorsum.
This last character is found in most or all Agathidium but is absent
in Anisotoma and allied genera of Anisotomini. I agree with
Wheeler that Stetholiodes is closely allied to, and possibly conge-
neric with, Agathidium. At present Stetholiodes appears to differ
from Agathidium only in having nine distinct punctate elytral striae,
rather than fewer or no striae, and in lacking an epistomal suture.
Further study of the large and diverse genus Agathidium is needed
to clarify the status of Stetholiodes.

I thank Mile. Nicole Berti of the Museum National d’Histoire
Naturelle, Paris, for loan of the holotype of Agathidiodes striati-
penne; and Fernando Angelini, Hermann Daffner, Stewart B. Peck
and my wife, Margaret K. Thayer, for commenting on the manus-
cript. Dr. Angelini has noted a recent collection of two males of S’.
striatipennis from Aru, Kashmir, October 1977, leg. H. Franz, now
in his collection and that of Dr. Franz.

Literature Cited

Fall, H. C.

1910. New Silphidae of the tribe Anisotomini. Can. Ent. 42: 4-8.

Portevin, G.

1926. Les Liodidae de l’lnde. Encycl. Ent. (B), Coleoptera 1: 75-83.

1944. Liodides nouveaux. Rev. Frang. Ent. 10: 168-169.

Wheeler, Q. D.

1981. Diagnosis and phylogenetic relationships of the monotypic genus Ste-
tholiodes (Coleoptera: Leiodidae). Ohio J. Sci. 81: 165-168.

FOSSIL TIGER BEETLES (COLEOPTERA: CICINDELIDAE):
REVIEW AND NEW QUATERNARY RECORDS

By Christopher D. Nagano1, Scott E. Miller2
and Alan V. Morgan

Introduction

Fossil Cicindelidae are extremely rare in the stratigraphic record,
probably due to the fragile nature of their exoskeleton. In this paper
we summarize previous records and comment on new finds, as well
as describe cicindelid specimens found in the southern California
asphalt deposits which were noted, but not identified, by Pierce
(1947a, 1947b).

Southern California Asphalt Deposits

Both the well known McKittrick asphalt deposit in Kern County,
California and the Rancho La Brea sequence, Los Angeles County,
California, have produced identifiable cicindelid specimens. Pierce’s
specimens are deposited in the Natural History Museum of Los
Angeles County (LACM). A fairly well-preserved specimen of Cicin-
dela haemorrhagica LeConte (LACM Invert. Paleo. hypotype 4944)
from the McKittrick asphalt deposit, retains complete markings on
the elytra, and the elytral pleura still show a blue coloration similar
to that of populations found along the sea coast of central San
Diego County, California. A mandible (LACM Invert. Paleo. hypo-
type 4945) also from McKittrick, is probably a cicindelid but family
placement is uncertain due to the poor condition of the specimen.
Both McKittrick specimens were collected by L. Bessom from W.
D. Pierce’s “site 4” (LACM Invert. Paleo. Loc. 260), at a depth of 4
feet (1.3m) (Pierce 1947b, Miller and Peck, 1979). This site has not
yet been radiocarbon dated, but a Cybister elytron taken from a
depth of 3 feet (l m) in the road cut at McKittrick has provided an

‘Natural History Museum of Los Angeles County, Los Angeles, California 90007.

2Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts
02138.

^Department of Earth Sciences, University of Waterloo, Waterloo, Ontario, Canada
N2L3G1.

Manuscript received by the editor May l, 1982.

339

340

Psyche

[Vol. 89

experimental sample for a 14C mass spectrometer run at Chalk River
Nuclear Laboratory, Ontario, Canada. The resultant age of ca 8,000
yr B.P. suggests that the samples are probably of Holocene age,
(Miller and Peck 1979, Morgan and Morgan 1980a) or of very late
Pleistocene age as stated by Pierce (1947b).

Three Cicindela specimens are known from Rancho La Brea; two
thoraces of C. haemorrhagica (RLP 8779E and RLP 9014E) and a
metasternum of C. oregona LeConte (RLP 9465E). RLP 8779E and
9465E are from Pierce’s “Bliss 29” material, which was collected in
1929 by W. Bliss from the sites of pits A, B, and C. The samples are
probably late Pleistocene in age, but this is questionable due to
unknown locality and possible contamination (Miller and Peck,
1979). RLP 9014E is from Pierce’s “Pit X”, which refers to mixed
material, lacking data, and of questionable age.

The Quaternary presence of C. haemorrhagica and C. oregona in
southern California is not unexpected. Both are presently widely
distributed in fresh and marine littoral habitats in western North
America. Although these two species are not narrowly restricted to
specific microhabitats, they are always found near permanent sour-
ces of water.

New Records of Fossil Cicindelids Elsewhere
in North America

Beside the above mentioned localities, a rekindled interest in the
examination of coleopterous faunas has recently revealed fragments
of cicindelids in a number of sites in the United States and Canada
(Fig. 1). All but one of these records are more recent than the last
review of North American fossil insects (Morgan and Morgan
1980b). The oldest specimen which is stratigraphically interpreted as
pre last interglacial (pre-Sangamon) is a partial elytral fragment
which is probably of the genus Omus from the Mountain View
-Dump site near Palo Alto, California (D. Adam pers. comm. 1978,
Morgan unpublished). In the Pacific northwest, a recently examined
site (Nelson and Coope, 1982) from Discovery Park, Fort Lawton,
Seattle, has produced the remains of Cicindela oregona LeConte. A
large and varied assemblage accompanies this find which is from
sediments previously radiocarbon dated at between 23,000 and
18,000 yr. B.P. The site pre-dates the last major (Vashon) ice
advance in the area and suggests a cooler climate with more open

1982] Nagano, Miller & Morgan — Fossil Tiger Beetles

341

vegetation, an interpretation similar to that made from another
18,000 year old coleopterous assemblage from Port Moody in
southern British Columbia (Miller, Morgan and Hicock, 1982).

In the central eastern section of the continent three sites post-
dating the retreat of Laurentide ice have produced cicindelid
remains. The Norwood site in Minnesota (Ashworth et al., 1981) is a
late-glacial kettle which has a sequence of silts overlain by peat
dated at 12,400±60 yr. B.P. (QL-1083). A left elytron of Cicindela
cf. C. sexguttata Fabricius was recovered from the upper silt, an
horizon characterized by a number of open ground beetle species. In
the Canadian province of Ontario two sites at Kitchener and
Brampton contain specifically identified cicindelids. The sites are
approximately equivalent in age to the Norwood locality (ca. 12,400
to 12,000 yr. B.P.) and both slightly post-date the last major (Port
Huron) ice readvance in the region. The Gage Street site. Kitchener,

342

Psyche

[Vol. 89

produced a solitary, well-preserved mandible of Cicindela repanda
Dejean from the basal level of a marl deposit (Schwert, 1978) found
in association with open-ground but largely boreal species. The
Brampton site near Toronto, is a kettle deposit from which a soli-
tary well-preserved right elytron and mandible fragment of Cicin-
dela limbalis Klug was recovered (Morgan and Freitag, 1982). Once
again the cicindelid fragments were associated with a fauna resident
today in open ground regions within the boreal zone (Morgan,
Morgan and Motz, 1982).

The presence of cicindelids in these early deposits of late Wiscon-
sinan sequences is not surprising. In all cases, with the exception of
the California examples, the tiger beetle remains are associated with
species which inhabit open ground situations. Undoubtedly the ice
merely forced many cicindelid populations southward at the time of
maximum advance and they remained there to successfully recolo-
nise sandy terrain after ice retreat. Tiger beetle remains also are
present due to the nature of the sediments; the very fine silts and
clays which are typical of most of these sequences is ideal for the
preservation of the extremely thin elytral chitin found in cicindelids.
In coarse sediments, or in sequences which are organic-rich, the
detritus would abrade, distort and fragment the remains to a degree
where most skeletal parts would become unrecognizable.

