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(V

PSYCHE

A Journal of Entomology

Volume 85

Editorial Board

Frank M. Carpenter, Editor
W. L. Brown, Jr.

E. O. Wilson
B. K. HOlldobler

P. J. Darlington, Jr.
H. W. Levi

Alfred F. Newton, Jr.
R. E. SlLBERGLIED

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. 84, no. 3-4, September-December, 1977: July 19, 1978

Vol. 85, no. 1, March, 1978: January 26, 1979

Vol. 85, no. 2-3, June-September, 1978: April 24, 1979

ISSN 0033-2615

PSYCHE

A JOURNAL OF ENTOMOLOGY

founded in 1874 by the Cambridge Entomological Club

Vol. 85

March, 1978

No. 1

CONTENTS

Webs of Miagrammopes (Araneae: Uloboridae) in the Neotropics. Y. D. Lubin,

W. G. Eberhard, and G. G. Montgomery 1

Revision of the South Temperate Genus Glypholoma Jeannel, with Four New
Species (Coleoptera: Staphylinidae: Omaliinae). Margaret K. Thayer and
Alfred F. Newton, Jr 25

Aggregations of M alios and Dictyna (Araneae, Dictynidae): Population
Characteristics. Robert R. Jackson and Sandra Smith 65

A Solitary Wasp that Preys upon Lacewings (Hymenoptera: Sphecidae,
Neuroptera: Chrysopidae). Howard E. Evans 81

Survey of Social Insects in the Fossil Record. Laurie Burnham 85

Parental Care in Guayaquila compressa Walker (Homoptera: Membracidae).

T. K. Wood 135

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1977-1978

President Gary D. Alpert

Vice-President JOHN A. SHETTERLY

Secretary ROBERT ROBBINS

Treasurer FRANK M. CARPENTER

Executive Committee MARTHA FISHER

Katherine Horton

EDITORIAL BOARD OF PSYCHE

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

ALFRED F. Newton, JR., Curatorial Associate in Entomology, Har-
vard 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. E. SlLBERGLIED, Assistant Professor of Biology, Harvard University
E. O. WILSON, Baird Professor of Science, Harvard University

PSYCHE is published quaterly by the Cambridge Entomological Club, the issues
appearing in March, June, September and December. Subscription price, per year,
payable in advance: $8.00 for United States and Canada, $9.50 for other countries.
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Checks and remittances should be addressed to Treasurer, Cambridge Entomo-
logical 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. Car-
penter, Biological Laboratories, Harvard University, Cambridge, Mass. 02138.

Authors are expected to bear part of the printing costs, at the rate of $24.50 per
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and for full page half-tones, $30.00 each; smaller sizes in proportion.

The September-December, 1977, Psyche (Vol. 84, Nos. 3-4) was mailed July 19, 1978

The Lexington Press, Inc., Lexington, Massachusetts

PSYCHE

Vol. 85

March, 1978

No. 1

WEBS OF MIA GRA MM OPES (ARANEAE: ULOBORIDAE)
IN THE NEOTROPICS

By Y. D. Lubin1, W. G. Eberhard2, and G. G. Montgomery1
Introduction

Uloborid spiders ( Uloborus sens, lat.) typically construct orb webs
composed of non-sticky threads (radii, frame threads, hub, and
temporary spiral) which support a sticky spiral made of cribellar
or hackled silk. Specialization of the web in the uloborid genus
Miagrammopes has involved the reduction of its structural com-
plexity together with changes in its operation as an insect trap. The
one described web of an unidentified species from Natal, South
Africa is reduced to a single horizontal capture thread (Akerman
1932). In this paper we describe the webs of six more species of
Miagrammopes and the prey capture behavior of the spiders, re-
vealing a substantial range of variation in simple web design within
the genus.

We studied M. simus on Barro Colorado Island, Panama Canal
Zone during the wet season of 1976. At no time was this species
common. In May and June, 1977, M, sp. 1 (ca. unipus) was studied
in a bamboo ( Guadua angustif olia) thicket in the Cauca valley near
Cali, Colombia where it occurred in abundance. In August, 1977,
M. intempus Chickering and M. sp. 2 were found in Valle, Colom-
bia. The former was common in some places on hanging moss on
exposed roots and low branches near the Rio Anchicaya at 400 m
elevation, while the latter was found in brush near the Rio Tulua
at 1100 m elevation. A small tree in a clearing on Finca La Selva

1 Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa, Panama
Canal Zone.

2Departamento de Biologia, Universidad del Valle, Cali, Colombia, and Smith-
sonian Tropical Research Institute.

Manuscript received by the editor April 16, 1978.

1

2

Psyche

[March

near Puerto Viejo, Heredia Province, Costa Rica, had substantial
populations of M. sp 3: M. sp. 4 was found on low vegetation in
January and February, 1978, in mid-elevation wet forest in Guatopo
National Park, Miranda State, Venezuela. Individuals of the last
four species were observed in the field on only one or two days each,
but in all cases more extensive observations had already been made
on the other species, and it was thus possible to make critical ob-
servations allowing comparisons among all six species. Miagram-
mopes sp. 1-4 appear to be either undescribed species or females of
species known only from males. Voucher specimens of these and
of the two previously described species are deposited in the Museum
of Comparative Zoology, Harvard University.

The Webs

M. simus

The web typically consisted of a single vertical capture thread
about 1 m long, attached above to a short, horizontal resting thread
strung under a leaf, and below to the ground or a leaf or twig (Fig.
la). The capture thread was covered with sticky, cribellar silk along
the central 50 to 60 percent of its length, and one or more very fine,
more or less horizontal threads often connected it to other supports.
Both end portions of the capture thread were non-sticky. For an
individual whose webs were measured periodically, the lengths of
sticky and non-sticky sections in new webs were (in cm; lengths of
sticky portions underlined): 20:50:30, 4:50:30, 6:52:34, 7:60:32,
and 7:60:34. One adult female which had been starved for seven
days made a web with two vertical capture threads and several thin,
non sticky lines between them.

One M. simus was seen laying sticky, cribellar silk on a non-
sticky, vertical thread which was already in place. The spider moved
slowly up the thread, combing out silk with legs IV until it was
about 5 cm below the resting thread, then ran up and assumed the
resting posture.

Individuals of M. simus rested under the horizontal thread and
held onto the broken end of the capture thread with one leg I and
one leg II, while the other legs held the resting thread (Fig. lb). Ten-
sion was exerted on the vertical capture thread both by pulling it up
with leg I and by backing up and pulling in the resting thread with
the fourth pair of legs. The spider which constructed a web with two

1978] Lubin, Eberhard, & Montgomery Miagrammopes 3

Figure 1. a) Typical web of Miagrammopes simus, showing the horizontal rest-
ing thread under a leaf, the vertical capture thread with sticky segment and thin,
non-sticky, horizontal threads; b) posture of M. simus as it holds its web and waits
for prey.

capture threads rested in essentially the same position; the leg I
holding the horizontal resting thread was in position to monitor
vibrations from the second capture thread.

When disturbed, or when hanging from a resting thread with no
capture thread present, M. simus assumed a stick-like, cryptic pos-
ture, orienting along the resting thread with the first and second
pair of legs held straight forward and the fourth pair held straight
behind. The small third pair of legs held the resting thread or the
substrate, but were pressed close to the body and did not break the
stick-like outline.

4

Psyche

[March

M. sp. 1 (ca. unipus )

The web of this species differed in that there was usually more
than one capture thread attached to a single horizontal resting thread
(Fig. 2). The average was 2.4 capture threads and some webs had
up to five (Table 1). There was no apparent relationship between
the number of capture threads in the web and the size of the spider
that constructed it. The capture threads were usually not perfectly
vertical and were often in different planes with angles of less than
90° between them. They were shorter and thinner than the capture
threads of M. simus and it was necessary to powder them with corn-
starch in order to count them. The horizontal resting thread was
always under a thin twig rather than a leaf, as in webs of M. simus.

In some webs of M. sp. 1 there were one or more very slack, non-
sticky, horizontal threads connecting the multiple capture threads.
Because of their looseness and their variable location and orienta-
tion, these lines were at first thought to be incidental (perhaps float-
ing threads made by other spiders), but their presence in many webs
of both this species and M. simus argues otherwise.

Web construction appeared to be similar to that of M. simus.
One spider was seen laying cribellar silk while moving up along a
vertical thread which was already in place. The spider advanced
slowly, combing out silk continuously with legs IV and attaching
it to the thread periodically with brisk dabs of the abdomen. Total
construction time for one capture thread was about 3 minutes.

At night, M. sp. 1 assumed a capture position similar to that of
M. simus, resting under the horizontal thread and holding a cap-
ture thread with legs I and II (Fig. lb). During the day it either
held the capture thread in the same way, or, more often, assumed
a more cryptic resting posture. The spider positioned itself near
one end of the resting thread which it broke and spanned with its
body. It held one end with one or both pairs of front legs, and then
pulled in the line behind it with the hind legs (and, occasionally, the

Table 1. Numbers of sticky capture threads in 66 webs of Miagrammopes sp. 1
(ca. unipus) and 22 webs of M. sp. 3.

Number of Capture Threads

Number of Webs

1

2

3

4

5

6 or more

M. sp. 1

15

21

22

6

2

0

M. sp. 3

4

5

3

3

6

1

1978]

Lubin, Eberhard, & Montgomery — Miagrammopes 5

Figure 2. Typical web of Miagrammopes sp. 1 (ca. unipus ) showing horizontal
resting thread under twig and three capture threads.

line in front of it with legs I). The result was to draw the spider
close to the twig. When adopting the cryptic posture, the spider
reached out briefly with legs II and III to pull itself closer to the
twig, then positioned legs II against legs I, holding the broken end
of the resting thread, and legs III against the sides of the abdomen.
In this position it was nearly invisible (see Fig. 3).

M. sp. 2

The web of one adult female was found in the morning (the spider
was without a web at 2100 the night before), and was similar to
some of the webs of M. sp. 1. The spider rested pressed to the
undersurface of a branch, at the end of a horizontal thread about
3 cm long that was strung under the branch (Fig. 3). She held the
broken end of the horizontal thread with one leg II and kept it tense
by pulling in the thread with her hind legs, as described for M.
simus and M. sp. 1. A single, vertical, capture thread (invisible
until powdered) was attached near the other end of the horizontal
thread. The lengths of the non-sticky and sticky portions of the
capture thread were 7:53:40.

6

Psyche

[March

Figure 3. Miagrammopes sp. 2 in cryptic posture as it feeds and holds the non-
sticky resting thread.

M. in temp us

Webs of this species were variable and most were different from
those of other Miagrammopes species. One mature female held
both a horizontal and a vertical sticky thread with her front legs,
and a single, short, non-sticky line with her rear legs (Fig. 4). A
second female also held two capture threads, but both were at an
angle rather than being either horizontal or vertical. The first spider
was induced to move forward along the horizontal thread several
times and her return to the waiting position was observed carefully
(Fig. 4). Each time she tensed the sticky threads by pulling them
in with her front legs; she did not move her hind legs. Another
individual, on a web which was similar except that the horizontal
thread did not appear to be sticky, held the sticky vertical thread
in the same way that M. simus held the capture thread and tensed
it by pulling thread with both front and hind legs. Still other in-
dividuals with single, horizontal, sticky threads (Fig. 5) failed to
pull in silk as they assumed the waiting position. One vertical
thread had several very fine, loose, horizontal lines attached to it,
similar to those shown in Fig. la for M. simus.

1978] Lubin, Eberhard, & Montgomery — Miagrammopes 7

Figure 4. Movements of a female M. intempus, illustrating how thread attach-
ments are manipulated. Letters mark spots on horizontal thread. The spider rested
(a) holding both sticky lines with its front legs, and a third, short, non-sticky line
with its hind legs at point * (the ends of the capture threads were not drawn as they
could not be seen). When lured out onto the horizontal sticky line, the spider car-
ried the vertical thread for some distance (b), then attached it to the horizontal line
and continued on (c). When she returned, she shifted the point of attachment of the
vertical thread again (d), then turned around and pulled in the line with her front
legs and resumed her original position (e). The shifts in attachment were extremely
rapid; the actual motions involved could not be followed, and the shifts were noticed
only by comparing thread positions before and after the spider passed by.

8 Psyche [March

M. sp. 3

The webs of this species were similar to those of M. sp. 1 in hav-
ing variable numbers of capture threads (Table 1). The sticky lines
were not all attached to a non-sticky line at one end, however, but
rather radiated in several directions from a more or less centrally
placed thread (Fig. 6). The spider rested on this thread, often break-
ing one of the capture threads and holding it as described for M.
simus (Figs, lb and 6). This position was also similar to that of
M. intempus in that the spider held a non-sticky line behind it and
a sticky line in front of it. In other cases the spider rested holding
only the non-sticky thread with both front legs. The sticky threads
differed from those of other species of Miagrammopes in being
relatively short (all less than 25 cm) and sticky all the way to the
lower end. The webs were found at night and were gone the next
morning.

M. sp. 4

Webs of M. sp. 4 had one or two capture threads (invisible until
powdered or sprayed with water), 20 to 40 cm long each. The cap-
tured threads were vertical or nearly vertical, but not necessarily
parallel or in the same plane. Of 9 spiders found during the day,
three had two capture threads each, three had a single capture
thread, and three had no capture thread. As in M. simus, the rest-
ing thread was generally under a leaf and often placed at an angle.
Spiders with capture threads rested with one leg I holding a vertical
thread (see Fig. lb) and adjusted the tension both by pulling in the
resting thread with legs IV and the capture thread with leg I.

Spiders without capture threads rested in a cryptic position sim-
ilar to that of M. simus. Often after going into the cryptic posture
(and particularly when disturbed), the spider bounced up and down
on the resting thread in a rhythmic motion reminiscent of rocking
motions of stick insects (Phasmidae). The significance of these
movements is not known.

Prey Capture

We observed in detail prey captures made by four M. simus, two
M. sp. 1, and one M. intempus. Insects that we gave to the spiders
as prey included fruitflies (2-3 mm long), moths (3-7 mm long),
and ants (3-5 mm long). In general, the sequences of prey capture
behavior were similar, but the spiders moved so rapidly that stop-

1978] Lubin, Eberhard, & Montgomery — Miagrammopes

9

Figure 5. Miagrammopes intempus female holding a single thread web. Note
the loose line just anterior to the tip of leg IV(a), and leg II holding the end of the
capture thread (b).

action analysis of video-recording was needed to permit adequate
analysis. Only M. simus was video-taped, using a SONY AV-3400
videorecorder and a macro lens. The descriptions below are based
mainly on analyses of these video-recordings.

Stage I: Prey detection — jerking the capture thread

When an insect was placed on the capture thread, the spider re-
sponded by jerking the thread, The spider quickly flexed her lower
leg I, which held the capture thread, and immediately extended it
again. The maximum distance travelled by the tip of the leg on an
upward jerk was 0.3 leg length (about 2.8 mm), and the quickest
jerks were accomplished in less than 1/60 second (the time span
of a single “frame” of the video-recording). It is tempting to think
that jerking functions in gauging the weight or size of the prey, as
seems to be the case in other uloborids (Eberhard 1969). Spiders
with multiple capture threads (both M. sp. 1 and M. simus) jerked
only the thread on which prey had been placed.

10

Psyche

[March

Figure 6. Miagrammopes sp. 3 on its web, as seen from below and slightly to the
side. The brighter threads are sticky (the web was not powdered). Note that the spider
has broken the end of the capture thread and holds it with the front legs bent to the
side in a manner similar to that shown for M. simus (Fig. 1 b).

Stage II: Entanglement of the prey — sagging the line

The spider sagged the capture thread by dropping the loose silk
it had pulled in with its hind legs, and perhaps also letting out
additional dragline. At almost the same time it manipulated the
capture thread with a series of complex movements of leg I (Fig. 7a)
which resulted in the prey being jerked rapidly back up and down
again (Fig. 7b). Whereas the jerks in stage I displaced a fruitfly
only 5-6 mm, sagging the capture thread caused the prey to drop
26-33 mm in less than 1/30 second. As the prey dropped, it was
often displaced sideways as much as 6 mm (due to air currents?).
Rapid and repeated sagging of the capture thread resulted in the
formation of one or more loops of silk that enveloped the prey.
Such loops were seen in the capture threads of both M. simus and
M. sp. 1.

The mechanism responsible for the formation of these loops is
not clear. One possible explanation is that, due to the relatively
higher air resistance and lower weight of the silk, the prey drops
more rapidly than the silk during a sag, and therefore falls into
the silk below it (Fig. 8a). An alternative explanation (Fig. 8b) is

1978] Lubin, Eberhard, & Montgomery — Miagrammopes 11

that, at the end of a sag, when the spider jerks the line up again,
the prey is “snapped” back upward and accelerated more than the
silk just above it so that it “runs into” the thread above it. The
second of these hypotheses is more appealing since 1) it would work
with non-vertical capture threads whereas the first would not, and
2) we saw two instances in which a loop clearly formed in the thread
just above the prey. In any event, the spider is somehow able to
entangle the prey from a distance by manipulating the capture
thread.

Stage III: Immobilization of prey-wrapping

After manipulating the capture thread to cause one or more sags,

(a) (b)

8

Figure 7. a) Movements of the tip of leg I of a female Miagrammopes simus as
she sagged the capture thread. Points are locations of the tip of leg I holding the
capture thread, taken from a video-taped sequence with “frames” 1/60 sec apart. In
frames 3-5 the tip of the leg remained in the same spot. In frames 9 and 1 1 the tip of
the leg was not visible; these points are not shown in the figure, b) Movements of a
prey on the capture thread while the thread is being sagged and jerked back up and
down, taken from a video-taped sequence (as above). Numbers refer to segments of
the path of movement of the prey on the line during consecutive 1/60 sec intervals.
Scale marker represents 10 mm.

12

Psyche

[March

the spider attached a dragline to the resting thread and moved rap-
idly down the capture thread, pulling in the capture thread and
wadding it up loosely with legs II as it moved. It touched the prey
one or more times with legs I, probably receiving tactile and chem-
ical clues as to the identity of the prey, and then turned 180° and
began wrapping. The wadded up capture thread was transferred
to legs III and wrapped onto the prey, probably thereby increasing
the effectiveness of the initial wraps.

While wrapping, the spider faced away from the prey, holding
the capture thread just above the prey with one leg I and the prey
itself with legs II and III. After 20-30 seconds of wrapping, the
spider cut the capture thread just above and below the prey. It then
rotated the prey package rapidly with legs II (and the palps?) while
continuing to wrap by pulling silk out from the spinnerets and
throwing it onto the prey with legs IV (rotation-wrapping in the
nomenclature of Robinson and Olazarri 1971). While wrapping
the prey, the spider spanned the gap between the two ends of the
capture thread, holding each end with one leg I as do other ulo-
borids (Marples 1962).

Stage IV: Transport of prey to the feeding site

The wrapped prey was transferred to the palps, and the spider
attached a dragline to the thread she had laid on her way down
and then to the broken end of the capture thread. After thus re-
pairing the web, she ran up to the resting thread, holding the prey
in the palps. Once on the resting thread, the spider transferred the
prey to the third pair of legs and again wrapped it. She wrapped
as described above, rotating the prey package with legs II while
hanging from the resting thread with legs I. After wrapping as long
as 5 minutes, the spider transferred the prey back to the palps,
turned facing away from the capture thread, and pulled the resting
thread with legs I as though testing the tension. She then turned
180° and resumed a resting posture with one leg I monitoring the
capture thread. As in other uloborids, the prey package was held
“overhead” in the palps and chelicerae while the spider fed (Fig 3)
and re-wrapped several times during the process of feeding. Feeding
often lasted an hour or more.

Variations in the prey capture sequence

We saw several modifications of the basic prey capture sequence
in M. simus. Small dolichoderine ants were rejected by a spider

1978] Lubin, Eberhard, & Montgomery — Miagrammopes 13

B

?

Y

1 'I

tit

l i

W

|

w

1 2

1 2 3 4 5

Figure 8. Two possible mechanisms which could result in prey becoming en-
tangled as a result of sagging behavior. A) The prey drops faster than the line
below it, and thus becomes entangled. B) The prey’s momentum, acquired when
the spider jerks the line up after a sag, causes it to become entangled in the line just
above it. This hypothesis depends on the thread below the prey being extensible.

on four occasions. Each time the spider jerked and sagged the cap-
ture thread several times, ran a short distance down the capture
thread, wadding it up as it went, and then cut the line above the ant
and ran back up to the resting thread. These ants were thus recog-
nized from a distance, perhaps by their strong alarm odor. After
an ant was rejected, the wadded-up section of the capture thread
was manipulated in the mouthparts for several minutes (feeding?),
then dropped. Rejection of prey thus resulted in destruction of the
capture thread. A new thread was often built within a few hours.

Three other ants, two Camponotus sp. and one Ectatoma sp.,
all about the same size as the spider (6-7 mm long), were attacked
successfully, but modifications of the capture sequence occurred in
all three trials. In two, the spider dropped the lower portion of the

14

Psyche

[March

capture thread after wrapping the prey instead of re-attaching it to
the dragline. In these trials, the spider did not rotate-wrap the prey,
but cut it out after the initial wrap and carried it directly back to
the resting thread. In all three trials, the ants were carried up to
the resting thread dangling from the spinnerets on a 1.5 to 2 cm
thread which was held with one or both legs IV. After reaching the
resting thread, the spider pulled the prey in with legs IV and rotate-
wrapped it.

Live moths of about the same length as the spider escaped readily
from the capture thread by fluttering down it, leaving behind a con-
spicuous trail of scales stuck to the cribellar silk. We observed four
complete prey capture sequences with moths and saw no major
modifications in prey capture behavior, such as those seen with
some araneids (Robinson 1969, Robinson et al. 1971). In three of
the trials, the spider discarded the remaining capture thread after
wrapping; as with the ants as prey, the rotate-wrap stage was
omitted from these captures.

These observations suggest that the decision to retain or discard
the remaining capture thread is made early in the attack sequence,
and is perhaps related to the size of the prey. If the capture thread
is to be abandoned, it may be advantageous for the spider to delay
rotation-wrapping until it reaches the resting thread, where it is less
exposed to visual predators. This explanation is not entirely satis-
factory, however, since if rotation-wrapping is not necessary at the
capture site (it would seem most necessary for just those large prey
for which it is omitted), it would seem advantageous to perform
all rotation-wrapping at the more protected resting thread.

Capture sequences with multiple prey

Capture of small prey such as fruitflies caused little damage to
the capture thread, because the repair of the thread left the remain-
ing sticky portion intact. When presented with a second or third
prey, the spider rushed down the capture thread holding the first
prey in its palps, and attacked the new prey in the usual manner.
Second prey were wrapped together with the first prey and carried
up to the resting thread in the palps in one large package, or
wrapped separately and carried up hanging from the spinnerets,
then wrapped with the first prey.

After only a few prey items were captured, the spider destroyed
the remaining capture thread by dropping the lower end of the

1978] Lubin, Eberhard, & Montgomery — Miagrammopes 15

thread after wrapping the prey instead of attaching it to the drag-
line. The capture thread was destroyed even when a substantial
portion remained undamaged, suggesting that the catching capa-
city of the thread does not limit the number of prey items the spider
will attack. Since Miagrammopes does not attach prey at the feed-
ing site (this is also true of Uloborus diversus — Eberhard 1967),
it is likely that the size of the prey package the spider can hold in
its palps limits it to capturing only a few insects in succession.

Prey capture in M. sp. 2, M. intempus, M. sp. 3, and M. sp. 4

Attack and prey capture behaviors of M. intempus, M. sp. 2,
M. sp. 3, and M. sp. 4 were similar to those described above, in-
volving dramatic sags of the capture thread as the spider ap-
proached the prey, wrapping of the prey at the capture site, and
continued wrapping after the spider returned to the resting thread.

One M. sp. 3 responded to a vibrating tuning fork held nearby
by quickly tightening the capture thread, either by pulling it in with
leg I or by pulling in the resting thread with leg IV. Four attacks
of M. intempus were observed, and in all cases the spider sagged
the capture thread before encountering the prey, then attacked it
by wrapping. One insect, an odorous pentatomid bug, was tapped
repeatedly with the front legs before being wrapped and discarded.
In one sequence it was possible to ascertain that the sticky capture
thread was wadded up as the spider approached the prey, and was
laid onto it as wrapping began.

Prey species captured

Prey taken from webs of an unidentified Miagrammopes sp. in
Bayano, Panama which constructed a web with a single capture
thread like that of M. simus included the following insects: 1 wasp,
1 winged ant, 2 nematocerous flies (1 psychodid), and 1 unidenti-
fied. An additional 29 prey collected as M. sp 1 fed on them included
14 winged ants of two species, 3 wasps, 2 nematocerous flies, 2 other
flies, 1 beetle, and 7 unidentified insects. Four flies were collected
as M. sp. 3 fed on them: 2 nematocerans of probably different fam-
ilies, a dolichopodid fly, and one acalyptrate. One small beetle was
taken from an immature M. sp. 4. These lists make it clear that the
spiders prey on a wide variety of insects, and are not specialists on
any one group.

16

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Reproduction

The egg sac and its web

The egg sacs of M. simus, M. sp. 1, M. sp. 3, and M. intempus
were tubular and elongate, two to four times the length of the
spider, and very similar in color to the adult female. The egg sacs
of M. simus and M. sp. 1 were brown, while those of M. sp. 3 and
M. intempus were lightly coated with green silk. The sacs were
thin-walled, with no fluffy silk inside, and the outlines of the eggs,
which were arranged in one or two rows, were clearly visible.

The females stayed by the egg sacs during the day, either in a
stick posture in line with the sac (Fig. 9) or holding one end of it
with leg I, as seen in some M. sp. 1. In these positions both the
spider and the egg sac were difficult to recognize; they looked like
a dead twig. One M. simus female remained with an egg sac con-
taining 52 eggs for 2 weeks in an outdoor cage. During this time
she did not construct a capture thread. One M. sp. 1, however,
nightly abandoned the daytime cryptic posture and laid several
more or less horizontal, radial lines, suspending the sac by one end
from the “hub” of this tiny web (Fig. 10). A single jagged loop of
sticky silk was laid and the spider rested under the hub. When a

Figure 9. Daytime posture of a Miagrammopes sp. 1 (ca. unipus) female with
an egg sac.

1978] Lubin, Eberhard, & Montgomery — Miagrammopes 17

Figure 10. Egg sac web spun at night by a Miagrammopes sp. 1 (ca. unipus)
female. The horizontal web is seen from above, with the tubular egg sac hanging
down from the “hub.” One end of the sticky “spiral” hung free and had probably
been connected to the radius just to its right. The spider rested near the hub, out
of contact with the egg sac.

small insect was placed on the sticky silk, the spider attacked and
fed on it. During the day this rudimentary orb was gone, and the
spider was back in the cryptic posture at the end of the egg sac.

Emergence of spiderlings

We observed emergence of spiderlings from one egg sac of M.
sp. 1. The spiderlings were first seen one evening easing themselves
through several ragged holes in the sac. They left behind, inside
the empty sac, empty egg shells each with a pink moulted skin stuck
to it. These second instar spiderlings (terminology of Hite, et al.
1966) were relatively inactive and stayed on the sac itself, holding
their anterior legs in an unusual position (Fig. 1 1). The next morn-
ing, they had all moulted again, and the cast skins remained on the
surface of the egg sac while the spiderlings wandered actively in the
vial, holding their legs normally. These spiderlings (third instar)
had fully developed cribella and calamistra.

18

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Uloborids do not have functional cribella until after their first
moult outside the egg sac (Wiehle 1931) and thus cannot produce
sticky silk as newly emerged second instar spiderlings and cannot
make functional, adult-type webs until after the second moult. Sec-
ond instar spiderlings of Uloborus spp. spin orb webs made of dense
sheets of fine threads, lacking the sticky spiral of the adult web
(Wiele 1931, Szlep 1961, Eberhard 1977a). Spiderlings of M. sp. 1
solve the same problem by going through the second moult on the
outside of the egg sac before dispersing; they thus have functional
cribella before spinning their first webs.

Discussion

The webs of the Miagrammopes species in this study are basically
similar in having one or a few simple, sticky capture threads that
are held under tension, sometimes with a few additional fine, non-
sticky threads attached to them. The spiders’ attack behaviors all
involve suddenly sagging the capture thread. Details of placement
of the capture and resting threads, and the spiders’ web tensing
behavior are variable among the species, and even to some extent
among individuals of some species. Two of the characteristics de-
scribed for M. sp. 1 appear to be unique among spiders — the
double moult of the young before leaving the egg sac, and the
special feeding web of the female near her egg sac.

The web of Miagrammopes species from Natal was similar to
some of the webs of M. intempus and M. sp. 3 in having a single
horizontal capture thread without a separate resting thread (Aker-
man 1932). The presence of additional fine threads attached to the
capture thread was not noted in webs from Natal, but they would
almost surely have gone unnoticed unless the webs were powdered.
Web construction behavior was similar in the Natal species. The
spider sat at one end of the completed capture thread, facing it;
the thread may have been broken with the spider bridging the gap
with its body (e.g. Marples 1962), but Akerman’s drawing shows
an intact line. The thread was held under tension by pulling it in
with legs IV as do all six species of this study and was also quickly
sagged when prey hit it. The single, horizontal, capture thread web
may represent a further simplification of an already simple web,
with a single sticky thread taking the place of both the horizontal
resting thread and the vertical capture thread.

1978] Lubin, Eberhard, & Montgomery — Miagrammopes 19

Figure 1 1. Typical leg positions of second instar Miagrammopes sp. 1 (ca. uni-
pus) as they rested on the outside of the egg sac.

Although uloborids in general seem to construct their webs in the
early morning (e.g. Eberhard 1972, Lubin and Eberhard unpubl.),
Miagrammopes are more variable. Thus, while M. simus, M. sp. 1,
and M. sp. 4 tend to have webs up early in the morning, they, as
well as the species from Natal (Akerman 1932) sometimes build at
other times, and M. intempus and M. sp. 3 commonly build in the
evening. An unidentified species in New Guinea which spins single,
horizontal threads also tends to build at night (Robinson and Rob-
inson 1974, M. Robinson, pers. comm.). Readiness to build at dif-
ferent times of the day might be expected in view of the rapidity
with which new webs can be made and the small investment of
material which they represent. The tendency to discard webs sup-
port this idea.

Kaston (1964) suggested that the reduced web of Miagrammopes
is derived from a Stegodyphus- type web (Eresidae) which consists
of irregularly spaced radii with connecting sticky threads. A web
similar to that of Sybota (Uloboridae) seems to us a more likely
precursor of a Miagrammopes- type web. Sybota producta (Sim.)

20

Psyche

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lays cribellar silk directly on the radii and frame threads of orb-
like webs which lack a sticky spiral (Wiehle 1931). The spider ap-
parently does not manipulate tensions in the web once it is built
(Wiehle 1931: Figs. 14 and 17). Such a web might conceivably
become reduced to a web like that of M. sp. 1 by loss of frame
and auxiliary spiral threads and reduction of the hub to a single
resting thread. On the other hand, the jagged pattern of the sticky
spiral found in the egg sac webs of M. sp. 1 suggests an affinity
with Uloborus (sens, lat.) or Hyptiotes (Uloboridae). Spiders of both
these genera commonly lay a jagged, sticky spiral on the periphery
of the orb (McCook 1889, Eberhard 1972, Eberhard, unpubl.). An
unidentified species of Uloborus (sensu strictu) builds an essentially
identical egg sac web (Eberhard, in prep.), and Uloborus diversus
also places sticky silk around its egg sacs (Eberhard 1969).

The most likely adaptive advantage of a single thread capture
web would seem to be its near invisibility to prey, since at least
some flying insects can detect and avoid webs (Bristowe 1941, Rob-
inson and Robinson 1970, 1973, Lubin 1973, Buskirk 1975, Eber-
hard in prep., Lahman and Zuniga in prep.). This is apparently
ruled out, however by the fact that at least two species (Af.sp.3 and
the New Guinea species) and perhaps a third ( M . intempus) usually
build their webs at night when visibility is probably unimportant.

Another possible advantage would be that predators using webs
as cues to the presence of prey would be unlikely to detect webs of
Miagrammopes. Some predators may use webs in this way, though
some are known not to (Eberhard 1970). The significance of the
very thin, slack lines attached to the capture threads remains even
more of a mystery.

The obvious disadvantage of a single thread capture web is the
low probability of a flying insect striking the web. Robinson and
Robinson (1976) suggested that the numerous nematocerous flies
which tend to rest on non-sticky spider threads might try to alight
on Miagrammopes capture threads and thus become entangled.
Indeed Akerman (1932) noted a number of “gnats” caught by the
Miagrammopes species in Natal. Some nematocerous flies were
among the prey collected in this study, but many other kinds of
small insects were collected as well. Certainly the webs of Mia-
grammopes are not specialized to the extent of exclusively or even
principally capturing nematocerous flies which alight on them. Some
other spiders with reduced webs use chemical attractants for spe-

1978] Lubin, Eberhard, & Montgomery — Miagrammopes 21

cific kinds of prey (Eberhard 1977b), but the wide variety of cap-
tured prey rules out this prey capture technique for Miagrammopes.

Summary

The webs of six species of Miagrammopes (Uloboridae) studied
in Panama, Colombia, Costa Rica, and Venezuela have only one
or a few sticky capture threads. Miagrammopes simus and M. sp. 2
have one vertical capture thread attached to a non-sticky, hori-
zontal resting thread. Miagrammopes sp. 1 (ca. unipus) builds
from 1 to 5 near-vertical capture threads, and M. intempus, M.
sp. 3, and M. sp. 4 use one or more capture threads that vary in
their spatial arrangement. Webs are pulled taut by pulling in silk
with either the front legs or the hind legs or both. The spiders as-
sume highly cryptic postures during the day as they rest on their
webs or near the egg sac.

Attack and prey capture behavior in all species involves rapid
jerking and sagging of the capture thread by the spider, resulting
(in at least two species) in the prey becoming entangled in one or
more loops of sticky thread before the spider arrives to attack.

Second instar spiderlings of M. sp. 1 do not disperse, but moult
a second time on the surface of the egg sac. Thus they construct
webs only after they have fully formed calamistra and cribella and
are capable of producing sticky silk. A mature female M. sp. 1
constructed nocturnal egg sac webs that were reminiscent of small
uloborid orb webs.

The adaptive advantage of the reduced web of Miagrammopes
is unclear. Many species of small insects are taken as prey and
chemical attractants do not seem to be used.

Acknowledgements

Brent Opell kindly identified the spiders and reviewed the manu-
script. Larry Kirkendall provided records of prey from Bayano,
Panama. We are grateful to the following institutions and persons
for their help and cooperation: Central Hidroelectica Anchicaya
and Dr. Victor Manuel Patino of the Jardin Botanico in Mate-
guadua (Colombia), Dr. Carlos E. Valerio of the Universidad de
Costa Rica, and the Ministerio de Recursos Naturales Renovables
(Venezuela). This work was supported by the Smithsonian Tropi-
cal Research Institute and the Comite de Investigaciones of the
Universidad del Valle.

22

Psyche

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References

Akerman, C.

1932. On the spider Miagrammopes sp. which constructs a single-line snare.
Ann. Natal Mus. 5: 83-88.

Buskirk, R.

1975. Coloniality, activity patterns and feeding in a tropical orb-weaving
spider. Ecology 56(6): 1314-1328.

Eberhard, W. G.

1967. Attack behavior of diguetid spiders and the origin of prey wrapping in
spiders. Psyche 74: 173-181.

1969. The spider Uloborus diversus and its web. Ph.D. Dissertation, Harvard
University, 214 pp.

1970. The predatory behavior of two wasps, Agenoides humilis (Pompilidae)
and Sceliphron caementarium (Sphecidae), on the orb weaving spider
Araneus cornutus (Araneidae). Psyche 77: 243-251.

1972. The web of Uloborus diversus (Araneae: Uloboridae). J. Zool., Lond.
166: 417-465.

1977a. The webs of newly emerged Uloborus diversus and of a male Uloborus
sp. (Araneae, Uloboridae). J. Arachnol. 4(3): 201-206.

1977b. Aggressive chemical mimicry by a bolas spider. Science 198(4322):
1173-1175.

Hite, J., W. Gladney, J. Lancaster, and W. Whitcomb

1966. Biology of the brown recluse spider. Agr. Expt. Sta. Univ. Ark. Bull.
711: 3-26.

Kaston, B. J.

1964. The evolution of spider webs. Am. Zool. 4: 191-207.

Lubin, Y. D.

1973. Web structure and function: the non-adhesive orb-web of Cyrtophora
moluccensis (Doleschall) (Araneae: Araneidae). Forma et Functio 6:
337-358.

Marples, B. J.

1955. A new type of web spun by the genus Ulesanis with the description of
two new species. Proc. Zool. Soc. Lond. 125: 751-760.

1962. Notes on spiders of the family Uloboridae. Ann. Zool. Agra 4: 1-11.

McCook, H. C.

1889. American Spiders and their Spinningwork. I. Webs and nests. Phila-
delphia: published by the author.

Robinson, M. H.

1969. Predatory behavior of Argiope argentata (Fabricius). Am. Zool. 9:
161-173.

Robinson, M. H. and J. Olazarri

1971. Units of behavior and complex sequences in the predatory behavior of
Argiope argentata (Fabricius) (Araneae: Araneidae). Smith. Contrib.
Zool. 65: 1-36.

Robinson, M. H. and B. Robinson

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

1978] Lubin, Eberhard, & Montgomery — Miagrammopes 23

1974. A census of web-building spiders in a coffee plantation at Wau, New
Guinea, and an assessment of their insecticidal effect. Trop. Ecol. 15:
95-107.

1976. A tipulid associated with spider webs in Papua New Guinea. Entom.
Mon. Mag. 112: 1-3.

Robinson, M. H., B. Robinson, and W. Graney

1971. The predatory behavior of the nocturnal orb web spider Eriophora
fuliginea (C. L. Koch) (Araneae: Araneidae). Rev. Per. Entom. 14(2):
304-315.

SZLEP, R.

1961. Developmental changes in the web-spinning instinct of Uloboridae:
construction of the primary-type web. Behaviour 27: 60-70.

Wiehle, H.

1931. Neue Beitrage zur Kenntnis des Fanggewebes der Spinnen aus den
Familien Argiopidae, Uloboridae, und Theridiidae. Z. Morph. Okol.
Tiere 22: 348 400.

REVISION OF THE SOUTH TEMPERATE GENUS

GLYPHOLOMA JEANNEL, WITH FOUR NEW SPECIES
(COLEOPTERA: STAPHYLINIDAE: OMALIINAE)*

By Margaret K. Thayer and Alfred F. Newton, Jr.

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

Introduction

The southern temperate silphid genus Glypholoma Jeannel was
transferred to the staphylinid subfamily Omaliinae by Newton
(1975), and Lathrimaeodes Scheerpeltz (originally placed in the
Omaliinae) was then synonymized with it. Newton (op. cit.) also
presented additional descriptive information, a new locality record,
and some discussion of the affinities of Glypholoma within the
Omaliinae. Since that time, four new species of the genus have come
to our attention, including one from Australia which greatly
enlarges the known range of Glypholoma (previously only parts of
Chile and Argentina). The discovery of these new species and the
availability of a wealth of material of the type species , pustuliferum
Jeannel, for detailed study led to our decision to revise the genus.

Methods

Measurements, made with an ocular micrometer in a Leitz
binocular dissecting microscope, are defined as follows:

Length: measured in lateral view from front of (closed) mandibles
to apex of abdomen (excluding genitalia if exserted, and attempting
to estimate “normal” degree of contraction of abdomen).

Width: maximum body width, across closed elytra at widest point
(usually near middle).

Head width: in dorsal view, maximum width including eyes.

Head length: measured along midline from anterior margin of
labrum to level of centers of ocelli, viewed perpendicular to line of
measurement.

*Published with the aid of a grant from the Museum of Comparative Zoology,
Harvard University.

Manuscript received by the editor October 6, 1978.

25

26

Psyche

[March

Antennal length: from constriction between scape and its basal
articulatory process to apex of last antennal segment.

Ocellar diameter: measured antero-posteriorly in dorsal view.

Pronotal width: maximum width.

Pronotal length: along midline from base to apex, viewed per-
pendicular to line of measurement.

Elytral width: same as Width, above.

Elytral length: measured along suture from apex of scutellum to a
line tangent to elytral apices.

Prosternal process length: measured along midline from a trans-
verse tangent to the anterior margin of the procoxal cavities to apex
of prosternal process.

Procoxal length: measured in ventral view between transverse
tangents to anterior and posterior faces of procoxae.

Mesosternal length: along midline from anterior margin to apex
of mesosternal process.

Mesosternal process length: along midline from transverse tan-
gent to anterior margin of mesocoxal cavities to apex of mesosternal
process.

Mesocoxal length: measured in ventral view between transverse
tangents to anterior and posterior margins of mesocoxal cavities.

Mesosternal procoxal cavity length: along midline from anterior
margin of mesosternum to a line connecting the posterior edges of
the cavities.

Metasternal length: along midline between extremities of inter-
coxal processes.

Metasternal antecoxal sutures: measured from lateral limit to
lateral limit.

Metasternal width: between posterolateral corners of metaster-
num.

Hind coxal length: lateral edge to posteromesal corner.

Hind femoral length: including trochanter, from posteromesal
corner of coxa to most distal point of femur.

Hind tibial length: along mesal side (not including any spines or
setae).

Hind tarsal length: from point of insertion of tarsus on tibia to
tarsal apex, not including claws or empodial setae. The dividing line
between the fourth and fifth hind tarsal segments was taken to be
the point of insertion of the fifth segment on the fourth.

1978] Thayer & Newton — Revision of Genus Glypholoma 27

Abdominal segments are numbered according to their morpho-
logical origin. (The first segment visible ventrally is therefore the
third segment.)

Mean length and width are given for each species, ± one standard
deviation.

Preparation of specimens for scanning electron microscope pic-
tures consisted of clearing heads, mouthparts, and prothoraces in
hot 1 N potassium hydroxide, critical-point drying all parts except
elytra, and coating with gold-palladium mixture. Cleared and
dissected specimens of pustuliferum and rotundulum and of the
aedeagi of other species were examined under dissecting and
compound microscopes.

Drawings were made with the aid of a camera lucida attachment
on a Leitz binocular dissecting microscope.

Acknowledgements

This study probably would not have come about were it not for S.
B. Peck’s extensive collecting in Chile and his kindly making this
material available to us; he later generously provided us with a
multitude of Australian specimens as well.

Specimens were borrowed from the following institutions (ab-
breviated in the text as indicated) and we extend our thanks to the
curators involved for their cooperation in lending specimens.

CAS California Academy of Sciences, San Francisco,
California, U.S.A. (D. H. Kavanaugh)

CNC Canadian National Collection, Ottawa, Ontario,
Canada (J. M. Campbell)

MCZ Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, Massachusetts, U.S.A.

NMVM National Museum of Victoria, Melbourne, Victoria,
Australia (A. Neboiss)

Specimens are also deposited in the following collections:

ANIC Australian National Insect Collection, Canberra,
A.C.T., Australia

ANMT A. F. Newton, Jr. and M. K. Thayer, Cambridge,
Massachusetts, U.S.A.

FMNH Field Museum of Natural History, Chicago, Illinois,
U.S.A.

SBP S. B. Peck, Ottawa, Ontario, Canada

28

Psyche

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Figs. 1-4. Glypholoma spp. 1. G. pustuliferum. 2. G. pecki, holotype. 3. G.
temporale, holotype. 4. G. tenuicorne, holotype. Scale lines = 1.0 mm.

1978] Thayer & Newton — Revision of Genus Glypholoma 29

The scanning electron microscope work done for this paper was
made possible by National Science Foundation grants BMS-7502606
(J. F. Lawrence, principal investigator) and BMS-74 12494 (SEM
operating grant) with the superb technical assistance of E. Seling.
We also thank L. H. Herman for calling our attention to the Klinger
and Maschwitz paper, H. S. Dybas for reading and commenting
upon the manuscript, and N. Hinnebusch for typing the manuscript.

Glypholoma Jeannel

Glypholoma Jeannel, 1962: 482; Newton, 1975: 53. Type species: Glypholoma
pustuliferum Jeannel, 1962: 483, by original designation and monotypy.
Lathrimaeodes Scheerpeltz, 1972: 58; (placed in synonomy by Newton, 1975: 54).
Type species: Lathrimaeodes pustulipenne Scheerpeltz, 1972: 59, by original
designation and monotypy.

Diagnosis: Separable from other known Omaliinae by the exca-
vate hind coxae, each elytron with eleven more or less distinct striae,
male genital segment with a small “button” internally at the anterior
end of sternite 9, and visible dorsal pleural-coxal articulation in the

Fig. 5. Glypholoma rotundulum. Scale line = 1.0 mm.

30

Psyche

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prothorax. In addition, the combined presence of all of the follow-
ing characters serves to distinguish Glypholoma from other Omali-
inae. (Some other genera share one or a few of these characters with
Glypholoma, but no other genus examined possesses all of these
characters as Glypholoma does.)

1 . Gular sutures widely separated, minimum separation equal to
0.12-0.18 times the head width.

2. Prothoracic pleural-sternal articulation present.

3. Procoxa with mesal articulating groove.

4. Mesosternal procoxal cavities long, 0.40-0.75 times the total
mesosternal length.

5. Hind femur relatively short, ranging from 0.92-1.05 times
hind coxal length.

6. Deflexed lateral portion of elytron short, only 0.64-0.67
times as long as total length of elytron (measured in lateral view).

7. Humeral margin of elytron serrulate.

8. Median lobe of aedeagus with membranous part of basal
bulb allowing dorsal-ventral instead of lateral-lateral contraction
and expansion.

9. Epistomal suture present and complete (although with or
without median stem).

Description: Ovoid (narrower posteriorly) to more or less oblong
in dorsal view, slightly to strongly convex dorsally in cross section.
Sparsely pubescent to nearly glabrous on dorsal surface, widely
spaced macrosetae on alternate intervals of elytra in a fairly
characteristic pattern (pattern varies some among the species; see
figs. 26, 29). Microsculpture lacking on dorsal surface of head,
pronotum, and elytra. Length 2. 1-3.5 mm, width 1.0- 1.5 mm.

Head capsule about as in fig. 58, lacking postocular ridge,
temples, and nuchal constriction except in temporale, which has
temples and a lateral nuchal constriction (fig. 57). Head about 1.8
times as wide as long, about 0.58 times as long as pronotal length,
with pair of distinct ocelli on dorsal surface (see especially fig. 6); no
anteocellar grooves or pits. Epistomal suture with internal rein-
forcing ridge present, angulate or arcuate, with or without median
stem. Antennae usually about 1 to 1.5 times as long as head width,
varying from filiform ( tenuicorne , about 2.2 times head width) to
having a moderately developed club of about three to five segments;
basal five to eight antennal segments glabrous except for a few

1978] Thayer & Newton — Revision of Genus Glypholoma 31

Figs. 6-9. Glypholoma pustuliferum. 6-7. Head, anterior oblique and ventral views, respectively. 8. Mandibles and
epipharynx, ventral view. 9. Tenth antennal segment. Scale lines: fig. 9 = 0.01 mm; others = 0.1 mm.

32

Psyche

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Figs. 10-13. Glypholoma pustuliferum. 10. Apical antennal segments. 11. Base
of fourth maxillary palpal segment, external view. 12. Hypopharynx and labium,
dorsal view. 13. Left maxilla, ventral view. Scale lines: fig. 1 1 = 0.01 mm; others =
0.1 mm.

1978] Thayer & Newton — Revision of Genus Glypholoma 33

scattered long setae. Apical antennal segments simple, without
apical gutters or invaginations (see figs. 9, 10), with fairly dense
short setae in addition to scattered long setae (see fig. 10, compare
basal segments, fig. 7).

Labrum transverse, narrowly rectangular to slightly bilobed
anteriorly, ventrally as in fig. 8. Mandible (of pustuliferum and
rotundulum, at least) without preapical teeth, with a medial setose
area about midway from base to apex and a well-developed molar
lobe (see figs. 8, 14-17). Molar lobe apparently articulating dorsally
and ventrally with mediobasal area of mandible proper. Maxilla
more or less as in fig. 13, the palp generally filiform with fourth
(apical) segment 2.5 to 5 times as long as third, a group of sensilla as
in fig. 1 1 on its dorsolateral surface near the base. Hypopharynx (of
pustuliferum and rotundulum, at least) about as in fig. 12. Labium
apparently bilobed, with three-segmented palps arising from separ-
ate palpigers; segments of palp subequal in length and width (see
fig. 7, also Newton, 1975, fig. 3). Mentum large and trapezoidal;
gular sutures separate, their minimum separation 0.12 to 0.18 times
the head width (see fig. 7).