In Table 1, we have attempted to compile known fossil Cicindeli-
dae records including those described in this paper, and we have
also commented, where appropriate, on some of the early identifi-
cations.

Table I: Known Fossil Cicindelidae1

343

1982] Nagano, Miller & Morgan — Fossil Tiger Beetles

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1982] Nagano, Miller & Morgan — Fossil Tiger Beetles

345

Acknowledgements

We would like to thank C. L. Hogue, P. C. Owen, C. A. Shaw
and E. C. Wilson (all of LACM), J. V. Matthews Jr. (Geological
Survey of Canada), A. Morgan (University of Waterloo), R. E.
Nelson (University of Seattle) and D. P. Schwert (North Dakota
State University) for providing specimens and data.

Literature Cited

Ashworth, A. C., Schwert, D. P., Watts, W. A. and H. E. Wright, Jr. 1981. Plant
and Insect fossils at Norwood in south-central Minnesota: A record of Late-
glacial succession. Quaternary Res. 16: 66-79.

Blair, K. G. 1927. Insect remains from oil sand in Trinidad. Trans. Ent. Soc.
Lond. 75: 137-141.

Bogachev, A. 1948. Fauna of the Binagady asphalt deposit: beetles-Coleoptera.
Trudy Estestvenno-Istoricheskii Muzei, Baku 1-2: 137-160. [in Russian].

Cockerell, T. D. A. 1920. Eocene insects from the Rocky Mountains. Proc.
U.S. Nat. Mus. 57: 233-260.

Coope, G. R. 1959. A Late Pleistocene insect fauna from Chelford, Cheshire.
Proc. Roy. Soc. (London) Ser. B. 151: 70-86.

Coope, G. R. 1977. Fossil coleopteran assemblages as sensitive indicators of cli-
matic changes during the Devensian (Last) cold stage. Phil. Trans. Roy. Soc.
(London) Ser. B. 280: 313-340.

Coope, G. R. 1979. The Carabidae of the glacial refuge in the British Isles and
their contribution to the post glacial colonization of Scandinavia and the North
Atlantic Islands, pp. 407-424 in Erwin, T. L. et al. (ed.), Carabid beetles: their
evolution, natural history, and classification. Junk, The Hague.

Coope, G. R. and J. A. Brophy. 1972. Late Glacial environmental changes indi-
cated by a coleopteran succession from North Wales. Boreas 1(2): 97-142.

Coope, G. R. and C. H. S. Sands. 1966. Insect faunas of the last glaciation from
the Tame Valley, Warwickshire. Proc. Roy. Soc. (London) Ser. B. 165:
389-412.

Horn, G. H. 1876. Notes on some coleopterous remains from the bone cave at
Port Kennedy, Pennsylvania. Trans. Amer. Ent. Soc. 5: 241-245.

Horn, W. 1906. Ueber das verkommen von Tetracha Carolina L. im preufsischen
bernstein und die phylogenie der Cicindela- arten. Deut. Ent. Zeit. pp. 329-336.

Horn, W. 1907. Brulle’s “ Odontochila aus dem baltischen Bernstein” und die
Phylogenie der Cicindeliden. (Col.). Deut. Ent. Zeit. pp. 461-466.

Kurten, B. and E. Anderson. 1980. Pleistocene mammals of North America.
Columbia Univ. Press.

Larsson, S. G. 1978. Baltic amber — a palaeobiological study. Entomonograph 1:
1-192.

Matthews, J. V., Jr. 1976. Insect fossils from the Beaufort Formation: Geologi-
cal and biological significance. Geol. Surv. Can., Paper 76-1B: 217-227.

346

Psyche

[Vol. 89

Matthews, J. V., Jr. 1977. Tertiary Coleoptera fossils from the North American
Arctic. Coleop. Bull. 31: 297-308.

Miller, R. F., Morgan, A. V. and S. R. Hicock. 1982. A pre-Vashon insect
assemblage from the Fraser lowland, British Columbia. Abs. Vol. VII, AMQUA
Conf. Seattle, Wash., 141.

Miller, S. E. and S. B. Peck. 1979. Fossil carrion beetles of Pleistocene Califor-
nia asphalt deposits, with a synopsis of Holocene California Silphidae (Insecta:
Coleoptera: Silphidae). Trans. San Diego Soc. Nat. Hist. 19: 85-106.

Morgan, A. V. and A. Morgan. 1980a. Beetle bits — the science of paleoentomol-
ogy. Geoscience Canada 7: 22-29.

Morgan, A. V. and A. Morgan. 1980b. Faunal assemblages and distributional
shifts of Coleoptera during the Late Pleistocene in Canada and the Northern
United States. Can. Ent. 112: 1 105-1 128.

Morgan, A. V. and R. Freitag. 1982. The occurrence of Cicindela limbalis
Klug (Coleoptera: Cicindelidae) in a late-glacial site at Brampton, Ontario.
Coleop. Bull. 36:105-108.

Morgan, A. V., Morgan, A. and J. Motz. 1982. Fossil insect assemblages from
the base of a late-glacial sequence near Brampton, Ontario. Prog, with Abs.
Geol. Assoc. Can. Annual Meeting, Winnipeg 67.

Nelson, R. E. and G. R. Coope. 1982. A pre-Vashon (Late Pleistocene) insect
fauna from Seattle, Washington. Abs. Vol. VII, AMQUA Conf. Seattle, Wash.,
146.

Osborne, P. J. 1972. Insect faunas of Late Devensian and Flandrian age from
Church Stretton, Shropshire. Proc. Roy. Soc. (London) Ser. B. 263: 327-367.

Osborne, P. J. 1980. The Late Devensian-Flandrian transition depicted by serial
insect faunas from West Bromwich, Staffordshire, England. Boreas 9: 139-147.

Pierce, W. D. 1947a. Fossil arthropods of California. 13. A progress report on
the Rancho La Brea asphaltum studies. Bull. So. Calif. Acad. Sci. 46: 136-138.

Pierce, W. D. 1947b. Fossil arthropods of California. 14. A progress report of
the McKittrick asphalt field. Bull. So. Calif. Acad. Sci. 46: 138-143.

Schwert, D. P. 1978. Paleoentomological analyses of two postglacial sites in
Eastern North America. Unpub. Ph.D. thesis, Univ. of Waterloo. 250 p.

Willis, H. L. 1967. Bionomics and zoogeography of tiger beetles of saline habi-
tats in the central United States. Univ. Kansas Sci. Bull. 47: 145-313.

PREDATION ON THE WESTERN HONEY BEE,
APIS MELLIFERA L., BY THE HORNET,
VESPA TROPICA (L.)

By Michael Burgett1 and Pongthep Akratanakul2

Hornets of the genus Vespa are recognized as efficient and devas-
tating predators of honey bees, especially in tropical and sub-
tropical biomes. Of the four species of honey bees in the genus Apis
only A. dorsata Fabr., the giant honey bee, appears free from attack
by hornets (Seeley et al. 1982). De Jong (1978) reviewed the records
of Vespa predation on A. mellifera and A. cerana Fabr. Matsurra
and Sakagami (1973) provided a detailed description on V. man-
darinia Smith attack behavior on A. mellifera in Japan.