Pronotum 1.5 to 1.8 times as wide as long, about 0.35 times as
long as elytra; widest point variable, from posterior corners to just
behind middle; with complete sharp lateral margins, explanate in at
least basal half; lacking lateral foveae with internal pillars. Post-
coxal process of pronotum acutely triangular, apparently a bar to
coxal flexation (see figs. 18, 20). Prosternum with or without
median longitudinal keel; prosternal intercoxal process extending
one-half to two-thirds of the (antero-posterior) length of the
procoxae. Procoxa with external longitudinal keel and mesal
transverse articulating groove (see figs. 18-21). Protrochantin ex-
posed, shorter than postcoxal pronotal process, dorsal pleural-coxal
(trochantinal-coxal) articulation visible, pleural-sternal articulation
present (the last possibly absent in tenuicorne), as in fig. 20.

Mesosternum 0.5 to 0.6 times as long as metasternum, with a
nearly acute process (except in rotundulum, see figs. 35, 37)
extending between the mesocoxae for four to seven tenths of their
length (figs. 34, 36). Mesosternal process not medially longitudinally
carinate. Anterior part of mesosternum with cavities for reception
of procoxae 0.4 to 0.75 times as long as whole mesosternum.

34

Psyche

[March

Figs. 14 17. Glvpholoma pustuliferum, right mandible. 14 15. Mesal view,
whole mandible and molar surface, respectively. 16-17. Dorsal view, whole mandible
and part of mola, respectively. Scale lines: figs. 14, 16 = 0.1 mm; figs. 15, 17 = 0.01 mm.

1978] Thayer & Newton — Revision of Genus Glypholoma 35

Figs. 18-21. Glypholoma pustuliferum. 18 19. Prothorax, ventrolateral view,
with and without procoxa and trochantin, respectively. 20. Right procoxa and
surrounding area, ventrolateral view, with coxa rotated anteriorly. 21. Prothorax
and head, anteroventral view, dpc = dorsal pleural-coxal articulation; g = mesal
articulating groove of coxa; k = external coxal keel; np = postcoxal process of
pronotum; ps = pleural-sternal articulation; sp = prosternal intercoxal process. Scale
lines = 0.1 mm.

36

Psyche

[March

Mesepisternal-mesepimeral suture present; mesosternal-pleural su-
ture present posteriorly, but disappearing anteriorly, apparently
with fusion of the two sclerites (except in tenuicorne, where it may
be complete). Metasternum with short anterior process meeting
mesosternal process between mesocoxae, short process with bifid
apex between metacoxae, and antecoxal sutures as in fig. 59
( temporale only) or 60. Metacoxae excavate, i.e. with posterior face
vertical and entire postero-ventral margin explanate (fig. 38).

Legs rather slender; tibiae only slightly, if at all, wider at apex
than at base, usually with some small spines on the external surface
(slightly fewer to slightly more than in fig. 23), meso- and metatibia
generally with more spines than protibia, the sexes usually similar
within each species. Hind coxa and femur subequal in length, hind
tibia about 0.9 times as long as femur, hind tarsus 0.5-0. 7 times as
long as tibia. All tarsi with five segments, hind tarsus with last
segment one-third to two-thirds as long as first four together.
Bisetose empodium between bases of tarsal claws (see fig. 39).

Elytra together about three-quarters as wide as long, each with
eleven punctate striae (most clearly delimited in the middle section,
more confused anteriorly and posteriorly) which are impressed
between the punctures except in tenuicorne; eleventh stria (adjacent
to epipleural keel) somewhat irregular. Macrosetae present on
alternate intervals, associated in some species with raised pustules
and/or spots; (except perhaps in tenuicorne?) intervals finely punc-
tate between macrosetae (see figs. 26, 27, 29). Elytron with epi-
pleural keel complete, serrulate in humeral region, intersecting with
upturned lateral edge of elytron at about two-thirds the distance
from elytral base to apex. Elytron covering abdomen through
segment 5 or 6, with apparent elytral-abdominal interlocking patches
on the internal elytral surface just antero-mesal to the confluence of
the epipleural keel and elytral edge (see figs. 30, 31) and on the
lateral area of abdominal segment 3 (see fig. 46). (Whole complex
seen in detail only in pustuliferum and rotundulum; elytral patch
present in all species except possibly tenuicorne; because of the
limited number of specimens available of three of the species, an
exhaustive search for the abdominal patches could not be made.)
Elytron with patch of small single or grouped teeth near apex of
ventral surface (as in figs. 30, 32, 33). Wings covering abdomen
through tergite 3 when folded, except in brachypterous individuals

1978] Thayer & Newton — Revision of Genus Glypholoma 37

Figs. 22-25. Glypholoma spp., prolegs. 22-24. G. pustuliferum; 22, male tarsus,
oblique ventral view; 23, male left tibia, posterior view; 24, female tarsus, oblique
ventral view. 25. G. rotundulum, male tarsus, oblique ventral view. Scale lines = 0.1

mm.

38

Psyche

[March

Figs. 26-29. Glypholoma spp. 26-28. G. pustuliferum, right elytron; 26, dorso-
lateral view; 27, detail of fig. 26, including two pustules; 28, strial puncture (detail of
fig. 26). 29. G. rotundulum, right elytron, dorsolateral view. Scale lines: figs. 26, 29 =
1.0 mm; fig. 27 = 0.1 mm; fig. 28 = 0.01 mm.

1978] Thayer & Newton — Revision of Genus Glypholoma 39

Figs. 30-33. Glypholoma spp., internal elytral surface. 30-32. G. pustuliferum;
30, overall view; 3 1 , detail of lateral area (“ 1 ” in fig. 30); 32, detail of apex (“a” in fig.
30). 33. G. rotundulum, same view as fig. 32. Scale lines: figs. 30, 31 = 0.1 mm; figs.
32, 33 = 0.01 mm.

40

Psyche

[March

of rotundulum, in which the wings only reach the apex of tergite 2
and are not folded. Distinct anal flap present on wings of pustuli-
ferum and rotundulum, probably in other species also. Folding
pattern of fully-developed hind wing similar to that illustrated for
Anthobium (=Eusphalerum) sorbi by Forbes (1926: fig. 33), the first
transverse fold being a hinge by which the costal margin is turned
about 90°. (In an individual, the hinges of the two wings usually
form slightly different angles.)

Abdomen with most of segment 8 and part of genital segment
exposed; first visible sternite is sternite 3. Tergites 4 or 5 to 8 fairly
well to well-sclerotized, spiracles located in tergites 4 or 5 to 8, in
membrane beside tergites 1 to 3 or 4; one pair of paratergites on
each of segments 3 to 6 or 7, may be partly or entirely fused with
sternites laterally (as almost complete fusion in fig. 47); tergite(s) 4
or 4 and 5 with paired patches of medially-directed microtrichia
which cover about half to nearly all of the surface of the tergites
(figs. 40-43); tergite 7 with an apical fringe (“palisade fringe” of
some authors) as in fig. 45. Intersegmental membranes with a brick
wall pattern of irregular plates as in fig. 44, some dorsal plates with
posterior teeth as shown. Sternite 2 extending up around sides of
abdomen slightly, but in tenuicorne appearing to be membranous,
its limits therefore not determinable. Sternites 2 and 3 with small
intimately associated intercoxal processes, more or less as in fig. 48,
sternite 3 (at least in pustuliferum and rotundulum) with a trans-
verse fold across its middle one-fourth to one-half about at posterior
margin of intercoxal process (this area not visible in other species
because of telescoping of abdomens). Sternite 3 without distinct
coxal cavities, metacoxae simply protruding parallel to sternite’s
surface; at least in some species with a ridge near basal margin and,
like sternites 4 to 5 or 6, a curved ridge just inside each lateral
margin. Sternite 8 with anterior median projection (see figs. 49,
61-65) associated with a gland system, similar to that described by
Klinger and Maschwitz (1977). In at least three species ( rotundulum ,
pecki, pustuliferum ) gland reservoir extending anteriorly as far as
anterior margin of segment 5.

Male: Peg setae apparently absent from trochanters, femora, and
tibiae; first four segments of protarsus slightly broadened, spatulate
setae in pairs on segments 1 to 3 of protarsus (figs. 22, 25), and
singly on first two (possibly three in pecki ) segments of mesotarsus

1978] Thayer & Newton — Revision of Genus Glypholoma 41

Figs. 34-37. Glypholoma spp., pterothorax. 34, 36. G. pustuliferum, ventral and
oblique lateral views, respectively. 35, 37. G. rotundulum, ventral and oblique lateral
views, respectively. Scale lines = 0.1 mm.

42

Psyche

[March

Figs. 38-41. Glypholoma spp. 38-40. G. pustuliferum; 38, metasternum and
right metacoxa, lateral view; 39, male left mesotarsus, ventral view; 40, abdomen,
dorsal view. 41. G. rotundulum, pterothorax and abdomen, dorsal view. t7 = tergite
7. Scale lines: figs. 38, 39 = 0.1 mm; figs. 40, 41 = 1.0 mm.

1978] Thayer & Newton — Revision of Genus Glypholoma

43

Figs. 42-45. Glypholoma pustuliferum, abdomen. 42-43. Microtrichial patch on tergite 5 (right side). 44. Intersegmental
membrane between tergites 5-6. 45. Apical fringe of tergite 7, oblique anterior view. Scale lines: fig. 42 = 0.1 mm; others =
0.01 mm.

44

Psyche

[March

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1978] Thayer & Newton — Revision of Genus Glypholoma 45

(see fig. 39). Genital segment with tergite 9 narrowly continuous
across dorsal midline, tergite 10 short, sternite 9 attenuate, fairly
well sclerotized in posterior one-third or so, only lightly sclerotized
anteriorly, with a small cylindrical protuberance on internal surface
near anterior end (see figs. 66-75). Aedeagus of general staphylinid
type (median lobe with basal bulb, small dorsal median foramen,
ventro-apical median orifice); parameral side of aedeagus facing
dorsally within abdomen; small basal piece present as a more or less
C- or U-shaped strap on dorsal side of basal bulb (often very lightly
sclerotized and difficult to see); basal bulb of median lobe lightly
sclerotized, with a membranous band around the abparameral side,
presumably allowing dorsal-ventral bellows-like contractions; in-
ternal sac with dense armature of short, fine spines; apex of median
lobe a finger-like projection, even with or slightly shorter than
apices of parameres; parameres with or without subapical setae (see
figs. 76-85).

Female: Pro- and mesotarsus without spatulate setae, as in fig. 24.
Genitalia (based on pustuliferum and rotundulum, except as noted)
about as in Newton, 1975, figs. 8-10: sclerotized spermatheca
apparently absent, sternite 9 present, tergite 9 divided into two
lateral sclerites, tergite 10 U-shaped; proximal gonocoxites separate,
about half as long as distal gonocoxites; in all species in which
females are known, distal gonocoxites separate, each bearing at its
apex a short stylus with 1 (or 2?) long apical seta(e) and one or more
subapical setae.

Immature stages unknown.

Range: Southern Chile and Argentina south to Tierra del Fuego;
Victoria, Australia; see maps, figs. 50, 51.

Key to the species of glypholoma jeannel

1. Antenna filiform (fig. 56); elytra without raised pustules; dorsal

surface uniform in color; body form as in fig. 4

G. tenuicorne n.sp.

Antenna with gradual apical club (figs. 52-55); elytra with or
without raised pustules; dorsal surface with various color
markings; body form otherwise (figs. 1-3, 5) 2

2. Elytra with raised pustules on alternate intervals (fig. 26) .3

Elytra without raised pustules (fig. 29) 4

46

Psyche

[March

3. Postocular lobe present (fig. 57); metasternal antecoxal sutures

long, about three-fourths the width of the metasternum (fig.

59) ; elytral pustules concolorous with surrounding area . .
G. temporale n.sp.

Postocular lobe absent (fig. 58); metasternal sutures short, no
more than about one-third the width of the metasternum (fig.

60) ; elytral pustules lighter in color than surrounding area
G. pustuliferum Jeannel

4. Antennal segment 11 about 4X as long as segment 10 (fig. 52);

mesosternal process a simple angulate projection between
mesocoxae (cf. figs. 34, 36); pronotum with lateral margins
explanate, dorsal surface uneven; dark median spot on ver-
tex of head, dark median streak on pronotum, no markings

on elytra G. pecki n.sp.

Antennal segment 11 not more than about 2.5X as long as
segment 10 (fig. 55); mesosternal process an elevated pentag-
onal protuberance between mesocoxae (figs. 35, 37); pro-
notum evenly convex, not broadly explanate laterally; head
and usually pronotum without markings, elytra with yellow
spots on alternate intervals, sometimes with additional pale
areas; usually brachypterous G. rotundulum n.sp.

Glypholoma pustuliferum Jeannel

Glypholoma pustuliferum Jeannel, 1962: 483; Newton 1975: 53-54; Szymczakowski,
1976: 424.

Lathrimaeodes pustulipenne Scheerpeltz, 1972: 59. (Placed in synonymy by Newton,
1975: 54.)

With the characters of the genus as described above.

Moderately convex dorsally, shape as in fig. 1, various shades of
brown in color: head darkest, reddish-brown to nearly black (with
area anterior to epistomal suture the darkest part), basal two-thirds
of elytra (except lateral margins) roughly concolorous with main
part of head except for much lighter yellow pustules on alternate
intervals; central area of pronotum similar to elytra or slightly
lighter, with pronotal and elytral margins yellow to yellowish-
brown, legs and dorsum of abdomen generally about the color of
the central pronotal area. Ventral body surface usually reddish-
brown, with lateral pronotal margins and deflexed portion of elytra

1978] Thayer & Newton — Revision of Genus Glypholoma 47

paler. Some specimens (teneral?) are yellowish to reddish-brown,
nearly unicolorous, with elytral pustules only slightly paler. Very
short setae (similar to that in fig. 28) sparsely distributed over body
surface, legs more densely setose, especially tibiae and tarsi. Length:
3.0 ± 0.2 mm (2. 6-3. 5); width: 1.3 ± 0.1 mm (1.1-1. 5) (N = 55).

Head capsule as in figs. 6, 7, and 58, epistomal suture with median
stem (not always visible on surface). Antennal length about 1.1
times head width, first six antennal segments without dense pubes-
cence, antennal club five-segmented, gradual (see fig. 54). Ocellar
diameter about 0.095 times head width. Labrum shallowly emar-
ginate anteriorly as in fig. 6. Maxillary palp with fourth segment
about three times as long as third, the two approximately subequal
in width,

Pronotum as in fig. 1, with several small shallow paramedian and
lateral depressions; lateral margins explanate. Prosternum deeply
pitted and rugose, but lacking microsculpture, and without median
longitudinal carina (see figs. 18-21). Mesosternum with dense
irregular microsculpture, cavities for reception of procoxae about
half the length of the mesosternum; mesosternal process fairly
narrow, blunt, extending seven-tenths of the way between meso-
coxae (see figs. 34, 36). Metasternum convex with faint median
longitudinal depression, surface without microsculpture but with
short setae in large punctures (see fig. 36), antecoxal sutures short,
extending only three-tenths the width of the metasternum (fig. 60).
Protibia setose, with several small spines on outer surface (fig. 23),
meso- and metatibia with slightly more spines. Hind tarsus six-
tenths as long as hind tibia, fifth segment of hind tarsus two-thirds
as long as first four together. Empodial setae subequal in length to
tarsal claws.

Elytral striae impressed between punctures, intervals punctate
and with short fine setae (in addition to macrosetae on alternate
intervals; see figs. 26, 27). Pale pustules located anterior and
adjacent to macrosetae. Angle formed by elytral apices at suture
about 180°. Wings fully developed, with anal flap.

Abdomen with tergites 4 to 8 well sclerotized, spiracles located in
tergites of these segments; paratergites separated anteriorly but not
posteriorly from sternites on segments 4, 5 and 6 (see fig. 47),
apparently not separated from sternites on segments 3 and possibly
7. Tergites 4 and 5 with paired patches of medially-directed

48

Psyche

[March

microtrichia as in figs. 40, 42, 43. Transverse fold on sternite 3 about
one-fourth width of sternite. Sternite 8 with anterior projection as in
figs. 49 and 61.

Male: Spatulate setae on protarsus as in fig. 22, three pairs on
first segment, two pairs each on second and third segments, and on
first two segments of mesotarsus as in fig. 39, three on first segment,
two on second, only on anterior half of mesotarsal segments; genital
segment and aedeagus as in figs. 66, 67, 84, 85.

Female: Genitalia as illustrated by Newton (1975, figs. 8-10).

Distribution: see map, fig. 50. (Includes records from the litera-
ture and those listed below.)

Types: Glypholoma pustuliferum Jeannel. Holotype (sex un-
known): CHILE: Aisen Province, Chile Chico [Museo Nacional de
Historia Natural, Santiago, Chile]; not seen. Paratype as listed by
Jeannel, 1962.

Lathrimaeodes pustulipenne Scheerpeltz. Holotype ( 6 ):
ARGENTINA: Rio Negro Province, El Bolson, Mt. Piltriquitron
[Hungarian Natural History Museum, Budapest, Hungary]; not
seen. Paratypes as listed by Scheerpeltz, 1972.

Material examined: CHILE: Malleco Prov.: 7 km W V. Portales,
1300 m, 23-24. XII. 1976, S. Peck (1) [SBP]; 15 km W V. Portales,
1650 m, 22-25.XII.1976, S. Peck (2) [SBP]; 15 km W Victoria, 200
m, 28-30. XII. 1976, S. Peck (2) [SBP]; 20 km E Manzanar (=30 km
E Curacautin, nr. Malalcahuello), 1100 m, 19-3 1 .XII. 1976, S. Peck
(106) [ANIC, ANMT, CNC, FMNH, MCZ, NMVM, SBP]; same
locality, 1300-1400 m, 19-21. XII. 1976, S. Peck (1) [SBP]; same
locality, no elev., 19-25. XII. 1976, S. Peck (16) [ANMT, CNC];
Talca Prov.: Alto Vilches, 1300 m, 10-13. XII. 1976, S. Peck (3)
[ANMT, SBP]; Nuble Prov.: Las Trancas, 70 km E Chilian, 1400 m,
13-17. XII. 1976 (some labeled 70 km SE Chilian 1300 m, 14-
17. XII), S. Peck (15) [ANMT, CNC, SBP]; Magallanes Prov.:
Punta Arenas, Feb. 1906, R. Thaxter (6) [MCZ]; Cautin Prov.: 22
km E Temuco, VJ-VII.1951, M. G. Smith (2) [CAS].

Habitat: Of the material examined, ecological data are available
for most of the specimens collected by Peck. Twenty-eight speci-
mens were collected at dung traps (three collections), 35 at carrion
traps (five collections), 60 from litter under carrion left for several
days (one collection), and one from mushrooms on a rotting stump
(with litter and moss; one collection). Most of these collections were

1978] Thayer & Newton — Revision of Genus Glypholoma 49

made in Nothofagus forest (also two specimens from Nothofagus-
Araucaria forest, one from Araucaria forest), and several of the
dung and carrion traps were set near streams (S. Peck, pers.
comm.). Scheerpeltz (1972), whose records are mainly small series,
reports this species (as Lathrimaeodes pustulipenne) primarily from
ground litter in forests of Nothofagus, Libocedrus, and other trees,
and also mentions one record from an old rabbit carcass in such a
forest. These beetles seem to live in forest litter, but are strongly
attracted to carrion and dung. Nothing is known about their feeding
habits.

Glypholoma temporale, new species

With the characters of the genus as described above.

Moderately convex dorsally, shape as in fig. 3, with central areas
of pronotum and elytra brown, their lateral margins and the elytral
apices yellowish, head and dorsum of abdomen between these two
colors, slightly reddish; venter of body darker than dorsum, legs
slightly paler than head. Head and pronotum with pale scattered
short fine setae, elytra glabrous between macrosetae, tibiae and tarsi
fairly densely setose. Length: 2.9 ± 0.1 mm; width: 1.3 ± 0.02 mm
(N = 6).

Head capsule as in fig. 57, epistomal suture angulate, lacking
median stem, temples and lateral nuchal constriction present.
Antennal length about 1.3 times head width, first four antennal
segments without dense pubescence, apical segments broader than
basal, but distinct club difficult to delimit (see fig. 53). Ocellar
diameter about 0.09 times head width. Labrum slightly emarginate
anteriorly. Maxillary palp with fourth segment about 2.5 times as
long as third, slightly wider at middle than third.

Pronotum as in fig. 3, with very shallow paramedian and lateral
depressions, lateral margins explanate. Prosternal surface finely
rugose, without median longitudinal carina. Mesosternum similar
to that of pustuliferum (figs. 34, 36), with microsculpture; cavities
for reception of procoxae about two-fifths the length of the
mesosternum, mesosternal process narrow, extending just over half
way between mesocoxae. Metasternum convex, lacking microsculp-
ture but with large punctures containing minute setae, antecoxal
sutures long, extending three-fourths the width of the metasternum

50

Psyche

[March

(fig. 59). Protibia setose, slightly fewer spines than in pustuliferum
(cf. fig. 23), meso- and metatibia with more spines than protibia.
Hind tarsus nearly seven-tenths as long as hind tibia, fifth segment
of hind tarsus half as long as preceding four together. Empodial
setae longer than tarsal claws.

Elytral striae impressed between punctures, intervals finely punc-
tate. Pustules (concolorous with surrounding area) present on
alternate intervals, anterior and adjacent to macrosetae. Angle
formed by elytral apices at suture slightly less than 180° (convex
posteriorly). Wings fully developed.

Abdomen with tergites 5 to 8 well-sclerotized, spiracles located in
tergites of these segments; paratergites clearly present and distinct
on segments 4 to 6, sutures between sternite and paratergites lacking
on segment 3 and (perhaps) posteriorly on segment 7. Tergites 4 and
5 with paired patches of medially-directed microtrichia, fairly
similar to those of pustuliferum (cf. fig. 40). Sternite 8 with anterior
projection as in fig. 63.

Male: Spatulate setae in pairs on protarsal segments 1-3, singly
on mesotarsal segments 1 and 2, similarly to fig. 39, but exact
number unknown; genital segment and aedeagus as in figs. 70, 71,
82, 83.

Distribution: see map (fig. 50).

Holotype: CHILE: Malleco Prov., 20 km E Manzanar, 19-
25. XII. 76 S. Peck, 1100 m, carrion + dung traps [MCZ].

Paratypes (2d, 3 9 ): Same locality and date as holotype; no
elevation or habitat data [ANMT, CNC, MCZ, SBP].

Habitat: In addition to the label data, the holotype was collected
in moist Nothofagus forest near a stream (S. B. Peck, pers. comm.).

Etymology: This species name refers to the well-developed tem-
ples on the head.

Glypholoma pecki, new species

With the characters of the genus as described above.

Moderately convex dorsally, shape as in fig. 2, unicolorous
yellowish-brown, except the following darker brown: spot on vertex
of head between ocelli, median longitudinal line on pronotum, last
three or four antennal segments, and third and fourth segments of

1978] Thayer & Newton — Revision of Genus Glypholoma 51

Fig. 50. Southern Chile and Argentina, showing locality records for South
American Glypholoma spp.: G. pustuliferum, dots; G. temporale, diamond; G.pecki,
square; G. tenuicorne, triangle.

52

Psyche

[March

maxillary palps. Dorsal surface of head, pronotum, and elytra
virtually glabrous except for elytral macrosetae. Length: 2.6 mm;
width: 1.2 ± 0.03 mm (only two specimens seen).

Head capsule similar to that of pustuliferum (fig. 58), but eyes
more prominent and epistomal suture lacking median stem. An-
tennal length about 1.3 times head width, first seven antennal
segments without dense pubescence, antennal club three-segmented,
apical segment four times as long as preceding segment (see fig. 52).
Ocellar diameter about 0.08 times head width. Labrum approxi-
mately rectangular (anterior margin straight). Maxillary palp with
fourth segment about 4.5 times as long as third, the two subequal in
width.