We observed the predation and ultimate destruction of a small A.
mellifera colony by V. tropica (L.) on the Kamphaeng Saen campus
of Kasetsart University, Nakorn Pathom, Thailand, during a four
day period in December 1981. The honey bee colony consisted of
four standard frames with a comb area of ca. 7,000 cm2 in a hive
body with a volume of 21 1. The colony entrance was restricted to
an area of ca. 3.5 cm2. The colony possessed one comb approxi-
mately one-half full of capped honey, two empty combs, one comb
with an active brood nest and an estimated 0.5 kg of worker bees
which occupied two combs. The brood nest was infested with the
parasitic brood mite Tropilaelaps clareae Delfinado and Baker.

Uninterrupted observations of hornet behavior at the colony were
conducted on December 21 and 23 for a total of 19 h and 50 min. To
facilitate the observations eight individual hornets were tagged on
their thoraces with color and number coded discs. Observations
were begun at 0730 h on the 21st and 0715 on the 23rd and con-
tinued until after 1700 h on both days. The ambient temperature
was 16°C at the start of observations on both days and reached a
maximum of 25° C by mid-afternoon.

We estimate that 25 to 35 hornets were involved in this predatory
episode. One of us (P. A.) first noted the presence of a few hornets at

'Department of Entomology, Oregon State University, Corvallis, Oregon 97331
department of Entomology, Kasetsart University, Kamphaeng Saen, Thailand
Manuscript received by the editor September 9, 1982.

347

348

Psyche

[Vol. 89

the colony some ten to 12 days prior to the 21st. This would corres-
pond to what Matsuura and Sakagami (1973) describe as the hunt-
ing phase for V. mandarinia. By the 21st the attack had escalated to
the slaughter phase where nearly two score of hornets were concen-
trating upon the now weakened honey bee colony.

The hornet attack was a campaign of slow attrition for the honey
bees. Usually two to five hornets would position themselves at the
colony entrance. They would engage any honey bee entering or
exiting the hive. The large hornets had no difficulty in seizing the
bees and would maul them with their strong mandibles. The hornets
would normally drop the disabled bees to the ground and only
rarely was a moribund bee observed to be eaten by a hornet. The
guard hornets would frequently position themselves in the entrance
passageway with only their abdomens visible to the observer. These
hornets would engage individual guard bees just inside the entrance
and after seizing a bee, quickly drag it out and drop it off the
landing board to the ground. A separate cadre of hornets would
enter the colony and position themselves on the comb containing
capped honey. This peripheral comb was without bees which were
concentrated on the brood comb. Honey scavenging hornets would
spend long periods of time within the colony uncapping honey stor-
age cells and engorging themselves on the contents. On the 21st the
average time spent by a hornet inside the colony was 22.9 ±17.7 min
(n = 64). Upon emerging from the hive the scavenger hornets were
frequently antennated by the guard hornets, and an exchange of
alimentary fluid would usually result.

Continuous observation at the colony was not conducted on
December 22. However, a one m2 piece of plywood was placed
directly in front of the hive to facilitate an estimate of adult honey
bee mortality. Between 0930 and 1415 h 119 dead honey bees had
been deposited on the plywood by guard hornets. At 1420 h the
colony absconded and within ten minutes had clustered on a small
shrub ca. 15 m north of the hive. For the remainder of the afternoon
hornets were observed for the first time exiting the hive with larvae
and pupae scavenged from the brood nest. At 1930 h on the 22nd we
reintroduced the swarm cluster and queen back into the hive.

Observations on the 23rd began at 0715 h. At 0834 h the colony

1982]

Burgett & Akratanakul — Apis mellifera

349

once again absconded. Several guard hornets were at the hive
entrance during the exodus of the bees. The hornets physically
engaged scores of worker bees, mauling and tossing them to the
ground. The queen was seen emerging from the hive at 0838 h. She
was immediately approached by a hornet which attacked her. She
was able to disengage herself from the hornet at the cost of the tarsi
from her left front leg. She flew to the branch of a small tree ca.
three m from the hive and the worker bees began clustering around
her.

With the abandonment of the hive the hornets began to concen-
trate on the undefended brood nest. From 0845 to 1705 h hornets
were observed on 109 occasions to exit the hive with brood as prey.
The time individual hornets spent inside the hive was significantly
shorter, ave. 13.1 ±1 1.3 min (n = 127), than on the 21st when bees
were present to mount a defense of the colony. The number of
foraging events by individual hornets showed a corresponding
increase with the departure of the bees. On the 21st, with the bees
present, eight marked hornets were observed to conduct 94 com-
plete forays upon the colony for an average of 10.6 ±6.4 trips per
hornet. On the 23rd eight marked hornets completed 171 trips for an
average of 21.4 ±6.0 trips per hornet.

An examination of the hive interior on December 24 revealed that
the hornets had completely removed all larvae and pupae from the
brood comb. The honey storage comb contained less than an esti-
mated 500 g of honey. Hornet traffic at the hive was considerably
reduced from the previous three days. Occasional observations of
the hive throughout the day revealed at most, six hornets still
engorging on the remaining honey.

Apis mellifera is an introduced species to Southeast Asia (Akra-
tanakul 1976). The colony we observed came from stock originally
imported from California in 1979. Absconding by A. mellifera in
temperate climates is an unusual phenomenon. However, abscond-
ing by other species of tropical Apis is a common defensive strategy
(Seeley et al. 1982). It is interesting to note that A. mellifera from
temperate origins still retains absconding as a defense mechanism in
the face of severe predation.

350

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[Vol. 89

REFERENCES CITED

Akratanakul, P.

1976. Honey bees in Thailand. Amer. Bee. J. 116: 120-121, 124, 126.

De Jong, D.

1978. Insects: Hymenoptera (ants, wasps and bees), pp. 138-157. In: Honey
Bee Pests, Predators and Diseases. R.A. Morse, Editor, Cornell Univ.
Press, Ithaca, NY.

Matsuura, M. and S. F. Sakagami

1973. A bionomic sketch of the giant hornet Vespa mandarinia, a serious pest
for Japanese apiculture. J. Fac. Sci. Hokkaido Univ. (Ser. Zool.) 19:
125-162.

Seeley, T. D., Seeley, R. H. and P. Akratanakul

1982. Colony defense strategies of the honeybees in Thailand. Ecol. Mono. 52:
43-63.

THE GUILD OF SAWGRASS-INHABITING ANTS
IN THE FLORIDA KEYS*

By Blaine J. Cole
Department of Biology,

University of Utah,

Salt Lake City, UT84112

A guild is a group of species using similar resources in a similar
manner (Root 1967). The guild of ants on which I report here uses,
as nest sites, the hollow stems of sawgrass (Cladium jamaicense).
The primary objectives of this study were to examine this guild for
the purposes of: 1. ascertaining the extent to which there is evidence
of competition for nest sites. 2. determining the extent to which
available nest sites are filled. 3. determining whether Solenopsis
picta (Emery) nests in association with other species.

Materials and Methods

I examined 119 dead, erect culms of sawgrass, Cladium jamai-
cense, on Sugarloaf Key in Monroe County, Florida. Data were
collected between 7/29/81 and 8/3/81. Each stem had seven or
more internodal regions. For the ants in sawgrass stems the
following data were recorded: the species present, the inside diame-
ter of the internodal segments occupied and the internodal segment
in which the colony was housed. For Pseudomyrmex pallida (F.
Smith) the number and location of queens were also recorded. The
inside diameter of a culm was measured with a micrometer to the
nearest 0.1 mm. The internodal segments were numbered with the
lowest segment numbered one. The numbered segments indicated
relative height on the culm. Due to individual variation in the height
of C. jamaicense, this does not translate directly into absolute
height.

Results

Out of 119 sawgrass culms examined, 34 (29%) did not have a
colony of any species. The occurrences of various species as well as

* Manuscript received by the editor September 10, 1982

351

352

Psyche

[Vol. 89

their co-occurrences with other species is given in Table 1. Pseudo-
myrmex pallida is by far the most frequent ant, found in 57 culms or
48% of the total. Tapinoma littorale (Wheeler) and Solenopsis picta
are each found in approximately 10% of the total.

The co-occurrence of series are also given in Table 1. Solenopsis
picta is found frequently with other species including P. pallida,
Zacryptocerus varians (F. Smith), and Camponotus planatus
(Roger). No other ant species co-occur with P. pallida. T. littorale is
the only species frequent enough to examine statistically. If P. pallida
and T. littorale assorted into culms independently of one another, the
expected number of co-occurrences would be 5.75. T. littorale never
co-occurs with P. pallida, a difference that is statistically significant
(X2 = 11.0, p< 0.001).

It is possible to calculate the probability that S. picta should be the
only species that co-occurs with P. pallida. This calculation can be
done independently of our knowledge that T. littorale is negatively
associated with P. pallida (p = 0.002) or contingent on our knowledge
of this relationship (p = 0.02). In either case, it is shown that if any
species occurs with P. pallida it is likely to be S. picta. It is not
possible to statistically demonstrate the stronger statement that S.
picta is positively associated with P. pallida. Indeed, this seems not to
be the case due to co-occurrence of 5. picta and other species.

If one combines the data of Z. varians, C. planatus, Leptothorax
allardvcei (Mann) and Pseudomyrmex elongatus (Mayr), one can
also demonstrate that this aggregate is negatively associated with P.
pallida (X2 = 12.0 p < 0.001). Due to the relative rarity of these
species, one cannot test each species individually. This result must
be considered tentative.

Table 2 gives some characteristics of the nests of the guild of
sawgrass inhabiting ants. The average inside diameter and standard
deviation of internodes occupied by P. pallida is calculated sepa-
rately for that subset of the colonies that occupy a single internode
and for that subset that occupy more than one internode. Nests of P.
pallida that occupy a single internode have an inside diameter of
2.38 mm. The inside diameter of internodes occupied by P. pallida
that are found in two internodes are 2.45 and 2.03 for the lower and
upper chamber respectively. The diameter of the single nest chamber
does not differ from that of the lower nest chamber of a P. pallida
colony that occupies two chambers (ts = 0.53, p > 0.5).

1982]

Cole — Sawgrass-Inhabiting Ants

353

Table 1.

Co-occurrence of Sawgrass

Ants.

Species found:

no

other

ant

P.p.

Co-occurring
T.l. S.p.

with:

Z.v.

c.p.

Total

Pseudomyrmex pallida

52

_

0

5

0

0

57

Tapinomla littorale

10

0

—

0

1

0

12

Solenopsis picta

2

5

0

—

3

1

11

Zacryptocerus varians

2

0

1

3

—

0

7

Camponotus plantus

3

0

0

1

0

4

Leptothorax allardycei

0

0

1

0

1

0

2

Pseudomyrmex elongatus

1

0

0

0

0

0

1

Unidentified spider

1

1

0

0

0

0

2

Nothing

—

—

—

—

—

—

34

Table 2

. Nest Characteristics of Sawgrass Ants

Species

Nest Characteristics

Inside Diameter Internode
mean (sdev, n) Occupied

# internodes
occupied/
culm

Pseudomyrmex pallida

sgl. chamber

2.38 (.41, 39)

3.7

1.4

lower

2.45 (.41, 13)

3.7

dbl. chamber

upper

2.03 (.37, 13)

4.7

Solenopsis picta

3.41 (1.04, 12)

2.2

1.2

Tapinoma littorale

2.45 (.63, 15)

4.3

1.15

Camponotus planatus

4.73 (.82, 7)

2.3

—

Zacryptocerus varians

3. 13 (.61, 8)

3.6

—

The average inside diameter is greatest for C. planatus (4.73),
surprisingly large for S. picta, which is such a minute ant, and
smallest for P. pallida (2.38, single chamber). The average diameter
of internodes occupied by P. pallida and T. littorale does not differ
significantly (t-test, ts = .70, p > 0.5).

As shown in Table 2 the internode occupied by the nest parallels
the results of internode diameter. Since larger internodes are lower

354

Psyche

[Vol. 89

on the culm, species that inhabit internodes with large diameter also
inhabit low internodes.

The number of internodes occupied per culm is given in Table 2
for P. pallida, S. picta, and T. littorale. P. pallida has a tendency to
occupy more internodes per culm ( 1 .4) than does either S. picta ( 1 .2)
or T. littorale (1.15).

In most P. pallida nests a queen was located. However, in 19% of
the nests a queen was not seen. It is conceivable that the queen could
have been overlooked in these nests. In the 46 nests in which a queen
was noted, 31 (67%) had a single queen, 8 (17%) had two queens, 6
(13%) and one had four queens. When multiple queens are found in
nests occupying multiple chambers, there is no tendency for the
queens either to be found in a single chamber or to disperse to
separate chambers. When a single queen is found in a nest occupy-
ing multiple internodes there is a tendency for the queen to occupy
the higher internode.

Discussion

The guild of sawgrass inhabiting ants is a collection of species for
which there is evidence that certain pairs of species compete for nest
sites and certain pairs of species do not. P. pallida and T. littorale
are strongly negatively associated. This pair of species was not
encountered inhabiting the same sawgrass culm. P. pallida and T.
littorale occupy internodes of similar physical characteristics (inside
diameter, and relative height on the culm). It is less likely that the
two species compete for an internode of particular character than
they compete for the space of an entire culm (Levings and Traniello
1981, Cole 1982).

P. pallida shows no evidence of competition for nest sites with S.
picta. The distributions of S. picta and P. pallida are independent of
one another. These two species are found in the same sawgrass culm
with S. picta occupying larger and lower internodes. There is little
evidence to suggest that S. picta is found in association with other
species of ants. It seems to be found frequently in association with
P. pallida simply due to the fact that P. pallida is common. S. picta
has been referred to as a thief ant which nests in close proximity to
other ants and specializes in stealing brood from them. In approxi-
mately half of the cases in which S. picta is found in a sawgrass culm

1982]

Cole — Sawgrass-Inhabiting Ants

355

with another species of ant, there is at least one intervening, empty
internode between S. picta and the other species.

Of the total sawgrass culms, 71% are occupied by at least one
species. Let C be the average probability that a species will colonize
a sawgrass culm and E be the average probability that a colony will
go extinct. Then the equilibrium fraction of sawgrass culms occu-
pied is C/ C+E = 0.7 1 . One can then obtain an estimate of the rate of
extinction relative to the rate of colonization as C = 2.5E.

If colonization takes place on an annual cycle, then one can esti-
mate that the average lifespan of a colony which becomes estab-
lished is about 2.5 years. This estimate assumes that the occupancy
of sawgrass culms is at equilibrium. In addition, data from several
species, each of which may not have the same demographic charac-
teristics, are combined. This is not as bad as it seems, however, due
to the fact that the bulk of the species’ occurrences are of P. pallida.
The estimate of average colony longevity is principally an estimate
based on P. pallida.

The inside diameter and position of the lower nest chamber of P.
pallida colonies that inhabit two internodes is comparable to the
inside diameter and position of the internode occupied by P. pallida
in a single nest chamber. This suggests that P. pallida move up to
occupy a second internode. The fact that the queen of P. pallida
tends to be found in the upper chamber suggests that the queen
moves into the newer, smaller or higher nest chamber.

The average inside diameter of sawgrass culms occupied by Z.
varians (3.13 mm) is not significantly different from the average
inside diameter of hollow stems of red mangrove occupied by Z.
varians (2.95 mm, Cole 1979, n = 1 14, ts = 0.94, p > 0.2). The major
workers of Z. varians are morphologically modified for passive col-
ony defense (Wilson 1976, Cole 1980). It is reasonable to suppose
that there is selective pressure of Z. varians to choose, as nest sites,
hollow stems that are of a suitable size to allow the major workers to
block off the stem and bulldoze out intruders.