Pronotum as in fig. 2, with shallow median and lateral depres-
sions, lateral margins explanate. Prosternum coarsely punctate with
median longitudinal carina. Mesosternum with dense microsculp-
ture except on mesosternal process, cavities for reception of pro-
coxae about half the length of the mesosternum, mesosternal
process similar to that of pustuliferum (cf. figs. 34, 36), extending
about two-thirds of the way between mesocoxae. Metathorax
convex, without microsculpture, coarsely punctate with minute
setae in punctures; antecoxal sutures short, extending one-fifth to
one-fourth the width of the metasternum. Male protibia with a few
external spines near apex, meso- and metatibia with a few more
spines, female tibiae with fewer spines than male. Hind tarsus about
two-thirds as long as hind tibia, fifth segment of hind tarsus three-
fifths as long as first four segments together. Empodial setae
subequal in length to tarsal claws.

Elytral striae impressed between punctures, elytral intervals finely
sparsely punctate between macrosetae on alternate intervals. Elytra
without pale spots or raised pustules. Angle formed by elytral apices
at suture slightly greater than 180° (concave posteriorly). Wings
fully developed.

Abdomen with tergites 5 to 8 well-sclerotized, spiracles located in
tergites of these segments (possibly 4 also); sutures between sternite
and paratergites apparently lacking on segments 3 to 7. Tergites 4
and 5 with paired more or less rectangular patches of medially-
directed microtrichia, less extensive than those of pustuliferum (cf.
fig. 40). Sternite 8 with anterior projection as shown in fig. 62.

1978] Thayer & Newton — Revision of Genus Glypholoma 53

Male: Spatulate setae in pairs on protarsus, number unknown,
singly on first two or three segments of mesotarsus, number
unknown; genital segment and aedeagus as in figs. 68, 69, 80, 81.

Distribution: See map (fig. 50).

Holotype ( 6 ): CHILE: Malleco Prov., 20 km E Manzanar, 1100
m, 19-25. XII. 1976, S. Peck, malaise trap [MCZ].

Paratype ( 9 ): Same data as holotype [CNC].

Habitat: Both specimens were collected in a malaise trap at the
edge of moist Nothofagus forest (S. B. Peck, pers. comm.), but this
species was absent from dung and carrion traps at the same locality
which produced over 100 pustuliferum and one temporale.

Etymology: This species is named after Dr. Stewart B. Peck, who
collected both types as well as most of the other specimens used in
this study.

Glypholoma tenuicorne, new species

With the characters of the genus as described above.

Slightly convex dorsally, shape as in fig. 4, unicolorous light
reddish-brown, head and pronotum dorsally with a few scattered
setae, elytra glabrous except for macrosetae, tibiae and tarsi less
densely setose than in other species. Length: 3 mm; width: 1.2 mm
(one specimen seen).

Head capsule generally similar to that of pustuliferum (fig. 58),
but epistomal suture arcuate and lacking median stem. Antennal
length about 2.2 times head width, first four antennal segments
without dense pubescence, club lacking (see fig. 56). Ocellar di-
ameter about 0.11 times head width. Labrum rectangular (anterior
margin straight). Maxillary palp with fourth segment about 3.5
times as long as third, the two subequal in width.

Pronotum as in fig. 4, with shallow median and anterolateral
depressions, lateral margins acute anteriorly, explanate posteriorly.
Prosternum minutely rugose and sparsely punctate, without median
longitudinal carina; pleural-sternal articulation possibly absent.
Mesosternum with dense microsculpture, cavities for reception of
procoxae about two-fifths the length of the mesosternum, meso-
sternal process similar to that of pustuliferum (figs. 34, 36),
extending half the distance between mesocoxae; mesosternal-pleural

54

Psyche

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suture possibly complete. Metasternum convex, without micro-
sculpture, more finely punctate than pustuliferum (cf. fig. 34) and
with most of the posterior half impunctate; antecoxal sutures short,
extending one-fourth the width of metasternum. Protibia without
spines externally, meso- and metatibia with only a few. Hind tarsus
seven-tenths as long as hind tibia, fifth segment of hind tarsus one-
third as long as first four segments together. Empodial setae
subequal in length to tarsal claws.

Elytral striae not impressed between punctures, intervals impunc-
tate except for macrosetae. Elytra without pale spots or raised
pustules. Angle formed by elytral apices at suture slightly greater
than 180° (concave posteriorly). Wings fully developed.

Abdomen with tergites 4 to 8 well-sclerotized, spiracles in tergites
of segments 4 (or possibly 3) to 8; paratergites clearly present and
distinct on segments 4 to 6, sutures between sternite and paratergites
lacking on segment 3. (Structure of segment 7 uncertain.) Tergites 4
and 5 with paired patches of medially-directed microtrichia, patches
slightly narrowed laterally but still occupying most of the width and
(medially) the length of both tergites. Sternite 8 with anterior
projection as in fig. 64.

Male: Spatulate setae in pairs on protarsal segments 1-3, number
unknown, present singly on anterior half of first two segments of
mesotarsus, number unknown; genital segment and aedeagus as in
figs. 72, 73, 76, 77.

Female unknown.

Distribution: See map (fig. 50).

Holotype ( d ): CHILE: Valdivia Prov., Corral, Dec. 1905, R.
Thaxter [MCZ].

Etymology: The species name tenuicorne refers to the long
slender antennae, apparently unique within the genus.

Glypholoma rotundulum, new species

With the characters of the genus as described above.

Very convex dorsally, shape as in fig. 5, reddish-brown to dark
brown or occasionally almost black, elytra with small yellow spots
on alternate intervals, a slightly larger yellow spot at each humerus,
and usually a still larger subapical yellow patch on each elytron.
Elytra sometimes with light areas laterally, apically, or basally (or

1978] Thayer & Newton — Revision of Genus Glypholoma 55

Fig. 51. Victoria, Australia, with dots showing locality records for Glypholoma
rotundulum.

various combinations of these), with the normal spots obscured in
these regions. Some individuals with very small yellow spots on the
pronotal margins, the maximum being an anterior paramedial pair,
a lateral pair, a posterior paramedial pair, and a posterior lateral
pair. Dorsal surface of body with minute to short setae (as well as
macrosetae on elytra), tibiae and tarsi fairly densely setose. Length:
2.5 ± 0.2 mm (2. 1-3.0); width: 1.2 ± 0.1 mm (1.0-1. 3) (N = 18).

Head capsule similar to that of pustuliferum (fig. 58), but
epistomal suture slightly arcuate medially, with short median stem.
Antennal length about 1.2 times head width, first seven antennal
segments without dense pubescence, antennal club three-segmented,
slightly more abrupt than in pustuliferum (see fig. 55). Ocellar
diameter about 0.08 times head width. Labrum slightly emarginate
anteriorly. Maxillary palp with fourth segment about five times as
long as third, its inner side slightly swollen so that fourth segment at
middle is about half again as wide as third segment.

Pronotum as in fig. 5, evenly convex except for a broad shallow
depression near each posterior angle, lateral margins acute anteri-
orly, explanate posteriorly. Prosternum coarsely punctate, with
microsculpture and median longitudinal carina. Mesosternum (see
figs. 35, 37) with dense microsculpture except on the raised pen-
tagonal mesosternal process, cavities for reception of procoxae
about three-fourths the length of the mesosternum, mesosternal

56

P svc he

[March

process extending about four-ninths of the way between mesocoxae.
Metasternum convex, without microsculpture but with large punc-
tures containing very short setae, antecoxal sutures short, extending
only one-fifth the width of the metasternum. Protibia setose, slightly
less spinose than that of pustuliferum, meso- and metatibia slightly
more spinose than protibia. Hind tarsus slightly less than six-tenths
as long as hind tibia, fifth segment of hind tarsus half as long as first
four together. Empodial setae longer than tarsal claws.

Elytral striae impressed between punctures, intervals minutely
punctate and with very short setae, in addition to the usual
macrosetae on alternate intervals (see fig. 29). Pale spots (but not
pustules) anterior and adjacent to macrosetae. Angle formed by
elytral apices at suture slightly greater than 180° (concave poster-
iorly). Wings greatly reduced in nearly all specimens seen, only
extending over tergite 2, and not folded at rest (see figs. 37, 41), very
rarely fully or almost fully developed; anal flap present.

Abdomen with tergites 4 to 8 well-sclerotized, spiracles located in
tergites of segments 5 to 8, at edge of tergite 4; paratergites clearly
present and distinct on segments 4 to 6, sutures between sternite and
paratergites lacking on segment 3; segment 7 with sutures between
sternite and paratergites fading out posteriorly. Tergite 4 with small
paired patches of medially-directed microtrichia (see fig. 41), tergite
5 with or without even smaller transverse patches of same near
posterior edge. Transverse fold on sternite 3 about one-half width of
sternite. Sternite 8 with anterior projection as in fig. 65.

Male: Spatulate setae on protarsus as in fig. 25, five pairs on first
segment, two pairs each on second and third segments, on first two
segments of mesotarsus, similarly to fig. 39, but with five on the first
segment and two on the second; genital segment and aedeagus as in
figs. 74, 75, 78, 79.

Female: Genitalia very similar to those of G. pustuliferum, distal
gonocoxites relatively slightly thicker.

Distribution: see map (fig. 51).

Holotype ( d ): AUSTRALIA: Victoria: Belgrave, F. E. Wilson,
22—5—29 / Fallen leaves/ F. E. Wilson Collection [NMVM],

Paratypes: AUSTRALIA: Victoria: Warburton, F. E. Wilson, 4080
feet/in tussocks/ F. E. Wilson Collection (3d) [NMVM]; Mt. Baw
Baw, summit, etc.. Mar. ’58, Darlington (Id) [MCZ]; Warburton,
Mt. Donna Buang, 1200 m, IV.28-V.7. 1978, carrion traps, Notho-
fagus forest, S. B. Peck (79 d , 104 9 ) [ANIC, ANMT, CAS, CNC,

Figs. 52 60. Glypholoma spp. 52-56. Right antenna, dorsal view; 52, G. pecki;
53, G. temporale; 54, G. pustuliferum; 55, G. rotundulum; 56, G. tenuicorne. 57-58.
Head, dorsal view; 57, G. temporale; 58. G. pustuliferum. 59-60. Metacoxae and
metasternal antecoxal sutures; 59, G. temporale; 60, G. pustuliferum. Scale lines =
0.5 mm; upper one applies to figs. 52-58, lower one to figs. 59-60.

Figs. 61-75. Glypholoma spp. 61-65. Abdominal sternite 8, ventral view; 61,
G. pustuliferum; 62, G. pecki; 63, G. temporale; 64, G. tenuicorne; 65, G. rotundulum.
66-75. Male genital segment, ventral and dorsal views, respectively; 66-67, G.
pustuliferum; 68-69, G. pecki; 70-71, G. temporale; 72-73, G. tenuicorne; 74-75, G.
rotundulum. b = internal cylindrical “button”; s9 = sternite 9; t9 = tergite 9; tlO =
tergite 10. Scale line = 0.5 mm.

1978] Thayer & Newton — Revision of Genus Glypholoma 59

Figs. 76-85. Glypholoma spp., aedeagus (in situ), dorsal and right lateral views,
respectively. 76-77, G. tenuicorne; 78-79, G. rotundulum; 80-81, G.pecki; 82-83, G.
temporale; 84-85, G. pustuliferum. bp = basal piece; mf = median foramen; ml =
median lobe; p = paramere. Scale line = 0.5 mm.

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FMNH, MCZ, NMVM, SBP]; Warburton, Acheron Gap, 750 m,
VI. 27-30. 1978, Ber.u.log bark Euc. + Notho., S. B. Peck (2 d , 6 9 )
[SBP]; same locality, IV.28-V.7.1978, carrion trap, Nothofagus
forest, S. B. Peck (4 d ) [ANMT]; Tara Valley N.P., 450 m,
V. 10-17. 1978, carrion traps, S.&J. Peck(4d ,89 ) [ANMT]; Bulga
N.P., 550 m, V. 10-17. 1978, carrion traps in Nothofagus ravine, S. &
J. Peck (11 d , 12 9) [ANIC, ANMT, CNC, SBP]; same locality,
V. 17. 1978 Ber. fungi on logs and stumps, S. & J. Peck (16,59)
[SBP]; Mt. Buffalo N.P., 1300 m, IV.22-26.1978 carrion trap, S. &
J. Peck (2d) [ANMT]; same locality, 500 m, IV. 24-27. 1978, wet
sclerophyll forest, creek: carrion, S. B. Peck (19) [MCZ]; Victoria?
/ex C. Oke Collection, no locality (2d, 2 9) [NMVM],

Habitat: Ecological data are available for all but five of the 252
specimens of this species seen. Of these, 225 specimens were
collected in carrion traps or at carrion (six collections), twelve from
fungi on logs and stumps (one collection), eight under bark of
Eucalyptus and Nothofagus logs (one collection), three in tussocks
(one collection), and one among fallen leaves (presumably loose
ground litter). Most of the specimens were collected in Nothofagus
forest, and in general Glypholoma rotundulum seems to occur in
habitats similar to those of G. pustuliferum, but nothing is known
about the feeding habits of this species, either. These beetles seem to
be notably cold-adapted. The largest collection (183 specimens) was
taken in carrion traps set in snow-covered ground (S. B. Peck, pers.
comm.), across which the flightless beetles must have walked.

Wing polymorphism: All specimens examined are brachypterous
except the three from Mt. Buffalo, the northernmost recorded
locality for the species. One of these specimens (d) has fully-
developed wings and the other two ( d9 ) have wings sharply
reduced beyond the costal hinge.

Etymology: The species name rotundulum (diminutive of rotun-
dum, round) refers to the great convexity and small body size of the
members of this species.

Discussion

We believe that the five species here included in Glypholoma form
a monophyletic group (in the strict, Hennigian sense) which can be
defined on the basis of these characters which we presume to be
uniquely derived or synapomorphic within Omaliinae: excavate

1978] Thayer & Newton — Revision of Genus Glypholoma 61

hind coxae with “retractile” hind femora, each elytron with eleven
more or less distinct striae, and male genital segment with a small
“button” internally near the anterior end of sternite 9. Some of the
species here included in Glypholoma have characteristics, such as
the filiform antenna of tenuicorne and mesosternal structure of
rotundulum, that might conventionally be used to justify the
erection of new genera. Because the south temperate omaliine fauna
is relatively poorly known, we feel that the use of a conservative
generic concept is best at present, since this will possibly reduce the
need for nomenclatural changes at the generic level as the full range
of variation within the south temperate fauna becomes better
known. A need for the creation of more genera may become
apparent eventually; judgment of this is best reserved until at least a
large proportion of the fauna (rather than the present fragments) is
known.

Although not enough information is available to allow formula-
tion of a complete phylogenetic hypothesis, a few comments can be
made concerning probable relationships among the species of
Glypholoma. Each species can be defined by an autapomorphous
trait, as follows: tenuicorne, elytral striae not impressed between
punctures; rotundulum, raised pentagonal mesosternal process;
pecki, elongate antennal segment 11; temporale, bulging temples;
and pustuliferum, coarsely punctured and rugose prosternum. All
species but tenuicorne have clavate antennae (instead of filiform,
which is presumably more primitive within Staphylinids) in com-
mon, and within this subgroup, pustuliferum and temporale share
the apparently derived condition of raised pustules on the elytral
intervals. No synapomorphies have been discovered yet which
clarify the relationships among these four species any further. The
availability of more material for detailed study of the three new
South American species described in the present paper, and the
discovery of immature stages or biological information for any
species, might allow the discovery of additional synapomorphies
and the development of a complete phylogenetic hypothesis.

The position of Glypholoma within the Omaliinae is somewhat
ambiguous, largely because relationships within the subfamily as a
whole are not very well worked out, especially the extra-Holarctic
representatives of the subfamily. A majority of the characters
distinguishing Glypholoma from most or all other Omaliinae are
characters exhibiting plesiomorphic states in Glypholoma. This

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Psyche

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suggests, at least, that Glypholoma is a relatively primitive member
of the subfamily and branched off early in the evolution of the
Omaliinae. The close relationship of Glypholoma to the Holarctic
Anthobium-group (viz., Anthobium, Camioleum, Deliphrum, Ma-
thrilaeum, Olophrum ) suggested by Scheerpeltz (1972) and Newton
(1975) mainly on the basis of general habitus does not seem to be
supported by more detailed evidence gathered to date. There are
several characters for which the members of the Anthobium-group
share what we believe are derived character states within the Oma-
liinae, and Glypholoma shows what we believe to be primitive char-
acter states. These characters appear in the Anthobium-group as
follows: presence of a sharp post-ocular ridge, absence of the episto-
mal suture, gular sutures very close together or partly fused, presence
of lateral foveae on the pronotum with internal pillars connecting
dorsal and ventral surfaces of pronotum, and female genitalia with
the proximal gonocoxites fused. We feel that this assemblage of dif-
ferences constitutes strong evidence against a close relationship be-
tween Glypholoma and the Anthobium- group. Unfortunately any
more precise statement concerning the placement and relationships
of Glypholoma must await future clarification of the evolution and
higher classification of the Omaliinae as a whole. In the meantime,
tarsal, palpal, and mandibular characters dictate placement of
Glypholoma in the heterogeneous tribe Anthophagini. If a family-
group name is eventually needed for Glypholoma, the name Gly-
pholomini Jeannel 1962 is available.

The presence of two of the species of Glypholoma (pustuliferum
and rotundulum) is strongly correlated with cool moist temperate
forests dominated by Nothofagus; the other three species have been
collected only within the Nothofagus zone of southern South
America. The fact that each of the latter species is known only from
a single small collection suggests that they may differ significantly
from rotundulum and pustuliferum in their biology or ecological
preferences. The disjunct southern temperate distribution of the
genus is similar to those of some other isolated and apparently
archaic groups of staphylinoids, some of whose ranges include New
Zealand as well as southern South America and Australia. The
occurrence of Glypholoma in New Zealand Nothofagus forests
would not be wholly unexpected.

1978] Thayer & Newton — Revision of Genus Glypholoma 63

References Cited

Jeannel, R.

1962. Les Silphidae, Leiodidae, Camiaridae et Catopidae de la Paleantarctide
Occidentale. Biol, de l’Amer. Aust. 1: 481-525.

Klinger, R. and U. Maschwitz

1977. The defensive gland of Omaliinae (Coleoptera: Staphylinidae). I. Gross
morphology of the gland and identification of the scent of Eusphalerum
longipenne Erichson. J. Chem. Ecol. 3(4): 401-410.

Newton, A. F.

1975. The systematic position of Glypholoma Jeannel, with a new synonymy
(Coleoptera: Silphidae, Staphylinidae). Psyche 82(1): 53-58.

SCHEERPELTZ, O.

1972. Wissenschaftliche Ergebnisse der Studienreise von Gy. Topal nach
Sudwest-Argentinien (Coleoptera: Staphylinidae). Folia Ent. Hung. 25,
Suppl., 268 pp., 5 pi.

SZYMCZAKOWSKI, W.

1976. Silphidae, Leiodidae, Catopidae et Colonidae (Coleoptera) du Parc
National du Nahuel Huapi en Argentine. Polskie Pismo Entom. 46:
423-438.

AGGREGATIONS OF MALLOS AND DICTYNA
(ARANEAE, DICTYNIDAE):
POPULATION CHARACTERISTICS

By Robert R. Jackson1 and Sandra E. Smith
North Carolina Division of Mental Health Services
Research Section
P.O. Box 7532

Raleigh, North Carolina 27611
Introduction

Generally web spiders are solitary animals. Usually only a single
spider occupies a single web, with the relatively common exception
of webs jointly occupied by a male-female pair or females with their
young. In most species, occupied webs are usually not connected to
other occupied webs by silk lines. However, aggregations of various
types and degrees occur in some species (reviewed by Buskirk, 1975;
Krafft, 1970; and Shear, 1970). These include cases in which
occupied webs are in close proximity (sometimes touching), but
occupied by single spiders, as well as cases in which single webs are
occupied by groups of spiders of all sex/age classes. The species
involved are sometimes referred to as “social spiders” (e.g., Burgess,
1978; Kullmann, 1972). At present relatively little is known about
the social adaptations of these spiders. Some of the first things we
would like to know about animals that occur in aggregations are the
size, composition and spacing characteristics of naturally occurring
groups. Data concerning these basic social characteristics will be
presented in this paper for several species of dictynid spiders.

The Dictynidae are a group of web-building cribellate spiders,
most of which have body lengths less than 5 mm. In the closely
related genera Mallos and Dictyna, there are species with different
types of social organization. Most are solitary; at least three are
communal, territorial; and at least one is communal, non-territorial.
Each individual web of a solitary species consists of a mesh in which
prey is captured and a nest within the mesh. Communal, territorial

Present address:

Department of Zoology, University of Canterbury, Christchurch 1, New Zealand.
Manuscript received by the editor June 10, 1978.

65

66

Psyche

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species (M. trivittatus Banks, Dictyna calcarata Banks, D. albopilosa
Franganillo) live in web complexes. Web complexes consist of
variable numbers of web units, each consisting of a nest and a mesh;
and web units are connected to each other by strands of silk in the
interstitial web. Evidently, web units are treated by the occupants as
defended territories (Jackson, 1978a). In the communal, non-
territorial species ( M . gregalis Simon), groups of spiders live in
communal webs which consist of surface sheets with holes leading
into the interior. The interior contains nests, tunnels and supporting
lines connecting the sheet with the substrate beneath (see Burgess,
1976; Diguet, 1909a, 1915). Communal webs are not divided into
defended units as in web complexes. More detailed information
concerning social organization, web characteristics, and habitats is
provided elsewhere (Jackson, 1978a).

Methods

DATA FROM NATURAL POPULATIONS

For census areas, locations were chosen which seemed to have
particularly high densities of spiders. The census areas for Mallos
niveus O. P. Cambridge were one-dimensional (horizontal lines).
All other census areas were rectangular. For D. calcarata and
Censuses 3-6 for M. trivittatus, complete web complexes of relatively
small size were used; and the boundaries of the census areas were
chosen to closely correspond to the edges of the web complexes. The
distance from each spider in the complex to the nearest conspecific
in the vicinity but not in the complex was greater than twice the
diameter of the complex in each of these census areas. For Census
No. 1 and 2 for M. trivittatus, each area censused was part of a large
web complex.

The corners of each census area were marked with pieces of tape.
Each web or web unit was completely searched, a process that
destroyed the web unit. The occupants of the units were recorded;
and a piece of tape, with a code number, was placed at the location
of each occupied nest before it was destroyed. The distance of each
of these nests to its nearest neighboring nest was measured.

The exact locations of spiders on the sheet and interstitial webs
proved excessively difficult to measure accurately. This was because
they were frequently obscured from view by silk and debris; and the
process of locating spiders inevitably disturbed them, leading to

1978]

Jackson & Smith — Mallos and Dictyna

67

changes in their positions before they could be recorded. Only the
component of the web (nest, mesh, interstitial web) occupied by the
spider could be recorded with confidence. When counting the
number of spiders per web unit, those in the interstitial web were
included in the record of the web unit to which they were in closest
proximity.

Spacing and density data were not collected for D. albopilosa.
The web complexes of this species were particularly difficult to
disassemble in the field because they were three-dimensional and
located on the leaves, stems and roots in dense growths of herbaceous
plants. Those of D. calcarata and M. trivittatus, in contrast, were
two-dimensional and usually built on more accessible flat surfaces.

At San Anton Falls (Morelos, Mexico), 23 occupied web units of
D. albopilosa were searched, and the occupants were recorded, with
no attempt to assess density or spatial relations. In addition to the
spiders in the census areas, the occupants of another 48 web units of
D. calcarata and 152 of M. trivittatus were recorded. Each of these
was in the general vicinity of the census areas.

Data for M. trivittatus were collected in June; those for M.
niveus, D. calcarata, and D. albopilosa were collected in September.
These species seem to feed predominantly in the late afternoon and
early evening (Jackson, 1977a). Data in this study were collected at
mid-day, when the spiders were less active.

LABORATORY STUDIES — Mallos gregalis

In the laboratory, M. gregalis in large communal webs were
maintained on a diet of houseflies (Musca domestica), provided at
approximately 5-day intervals. Temperature was maintained at
approximately 24° C, and the light cycle was approximately 13 L:
11D. These large colonies were begun from fewer than 200 spiders
collected by Burgess (see Burgess, 1976) in Guadalajara, Mexico, 3
yr previous. The webs in the laboratory were on plants and other
objects, and they were not enclosed.

Two census methods were used. 1. Small discrete communal webs
and relatively discrete portions of larger communal webs were
completely disassembled in the spring and early summer. All occu-
pants were recorded; and the condition, structure, and size of the
webs were recorded. 2. The surface of a large communal web, at
least 2 years old, was censused with 30cm transects in the fall. Each
transect consisted of a string laid haphazardly onto the web, and all

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spiders within 1 mm of the string were counted. The web was
arbitrarily divided into 3 sections. Two transects were placed onto
each section on each census-day; and there were 1 1 census days,
each 2 to 3 days apart. A “census” was defined as the sum of the
spiders found on the two transects on a given section on a given
census-day.

Census Areas

Mallos niveus

Three aggregations of M. niveus were censused in Guanajaato
(Guanajaato, Mexico), each under a different window sill on
exterior walls of buildings in the city. The webs were arranged in
horizontal straight lines, and the length of the aggregation was
defined as the distance between the first web at one end and the last
web at the other end (Census Area No. 1, 54 cm; No. 2, 87 cm; No.
3, 273 cm). The distance to the nearest conspecific not in the
aggregation was greater than length of the aggregation in each case.

Mallos trivittatus

Census Areas Nos. 1, 2, and 3 were at East Turkey Creek in the
Chiracahua Mountains (Arizona, USA). The creek passes through a
1.8 m diameter metal culvert, with an interior surface area of 88m2.
Almost the entire interior of the culvert was covered by a single
enormous web complex. The bottom of the culvert was covered by
water; and in the immediate vicinity of the water, web units were
scarce. Subtracting these parts of the culvert, the web complex was
conservatively estimated to be 79 m2 in area. Census Areas Nos. 1
and 2 were each inside the culvert, within 2 m of opposite entrances
and 1.5 m above the bottom of the culvert. Census Area No. 1 was
56 cm X 50 cm; No. 2, 60 cm X 56 cm. Census Area No. 3 was a web
complex (54 cm X 42 cm) 30 m from the culvert, on a large boulder
(ca. 2 m high) beside the creek. The boulder was overhanging the
ground, forming a shallow cave (ca. 1 m from ground to top; 0.5 m
from entrance to back). The web complex was on the upper surface
of the cave.

Census Area No. 4 (56 cm X 32 cm) and No. 5 (56 cm X 56 cm)
were web complexes on rock ledges near Estes Park (Colorado,
USA). Each was 1 to 2 m above the ground. Census Area No. 6 was
a web complex (124 cm X 34 cm) on the upper surface of a shallow

1978]

Jackson & Smith — Mallos and Dictyna

69

cave (ca. 1 m high and 1 m deep) made by an overhanging rock ledge
in Big Thompson Canyon (Colorado, USA).

Dictyna calcarata

Census Area No. 1 (24 cm X 24 cm) and No. 2 (54 cm X 46 cm)
were web complexes on the external walls of buildings in Chapala
(Jalisco, Mexico).

Results

POPULATION SIZE AND DENSITY

Densities for M. trivittatus and D. calcarata were relatively
similar (Table 1). Since the M. niveus aggregations were linear (one-
dimensional) it is problematic to compare densities in this species
with the densities found in the other species. The webs of M. niveus
in the census areas were oriented such that the shorter distance
across each was vertical. The areas given in Table 1 were obtained
by multiplying the length of the aggregations by 3 cm, the mean
widths of M. niveus webs found on buildings (Jackson, 1978a). M.
gregalis tends to live in higher densities. The webs censused by
Method No. 1 (Table 2) had a mean density of 0.5 ± 0.66 spiders per
cm2 and 0.2 ± 0.14 per cm3.

Based on many hours spent in the culvert at E. Turkey Creek, it
seems that the densities obtained in Census Nos. 1 and 2 for M.
trivittatus were representative of the large complex in the culvert.
Using the mean of the two densities for spiders (14.23 per 1000 cm2),
webs (9.79 per 1000 cm2), and occupied webs (8.79 per 1000 cm2) as
estimates for the densities for the complex as a whole, this web
complex contained 10,200 spiders, 7,700 webs, and 6,500 occupied
webs. This was by far the largest web complex of this or any other
species in this study.

Web units of M. trivittatus tend to be 5 cm X 4 cm; those of D.
calcarata tend to be 3 cm X 2 cm; and webs of M. niveus on walls of
buildings tend to be 4 cm X 3 cm (Jackson, 1978a). In the culvert on
E. Turkey Creek, the interstitial area between web units tended to be
quite small, since the mean for nearest neighbor distance was not
much greater than the diameter of web units (Table 1). In all other
censused web complexes, web units tended to be 2 to 4 web unit
diameters apart; and aggregated M. niveus webs tended to be 2 to 4
web diameters apart also.

70

Psyche

[March

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Jackson & Smith — Mallos and Dictyna

71

Some of the larger communal webs of M. gregalis in the
laboratory had surface areas and volumes estimated to be as large as
1300 cm2 and 1500 cm3. If the densities in these large webs were
comparable to those in smaller webs, they can be estimated to have
contained 300 to 700 spiders. Based on the descriptions of Diguet
(1909a, b) and Burgess (1976), 4 m2 will be used as an approximate
estimate for the surface area of larger communal webs on trees in
Mexico. If densities in nature are comparable to those in the
laboratory then these very large webs may have contained as many
as 20,000 spiders.

WEB OCCUPANTS

Solitary species, including M. niveus , usually occurred one spider
per web (Jackson, 1978a), and this was true of all censused webs in
the aggregations of M. niveus. However, each occupied web unit of
the three communal, territorial species usually contained a set of
several spiders. These could be almost any combination of females,
males, and immatures of varied sizes (Table 3), except that web
units were never shared by two adults of the same sex. In the only
instances in which male-male or female-female pairs were seen
together in the same web units, aggressive interactions took place,
after which one individual departed. In contrast, large numbers of
M. gregalis belonging to all sex/ age classes shared the same
communal webs without aggressive behavior or cannibalism.

DISTRIBUTION OF WEB OCCUPANTS

Large proportions of each sex/ age class of M. trivittatus occupied
the nests (Tables 3 and 4); but adult females were especially prone to
be inside nests, and there was a preponderance of immatures in the
interstitial web. These data were consistent with more casual
observations of M. trivittatus in other locations and of the other
communal, territorial species. During the day, M. niveus and other
solitary species were most often, but not always, inside nests. All M.
niveus in censuses were inside nests.

Many hours of casual observation in the laboratory gave the
impression that adult males and immatures of M. gregalis were
more prone to be on the outer surfaces of webs, and females were
more prone to be in the interior. (Data related to this will be
presented below.)

72

Psyche

[March

Table 2. Data from complete disassembly of 19 M. gregalis webs or subunits of
webs (census Method No. 1, see text). Area and volume are estimates, since borders
of webs were irregular. Small immature: ca. 1 mm in body length (color: uniformly
brown). Large immature: body length ca. 2 mm (gray markings present on body).
Adult females: ca. 3 mm. Adult males: ca. 3 mm. Web class A: 3-D and lacking
evidence of deserted areas. B: 3-D and containing conspicuous deserted areas.
C: 2-D and lacking conspicuous deserted areas. D: 2-D (but not as flat as C) and
containing conspicuous deserted areas. E: 3-D, small discrete web on irregular
substrate. A-D: census areas were subunits of large webs, except for some of C
that were discrete webs. 3-D: web occupies relatively much space in each of 3 dimen-
sions. 2-D: web confined predominately to 2 dimensions (“flat”). “Deserted” areas:
no longer occupied by M. gregalis, not substantially adhesive, and covered by fly
carcasses and dust. Note: as population size increases, composition of population
shifts toward decreasing percentages of females and increasing percentages of im-
matures. Kendall rank correlation (Sokal and Rohlf, 1969) for increasing percent-
ages of small immatures with increasing population size (P < 0.001).

Popu-

Area

Volume

lation

(cm2)

(cm3)

1

144

144

2

144

144

3

105

300

4

150

300

5

144

144

6

75

150

7

70

150

8

70

350

9

150

150

10

70

350

11

144

144

12

70

350

13

64

320

14

150

600

15

155

155

16

100

300

17

400

800

18

45

225

19

150

600

Sex/

Web

Popu-

lation

Adult

Class

Size

Female

C

5

60.0

C

7

71.4

c

9

0.0

D

10

90.0

D

10

30.0

B

11

27.3

B

21

19.0

A

23

65.2

B

25

4.0

A

27

48.2

C

31

12.9

E

34

11.7

B

40

27.5

A

47

23.4

D

53

32.1

A

74

17.5

A

82

9.7

A

114

7.0

A

288

9.3

Age Class Composition
(%) of Population

Large

Small

Adult

Imma-

Imma-

Male

ture

ture

0.0

20.0

20.0

0.0

14.2

14.2

11.1

44.4

44.4

10.0

0.0

0.0

0.0

10.0

60.0

0.0

27.3

45.4

0.0

19.0

61.0

17.4

17.4

0.0

4.0

8.0

84.0

18.5

7.4

25.9

0.0

0.0

87.0

5.8

32.3

50.0

0.0

40.0

32.5

6.3

21.3

48.9

0.0

20.7

47.2

1.3

22.2

55.4

2.4

7.3

80.4

0.0

12.3

80.4

2.0

9.7

78.8

1978]

Jackson & Smith — Mallos and Dictyna

73

SEX/AGE CLASS RATIOS

Based on the spiders in the two censuses in the culvert at E.
Turkey Creek, this large web complex was estimated to have had a
population consisting of 27% adult females, 9% adult males, and
64% immatures. The adult sex ratio (Male: Female) was approxi-
mately 1:3. Based on the censuses at Chapala, populations of D.
calcar ata in web complexes on buildings were composed of 14%
adult females, 5% adult males, and 81% immatures; and the adult
sex ratio was 1:3, as for M. trivittatus. No males were found in the
censuses of M. niveus. Generally only small numbers of males of the
solitary species were found, and these were predominantly ones in
webs with females (Jackson, 1977b).

Based on the 19 censuses of M. gregalis by Method No. 1
(disassembly), population composition was 29.8 ± 25.49% adult
females, 4.1 ± 5.97% adult males, 17.6 ± 12.17% large immatures,
and 48.2 ± 27.41% small immatures. Immatures comprised a greater
proportion of the spiders in webs with large population size (Table
2). The average adult sex ratio was approximately 1:7. For the 33
transect censuses (Method No. 2), population composition was 12.3
± 7.44% adult females, 3.0 ± 3.85% adult males, 38.0 ± 12.96% large
immatures, and 46.2 ± 13.11% small immatures (adult sex ratio, ca.
1:4), with a total of 16.8 ± 2.34 spiders occurring in each census. The
three sections of the web were comparable.

Discussion

AGGREGATION SIZE AND DENSITY —

EVOLUTIONARY CONSIDERATIONS

The occurrence in certain areas of very large populations of some
species, especially M. gregalis and M. trivittatus, and the great
densities of spiders within these populations contrast markedly with
the majority of Dictyna and Mallos species. Most species are
solitary, with individuals living in relatively widely spaced indi-
vidual webs on stems and leaves of vegetation (Jackson, 1978a); and
although density censuses were not carried out for these, our
impression is that an area of 50 to 100 m2 in a population of a
solitary species would usually contain fewer than a dozen indi-
viduals. In contrast, the largest web complex of M. trivittatus
occupied approximately this much space and contained more than
10,000 individuals.

74

Psyche

[March

A related comparison can be made by looking at the amount of
web area or volume per individual spider for different species. There
is a trend toward decreasing web surface per spider as one goes from
solitary to communal species. Individual webs of solitary species
tend to have a surface area of 35 cm2 (Jackson, 1978a). Since each
web unit in web complexes of communal, territorial species tends to
contain 2 or 3 spiders (Table 3), and since web units tend to have a
surface area of 4 to 21 cm2 (Jackson, 1978a), there tends to be 1 to 7
cm2 of surface area per spider in these species. In the laboratory, in
communal webs of M. gregalis, there tends to be a surface area of
2.1 cm2 and a volume of 6.4 cm3 of web per spider. (Volume was not
calculated for other species since their webs tended to be largely 2-
dimensional; Jackson, 1978a.)

The influence of prey availability on these trends deserves atten-
tion. Diptera of the same approximate size relative to the spider
seem to predominate in the diet of species of Dictyna and Mallos

Table 3. Percentages of web units (in web complexes) occupied by different sex/
age class combinations. Data based on nests alone ( M . trivittatus ) given in paren-
theses. More than one immature sometimes occurred together in same web unit.
More than one adult of same sex never occurred together. N: No. of web units
sampled, including ones in Table 1 plus others (ones not occupied by dictynids
excluded).

M. trivittatus

D. calcarata

D. albopilosa

Females'

33.33 (34.15)

30.30

39.13

Males'

7.58 (4.07)

24.24

21.74

Immatures'

84.09 (75.61)

77.24

65.22

Female Only

14.39 (20.33)

7.58

21.74

Male Only

0.76 (1.63)

9.09

8.70

Immatures Only

61.36 (61.79)

56.06

47.83

Female plus Immatures Only

16.67 (13.82)

12.12

8.70

Female plus Male Only

0.76 (0.00)

9.09

8.70

Male plus Immatures Only

4.55 (2.44)

7.58

8.70

Female, Male, and Immatures

1.52 (0.00)

1.52

0.00

No. of Spiders per Web Unit2

3.1 ± 4.83 (20)

1.6 ± 0.91 (4)3

1.3 ± 0.47 (2)3

N

132 (123)

66

23

'Regardless of other sex/ age classes present.

2Mean ± S.D. (maximum).

3Larger maxima were seen during more casual observations.

1978]

Jackson & Smith — Mallos and Dictyna

75

(Jackson, 1977a). Thus large prey populations would seem neces-
sary to support some of the larger aggregations of dictynids. Diguet
(1909a, b, 1915) and Burgess (pers. comm.) noted numerous adult
Diptera in the vicinity of M. gregalis webs in Mexico during the
rainy season. Studies are needed to determine whether Diptera
populations are larger and/or more predictable in habitats con-
taining communal webs of M. gregalis compared to neighboring
areas without communal webs. Great numbers of Diptera were
noted in the metal culvert in Arizona that contained the enormous
web complex of M. trivittatus. Another consideration in this case is
that flies entering the culvert may be especially vulnerable to
capture, since inside the culvert they were almost completely
surrounded by web. Great masses of nematocerous flies were active
in the vicinity of web complexes of D. calcarata in Chapala,
emerging from nearby Lake Chapala in the late afternoon and early
evening. Diptera were also numerous in the vicinity of web com-
plexes of D. albopilosa at San Anton Falls. In general, wherever
there were web complexes, there were also numerous Diptera.

However, the question of why some species live in large, dense
populations, while others do not, cannot be answered simply on the
basis of prey densities in the habitats of different species. Riechert
and Tracy (1975) have shown relationships between density within
spider populations and prey availability. However, they found prey
availability to depend not only on the absolute abundance of prey
but also on factors that influence how the spider experiences prey
abundance, especially its thermal relations with its environment.

Another consideration is that species of Dictyna and Mallos with
differing aggregating tendencies may occur side-by-side in the same
habitats (Jackson, 1978a). For example, M. niveus were found in
individual webs on the same trees with web complexes of M.
trivittatus in the Chiracahua Mountains. It would seem that dictynid
species that routinely occur in aggregated states (communal, terri-
torial and communal, non-territorial) and those that generally live
more widely dispersed (solitary) are somehow adapted to exploit
different sets of resources, but we have no clear insights at this time
concerning what these different resources might be.

The adaptive advantages and disadvantages for animals related to
living in groups have been subjects of considerable interest in recent
years (see Wilson, 1975). For a review of ideas concerning spiders,

76

Psyche

[March

Table 4. Distribution of sex/ age classes of M. trivittatus within web complexes.
Based on occupied web units. No. of spiders: means ± S.D. Females, Immatures,
Males: percentages of the total found for each that were in each part of the web
complex. Total: Number of spiders — mean ± S.D. for spiders — per web unit
(spiders in interstitial web included with nearest web unit); Females, Immatures,
Males — percentage of total number of spiders belonging to each sex/ age class.

Number of
Spiders

Females

Males

Immatures

Inside Nest

1.5 ± 1.88

93.33

50.00

52.49

Inside Mesh

Less than 2 cm from Nest

0.1 ± 0.47

6.67

30.00

4.32

More than 7 cm from Nest

0.1 ± 0.38

0.00

20.00

5.98

Inside Interstitial Web

0.9 ± 2.19

0.00

0.00

37.21

Total

3.1 ± 4.83

12.64

2.81

84.55

see Buskirk (1975) and Lubin (1974). With the dictynids, we need to
compare the varied species with respect to the importance of each
potential advantage and disadvantage, given the nature of the
resources each species exploits.

PHENOLOGY

Most Dictyna and Mallos probably have an annual life cycle in
nature, with adult females and males present during spring, summer
and/or fall (Chamberlin and Gertsch, 1958). The mating season for
Mallos species generally seems to be later than that of Dictyna
species, although the season for some species may last many
months. Since both adult males and adult females were found,
evidently each census was carried out during the mating season of
the species involved, although not necessarily during the peak of the
season for all species. No doubt, if censuses had been undertaken at
different times of the year, different ratios of each sex/ age class
would have been found for each species.

Seasonal changes in the ratios of the different sex/ age classes in
laboratory populations of M. gregalis apparently occur, although
data have not been collected. Each sex/age class was found
throughout the year, but males were less numerous in winter than in
other seasons. The differences in adult sex ratios in the two types of
censuses of M. gregalis may be a reflection of the tendency, noted
during casual observation, for females to predominate in the
interior of webs, with males predominating on the exterior. How-

1978]

Jackson & Smith — M alios and Dictyna

77

ever, another factor that must be considered is that the two types of
censuses were made at different times of the year, and differences
may be influenced by phenology.

CAUSES OF DISTRIBUTION PATTERNS WITHIN WEBS

In all species, eggs tend to be oviposited in the vicinity of the
nests, and the tendency for females to be found in nests is probably
at least partly related to this. Also, nests may be optimal resting sites
with respect to protection from predators and parasites (Jackson,
1978a). If obtaining space within a nest is accompanied by aggres-
sive behavior, which seems likely in the communal, territorial
species, then females may have an advantage related to their larger
size. Females in each species have the largest body size of any
sex/ age class. Compared to males, females may be more sedentary;
and a stronger tendency to occupy nests may be related to this. This
might also apply to comparison of females with immatures if the
immatures are the dispersal phase in the life histories of these spe-
cies. Also, by virtue of their smaller size, young immatures may be
safer from predation on the mesh or interstitial web than females,
since they might take refuge under strands of silk, particles of
debris, etc., which are too small to be effective for females. As a
result, there might be lesser selection pressure against immatures
that remain outside nests compared to larger females that remain
outside nests.

Since females seem to be prone to be in the interior of webs of M.
gregalis, we might expect males to spend considerable time search-
ing for and courting females in the interior of webs. Instead, more
males seem to be on the exterior surfaces of the webs. In M. gregalis,
unlike M. trivittatus and D. calcarata, the presence of females and
silk spun by females are not releasers of courtship behavior. Instead,
males seem to have an advertising routine, as part of their daily
activity pattern, in which they perform behavior referred to as
“pluck-walking.” Females seem to indicate their receptivity to
pluck-walking males by failing to run away (Jackson, 1978b). We do
not know the factors that determine female receptivity, but perhaps
receptive females are more likely to be on the outer surfaces of webs.
For example, females might be unreceptive near the time of
oviposition, and oviposition takes place in the interior of the web.
Although the difficulty of observing behavior of spiders in the
interior of webs should be kept in mind, it is of interest that the

78

Psyche

[March

majority of observed instances of pluck-walking involved males on
the surfaces of webs.

We have no information at this time concerning how new
communal webs are formed by M. gregalis. One possibility is that
new webs are founded by single adult females or by groups of
several adult females. In the laboratory web censuses (method 1),
the webs with fewer spiders (predominantly adult females) may
have been ones that were newly founded. Those with more spiders
and a greater proportion of smaller immatures may have been older
webs in a period of rapid growth. Further study, especially in the
natural habitats of these spiders, is necessary. Information is also
lacking on the formation of web complexes by M.trivittatus and D.
calcarata.

CAUSES OF SKEWED SEX RATIOS

The causes of the relative rarity of males in all species investigated
are unknown. We do not know the sex ratios at hatching. If maternal
investment in progeny of the two sexes is equal, we would expect a
1:1 ratio (Fisher, 1930; but see Hamilton, 1967; Trivers and Willard,
1973). There are a number of factors that might skew the adult ratio
in favor of females even if the ratio at hatching is 1:1. Earlier studies
(Jackson, 1978b) suggested that adult males of solitary and com-
munal, territorial species are relatively nomadic, expending con-
siderable time and energy wandering about searching for females.
As a result, a sizeable proportion of the males in populations of
these species might not be found in censuses of webs. Also, the
nomadic character of males might subject them to earlier mortality
from predation, starvation, and other factors. Shorter male longev-
ity would skew the adult sex ratio in favor of females. However,
mortality factors such as predation would seem less important for
M. gregalis populations in the laboratory; yet the sex ratio was
skewed in favor of females here also, suggesting that mortality
factors of a more intrinsic nature might be involved. Females of each
investigated species oviposited several batches of eggs over a period
of time. In contrast to females, males may be adapted to a lifestyle
that emphasizes courtship, mating, and searching for females, in
conjunction with greater vagility and smaller size, at the expense of
maintenance functions that would serve to prolong survival.

1978]

Jackson & Smith — M alios and Dictyna

79

Acknowledgements

For comments on the manuscript, we thank P. N. Witt, and M. C.
Vick. Special thanks go to W. J. Gertsch for his assistance in the
identification of spiders. C. E. Griswold, P. S. Jackson, and V. D.
Roth are gratefully acknowledged for assistance in the field. The
assistance of the Southwestern Research Station of the American
Museum of Natural History is gratefully acknowledged. This work
was supported in part by the North Carolina Division of Mental
Health Services, Research Section and by N.S.F. grant number
BMS 75-09915 to P. N. Witt.

Summary

Three types of social organization and three corresponding types
of webs occur in the Dictynidae: solitary (individual webs); com-
munal, territorial (web complexes); communal, non-territorial
(communal webs). Solitary (M. niveus ) and communal, territorial
(M. trivattatus, D. calcarata, and D. albopilosa) species were
censused in nature; for M. gregalis (communal, non-territorial),
free-living populations in communal webs in the laboratory were
censused. Web surface area per spider decreases as one goes from
solitary (ca. 35 cm2) to communal, territorial (ca. 10 cm2) to
communal, non-territorial (ca. 2 cm2) species. Very large popula-
tions may occur on single web structures of communal species
(estimates of maxima: M. trivittatus, 10,000 spiders; M. gregalis,
20,000). There is a tendency for adult females to occupy nests and
the interior of webs, with males and immatures occupying the
exterior and the interstitial areas. Sex ratios (male:female) are
skewed in favor of females (M. gregalis 1:7; communal, territorial
species, 1:3; males of solitary species are infrequently found).

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Psyche

[March

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1977b. Web sharing by males and females of dictynid spiders. Bull. Brit. Arach.
Soc. 4: 109-112.

1978a. Comparative studies of Dictyna and Mallos (Araneae, Dictynidae): I.

Social organization and web characteristics. Rev. Arach. 1: 133 164.
1978b. Male mating strategies of dictynid spiders with differing types of social
organization. Symp. Zool. Soc. London (In Press).

Krafft, B.

1970. Contribution a la biologie et a l’Ethologie d 'Agelena consociata Denis
(Araignee sociale de Gabon). Premiere Partie. Biol. Gabon. 6: 197-301.
Kullmann, E.

1972. Evolution of social behavior in spiders (Araneae; Eresidae and
Theridiidae). Amer. Zool. 12: 419 426).

Lubin, Y. D.

1974. Adaptive advantages and the evolution of colony formation in Cyrto-
phora (Araneae: Araneidae). Zool. J. Linn. Soc. 54: 321-339.

Riechert, S. E. and C. R. Tracy.

1975. Thermal balance and prey availability: bases for a model relating web-
site characteristics to spider reproductive success. Ecology 56: 265 284.