This research supported, in part, by a grant from Sigma Xi.

References

Cole, B. J. 1979. Assembly of mangrove ant communities. Ph.D. dissertation.

Princeton University, vi + 123 pp.

356

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[Vol. 89

Cole, B. J. 1980. Repertoire convergence in two mangrove ants, Zacryptocerus
varians and Camponotus (Colobopsis) sp. Insectes Sociaux 27: 265-275.

Cole, B. J. 1982. Assembly of mangrove ant communities: patterns of geographi-
cal distribution. J. of Anim. Ecol. (in press).

Levings, S. C. and J. F. A. Traniello. 1981. Territoriality, nest dispersion and
community structure in ants. Psyche 88: 265-319.

Root, R. B. 1967. The niche exploitation pattern of the blue-gray gnatcatcher.
Ecol. Monogr. 37: 317-350.

Wilson, E. O. 1976. A social ethogram of the neotropical arboreal ant, Zacryp-
tocerus varians (Fr. Smith) Anim. Behav. 24: 354-363.

<x

DEFENSIVE SPRAY MECHANISM OF A SILPHID BEETLE
(NECRODES SURINAMENSIS)*

By Thomas Eisner and Jerrold Meinwald
Section of Neurobiology and Behavior,
and Department of Chemistry,

Cornell University, Ithaca, NY 14853

Introduction

Although much has been learned about chemical defenses of bee-
tles in recent years (Weatherston and Percy, 1978), few studies have
been made of Silphidae, the family that includes the largest carrion
beetles. As is known to anyone who has collected these insects,
many silphids respond to disturbance by emitting a nauseatingly
malodorous ooze from the anus. The fluid is said to be strongly
alkaline in some species, and rich in ammonia (Schildknecht and
Weis, 1962). In Silpha, a gland had been noted that opens into the
rectum (Dufour, 1826; Leydig, 1859), but no chemical work had
been done to determine whether specific defensive chemicals in the
anal effluent might stem from the gland.

Personal observation had told us that one silphid, the so-called
red-lined carrion beetle, Necrodes surinamensis, might be unusual.
First, the beetle seemed able to eject its anal fluid as a spray rather
than an ooze, which no other silphid had been reported to do, and
second, the fluid gave an acidic test on indicator paper and had a
stench that was overlain by a distinct aromatic fragrance.

We have now studied N. surinamensis in some detail. Chemical
work, carried out in collaboration with others, led to the isolation of
several fatty acids and terpenoid compounds, present in the spray
and produced by a special rectal gland. An account of these chem-
ical findings, which are summarized in Figure 1, will be published
elsewhere. We here give details of the beetle’s defensive behavior,
plus a brief description of the gland, and data on the beetle’s unac-
ceptability to predators.

♦Paper No. 72 of the series Defense Mechanism of Arthropods. Paper No. 71 is
Eisner, T. and Nowicki, S., Science 219, 185 (1983).

Manuscript received by the editor October 12, 1982.

357

358

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[Vol. 87

Procedures and Results

Necrodes surinamensis is a large beetle, about 2 cm in average
body length. It occurs throughout the United States, east of the
Rocky Mountains. It is mostly taken at carcasses. We collected
large numbers at baits (dead fish and chickens) in the environs of
Ithaca, New York, and on the grounds of the Archbold Biological
Station, Lake Placid, Florida. They were maintained in the labora-
tory on commercial meat-based dog food preparations. Observa-
tions were made mostly on freshly captured specimens.

Spray ejection was studied by techniques previously used with
other chemically protected arthropods (Eisner, 1958). The beetles
were individually fastened with wax to tethers and placed in normal
stance upon sheets of indicator paper (filter paper presoaked in red
alkaline phenolphthalein solution, blotted off to near dryness just
before use). They were then subjected to simulated attack by pinch-
ing some of their appendages with forceps or briefly touching parts
of the body with a hot spatula. Their responses were immediate. No
sooner had a stimulus been applied than they revolved the abdomi-
nal tip, which projects free beyond the posterior margin of the ely-
tra, aimed it toward the site stimulated, and sprayed. As evidenced
by the pattern of white spots induced by the acid fluid on the indica-
tor paper, the discharges were accurately directed (Fig. 2A, B). The
site of emission of the spray was clearly noted to be the anus. The
abdominal tip is essentially a revolvable emplacement for the anal
nozzle. It can be pointed in all directions, even anteriorly over the
beetle’s own back (Fig. 2C-F). Regions of the body stimulated were
always noticeably wetted by the spray. Beetles that had remained
undisturbed in confinement for several days, and were tethered
without being caused to discharge (they were kept refrigerated dur-
ing the tethering procedure), proved capable of spraying repeatedly,
even in quick succession if a rapid sequence of stimuli was applied.
The number of discharges (x ± sd) that could be elicited from such
beetles was 4.9 ± 1 . 1 (N = 5 females + 3 males). Only direct contact
elicited discharges. The beetles never sprayed in response to move-
ment or tapping nearby.

The rectal gland, which is identical in both sexes, was readily
exposed by dissection. It consists of a tubule and a sac (Fig. 3). The
tubule lies free in the hemocoel, is long and narrow (actual meas-
urement in a female = 18 X 0.2 mm) and closed at its distal end. It
opens proximally into the bladder-like sac, which itself opens by

1982] Eisner & Meinwald — Defensive Spray Mechanism 359

Aliphotic Acids

fj.q per Beetle

Caprylic acid CH3(CH2)6 C02H 25

Capric acid CH3(CH2)8C02H 5

c/s-3-Decenoic acid CH3(CH2)5 CH =CHCH2C02H 5

c/s-4-Decenoic acid CH3(CH2)4CH =CH(CH2)2C02H 5

Terpene Alcohols

Lavandulol

a-Necrodol

)3-Necrodol

4

14

3

Fig. 1. Substances isolated and characterized from the rectal gland of Necrodes
surinamensis. The two terpene alcohols, a-necrodol and 0-necrodol, are new natural
products; m-3-decenoic acid and m-4-decenoic acid have not previously been
reported from an insectan source. Details of the chemical procedures will be pub-
lished elsewhere.

way of a narrow neck into the rectum. The tubule is surrounded by a
loose meshwork of muscle fibers, clearly identifiable as such in
whole mounts of the gland viewed by transmitted polarized light.
Comparable compressor muscles, arranged in a thick layer, envelop
the sac. The entire gland has an inner lining of membranous cuticle,
which was readily isolated by treatment of the gland with 10%
aqueous potassium hydroxide, and was shown to be continuous with
the cuticular lining of the hindgut. In freshly dissected preparations,
both parts of the gland were seen to be filled with clear fluid. The
hindgut, in contrast, was usually replete with opaque fecal paste.

The compounds listed in Figure 1 had been shown to be present
both in extracts of isolated glands and in samples of the spray itself.
None were present in more than trace amounts in extracts of the
region of the hindgut anterior to the glandular junction. It seemed
reasonably certain, therefore, that the fatty acids and terpenes are
products of the gland rather than the enteron. This conclusion was
further supported by circumstantial evidence. Fluid squeezings from
isolated glands, unlike squeezings from the hindgut, gave acidic spot

360

Psyche

[Vol. 87

Fig. 2. A-B, Aimed discharges elicited by pinching a left midleg (A) and left
hindleg (B) of Necrodes with forceps. The spray pattern is visible on phenophthalein
indicator paper. C-F, Directional aiming movements of the anal turret of Necrodes.
Note that the abdominal tip is accurately pointed toward the site of application of the
stimulus: (C) tibia of midleg, (D) tarsus of midleg, (E) tibia of hindleg, pinched with
forceps; (F) back of beetle touched with hot spatula.

tests on phenolphthalein indicator paper and had the recognizable
terpenoid fragrance of the spray.