Shear, W. A.

1970. The evolution of social phenomena in spiders. Bull. Brit. Arach. Soc. 1:
65 76.

SOKAL, R. R. AND F. J. ROHLF.

1969. Biometry. Freeman, San Francisco.

Trivers, R. L. and D. E. Willard.

1973. Natural selection of parental ability to vary sex ratio of offspring.
Science (N.Y.) 179: 90-92.

Wilson, E. O.

1975. Sociobiology. Belknap, Cambridge.

A SOLITARY WASP THAT PREYS UPON LACEWINGS
(HYMENOPTERA: SPHECIDAE;

NEUROPTERA: CHRYSOPIDAE)*

By Howard E. Evans

Department of Zoology and Entomology,

Colorado State University, Fort Collins, Colorado 80523

The tendency of solitary wasps to restrict predation to members
of specific groups of arthropods is well known. There are several
hundred records for diverse species of Bembix, for example, which
suggest that all Northern Hemisphere species of this genus prey only
upon Diptera. This includes 16 of the 19 species occurring north of
Mexico; the remaining three have remained unstudied. Two of
these, gillaspyi Evans and Matthews (1968) and frommeri Bohart
(1970) were described only recently, while the third, stenebdoma
Parker (1917) was described long ago but has remained one of the
most infrequently encountered members of the genus. Although
stenebdoma was placed in the belfragei species-group by Evans and
Matthews (1968), largely on the basis of the partially preserved
ocellar lenses, the species fits poorly in that group on most features
and in fact is unique in the genus in having the first intercubital vein
straight rather than angulated. Evidently this species is also unique
in the genus, indeed among all digger wasps, in that the prey consists
of lacewings (Chrysopidae).

The use of Neuroptera is not, however, unknown among Bembi-
cini. Evans and Matthews (1973) presented a record of the Aus-
tralian species Bembix wilcannia Evans and Matthews preying upon
adult ant lions (Myrmeleontidae), and Alcock (1975) found Xeros-
tictia longilabris Gillaspy preying upon adult ant lions as well as
flatid buds in southern Arizona. Thus the record of Bembix
stenebdoma using Chrysopidae seems less unusual than it would
have seemed a few years ago. Why this species appears so rare when
it has apparently entered a new and unexploited adaptive zone
remains a mystery.

* Manuscript received by the editor October 10, 1978.

81

82

Psyche

[March

My observations on B. stenebdoma involve a single female seen
plunging into an open hole in a small draw 1 .5 km SW of Bernardo,
Socorro Co., New Mexico, at 1000 hours on 2 August 1978. The soil
in this area was coarse, stony, and hard, with sparse desert
vegetation, but the draw was sandy and bordered by Croton plants
which were in bloom and attracting many bees and wasps. Despite
several hours of intensive collecting in this area by Kevin M. O’Neill
and myself, we took no other specimens of B. stenebdoma.

This female was captured as she left the nest, and the nest was
excavated. Burrow diameter was about 8 mm, and there was no
obvious mound of soil at the entrance. The burrow was straight and
oblique, 30 cm long, terminating in a single cell at a vertical depth of
20 cm. The cell was horizontal, 10 mm in diameter by 20 mm in
length. It contained 10 paralyzed lacewings, all lying on their sides
with their heads facing the entrance to the cell. The egg of the wasp
had been laid on the side of one of the lacewings deep in the cell. The
egg was elongate and curved, measuring 2.2 mm in length; it was

Figure 1 . Prey and egg of Bembix stenebdoma. The cell contents have here been
removed to an artificial cell and the lacewing bearing the egg moved to the top of the
pile for purposes of photography.

1978] Evans — Solitary Wasp that Preys on Lacewings

83

attached between the middle and hind coxae of the prey and
extended over the wing base and well above the dorsum (Fig. 1).

This egg position is similar to that of many Bembix which prey
upon flies. That there were 10 prey in the cell, with the female still
provisioning, indicates that this species mass-provisions its cells.
Mass provisioning is uncommon in Bembix, B. hinei Parker being
the only other North American species for which this has been
reported (Evans, 1957).

The lacewings proved to belong to three species: Eremochrysa
tibialis Banks (2), E. punctinervis (McLachlan) (5), and Chrysoperla
comanche (Banks) (3) (det. Phillip A. Adams). Dr. Adams writes
that all three species are common in the Southwest. Although some
Chrysopidae release odorous substances from thoracic glands, this
is not notably true of members of these genera. It will be of interest
to discover if B. stenebdoma consistently uses lacewings in other
areas and whether predation is restricted to species of Eremochrysa
and Chrysoperla.

Acknowledgments

This paper is part of a study of the comparative behavior of
solitary wasps, supported by the National Science Foundation,
grant BNS76-09319. The assistance of Dr. Phillip A. Adams of
California State University, Fullerton, in identifying the Chrysopi-
dae is gratefully acknowledged.

References

Alcock, J.

1975. The behavior of some bembicine wasps of southern Arizona (Hymen-
optera: Sphecidae, Microbembex, Glenostictia, Xerostictia). Southwest-
ern Nat. 20: 337-342.

Bohart, R. M.

1970. New species, synonymy and lectotype designations in North American
Bembicini. Pan-Pac. Ent. 46: 201-207.

Evans, H. E.

1957. Studies on the Comparative Ethology of Digger Wasps of the Genus
Bembix. Comstock Publ. Assoc., Ithaca, N.Y. 248 pp.

Evans, H. E., and R. W. Matthews

1968. North American Bembix, a revised key and suggested grouping. Ann.
Ent. Soc. Amer. 61: 1284-1299.

Evans, H. E., and R. W. Matthews

1973. Systematics and nesting behavior of Australian Bembix sand wasps.
Mem. Amer. Ent. Inst., no. 20. 387 pp.

84

Psyche

[March

Parker, J. B.

1917. A revision of the bembicine wasps of America north of Mexico. Proc.
U.S. Nat. Mus. 52: 1-155.

SURVEY OF SOCIAL INSECTS
IN THE FOSSIL RECORD*

By Laurie Burnham
Museum of Comparative Zoology,

Harvard University,

Cambridge, Massachusetts 02138, U.S.A.

Biologists have long been intrigued by the complex social systems
of various insects. Despite a voluminous literature dealing with the
evolution of these systems, immense gaps remain in our understand-
ing of insect sociality. Several theories have been proposed to
explain the evolution of social behavior in certain groups of insects
(e.g., Hamilton, 1964), but none consider this problem with respect
to geological time. The present paper does so by examining the
fossil record for clues not only on the antiquity of sociality, but also
on the nature of these early social insects. Included in this survey are
those insects recognized as eusocial: the Isoptera, and three super-
families of the Hymenoptera: Vespoidea, Formicoidea, and

Apoidea.

ISOPTERA

The termites are remarkable in two regards: 1) as a group, they
are fully eusocial, exhibiting a wide range of behavioral modifica-
tions and sophistications, and 2) their record in the geological past,
although sparse, is highly indicative of an Early Mesozoic origin.
This latter point is of particular significance if one considers
sociality among insects as a pinnacle of evolutionary success.
Wilson (1971, p. 1) states that “[insect societies] best exemplify the
full sweep of ascending levels of organization, from molecule to
society.” The possibility that termites evolved a social organization
as far back in geological time as the Jurassic (roughly 190 million
years ago) is of great interest, particularly when attempting to draw
parallels with the evolution of sociality in the Hymenoptera, a group
phylogenetically very remote from the termites.

* Manuscript received by the editor July 7, 1978.

85

86

Psyche

[March

Five of the six families1 of termites recognized by Emerson (1955)
have a fossil record extending at least as far back as the Tertiary. In
1967, Cretatermes carpenteri (Hodotermitidae) was found in an
Upper Cretaceous deposit in Labrador (Fig. 1), a discovery which
immediately placed the origin of the Isoptera no later than the
Mesozoic — an extension of 45 million years from previously
known specimens. In addition, the advanced phylogenetic position
of Cretatermes provides evidence for a much earlier origin of the
order than has formerly been recognized (Emerson, 1967).

An examination of various fossil localities reveals a widespread
termite fauna during the Tertiary Period (Table 1). The Termitidae
are found in Miocene deposits of California and Germany; the
Rhinotermitidae, Hodotermitidae, and Kalotermitidae are found at
various Tertiary deposits throughout the United States and Europe;
and the Mastotermitidae have the most widespread Cenozoic
distribution of all, having been found at localities in the United
States, Europe, South America, and Australia. This latter finding is
highly intriguing because the family Mastotermitidae today has but
one species, Mastotermes darwiniensis, which is restricted to north-
ern Australia.2 Emerson (1955) postulates that this widespread

'The sixth family is the Serritermitidae — an aberrant taxon known from only one
species.

2A look at past climatic shifts provides additional insight into the redistribution of
the termites, particularly with respect to the Mastotermitidae, now solely restricted to
Australia. Reconstructions of paleo-climatic patterns may be made fairly accurately
on the basis of floral analyses (Reid and Chandler, 1933). The presence of Sequoia
stumps in the Florissant Shales of Colorado provides evidence for warmer tempera-
tures during the Oligocene (Emerson, 1969). Tiffney (1977) postulates on the basis of
fossil angiosperm assemblages that temperatures in New England during the
Oligocene were much more equable than at present — the temperatures ranging from
26° C to 9° C in contrast to today’s 21° C to -10° C. Furthermore, extended frosts and
hard freezes were unknown. In the more tropical climate of the Oligocene, colony
activities were presumably carried out year round in a relatively warm, moist
environment, explaining the widespread distribution of the Mastotermitidae during
the Lower to Middle Tertiary. By the Late Miocene or Early Pliocene, the earth’s
climate began shifting towards cooler temperatures with the rising level of the
continental land masses and increasingly large polar ice caps. My hypothesis is that,
unable to adapt to an increasingly colder climate, and possibly to a concomitant
change in predator pressures, the Mastotermitidae began to die out during the
Tertiary. And, because at this time the Termitidae were undergoing tremendously
successful radiation in Africa and South America, the Mastotermitidae became
geographically restricted to northern Australia, represented today by only one relict
species, Mastotermes darwiniensis.

1978]

Burnham — Social Insects in Fossil Record

87

Figure 1. Cretatermes carpenteri Emerson from lower part of Upper Cretaceous
of Labrador. Note humeral suture at wing base. Original photograph of holotype in
Princeton Museum. Length of wing, 7.5 mm.

geographical distribution provides strong evidence to support a
Mesozoic origin of the order. He argues (1975) that the breakup of
the united land mass Pangaea in the Permian or Lower Triassic
must have occurred subsequently to the origin of the Isoptera to
explain their distribution in the southern and northern continental
land masses and that all five families must have been present in the
Late Mesozoic to explain their diversity and distribution by the
Tertiary.

In 1971 he looked at a variety of primitive and derived characters
of each family and analyzed the geographical distribution of the
groups, using plate tectonics to provide the following estimates on
the geological origin of the families:

Mastotermitidae — possibly Early Mesozoic.

Hodotermitidae — Triassic, or Early Jurassic before the breakup
of southern continents.

Kalotermitidae — mid-Jurassic, or Lower Cretaceous, before the
separation of Africa and South America.

Rhinotermitidae — Late Jurassic, Early Cretaceous.

Termitidae — Cretaceous.

Because termites are such poor fliers and do not mate until the
adults have cast their wings, he considers water gaps of more than
50 miles capable of preventing termite dispersal.

While I am supportive of the theory that places great importance
on the role of a unified land mass in animal dispersal, I do not agree
that this can effectively be used to date the origin of the Isoptera.

TABLE 1 ISOPTERA IN THE FOSSIL RECORD.

88

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Kalotermes tristis (Cockerell) Burma Emerson, 1969

Kalotermes nigritus Snyder Chiapas, Mexico Snyder, 1960

TABLE

ISOPTEF

[•HE FOSSIL RECORD.

CRETACEOUS

Hodotermitidae

Hodotermitidac

Ter mops is mallaszi Pongracz
OLIGOCENE
Mastotermitidae

* Mioiermes insignis (Hecr)

•Uioicrmt-. •.pciiahiln (Hceri
Mastoiermes bournemouthensis von Rosen
Mastoiermes heeri (Goppert)

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Kalotermitidae s dd )

•Electrotermes giradi (Giebel)

* Electrotermes affinis (Hagen)

Kalotermes rhenanus Hagen
•Eotermes grandaeva Statz

Oeningen. Germany
Oeningen, Germany
England

Schlesien, Germany
England

Florissant, Colorado

Brazil

Radoboj, Croatia
Radoboj, Croatia
Radoboj, Croatia
Radoboj, Croatia
Radoboj, Croatia
Wiirttemberg, Germany

Calico, California

Burma

Burma

III iiiiiiii ilifl ill111 iliill UUl

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1978]

Burnham — Social Insects in Fossil Record

91

Simpson (1952) has made some insightful remarks on the matter.
He contests the premise that if a given group of organisms requires a
land connection, then disjunctive areas occupied by the group must
have been once connected by continuous land. His contention is
that there is no group of organisms that cannot be dispersed over
water. Given a probability of only one chance in a million that an
organism can cross a stretch of water, when geological time is
considered the chance that the event will actually take place (over
tens of millions of years) becomes significantly greater. It is further
argued that successful colonization is dependent on successful
invasion and the ability of the intruder to compete with existing
species. Chances for survival are much higher when there are
numerous, simultaneous arrivals of individuals.

In my opinion, the termites support such reasoning, and this can
be argued in several ways. Firstly, termites are relatively light-
bodied, winged insects. Studies by Simberloff and Wilson (1969)
and Glick (1933) on the repopulation of an island by wind trans-
ported insects strongly support the possibility that termites are
capable of being carried considerable distances in the upper atmos-
phere. Furthermore, because termites swarm in such large numbers
prior to reproduction, a reasonable possibility exists that they will
be dispersed to a new habitat as either a group or at least as a
male/ female pair. A wind current strong enough to blow one
individual into the upper atmosphere should be equally capable of
carrying multiple individuals, and, according to windflow, of trans-
porting them in the same directional pathway.

Secondly, termites are ideally suited to dispersal over large bodies
of water via floating logs. The more primitive families construct
their extensive nesting colonies in wood and logs; as a consequence,
it is entirely plausible that a dead tree falling into a body of
circulating water could be carried extended distances. Furthermore,
this mode of transportation provides the termites with a source of
food during their sojourn, and travel en masse obviates the prob-
lems of reproduction upon arrival. In addition, as Simpson points
out, the larger the number of individuals, the more likely it is that
they will be successful competitors in the new habitat. I am not
presenting this as evidence that the termites did not evolve while the
earth’s land masses were still contiguous, but am merely pointing
out the problems in arguing that land dispersal was essential for
termites.

92

Psyche

[March

The Isoptera exhibit strong affinities to the Blattodea; evidence
linking the two groups to a common ancestor is well marked
between the Mastotermitidae, an archaic termite family, and the
Cryptocercidae, a family of generalized cockroaches. This theory of
common ancestry is supported by several comparative morphologi-
cal and behavioral studies (Emerson, 1965; McKittrick, 1965;
Ahmad, 1950; Cleveland, 1934; Hill, 1925). McKittrick (1965) goes
so far as to incorporate both groups into the Dictyoptera, an order
which also includes the Mantodea. The gut fauna, female genitalic
structures, anal expansion of the hind wing, morphology of the
proventriculus, and deposition of eggs in ootheca-like masses are
much alike in Mastotermes and Cryptocercus. Furthermore, both
groups inhabit similar habitats. As a consequence, termites have
often been referred to as merely social cockroaches. This degree of
relatedness becomes immediately interesting in view of the extensive
geological record of the cockroaches.

Fossil cockroaches are first found in deposits from the Upper
Carboniferous, which makes them among the oldest insects known.
Furthermore, they comprise 80 percent of the fossil insect fauna
during that period (Carpenter, 1930) — an indication that they have
not only existed, but have flourished, for three hundred million
years. If the similarities between termites and cockroaches are
indeed the result of monophyletic, rather than convergent or
parallel evolution, one might speculate on a much earlier origin for
the Isoptera than is shown by the fossil record.

McKittrick (1965) admits that the flagellate gut fauna essential
for cellulose digestion in both groups may have arisen independ-
ently in each; however, she believes that the similarities in two
important morphological characters, the female genitalia and the
dental belt of the proventriculus, represent primitive characters and
are therefore indicative of a common origin for Mastotermes and
Cryptocercus. On the other hand, Tillyard (1926, 1936), Cleveland
(1934), Imms (1919), Carpenter (personal communication), among
others, believe that the termites were derived from more ancient
stock and may have evolved during the Late Paleozoic. Hamilton
(1978) supports the view that social termites arose from “roach-like
ancestors” in the habitat of dead phloem, and suggests that the
invasion of Cryptocercus into the same type of habitat was inde-
pendent of the ancestral termite. The possibility of termite “evolu-

1978]

Burnham — Social Insects in Fossil Record

93

tion under bark” seems immensely feasible; not only is isolation
(and, hence, inbreeding) possible, but selective pressures leading to
dependence on a cellulose diet would also be high. It seems an
excellent explanation for the early separation of the termites and
cockroaches from a common protorthopteran (protoblattoid) an-
cestor as long ago as the Late Paleozoic. More definite conclusions
on the origin of the Isoptera must wait until termites or termite-like
insects have been found in pre-Cretaceous strata.

HYMENOPTERA

The Hymenoptera belong to the major subdivision of the Insecta
known as the Endopterygota. There are no clues elucidating the
nature or precise age of the earliest endopterygote insects, but the
fossil record does provide insight into the history of the group as a
whole. Representatives of two endopterygote orders, Neuroptera
and Mecoptera, are found as far back as the Early Permian, some
280 million years ago. This occurrence suggests an origin of the
Endopterygota approximately 100 million years after the origin of
the true insects.3

The earliest known Hymenoptera have been found in Triassic
beds of Central Asia (Rasnitsyn, 1964) and Australia (Riek, 1955).
These fossils establish a minimum age for the order of about 220
million years. All the specimens known from this period belong to
the suborder Symphyta, and surprisingly enough belong to the
existing family Xyelidae.

A major advance in the evolution of the Hymenoptera occurred
with the development of a constriction between the first and second
abdominal segments; this presumably had the selective advantage of
increasing the flexibility of the abdomen, important for both
oviposition and defense. Hymenoptera which possess this adapta-
tion, a diagnostic character of the suborder Apocrita, are first
known from Upper Jurassic deposits of Central Asia (Rasnitsyn,
1975, 1977). These specimens have been assigned to the more
primitive division of the Apocrita known as the Terebrantia or

3The oldest known insects, found in Upper Carboniferous deposits, comprise 11
orders and include the Apterygota (Thysanura), Paleoptera and Exopterygota. It
should be noted that here the use of the term insect does not include the Collembola,
Protura or Diplura.

94

Psyche

[March

Parasitica; the other division within this suborder is the Aculeata.4
Members of the latter are characterized by modifications of the
ovipositor that have enabled its use not only for oviposition, but
also as a transport vessel for defensive and prey-paralyzing com-
pounds. This structure unquestionably plays an important role in
colony defense and might provide an explanation for the restriction
of eusociality within the Hymenoptera to the Aculeata.

The oldest known aculeate hymenopteron, Cretavus sibericus, was
discovered in an Upper Cretaceous (Cenomanian) deposit in Siberia
in 1957. Although placed by Sharov (1962) in an extinct superfamily
Cretavidea, related to the Scolioidea, it has recently been transferred
to the existing family Mutillidae by Rasnitsyn (1977, p. 109). Since
1967, species representing 10 families and 19 genera of aculeate
Hymenoptera have been found in Upper Cretaceous deposits in
Central Asia (Rasnitsyn, 1977) (Table 2). Evans (1966) believes that
such diversity by the Late Cretaceous is indicative of an earlier origin
and postulates that the group may have evolved during the Jurassic.
However, it must be pointed out that the Cretaceous is one of the
longer periods in the earth’s history, having a duration of roughly 70
million years, and may have been of sufficient length to account for
such diversification.

Vespoidea

Included in this group are the three families considered to be “true
wasps”: The Masaridae and Eumenidae, both of which are solitary,
and the Vespidae, where one finds behavioral modifications ranging
from subsocial to highly advanced eusocial (Richards, 1953, 1971).
It is the Vespidae, by virtue of their sociality, with which I am
primarily concerned in this paper.

There are many gaps in our record of the early social wasps and of
the Vespoidea in general. Most striking, perhaps, about the fossil
record of the wasps is their lack of representation (see Table 3). The

4The classification of the Aculeata has recently undergone a major revision by D. J.
Brothers (1975), in which the seven previously recognized superfamilies (Bethyloidea,
Scolioidea, Pompiloidea, Formicoidea, Vespoidea, Sphecoidea, and Apoidea) are
now combined into three: the Bethyloidea, Sphecoidea (subdivided into the Spheci-
formes and Apiformes), and Vespoidea (subdivided into the Vespiformes and
Formiciformes). However, since this revised classification has not been generally
accepted in its entirety, I am employing here the more conventional classification
(sensu Riek, 1970; Richards, 1971).

1978]

Burnham — Social Insects in Fossil Record

95

Table 2. Genera of aculeate Hymenoptera known from Cretaceous deposits
(based on Rasnitsyn, 1977, and Evans, 1973). All genera are extinct.

SCOLIOIDEA

Mutillidae

Cretavus

Sharov, 1962;

Rasnitsyn, 1977

7SCOLIOIDEA

Scolioidae

Angarosphecidae

Falsiformicidae

Oryctopterus

Angarosphex

Falsiformica

Rasnitsyn, 1977
Rasnitsyn, 1977
Rasnitsyn, 1977

7SCOLIOIDEA-BETHYLOIDEA

?Scolebythidae

Cretabythus

Evans, 1973

BETHYLOIDEA

Bethylidae

Archaepyris

Celonophamia

Evans, 1973
Evans, 1973

Cleptidae

Procleptes

Hypocleptes

Protamisega

Evans, 1969
Evans, 1973
Evans, 1973

Dryinidae

Cretodryinus

Rasnitsyn, 1977

POMPILOIDEA

Pompilidae

Pompilopterus

Rasnitsyn, 1977

FORMICOIDEA

Formicidae

Sphecomyrma

Cretomyrma

Paleomyrmex

Wilson and Brown, 1967
Rasnitsyn, 1977
Rasnitsyn, 1977

SPHECOIDEA

Sphecidae

Lisponema

Pittoecus

Evans, 1969
Evans, 1973

7SPHECOIDEA

?Sphecidae

Archisphex

Taimyrisphex

Evans, 1969
Evans, 1973

VESPOIDEA

Masaridae

Curiovespa

Rasnitsyn, 1975

96

Psyche

[March

absence of Vespidae from Baltic Amber (Lower Oligocene) and
other fossil resins, in which ants are abundant, is probably due to
their relatively large size, which reduces the likelihood of their
entrapment in the sticky tree resin. Spradbery (1973, p. 316),
attributes their scarcity in sedimentary deposits to “the behavioral
characteristics and paper nest structures which do not lend them-
selves to fossilization.” As with any other fossil, the absence of an
insect in the paleontological record provides no proof as to its actual
occurrence in the past; one can only reconstruct and evaluate
paleofaunas on the basis of those organisms that are represented.
Therefore, it is conceivable that wasps were present earlier than the
record indicates, but that conditions conducive to their preservation
were lacking. The following does, however, provide information on
the diversity of the group as we know it.

Cretaceous

The earliest record of the Vespoidea extends back to the Upper
Cretaceous (Turonian). Two species of vespoid wasp have been
found in a deposit of this age in the USSR — both assigned to
the genus Curiovespa (Rasnitsyn, 1975). Unfortunately, nothing is
known about the body structure of these insects but on the basis of
their wing venation they are placed in the family Masaridae. The
presence of two distinct species in the same deposit suggests that some
diversification of the Vespoidea had taken place as early as the Upper
Cretaceous, although nothing is known about the morphological
character of these early wasps.

Paleocene

No Vespoidea from this period are known.

Eocene

The Eocene beds of Green River have yielded a surprisingly
diverse assemblage of aculeates, but most of these belong to the
Terebrantia or Sphecoidea; the only vespoid recovered from this
deposit, Didineis solidescens, is of uncertain systematic position
(Evans, 1966, p. 393). Scudder (1890) described this specimen as a
sphecid of the subfamily Nyssoninae. However, Evans (1966)
examined the type and concluded that it did not belong to the family
Sphecidae, but was probably a eumenid, and tentatively assigned it
to the genus Alastor.

1978]

Burnham — Social Insects in Fossil Record

97

Figure 2. Vespoid wasp from Eocene of British Columbia. Original photograph
of specimen in Royal Ontario Museum, Toronto. Length of forewing, 12 mm.

Piton (1940), in a thesis on the Eocene fossil beds of Menat,
France, described an assemblage of Vespoidea found in this sedi-
mentary deposit. However, because the six specimens he described
are all assigned to extant genera, and do not show the characters
essential for such generic designation, Piton’s taxonomic determina-
tions are perforce questionable. Particularly dubious is his place-
ment of one specimen in the family Vespidae, genus Polistes.
Because the morphological features necessary for accurate taxo-
nomic placement are obscured in this fossil, I prefer to place it in
Vespoidea incertae sedis. The remaining five specimens are assigned
to the Eumenidae incertae sedis.

Another vespoid species was recently recovered from a Middle
Eocene deposit in British Columbia (M. V. H. Wilson, 1977). Al-
though not formally described, the fossil clearly shows the charac-
teristic venation of the vespoid complex (see Fig. 2), but could be
either a vespid or a eumenid. Of course, one has no way of stating

98

Psyche

[March

with certainty that these early vespids were social. Within the
Vespidae, divisions into subfamily and tribe are based primarily on
behavioral rather than morphological characters. Furthermore, the
morphological differences between the castes in any given species
are often not obvious in the preserved fossils.

Oligocene

True vespids are first found in the Upper Oligocene shales of
Florissapt, Colorado and Rott, Germany, two highly productive
fossiliferous deposits. These beds and other various localities listed
in Table 3 have turned up an assemblage comprised of four genera
and 14 species. It is quite remarkable that three of the four genera
represented are extant and this supports the possibility that the
Vespidae were essentially modern by the Oligocene. Furthermore,
the diversification of taxa suggests a much earlier origin for the
family than is evidenced by the fossil record.

Miocene

Scarcely any Vespidae are known from the Miocene, although
this is most likely due to the overall dearth of deposits from this
epoch. One vespid has been described from a deposit in Germany.
This is Polistes kirbyanus and clearly belongs to the subfamily
Polistinae. Other wasps from Miocene deposits have yet to be
discovered, but one can assume that the wasp fauna of this age
would be barely distinguishable from the wasp fauna of today.

Formicoidea

The following review of the fossil history of the Formicidae
provides important information on their dominance, distribution,
and supposed habits during the Mesozoic and Cenozoic eras. In
contrast to the Vespoidea, ants are the most abundant insects in
Tertiary formations. This may be attributed to their foraging
behavior on and around trees, which enhances their chances of
preservation in amber. A rough total of 20,000 specimens represent-
ing some 200 species of ants has been studied (Table 4); this massive
amount of work far exceeds the paleontological investigations
carried out on any other family of insects. Several comprehensive
monographs on the subject have been written, including The Ants of
the Baltic Amber (Wheeler, 1914), and The Fossil Ants of North
America (Carpenter, 1930), which are drawn on extensively in the
following pages.

1978]

Burnham — Social Insects in Fossil Record

99

Cretaceous

The Cretaceous Period has, without question, provided more
information on the early evolution of the ants than any other
period, primarily because of the discovery in 1967 of two perfectly
preserved worker ants in a New Jersey amber deposit. No doubt
exists as to the primitive nature of these Cretaceous ants — both are
members of the same species, Sphecomyrma freyi Wilson and
Brown, and possess a mixture of wasp and ant characters. The
petiole is distinctly ant-like, although the mandibles, which are short
and bidentate, are very wasp-like (see Fig. 3A). A new subfamily,
Sphecomyrminae, was named to accommodate S. freyi (Wilson,
Carpenter, and Brown, 1967), and is considered ancestral to all
known formicid subfamilies (see Taylor, 1978).

Since the discovery of Sphecomyrma, several other Cretaceous
ants have been found, and these provide strong evidence that the
family was widespread during this period. Dlussky (1975) described
two new genera and three species, Cretomyrma arnoldii, C. uni-
cornis, and Paleomyrmex zherichini (from a Late Cretaceous amber
deposit in Yantardak, USSR) which he assigned to the Sphecomyr-
minae. It is of interest that the type of P. zherichini is the first
winged male ant to be found in a Cretaceous deposit and provides
the only indication of wing venation in the Sphecomyrminae (Fig.
3B). The figured specimen of Cretomyrma unicornis raises doubts
as to its position in the Formicidae for it is a badly mangled, poorly
preserved specimen and might be better assigned to Hymenoptera
incertae sedis.5 A fifth specimen, apparently a worker, has recently
been discovered in the Cretaceous amber of Manitoba, Canada.
Although not yet described, it undoubtedly belongs to the subfamily
Sphecomyrminae (Wilson, personal communication).

Paleocene

No ants from the Paleocene are known, undoubtedly because so
few fossiliferous beds containing insect remains from this epoch

5Dlussky (1975) also described several other “ants” which were found in Upper
Cretaceous deposits in the Kzyl-Zhar of Russia. Three genera (3 species) were placed
in the subfamily Ponerinae: Petropone petiolata, Cretopone magna, and Archaeo-
pone kzylzharica. These are all fragmentary specimens, and, as figured by Dlussky,
present no characters which would place them unequivocally in the Formicidae. They
much more obviously belong in Hymenoptera incertae sedis, as does Dolichomyrma
longiceps from the Upper Cretaceous of Kzyl-Zhar, which Dlussky put into
Formicidae incertae sedis.

TABLE 3. VESPOIDEA IN THE FOSSIL RECORD.

100

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1978]

Burnham — Social Insects in Fossil Record

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TABLE 3. VESPOIDEA IN THE FOSSIL RECORD.

§

Kazakh, U
Kazakh, U

Evans,

Piton,

Florissant, Colorado
Florissant, Colorado
Rott, Germany

lorissantia Cockerell
cudderi Cockerell
■elecia Cockerell
'tallica Cockerell
vilsoni Cockerell

Polisi

rius Theobald
Polisles signata Statz
I Poly bia anglica Cockerell
Polybia oblita Cockerell
Vespa bilineata Statz
Vespa cordifera Statz

Florissant, Colorado
Baltic Amber
Florissant, Colorado

Rott, Germany
Isle of Wight, England
Isle of Wight, England
Rott, Germany
Rott, Germany

Bequacrt, 1930
Bequaert, 1930
Bequaert, 1930
Bequaert, 1930
Cockerell, 1909b
Cockerell, 1914
Theobald, 1937
Statz, 1936
Cockerell, 1921a
Cockerell, 1921b
Statz, 1936
Statz, 1936
Statz, 1936

Oeningen, Germany
Parschlug, Germany
Radoboj, Croatia

Cockerell, 1914
Heer, 1849
Heer, 1867

min

102

Psyche

[March

Figure 3 A. Sphecomyrma freyi Wilson and Brown from the lower part of Upper
Cretaceous of New Jersey. Drawing of holotype worker in Museum of Comparative
Zoology, modified from Wilson, Carpenter, and Brown (1967). Length of body, 3.5
mm.

Figure 3B. Paleomyrmex zherichini Rasnitsyn from the lower part of the Upper
Cretaceous of U.S.S.R. Drawing of holotype male in Paleontological Institute,
Moscow, from Rasnitsyn, 1977. Length of body, 5.4 mm.

have been discovered. Mention is made by Brues (1936) of a piece of
petrified wood containing what he considers ant borings, highly
resemblant of borings made by Camponotus today. Although there
is no clear-cut evidence that these borings represent Camponotus
activity, or insect activity of any kind, it is conceivable that
Camponotus was present in New Mexico during the Paleocene;
several species have been dscribed from the Florissant Shales,
Colorado (Upper Oligocene), and one from the Baltic Amber

1978]

Burnham — Social Insects in Fossil Record

103

(Lower Oligocene). In addition, it must be remembered that the
Paleocene did not begin for at least 40 million years after the
appearance of Sphecomyrma freyi.

Eocene

Very few fossil ants have been found in deposits of this age, and
the determinations of many of these ant species are in doubt.
Scudder (1877, 1878) described four supposed ants from the Green
River formation, and five ants (1877) from the Quesnel Beds in
British Columbia. Generic identifications on all of these fossils are
to be considered dubious at best, and more likely incorrect (Car-
penter, 1930).

In 1920, two species, Oecophylla bartoniana and Formica heter-
optera, were described by Cockerell from an Eocene deposit in
Bournemouth, England. Wheeler (1928) considered these ants for-
micines, but because the descriptions were based on wing fragments,
he questioned their generic determinations. Similarly, Cockerell’s
Formica eoptera (1923a) from the Eocene of Texas is of uncertain
position at both the generic and subfamily levels. Archimyrmex
rostratus (Cockerell, 1923b) from the Eocene shales of Colorado is
probably a myrmicine (Carpenter, 1930), and is the only Green
River ant that can be placed with any certainty in a subfamily.
Carpenter (1929) described Eoponera berryi from the Wilcox
formation of Tennessee, and placed this ant in the subfamily
Ponerinae. He suggests that it may be closely allied to the Neotrop-
ical genus Dinoponera. This is of interest because Eoponera berryi
is the oldest known ant (Lower Eocene) to be assigned to a living
subfamily of Formicidae.

Wilson (personal communication) mentions the recent discovery
of three ants in a Middle Eocene amber deposit near Malvern,
Arkansas, each belonging to a different subfamily. One belongs to
the Dolichoderinae, genus Iridomyrmex; one is a formicine closely
allied to the genus Paratrechina, and considered a relatively primi-
tive, or “typical euformicine”; the last is a new genus of myrmicine,
unique by virtue of its inflated postpetiole. These ants have yet to be
formally described but they are nevertheless of paramount interest.
The presence of these subfamilies in North America in the Eocene is
strongly suggestive of their rapid evolution and dispersal during the
Paleocene and perhaps during the Cretaceous.

104

Psyche

[March

O/igocene

The Baltic Amber is, most certainly, the best studied of all
Tertiary insect deposits, and has revealed a great deal about the
nature and diversity of Oligocene ants.6 As of 1928, 1 1,71 1 ants (93
species) were examined from this deposit. Of this number, 1461 were
studied by Mayr (1868); 690 by Andre (1895); and 9,560 by Wheeler
(1914, 1928).

An examination of the ant fauna reveals wide representation at
the subfamily and generic levels. All extant subfamilies of Formici-
dae are found in the amber with the exception of the Dorylinae and
Leptanillinae. The absence of the Dorylinae is probably not due to
selective exclusion on the part of the amber, but more likely
indicates their absence from that part of the European continent
during the Oligocene. Wheeler (1914) speculates that the foraging
behavior of doryline ants should readily lead to entrapment in tree
resin, but, in all probability, this group was then, as it is now,
confined to the tropics. It is not surprising that the Leptanillinae are
absent from the Baltic Amber; this is a small subfamily once
considered a tribe of the Dorylinae, consisting of one genus and a
few species; and although pantropical is hypogaeic and rarely
encountered.

The Dolichoderinae and Formicinae together constitute 97 per-
cent of all specimens and evidence indicates that these amber ants
were already extraordinarily specialized. Workers of Iridomyrmex
goepperti were found in a piece of amber (originally in the Konigs-
berg collection) with several aphids. On the basis of this discovery,
Wheeler (1914) concludes that Homoptera were attended by ants
then much as they are today. The finding of several genera of
paussid beetles (e.g., Arthropterus, Cerapterites and Eopaussus) in
the Baltic Amber (Wasmann, 1929) suggests that myrmecophiles
were established at this time. Perhaps most remarkable of all was
the discovery of two Lasius schiefferdeckeri workers — each found
with a mite attached to the base of the hind tibia, in precisely the

6Because the Baltic Amber was secondarily deposited in a clay bed of Lower
Oligocene age, it is necessarily older than the glauconitic sand (“blue-earth” clay) in
which it lies. How much older is uncertain. In some published accounts it is referred
to as Eocene. However, since the composition of the Baltic Amber ant fauna is very
similar to that of the Florissant Shales and other bona fide Oligocene deposits, I am
following Zeuner (1939, p. 26) in referring to the amber as Lower Oligocene.

1978] Burnham — Social Insects in Fossil Record 105

same position on each. This demonstrates almost certainly that by
the Lower Oligocene mites had acquired distinct preferences for
attachment on specific regions of their host’s integument.

Almost as valuable as the Baltic Amber in providing a large and
diverse assemblage of fossil ants is the Upper Oligocene deposit in
Florissant, Colorado, studied by Carpenter (1930). The ant fauna of
this deposit is strikingly similar to that of the Baltic Amber in many
respects. It is interesting to note that roughly the same percentage of
extant genera is found in both places; in the Florissant Shales this
figure is given as 60 percent (Carpenter, 1930), in the Baltic Amber
56 percent (Wheeler, 1914). Iridomyrmex is clearly a dominant
genus in the Baltic Amber, and although not so common in the
Florissant Shales, a closely allied genus, Protazteca, comprises more
than 25 percent of all specimens (Brown, 1973).

Another similarity between the two deposits is the relative
percentages of the various subfamilies. As in the amber, the
Dolichoderinae are predominant, comprising 60 percent of the total
number of ants. The Formicinae comprise another 25 to 30 percent,
and the Myrmicinae in each deposit are represented by five percent
or less of the total specimens. This suggests that the ant fauna in the
northern hemisphere was essentially homogenous during the Oli-
gocene.

The remaining deposits of Oligocene age from which ants have
been described are of relatively minor importance. Most of the
specimens are fragmentary and the determinations dubious; never-
theless, a mention of them is certainly necessary. Specimens from
Gurnet Bay, Isle of Wight, England, have been studied by Cockerell
(1915) and Donisthorpe (1920). Cockerell described eight species of
ants from this deposit but, because his generic determinations are
based chiefly on highly variable measurements of wing fragments,
they are of dubious significance. Donisthorpe examined a total of
eight genera and fourteen species belonging to the subfamilies
Ponerinae, Dolichoderinae, and Formicinae. Surprising is the large
number of Oecophylla workers recovered (245); this genus is now
restricted to Africa, India, and Australia, and is much more
numerous in the Gurnet Bay deposit than in the Baltic Amber or
Florissant Shales. This might be due to the difference in latitude
between the deposits which would account for a warmer climate at
Gurnet Bay later into the Tertiary than at the more northern
deposits.

106

Psyche

[March

Another Lower Oligocene deposit which has provided beautifully
preserved fossil ants is Aix-en-Provence, France. Several species
have been described by Theobald (1937), who recognized four
subfamilies: Myrmicinae (1 species); Ponerinae (1 species); Doli-
choderinae (1 genus, 2 species); and Formicinae (3 genera, 9
species). Also described by Theobald (1937) is an Oligocene collec-
tion from Haut-Rhin, France, in which he recognizes the same four
subfamilies (16 genera, 34 species). This fauna is very similar to that
found in the Baltic Amber; in fact, Theobald has found five species
which he considers identical to species in the Baltic Amber. In a
deposit in Gard, France, Theobald (1937) describes two species, one
a myrmicine, the other a dolichoderine.

Meunier (1917) has described four ant species from an Upper
Oligocene deposit in Rott, Germany. These have been assigned to
three genera: Formica, Ponera, and Myrmica. The specimens are
well-preserved, as may be seen in Meunier’s photographs, but his
generic determinations are questionable.

In 1957, two female reproductives of the same species were
discovered in an Upper Oligocene deposit in Argentina. The authors
described the species as Ameghinoia piatnitskyi and placed it in the
subfamily Ponerinae (Viana and Haedo-Rossi, 1957). E. O. Wilson
(personal communication) is highly sceptical of the placement of A.
piatnitskyi in the Ponerinae, and thinks that it is very clearly a
myrmeciine. This is quite extraordinary because no other fossil ants
have been recovered from South America, and more importantly, if
Wilson is correct, this is the first indication that the Myrmiciinae
were so widespread by the Oligocene.

Miocene

The deposits of Miocene age which have provided the greatest
number of ant specimens have been the Oeningen beds in Germany,
and the Radoboj formation in Croatia. Approximately 60 species of
ants from these places were described by the Swiss myrmecologist
Heer (1849, 1856, 1867), but his generic assignments are necessarily
questionable in terms of present-day concepts of a formicid genus.
Regrettably, the type specimens which are essential to a revision of
this fossil fauna are believed to be lost.

1978]

Burnham — Social Insects in Fossil Record

107

A few species were described by Emery (1891) in Sicilian ambej,
presumed to be Miocene, but these, like the specimens studied by
Heer, are of questionable generic position.7

Another Miocene amber deposit has been found in Chiapas,
Mexico, from which some one hundred ants have been recovered.
Unfortunately, the majority of these are fragmentary, or otherwise
too poor for determination. The assemblage does, however, suggest
that the ant fauna in Mexico during the Miocene was essentially the
same as might be found in that region today (Brown, 1973).

Fujiyama (1970) described a single ant from the Chojabaru
formation in Japan (middle Miocene) which he named Aphaeno-
gaster axila, thought to be closely allied to the subgenus Dero-
myrma. This is not particularly unusual inasmuch as Aphaenogaster
is a world-wide genus, and several species are found in Japan today.

Perhaps the most interesting of all Miocene material is an ant
colony of Oecophylla leakeyi found in Kenya (Wilson and Taylor,
1964). This is the first record of an actual, although fragmented, ant
colony and contains a total of 366 specimens: 197 larvae, 105 worker
pupae, and at least 64 workers. No Nearctic fossils of Oecophylla
are known, but the species is well represented in European Tertiary
deposits. Wilson and Taylor suggest on the basis of these fossil
specimens that Oecophylla is a morphologically stable paleotropical
genus which has persisted through most of the Tertiary with very
little specialization.

Apoidea

The Apoidea form an interesting complex of social insects. Unlike
the other social insect groups that are consistent in their degree of
social achievement at the ordinal level (Isoptera), family level
(Formicidae), and virtually the subfamily level (Vespinae), the
Apoidea present a wide spectrum of social behavior at the generic
level. Evidence suggests that eusociality has arisen in the bees at
least eight times (Michener, 1962; Wilson, 1971), which may explain
this variance. Nevertheless, it is noteworthy that of roughly 20,000
existing species of bees only a small minority are thought to be
presocial and eusocial (Wilson, 1971). Why sociality in the Apoidea

7These generic determinations are currently being reviewed by Dr. W. L. Brown, Jr.

TABLE 4. FORMICOIDEA IN THE FOSSIL RECORD.

108

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Aphaenogaster sommerfeldti Mayr Baltic Amber Wheeler, 1914

Aphaenogaster oligocenica Wheeler Baltic Amber Wheeler, 1914

TABLE

FORMICOIDEA IN THE FOSSIL RECORD.

CRETACEOUS

Sphccomyrminac ^ ^

1*Sphecomyrma sp.

•Creiomvrma arnoldii Dlussky
•Cretomyrma unicornis Dlussky
• Paleomyrmex zherichini Dlussky
EOCENE

yT*ArMmyrmex rostratus Cockerell

Oecophylla bartoniana Cockerell
Formica eoptera Cockerell
, Formica heieropiera Cockerell

l Parat rechina sp.

*Eoponera berryi Carpenter
Dolichoderinae

Taymyr, U.S.S.R.
Taymyr, U.S.S.R.
Taymyr, U.S.S.R.

Bournemouth, England
Texas, U.S.A.
Bournemouth, England

r, 1929

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* Electoponera dubia Wheeler

Euponera calcarea Theobald
Euponera succinea (Mayr)

Euponera crawleyi Donisthorpe
Euponera globivenlris Theobald

Aphaenogasier mayri Carpenter
Aphaenogasier donislhorpei Carpenter
Aphaenogasier maculipes Theobald
Aphaenogasier maculata Theobald

Aphaenogasier oligocenica Wheeler

Florissant. Colorado
Baltic Amber
Baltic Amber

Baltic Amber
Baltic Amber

Haut-Rhin, Germany

Isle of Wight, England

Haut-Rhin, Germany

Baltic Amber

Isle of Wight, England

Rott, Germany

Isle of Wight, England

Rott, Germany

Baltic Amber

Isle of Wight. England

Isle of Wight, England

Florissant, Colorado

Florissant, Colorado
Florissant, Colorado
Haut-Rhin, Germany

Donisthorpe, 1920

Meunicr, 1923
Cockerell, 1915
Meunier, 1917
Wheeler, 1914
Donisthorpe, 1920
Donisthorpe, 1920

Carpenter, 1930

Carpenter, 1930
Carpenter, 1930
Theobald, 1937
Theobald, 1937
Wheeler, 1914
Wheeler, 1914

TABLE 4. (CONTINUED)

110

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Dolichoderus coquandi Theobald Haut-Rhin, Germany Theobald, 1937

TABLE 4. (CONTINUED)

Baltic Amber
Baltic Amber

Baltic Amber

Wheeler, 1914
Wheeler. 1914

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Haut-Rhin, Germany
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Carpenter, 1930
Carpenter, 1930

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d, 1937

OLIGOCENE Dolichoderinae (continued)

112

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Rhopalomyrmex pygmaeus Mayr Baltic Amber Wheeler, 1914

Dimorphomyrmex theryi Emery Baltic Amber Wheeler, 1914, 1929

Dimorphomyrmex mayri Wheeler Baltic Amber Wheeler, 1914

TABLE 4. (CONTINUED)

Iridomyrmex breviantennis Theobald
Iridomyrmex florissantius Carpenter
Iridomyrmex obscurans Carpenter

Iridomyrmex samlandicus Wheeler
Iridomyrmex oblongiceps Wheeler
Proiazieca elongata Carpenter
Protazteca quadraia Carpenter

Liomeiopum miocenicum Carpenter
Liometopum oligocenicum Wheeler
Liomeiopum scudderi Carpenter
Elaeomyrmex gracilis Carpenter
Elaeomyrmex coloraderisis Carpenter
Asymphylomyrmex balticus Wheeler
Pilyomyrmex tornquisii Wheeler
Miomyrmex impacius (Cockerell)
Miomyrmex striatus Carpenter

Formicinae

Plagiolepis klinsmanni Mayr
Plagiolepis kuenowi Mayr
Plagiolepis squamifera Mayr
Plagiolepis singulars Mayr
Plagiolepis soliiaria Mayr

Haul-Rhin, Germany
Florissant, Colorado
Florissant, Colorado

Baltic Amber
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Baltic Amber
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado

Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber

Wheeler, 1914
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Wheeler, 1914
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Wheeler, 1914
Wheeler. 1914
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930

Wheeler, 1914

Wheeler, 1914 >3

Wheeler, 1914 ;

Wheeler, 1914
Wheeler, 1914, 1929
Wheeler, 1914

OLIGOCENE Formicinae (continued)

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Burnham — Social Insects in Fossil Record

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Cpmponotus mengei Theobald Haut-Rhin, Germany Theobald, 1937

TABLE 4. (CONTINUED)

OLIGOCENE Formicinac (continued)

Oecophylla brischkei Mayr
Oecophylla brevinodis Wheeler
Oecophylla megarche Cockerell
Oecophylla aiavina Cockerell
Oecophylla perdila Cockerell
Prenolepis henschei Mayr
Prenolepis pygmaea Mayr
Lasius schiefferdeckeri Mayr

Lasius pumilus Mayr

Lasius epicenirus Theobald

Lasius chambonensis Piton and Theobald

Baltic Amber
Haut-Rhin, Germany
Haut-Rhin, Germany
Haut-Rhin, Germany
Baltic Amber
Baltic Amber
Haut-Rhin, Germany

Baltic Amber
Isle of Wight, England
Isle of Wight, England
Isle of Wight, England

Aix-en-Provence, France
Lac Chambon, France
Ukraine, U.S.S.R.