Examination of fresh spray ejected by Necrodes on glass showed
occasional presence of opaque material in the discharged fluid, sug-
gesting that the secretion may sometimes be expelled with admix-
ture of fecal paste. Since the glandular contents are forced to the
outside by way of the rectum, such admixture may occur whenever
the pathway of secretory egress is blocked by enteric matter. Two
fatty acids not listed in Table 1, stearic acid and palmitic acid, were

1982] Eisner & Meinwald — Defensive Spray Mechanism 361

Fig. 3. Diagram of Necrodes surinamensis showing the position of the rectal
gland (tb = tubule; sc = sac) relative to the hindgut (hg).

362

Psyche

[Vol. 87

identified as occasionally present in the spray. Neither was detected
with consistency or in substantial amounts in extracts of the gland,
but they were always present in extracts of the hindgut. Their occur-
ence in the spray may be a further indication that rectal contents are
sometimes ejected with the secretion.

Laboratory tests done with formicine ants (Formica exsectoides )
and Swainson’s thrushes (Catharus ustulatus) demonstrated that
Necrodes is well protected against such predators. The tests with
Formica involved presenting individual tethered Necrodes to groups
of 10 ants in small glass enclosures. The ants attacked immediately,
by clamping onto the beetles with their mandibles, in response to
which the beetles revolved their abdominal tip and sprayed. As was
particularly clear from the patterns of droplets sometimes visible on
the bottom of the enclosures, the discharges were accurately aimed
toward the ants. These usually released their hold quickly and fled.
At varying intervals thereafter they engaged in intensive cleansing
activities, which seemed all the more protracted when the ants had
been heavily contaminated with spray. Five beetles were exposed to
ants in this fashion for 30 min. each. None received noticeable
injury.

The tests with the thrushes followed a protocol previously used
with these birds in experiments with other chemically protected
insects (Eisner et al., 1978). Necrodes were offered together with
mealworms (larvae of Tenebrio molitor, which served as edible con-
trols) to 3 individually caged birds (all males), in 3 daily feeding
sessions per bird. Mealworms outnumbered Necrodes 2 to 1. The
insects were offered one at a time, up to a total of 14-15 per session.
Sequence of presentation was such that each series of 3 consecutive
items consisted of two mealworms and one randomly placed
Necrodes. Each item was left with a bird until it was eaten, or for a
maximum of 2 min. Fate of prey was scored as follows: eaten ( E , if
the insect was ingested after having been pecked no more than 3
times); eaten with hesitation {EH, if the insect was eaten after having
been pecked more than 3 times); rejected ( R , if the insect was
ignored after having been pecked one or more times); not touched
{NT, if the insect was not contacted by the bird during the 2 min. of
presentation). Insects not touched at the end of a feeding session
were not tallied, since such avoidance might have been due to satia-
tion of the bird.

1982] Eisner & Meinwald — Defensive Spray Mechanism 363

The results, lumped for the 9 feeding sessions with the 3 thrushes,
are shown in Figure 4. It is clear that the birds rated Necrodes
distinctly undesirable relative to mealworms. While the latter were
all eaten outright, 74% of Necrodes were either rejected or left
untouched. The 26% that were eaten were only taken after repeated
peckings. A special point was made to check the rejected Necrodes
for injury. None was found to bear any, and all were live when
examined several days later. Although it proved impossible to
determine with certainty whether Necrodes always sprayed when
pecked or grasped by a bird, in some cases there was evidence that
discharges had occurred. Streaks of spray occasionally made their
appearance on the glossy floor of the cage during an attack, or birds
shook their heads violently after seizing a beetle, as we have repeat-
edly seen captive thrushes do when attempting to take insects that
spray (e.g. carabid beetles).

E EH R NT

FA TE

Fig. 4. Fate of Necrodes surinamensis and mealworms fed to three Swainson’s
thrushes; E = eaten; EH = eaten with hesitation; R = rejected; NT = not touched.
Details in text.

364

Psyche

[Vol. 87

Discussion

The discovery of a chemical defense mechanism in an insect
should come as no surprise, since such mechanisms are extra-
ordinarily widespread among arthropods. Moreover, many insects,
including a multiplicity of beetles, termites, ants, earwigs, cater-
pillars, and phasmids, eject their defensive secretions in the form of
accurately directed jets. Necrodes is anomalous in that it expels its
aimed secretory discharges from the anus. Other beetles that spray,
such as Carabidae, also discharge from the tip of the abdomen and
may aim their ejections by movement of the abdominal tip (e.g.
Eisner, 1958), but their glands are integumental and open beside the
anus on the body wall itself. Necrodes is further unusual in that it
has only one gland. Exocrine defensive glands in beetles commonly
occur in pairs.

It seems reasonable to presume that the gland of Necrodes arises
developmental^ as an outpocketing of the rectum. Other rectal
glands in Silphidae, such as that of Silpha, are doubtless homolo-
gous to that of Necrodes. We feel this to be so despite some differ-
ences in gland morphology [In Silpha americana the lateral tubule is
reduced to a short elaborately subdivided diverticulum (Alsop,
1970)]* and in gland chemistry ( Silpha americana, as we shall report
elsewhere, produces steroids in its gland). While in the absence of
histological work little can be said about the function of the two
parts of the Necrodes gland, the strongly muscled condition of the
sac suggests that it might serve as the reservoir from which secretion
is expelled for the discharge. The tubule might be strictly secretory.

It seems clear from the tests with ants and birds that Necrodes is
relatively unacceptable to such predators. But to what extent this is
attributable to the glandular components of the spray, or to enteric
additives of the spray, or even to entirely different factors (carrion
contamination of the beetle’s body?) remains to be seen. The secre-
tion, no doubt, plays a defensive role, but the other factors may
amplify the effect. It is interesting in this connection that another
common inhabitant of carrion, the staphylinid beetle Creophilus

*Dufour ( 1826) writing of Silpha littoralis, speaks of a rectal gland with a “vaisseau
secreteur” almost as long as the body, suggesting that he was dealing with a gland
similar to that of Necrodes.

1982] Eisner & Meinwald — Defensive Spray Mechanism 365

maxillosus, also mixes intestinal fluid with the secretion of its defen-
sive glands (Jefson et al., 1983). A diet of carrion, one might imagine,
could render an insect’s enteric contents potently deterrent. The
ammonia reportedly present at high concentrations in the anal
effluent of some silphids (Schildknecht and Weis, 1962) is probably
derived from decaying ingested animal protein and may well serve
for defense. To us at least, the odor of the intestinal fluid discharged
by many carrion insects upon handling, or for that matter the odor
of the insects themselves, is repugnant. The fragrance emitted by
Neerodes after a discharge is transient, and certainly does not mask
the intrinsic stench of the animal.

While it would have been desirable to test the various secretory
components of Neerodes for repellency, this proved impossible due
to lack of sufficient synthetic quantity of a-necrodol and (3-
necrodol, the two most interesting novel compounds in the mixture.
It seems likely, however, that these terpenes are deterrent to insects.
They are cyclopentanoid compounds, of which many are known to
occur in the defensive glands of insects and in plants (Nakanishi et
al., 1974), and some are provenly repellent to insects (Eisner, 1964;
Smolanoff et al., 1975; Meinwald et al., 1977; Jefson et al., 1983).
Fatty acids have also been reported from other arthropodan defen-
sive glands. They may themselves be deterrent, and may also serve
as surfactants. As part of a spray they may promote spread and
penetration of droplets on target, a role that has been demonstrated
for caprylic acid in whip scorpion secretion (Eisner et al., 1961). The
fatty acids of Neerodes may have a similar function, and may also
facilitate the mixing of the apolar glandular material with the large-
ly aqueous enteric fluid when the two are discharged together. Two
of the Neerodes fatty acids, m-3-decenoic acid and ns-4-decenoic
acid, have not previously been identified from an insectan source.
The apparent enteric, rather than glandular, origin of stearic and
palmitic acid should come as no surprise, since these fatty acids are
major components of animal fats and hence likely to be ingested by
Neerodes with carrion.

Only speculation can be offered to account for the presence of
lavandulol in the Neerodes spray. The substance has not previously
been reported from insects, although it is known from plants as a
major component of lavender oil (Karrer, 1958). We suspect the
compound to be repellent to insects, as some low molecular terpenes

366

Psyche

[Vol. 87

are known to be, which if true would provide some explanation for
the presence of the substance in plants. An increasing number of
compounds known previously only from plants is being isolated
from the defensive glands of insects. In our judgment the very
occurrence of such compounds as defensive agents in animals sug-
gests that they may (sometimes at least) fulfill a similar function in
plants.

Carrion insects, often crowded in their food source, undoubtedly
interact in subtle competitive ways. To what extent Necrodes, or for
that matter any other chemically protected carrion insect, makes use
of its defensive glands in such interactions, remains an intriguing
unknown.

Summary

When disturbed, the carrion beetle Necrodes surinamensis (family
Silphidae) ejects jets of fluid from the anus. The abdominal tip,
which projects beyond the posterior margins of the elytra, serves as
the revolvable turret by which the ejections are aimed. Only contact
stimulation elicits discharges. The fluid is primarily of glandular
origin but may contain admixed enteric matter. The gland, which
consists of a tubular portion and a vesicular sac, opens into the
rectum itself. Chemical work (to be reported elsewhere) has shown
the secretion to contain two novel cyclopentanoid compounds (a-
necrodol and /3-necrodol) as well as lavandulol and several fatty
acids. Two of the fatty acids, n's-3-decenoic acid and m-4-decenoic
acid, were not previously known from insects.

Acknowledgements

Study supported in part by NIH Grants (AI02908 and AI 12020)
and Hatch Grants (NYC-191406 and NYC-191409). We thank
Karen Hicks, Maura Malarcher, and Maria Eisner for excellent
technical help, and the staff of the Archbold Biological Station for
personal and professional generosity. Vivian Eisner did the drawing.
Brady Roach, our principal collaborator on the chemistry of
Necrodes, helped in the unpleasant task of collecting the beetles.

1982] Eisner & Meinwald — Defensive Spray Mechanism 367

References Cited

Alsop, D. W.

1970. Defensive glands of arthropods: comparative morphology of selected
types. Ph.D. Thesis, Cornell University, Ithaca, NY

Dufour, L.

1826. Recherches anatomiques sur les carabiques et sur plusieurs autres
Insectes coleopteres. Ann. Sci. Nat. 8: 5-19.

Eisner, T.

1958. The protective role of the spray mechanism of the bombardier beetle,
Brachynus ballistarius Lee. J. Ins. Physiol. 2: 215-220.

Eisner, T.

1964. Catnip: its raison d’etre. Science 146: 1318-1320.

Eisner, T., J. Meinwald, A. Monro, and R. Ghent.

1961. Defense Mechanisms of Arthropods-I. The composition and function of
the spray of the whipscorpion, Mastigoproctus giganteus (Lucas) (Arach-
nida, Pedipalpida). J. Ins. Physiol. 6: 272-298.

Jefson, M., J. Meinwald, S. Nowicki, K. Hicks, and T. Eisner.

1983. Chemical defense of a rove beetle (Creophilus maxillosus). J. Chem.
Ecol. (in press).

Karrer, W.

1958. Konstitution und Vorkommen der organischen Pflanzenstoffe (exclusive
Alkaloide). Basel: Birkhauser.

Leydig, F.

1859. Zur Anatomie der Insekten. Archiv Anat. Physiol, wissen. Med.: 33-89,
149-183.

Meinwald, J., T. H. Jones, T. Eisner, and K. Hicks.

1977. New methylcyclopentanoid terpenes from the larval defensive secretion
of a chrysomelid beetle (Plagiodera versicolora). Proc. Nat. Acad. Sci.
74: 2189-2192.

Nakanishi, K., T. Goto, S. Ito, S. Natori, and S. Nozoe (eds.)

1974. Natural Products Chemistry, Vol. 1. New York: Academic Press, pp.
48-59.

SCHILDKNECHT, H. AND K. H. WEIS.

1962. Uber die chemische Abwehr der Aaskafer. XIV. Mitteilung liber
Insektenabwehrstoffe. Z. Naturforschg. 17b: 452-455.

Smolanoff, J., A. F. Kluge, J. Meinwald, A. McPhail, R. W. Miller, K. Hicks,
and T. Eisner.

1975. Polyzonimine: a novel terpenoid insect repellent produced by a milliped.
Science 188: 734-736.

Weatherson, J. and J. E. Percy.

1978. Venoms of Coleoptera. In S. Bettini (ed.) Arthropod Venoms, Hand-
book of Experimental Pharmacology, Vol. 48. Berlin: Springer-Verlag,
pp. 511-554.

PSYCHE

A Journal of Entomology

Volume 89
1982

Editorial Board

Frank M. Carpenter, Editor P. J. Darlington, Jr.
W. L. Brown, Jr. H. W. Levi

E. O. Wilson Alfred F. Newton, Jr.

B. K. HOlldobler M. D. Bowers

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. 88, nos. 3-4, for 1981, May 28, 1982
Vol. 89, nos. 1-2, for 1982, December 17, 1982

PSYCHE

INDEX TO VOLUME 89, 1982

INDEX TO AUTHORS

Adams , Phillip A. and J. Allan Garland. A Review of the Genus Mallada in the
United States and Canada, with a New Species (Neuroptera: Chrysopidae).
239

Akratanakul, Pongthep. See Burgett, Michael

Akre. Roger D. See Reed, Hal C.

Alloway, Thomas M., Alfred Busehinger, Mary Talbot, Robin Stuart, and Cynthia
Thomas. Polygyny and Polydomy in Three North American Species of the
Ant Genus Leptothorax Mayr ( Hymenoptera:Formicidae). 249

Blomquist, Gary J. See How ard, Ralph W.

Burgett, Michael and Ponghep Akratanakul. Predation on the Western Honey Bee,
Apis meUifera L., by the Hornet, Vespa tropica (L.). 347

Busehinger, Alfred. Leptothorax faberi n.sp., an Apparently Parasitic Ant from
Jasper National Park, Canada (Hymenoptera:Formicidae). 197

Busehinger, Alfred. See Allow'ay, Thomas M.

Carpenter, Frank M. Dedication: Joseph C. Bequaert. 1

Cole, Blaine J. The Guild of Sawgrass-Inhabiting Ants in the Florida Keys. 351

Coles de Negret, Helen R. and Kent H. Redford. The Biology of Nine Termite
Species (Isoptera: Termitidae) from the Cerrado of Central Brazil. 81

Darling, Christopher D. Description of a New Species of Krombeinius ( Hymenop-
tera: Perilampidae) from the Philippines, and the Phylogenetic Relationships of
the Genus. 307

Dow ning, H. A. and R. L. Jeanne. A Description of the Ectal Mandibular Gland in
the Paper Wasp, Polistes Juscatus (Hymenoptera: Vespidae). 317

Eberhard, W. G. See Lubin, Y. D.

Engel-Siegel, Hiltrude. See Holldobler, Bert

Eisner, Thomas, and Jerrold Meinwald. Defensive Spray Mechanism of a Silphid
Beetle ( Necrodes surinamensis). 357

Garland, J. Allan. See Adams, Phillip A.
Garling, Lyn. See Otis, Gard W.

371

Holldubler, Ben. Raiding Behavior and Prey Storage in Cerapachys (Hymenop-
tera: Formicidae). 3

Holldubler, Bert and Hi/trude Engel-Siegel. Tergal and Sternal Glands in Male
Ants ( Hymenoptera: Formicidae). 113

How ard, Ralph W., C. A. McDaniel, and Gary J. Blomquist. Chemical Mimicry as
an Integrating Mechanism for Three Termitophiles Associated with Reticuli-
termes virginicus (Banks). 157

Jeanne, R. L. See Downing, H. A.

Kitnsey, Lynn S. Parataruma, a New Genus of Neotropical Crabronini (Hymenop-
tera: Sphecidae). 169

Levi, Herbert W. and Deborah R. R. Smith. A New Colonial Anelosinius Spider
from Suriname (Araneae: Theridiidae). 275

Levi, H. W. See Lubin, Y. D.

Lubin, Y. D., B. D. Opell, W. G. Eberhard, and H. W. Levi. Orb Plus Cone-webs in
Uloboridae (Araneae), with a Description of a New Genus and Four New
Species. 29

Malaret, Luis. See Otis, Gard W.

Mansour, Fadel. See Ross, John

McDaniel, C. A. See Howard, Ralph W.

McGinley, Ronald J. See Otis, Gard W.

Meinwald, Jerrold. See Eisner, Thomas

Miller, Scott E. See Peck, Stewart B.

Miller, Scott E. See Nagano, Christopher D.

Mockford, Edward L. Redescription of the Type Species of Myopsocus, M.
unduosus (Hagen), and Resulting Nomenclatural Changes in Genera and
Species of Myopsocidae (Psocoptera). 21 1

Morgan, Alan V. See Nagano, Christopher D.

Nagano, Christopher D., Scott E. Miller and Alan V. Morgan. Fossil Tiger Beetles
(Coleoptera: Cicindelidae): Review and New Quaternary Records. 339

New ton, Alfred F., Jr. Agathidiodes Portevin, New Synonym of Stetholiodes Fall
(Coleoptera: Leiodidae: Anistomini). 337

Obin, Martin S. Spiders Living at Wasp Nesting Sites: What Constrains Predation
by Mud-Daubers? 321

Opell, B. D. See Lubin, Y. D.

Otis, Gard W., Ronald J. McGinley, Lvn Garling, and Luis Malaret. Biology and
Systematics of the Bee Genus Crawfordapis (Colletidae, Diphaglossinae). 279

Peck, Stewart B. The Life History of the Carrion Beetle, Ptomascopus morio and
the Origins of Parental Care in Necrophorus (Coleoptera, Silphidae, Nicrophini).
107

372

Peck, Stewart B. and Scott E. Miller. Type Designations and Synonymies for
North American Silphidae (Coleoptera). 151

Redford, Kent H. See Coles de Negret.

Reed. Hal C. and Roger D. Akre. Morphological Comparisons between the Obli-
gate Social Parasite, Vespula austraica (Panzer), and Its Host, Vespula acadica
(Sladen). 183

Richman, Davis B. See Ross, John

Ross, John, Davis B. Richman, Fade I Mansour, Anne Trambarulo, and W. H.
Whitcomb. The Life Cycle of Heteropoda venatoria (Linnaeus) (Araneae:
Heteropodidae). 297

Smith, Deborah R. R. See Levi, Herbert W.

Stuart, Robin. See Allowav, Thomas M.

Talbot, Mary. See Allowav, Thomas M.

Tepedino, V. J. See Torchio, P. F.

Thomas, Cynthia. See Allowav, Thomas M.

Torchio. P. F. and V. J. Tepedino. Parsivoltinism in Three Species of Osmia
Bees. 221

Thorne, Barbara L. Termite-Termite Interactions: Workers as an Agonistic Caste.
133

Trambarulo, Anne. See Ross, John

Traniello, James F. A. Population Strcuture and Social Organization in the
Primitive Ant, Amblyopone pallipes (Hymenoptera: Formicidae). 65

Whitcomb, W. H. See Ross, John

INDEX TO SUBJECTS

All new genera, new species and new names are printed in capital type.

Agathidiodes, new synonym, 337
Amblyopone pallipes, 65
Anelosimus saramacca, 275
Ant larvae, 175
Ants inhabiting saw-grass, 351
Armitermes, 81, 133
Bequaert, Joseph C., 1

Biology and systematics of Crawforda-
pis, 279

Biology of nine termite species, 81
Cerapachvs, 3

Chemical mimicry by termitophiles, 157
Colonial spider, 275

Communication, raiding behavior, and
prey storage in Cerapachvs, 3

CONIFABER PARVUS, 52
Cornitermes, 81
Cortaritermes, 81
Crabronini, 169
Crawjordapis, 279
Cyclogaster, 25

Dedication: Joseph C. Bequaert, 1

373

Defensive spray mechanism of a silphid
beetle, 357

Description of ectal mandibular gland in
Polistes, 317

Description of a new species of Krom-
beinius, 307

Designation of a type-species for Cvc-
logaster, 25

Ectal gland in Polistes, 317
Fossil tiger beetles, 339
Grigiotermes, 81

Guild of sawgrass-inhabiting ants, 351
Heteropoda venturia, 297
Krombeinius sauion, 313
Leptothorax faberi, 137

Leptothorax, polygyny and polydomy,
249

Life cycle of Heteropoda ventoria, 297

Life history of Japanese Carrion Beetle,
107

Mai la da luctuosus, 246
Mallada macleodi, 239
Mallada perfectus, 244
Mallada sierra, 245

Morphological comparisons between
Vespula austraica and V. acadica, 183

Mud-daubers, 321
Myopsocidae, 21 1

Myopsocus unduosus, redescription,
212

Nasutitermes, 81
Nasutitermes corniger, 133
Necrocharis, synonymy
Necrodes surinamensis, 357
Necrophilus, type designations, 1 5 1
Necrophorus, type designations, 151

New colonial spider, 275

Nicrophorus, 107, 151

Orb plus cone-webs in Uloboridae, 29

Orthognat hotermes, 8 1

Osmia, 212

PARATARI MA LEC'LERCQI, 170
PARATARUMA TROPICAUDA, 173
Parental care in Nicrophorus, 107
Parsivoltinism in Osmia bees, 212
Polistes fuscatus, 317

Polygyny and Polydomy in Leptothor-
ax, 249

Population structure and social organi-
zation of Amblyopone pallipes, 65

Predation on Apis mellifera, 347
Procornitermes, 81
Ptomascopus morio, 107

Redescription of type species of Myop-
socus, 2 1 1

Reticulitermes virginicus, 157
Review of the genus Mallada, 239
Saw-grass inhabiting ants, 351
Silpha, type designations, 151

Silphidae, type designations and synon-
ymies, 151

Spiders living at wasp nesting site, 321
Stetholiodes, synonymy, 337
Sternal glands in male ants, 1 13
Syntermes, 81

Supplementary studies on ant larvae:
Formicinae, 175

Tergal and sternal glands in male ants,
113

Termite-termite interactions, 133
Termites, 81, 133

374

Termitophiles associated with Reticuli-
termes, 157

Tiger beetles, fossil, 339

Type designations and synonymies for
Silphidae, 151

Uloboridae, new species, 29
Uloborus ABOLINEATUS, 57

Uloborus bispiralis, 59
Uloborus corn us, 53
Velocitermes, 8 1
Vespa tropica, 347
Vespula, 183
Western honey bee, 347

375

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