Baltic Amber
Baltic Amber
Baltic Amber
Florissant, Colorado

Wheeler, 1914
Theobald, 1937
Theobald, 1937
Theobald, 1937
Wheeier, 1929

Theobald, 1937
Wheeler, 1914
Wheeler, 1914
Donisthorpe, 1920
Cockerell, 1915
Cockerell, 1915

Wheeler, 1914
Wheeler, 1914
Wheeler, 1914
Theobald, 1937
Piton and Theobald, 1935
Zalessky, 1949
Wheeler, 1914
Wheeler, 1914
Wheeler, 1914
Wilson, 1955
Cockerell, 1921c

Wheeler, 1914
Wheeler, 1914
Wheeler, 1914
Wheeler, 1914
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Piton and Theobald, 1935
Piton and Theobald, 1935

and Theobald, 1935

Theobald, 1937

Wheeler, 1914
Theobald, 1937

Psyche

OLIGOCENE Formicinae

116

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Lasius martynovi Popov Caucasus, U.S.S.R. Popov, 1933

Oecophylla leakeyi Wilson and Taylor Kenya Wilson and Taylor, 1964

1978]

Burnham — Social Insects in Fossil Record

117

is so highly polyphyletic remains unanswered, and is a problem
unlikely to be resolved by the geological past.

However, the fossil record does provide intriguing information on
the evolution of the bees and indicates that their sociality may well
have been established prior to the Oligocene. The following survey
of the fossil Apoidea is indicative of the diversity of bees which have
been found (Table 5). Those species which were described by early
19th century entomologists (Latreille, Heer, Heyden, etc.) are
excluded from this coverage because these were uniformly assigned
to modern genera.8 Cockerell (1909) claims that most of these
species actually belonged to quite different and extinct genera.

Oligocene

The earliest bees in the fossil record are found in the Baltic
Amber, of Lower Oligocene age. The bees in this deposit are well-
diversified (Zeuner and Manning, 1976), and the most prevalent
apoid genus in the amber, Electrapis, is thought to have been social.
Cockerell (1909) based this conclusion on the occurrence of many
specimens of Electrapis meliponoides crowded together in a small
piece of amber, a suggestive but certainly not conclusive deduction.
Zeuner (1944, 1951), however, believed Electrapis to be social based
on its pollen collecting apparatus. The extent to which social
behavior was developed in this genus nevertheless remains a matter
of conjecture. Electrapis is considered by some to be directly
ancestral to the highly eusocial Apis, although Kelner-Pillault
(1974) disagrees with this relationship. She suggests that Electrapis
is actually a long extinct genus which possessed many primitive
characters and represents an evolutionary side-line of the Apoidea.
Both hypotheses are highly conjectural.

The presence of long-tongued bees such as Electrapis suggests
that the Baltic Amber bees were rather specialized. Tongue structure
is assumed to have evolved in response to various morphological
changes (i.e., longer corollas) which took place during the evolution
of the angiosperms (Michener, 1974). Short-tongued bees such as
the colletids are considered the more primitive members of the
Apoidea and are representative of bee radiation that occurred at a
time when most of the angiosperms had shallow flowers (Michener,
1974).

For a listing of these specimens, see Zeuner and Manning (1976).

118

Psyche

[March

In Late Oligocene deposits, the Apoidea are extremely well
represented. Six major families of bees are known from this epoch:
Halictidae, Andrenidae, Melittidae, Megachilidae, Anthophoridae,
and Apidae. A total of 29 genera are represented, many of which are
extant. Several specimens belonging to Chalcobombus and Bombus
are described from deposits in both Europe and North America
suggesting widespread radiation of this specialized group of bees by
the Early Oligocene. In the Late Oligocene, bees very similar to Apis
mellifera are found. Manning (1952) feels that some species from the
Rott Shales possess almost all the necessary characters for place-
ment in the genus Apis (Fig. 4). Moreover, in the Dominican Amber
of Oligocene-Miocene age, several Trigona workers are found,
providing convincing proof that social behavior was well established
at this time (Michener, 1974).

Figure 4. Apis henshawi Cockerell from Upper Oligocene of Rott, Germany.
Original photograph of holotype in Museum of Comparative Zoology. Length of
body, 15 mm.

1978]

Burnham — Social Insects in Fossil Record

119

Miocene

By the Miocene, the bee fauna is essentially modern. In Chiapas
Amber from Mexico, bees have been discovered that are so similar
to an existing Neotropical species that they have been assigned to
the same subgenus, Trigona ( Nogueirapis ), and are scarcely different
at the specific level (Wille, 1959). Fujiyama (1970) mentions the
discovery of a fossil bee in a Japanese Miocene deposit and states
that, “There is no room for doubt that this is a species of honeybee.”

A review of the fossil record reveals the following about the
evolution of the bees. 1) We know that the Early Oligocene fauna is
a mixture of primitive and advanced genera, although it appears to
be dominated by fairly advanced species. By the end of this epoch,
the fauna is modern in overall character. 2) We know that sociality
had clearly arisen by the end of the Oligocene, and possibly much
earlier. And 3) by the Miocene, the bees were virtually indistinguish-
able from the bees of today. Six families of bees are represented in
the Oligocene: including the phylogenetically advanced Apidae with
six genera and 22 species. Such diversity of relatively advanced bees
is indicative of either a much longer history of the group than is
evidenced by the fossil record, or a fairly short history characterized
by the rapid speciation and explosive radiation of the group.

The bees are clearly derived from the spheciform wasps, al-
though nothing is known about the nature of this sphecid ancestor
(Wilson, 1971; Michener, 1974). In 1964, just prior to his death, F.
J. Manning was investigating a sphecid from the Jurassic beds of
Lerida Province, Spain, which “he thought might be (or be closely
related to) the ancestor of the bees” (Zeuner and Manning, 1976, p.
155). This would be an astounding find if true, and it is unfortunate
that nothing more is known — either about the specimen or about
Manning’s reasons for thinking it ancestral to the bees.

The distinction between the Sphecoidea and the Apoidea is
sufficiently subtle as to make determinations of fossil compressions
extremely difficult. The presence of plumose hairs and enlarged
basitarsi, characters which are important apoid features, rarely
survive preservation unless the insect is preserved in amber.

The origin of the bees remains a subject of much speculation. It is
believed that “insect-plant interactions played a key role in the
origin of the angiosperm flower and component structures” (Hickey

120

Psyche

[March

and Doyle, 1977, p. 92). Conversely, angiosperms have been
instrumental to the evolutionary success of the Apoidea. On the
basis of the evolutionary dependence of the two groups, can
anything be said about their relationship in geological time? Two
possibilities present themselves: 1) the angiosperms evolved first and
were initially wind pollinated9 or pollinated by arthropods other
than Hymenoptera (e.g., Coleoptera, Diptera, Thysanoptera, pos-
sibly spiders); and 2) the first bees evolved from sphecid wasps prior
to the origin of the angiosperms by adapting themselves to feeding
on pteridosperm pollen or reproductive organs.

A closer look at these possibilities is warranted. Coleoptera and
Diptera are found in the fossil record at least by the Triassic. This
supports the argument that they could have served as vectors for
dispersal of angiosperm pollen. The question arises, if these insects
were capable of performing essential roles as pollinators, why didn’t
angiosperms arise earlier in the Mesozoic than the Cretaceous?
Regal (1977) suggests that the limiting factor to angiosperm dis-
persal was the presence of seed-carrying birds and mammals. He
argues that this method of seed dispersal, acting in conjunction with
insect pollination, provided the selective advantages behind the
subsequent primary radiation of the angiosperms. This is a sound
argument, but says little about the insects which may have been
pollinating these early plants. It would seem that successful dispersal
of flowering plants is dependent on efficiency at two levels —
pollination and seed dispersal. The explosive radiation of the
angiosperms during the Cretaceous indicates that the more special-
ized insect pollinators, the bees, may have been present in order to
explain this success.

This might support the possibility that pollen collecting bees had
already evolved by the time the first angiosperms appeared. Accord-
ing to Wilson (1971, p. 75), the “Apoidea can be loosely character-
ized as sphecoid wasps that have specialized in collecting pollen
instead of insect prey as larval food.” The possibility, however
speculative, exists that bees evolved in response to the food source
presented by the pteridosperms but subsequently abandoned this
resource when the angiosperms appeared. Certainly one way of
accounting for the explosive radiation of the angiosperms would be

9Stebbins (1970, p. 323) suggests that the earliest angiosperms were not wind
pollinated.

1978]

Burnham — Social Insects in Fossil Record

121

TABLE 5. APOIDEA IN THE FOSSIL RECORD.10

Geological Age

Locality

EOCENE

?Apidae

Probombus hirsutus Piton

Menat, France

OLIGOCENE

Halictidae

*Cyr tapis anomalis Cockerell

Florissant, Colorado

Halictus ruissatelensis Timon-David

Marseille, France

Halictus florissantellus Cockerell

Florissant, Colorado

Halictus miocenicus Cockerell

Florissant, Colorado

Halictus scudderiellus Cockerell

Florissant, Colorado

Andrenidae

Andrena wrisleyi Salt

Baltic Amber

Andrena clavula Cockerell

Florissant, Colorado

Andrena grandipes Cockerell

Florissant, Colorado

Andrena hypolitha Cockerell

Florissant, Colorado

Andrena lagopus Latreille

Florissant, Colorado

Andrena percontusa Cockerell

Florissant, Colorado

Andrena sepulta Cockerell

Florissant, Colorado

* Lithandrena saxorum Cockerell

Florissant, Colorado

* Pelandrena reduct a Cockerell

Florissant, Colorado

* Libellulapis antiquorum Cockrell

Florissant, Colorado

* Libellulapis wilmattae Cockerell

Florissant, Colorado

Melittidae

*Ctenoplectrella dentata Salt

Baltic Amber

*Ctenoplectrella viridiceps Cockerell

Baltic Amber

*Ctenoplectrella splendens Kelner-Pillault

Baltic Amber

*Glyptapis fuscula Cockerell

Baltic Amber

*Glyptapis mirabilis Cockerell

Baltic Amber

*Glyp tapis neglecta Salt

Baltic Amber

*Glyptapis reducta Cockerell

Baltic Amber

*Glyp tapis reticulata Cockerell

Baltic Amber

Melitta willardi Cockerell

Baltic Amber

Megachilidae

Anthidium mortuum (Meunier)

Rott, Germany

Anthidium exhumatum Cockerell

Florissant, Colorado

Anthidium scudderi Cockerell

Florissant, Colorado

* Dianthidium tertiarium Cockerell

Florissant, Colorado

* Lithanthidium pertriste Cockerell

Florissant, Colorado

Heriades bowditchi Cockerell

Florissant, Colorado

Heriades halictinus Cockerell

Florissant, Colorado

Heriades laminarum Cockerell

Florissant, Colorado

Heriades mersatus Cockerell

Florissant, Colorado

10See Zeuner and Manning (1976) for reference citations.

122

Psyche

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TABLE 5. (CONTINUED)

Geological Age

OLIGOCENE Megachilidae (continued)
Heriades mildredae Cockerell
Heriades priscus Cockerell
Heriades saxosus Cockerell
Megachile praedicta Cockerell
Osmia carbonum Heyden
Anthophoridae

Ceratina disrupta Cockerell
Xylocopa friesei Statz
Tetralonia berlandi Theobald
Anthophora melfordi Cockerell

* Anthophorites gaudryi Oustalet

* Protomelecta brevipennis Cockerell
Apidae

* Chalcobombus hirsutus Cockerell

*Chalcobombus humilis Cockerell

* Chalcobombus martialis Cockerell
Bombus florissantensis (Cockerell)

*Sophrobombus fatalis Cockerell
Trigona dominicana Wille and Chandler
Trigona eocenica Kelner-Pillault

* Elec tr apis apoides Manning

*Electrapis meliponoides (Buttel-Reepen)

* Electrapis indecisus (Cockerell)

*Electrapis tristellus (Cockerell)

* Electrapis palmnickenensis (Roussy)

* Electrapis minuta Kelner-Pillault

* Electrapis bombusoides
Electrapis proava (Menge)

Electrapis tornquisti Cockerell
Apis cuenoti Theobald

Apis he ns ha wi Cockerell

Apis henshawi dormitans (Cockerell)

Apis henshawi kaschkei (Statz)

Apis aquitanensis de Rilly

Locality

Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Rott, Germany

Florissant, Colorado
Rott, Germany
Gard, France
Florissant, Colorado
Corent, France
Florissant, Colorado

Baltic Amber

Baltic Amber

Baltic Amber

Florissant, Colorado

Baltic Amber

Dominican Amber

Baltic Amber

Baltic Amber

Baltic Amber

Baltic Amber

Baltic Amber

Baltic Amber

Baltic Amber

Baltic Amber

Baltic Amber

Baltic Amber

Cereste, France

Rott, Germany

Rott, Germany

Rott, Germany

Aix-en-Provence, France

1978]

Burnham — Social Insects in Fossil Record

123

TABLE 5. (CONCLUDED)

Geological Age

Locality

MIOCENE

Halictidae

Halictus schemppi (Armbruster)

Randeck, Germany

Andrenidae

Andrena primaeva Cockerell

Oeningen, Germany

Megachilidae

Lithurge adamitica (Heer)

Oeningen, Germany

Megachile amaguensis Cockerell

Siberia, U.S.S.R.

Osmia antiqua Heer

Oeningen, Germany

Osmia nigra Zeuner and Manning

Oeningen, Germany

Anthophoridae

Xylocarpa jurinei (Heer)

Oeningen, Germany

Xylocopa hydrobiae Zeuner

Biebrich, Germany

Xylocopa senilis Heer

Oeningen, Germany

* Anthophorites thoracica Heer

Radoboj, Croatia

* Anthophorites longaeva Heer

Radoboj, Croatia

* Anthophorites mellona Heer

Oeningen, Germany

* Anthophorites titania Heer

Oeningen, Germany

* Anthophorites tonsa Heer

Oeningen, Germany

* Anthophorites veterana Heer

Oeningen, Germany

Apidae

Bombus abavus Heer

Oeningen, Germany

Bombus proavus Cockerell

Latah, Washington

Trigona succini (Tosi)

Sicilian Amber

Trigona sicula (Tosi)

Sicilian Amber

Trigona silacea Wille

Chiapas, Mexico

Trigona devicta Kerr and Maule

Burma Amber

Apis armbrusteri armbrusteri Zeuner

Wiirttemburg, Germany

Apis armbrusteri scharmanni (Armbruster)

Wiirttemburg, Germany

Apis armbrusteri scheeri (Armbruster)

Wiirttemburg, Germany

Apis armbrusteri scheuthlei (Armbruster)

Wiirttemburg, Germany

Apis catanensis avolii Roussi

Sicilian Amber

Apis melisuga (Handlirsch)

Italy

* Extinct genera.

124

Psyche

[March

the explanation that the insect pollinators so important to their
success were pre-adapted as pollination vectors. It is interesting to
note at this point that bees have been observed foraging on conifer
pollen in areas where other food resources are scarce. Ray Angelo
(personal communication, May, 1978) reports observing Colletes
sp. foraging in high numbers on Juniperus virginiana pollen cones.
This is noteworthy in two respects: 1) this conifer is the only readily
available pollen source in the particular habitat where observations
took place (Concord, Mass.), and 2) the bees foraging on the tree
are members of the primitive bee family Colletidae. This suggests
that they are generalized enough to have retained the ability to
forage on gymnosperm pollen. Nevertheless, the hypothesis that
bees evolved before the advent of the angiosperms is highly
speculative, and remains a difficult theory to prove. The possibility
of a pre-angiosperm origin for the bees implies that the Apoidea,
and possibly sociality in the Apoidea, may be older than indicated
by the fossil record. An inherent problem, of course, is whether or
not these early bees would be recognizable as such, or would be
mistaken for sphecid wasps. The discovery of additional Cretaceous
amber might well provide valuable insight into this problem.

SUMMARY

Wheeler writes in his 1928 book, “from the lowest to the highest
forms in the series, all animals are at some time in their lives
immersed in some society.” It is the elaboration or evolution of
these habits that leads to the eusocial behavior found in the Isoptera
and certain groups of the Hymenoptera. The preceding account has
examined insect sociality from a paleontological perspective in the
hope that it will provide insight into the antiquity of this behavioral
phenomenon. In addition, it has provided information on certain
aspects of the evolution of the four major groups of social insects.

The Isoptera are highly eusocial at the ordinal level and evidence
suggests an ancient origin for the group. The oldest fossil termite
known is from a Late Cretaceous deposit in Canada. The presence
of a distinct humeral suture at the wing base indicates that social
behavior was developed in the Isoptera at this time. It is further-
more presumed that the termites arose in the early Mesozoic or
possibly earlier, and from “protoblattoid” or blattoid stock. The
hypogaeic lifestyle of most termites is not conducive to their

1978]

Burnham — Social Insects in Fossil Record

125

preservation as fossils and this may explain their absence in pre-
Cretaceous deposits.

The first Hymenoptera appear in the Triassic and belong to the
primitive family Xyelidae (Symphyta). Social Hymenoptera are not,
however, found in the fossil record until the Upper Cretaceous. The
ant species discovered in deposits of this age are more primitive than
any now existing and have been of paramount importance in our
understanding of ant phylogeny. By the mid-Tertiary, the ant fauna
was extremely diverse; by the Miocene, the genera were essentially
modern, and geographic distribution of the ants was apparently
similar to that of today.

The Vespoidea although not very numerous in fossil deposits,
have been found as far back as the Late Cretaceous, represented by
one specimen assignable to the Masaridae. The presence of several
vespoids in Eocene deposits strongly supports the possibility that
social wasps evolved during the Late Cretaceous or Early Paleocene.

Apoidea extend into the fossil record only as far as the Oligocene,
although it is speculated that they may have evolved much earlier.
This is suggested by the fact that the bee fauna was essentially
modern by the end of the Oligocene and also because the inter-
dependence of angiosperms and bees suggests a co-evolutionary
relationship beginning sometime in the Cretaceous.

Any discussion of sociality in the geological past must necessarily
involve a certain amount of speculation. Morphological characters
play an essential role in the analysis of an insect’s social status, an
example of this being the presence of the humeral suture in
Cretatermes. In those social insect groups possessing very little
morphological variation between castes, recognition of such social
distinctions in the fossils is virtually impossible. It is generally
assumed that extinct species belonging to extant genera possessed a
similar type of social behavior in the past as is exhibited by the
group today. To speculate further about the social habits of fossil
insects is simply not possible. The mechanisms behind the evolution
of eusociality in the insects remain unknown, yet the success of this
form of social behavior is unquestioned. Only the recovery of
additional material will provide evidence to further elucidate our
understanding of the paleontological record of these insects.

As the record now stands, it is possible to state with a fair degree
of certainty that insect sociality had evolved by the middle of the
Cretaceous and perhaps much earlier.

126

Psyche

[March

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Figure 5. Geological time scale showing the distribution of twenty social insect families found in the fossil record. Dotted lines
represent age extensions based on specimens of questionable taxonomic assignment.

1978]

Burnham — Social Insects in Fossil Record

127

ACKNOWLEDGEMENTS

This study was originally intended as a brief survey of the social
insects in the fossil record but underwent rapid expansion shortly
after its initiation. This is partly due to my burgeoning interest in the
subject matter, partly due to the vast amount of material requiring
my attention, and partly due to the stimulation and encouragement
received from friends in the academic community at Harvard. It is
to the following friends that I extend my thanks and appreciation:
K. M. Horton, Paul Strother, Robert E. Silberglied, Kenneth
Miyata, and N. E. Woodley. Special thanks are given to F. M.
Carpenter for his continuous guidance and advice, for his patience
as my photographic assistant; and most of all, deep appreciation
is extended to him for providing the inspiration integral to the
success of this study. The Royal Ontario Museum, Toronto, is
gratefully acknowledged for the loan of the Eocene vespoid. In
addition, partial financial support is acknowledged to National
Science Foundation Grant DEB 78-09947 — F. M. Carpenter,
principal investigator.

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Zeuner, F. E. and F. J. Manning

1976. A monograph on fossil bees (Hymenoptera: Apoidea). Bull. Brit. Mus.
(Natur. Hist.) 27(3): 151-268.

PARENTAL CARE IN GUAYAQUILA COMPRESSA
WALKER (HOMOPTERA: MEMBRACIDAE)*

By T. K. Wood

Box 1224, Wilmington College, Wilmington, Ohio 45177
Introduction

Egg guarding or parental care is common in the Heteroptera. In
the Hemipteran families studied (Bequaert 1925, Eberhard 1975,
Melber and Schmidt 1975, 1977), most females desert their offspring
before they reach maturity. Typically, parental investment in off-
spring is provided by females, but in the families Reduviidae
(Ralston 1977) and Belostomatidae (Smith 1976), they are replaced
by males. In the Homoptera, only females in the Membracidae
(Hinton 1976, Wood 1974, 1976a, b, 1977a) and closely related
families (Brown 1975) provide parental care of offspring. My studies
(unpublished) and those by Hinton (1977) indicate female parental
care is common in this family, particularly in the new world tropics.

My studies of 3 membracid species provide evidence for 2 major
types of parental care in the family. In the 1st type, exemplified by
Umbonia crassicornis Amyot and Serville and Platycotis vittata F.,
females remain on eggs until hatch and make a series of feeding slits
for nymphs in the branch of the host plant. First instars move off
the egg mass and aggregate along these slits with the female
positioned below. Parent females actively maintain aggregated
nymphs and defend them from potential predators such as adult
coccinelids. Successful maturation in the field depends on both
nymphs and the parent female remaining together on the same
branch until offspring become adults (Wood 1974, 1976a, b).

Entylia bactriana Germar exemplifies the 2nd type of parental
investment where the role of the parent female is reduced to the
protection of eggs and the 1st two instars. Presumably, reduction of
female investment is brought by mutualistic ant associations in this
species. Although females are capable of protecting eggs and 1st to
2nd instar offspring, protection is enhanced if ants are in attendance.
When females desert 1st to 2nd instars, nymphal maturbation in the
field depends on protection provided by ants (Wood 1977a).

* Manuscript received by the editor July 26, 1978.

135

136

Psyche

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Haviland’s (1925) observations suggested females of Guayaquila
compressa Walker may protect eggs. Although detailed observa-
tions were not reported, she clearly indicates females relocate
offspring and that nymphs readily disperse when disturbed. Such
reports require detailed confirmation before attempting a systematic
analysis of the types of parental investment within Membracoidea.

Methods

The study site was a lowland wet forest at Finca, LaSelva, a field
station of the Organization for Tropical Studies in the province of
Heredia in Costa Rica. G. compressa was always found in succes-
sional or edge areas along the forest on the following host plants:
Alchornea sp., Pterocarpus officinalis, Theobroma cacao, Eu-
phorbiaceae and an unidentified vine. Branches or leaves with
insects were marked and observations were made daily for 2 to 16
days. Observations of 17 different females on eggs or with nymphs
were made during 2 separate trips in August 1976 and 1977.
Extensive attempts to increase numbers were made, but individuals
tended to be on isolated trees some distance from each other.

Results

Females on egg masses — Eggs are deposited by females in masses
surrounded or embedded in a sticky, white matrix on top of plant
tissue (Figs. 1 and 2). Two egg masses contained 78 and 88 eggs.
Ovipositional sites varied; 6 females placed eggs on the underside of
mature leaves on top of the main midrib, while 4 others placed egg
masses on branches 6 to 12 inches from the apical meristem.
Females sat on egg masses and usually faced the leaf petiole or the
apex of the branch. In 1 egg mass, eggs hatched 12 days after
deposition, while 9 others were observed for 7 to 10 days before eggs
hatched. All egg masses which hatched had females present, while 1
egg mass where the female was removed failed to hatch. A portion
of this egg mass without a female was damaged (as if eaten by a
predator) the following day and developed mold growth during the
subsequent 6 days of observation. This egg mass, deposited within 3
days of an egg mass which hatched, was followed long enough that
if there were viable eggs, they should have hatched.

1978] Wood — Parental Care in Guayaquila compressa

137

Figure 1 . Photograph of female G. compressa on egg mass deposited on plant
tissue on the underside of leaf (X7).

Females on eggs are sensitive to disturbances and will fly off egg
masses. In several cases, touching the branch or leaves caused flight,
while other females had to be repeatedly poked with a pencil before
taking flight. Six females were disturbed to the point of flight in 12
separate trials. Some of these females simply dropped from the egg
mass, but most flew off, landing on plants 5 to 15 feet away or made
a circular flight back to the host. In all 12 trials, these 6 females
found their way back to egg masses. In 5 trials, females returned
within a 24-hour period, while in the remaining 7 trials they
returned within 5 to 80 min. Females which could be observed
often moved to several plants before locating the tree with the egg
mass. Once on the host, they walked up and down branches until
finding an egg mass. As they approached egg masses, females
appeared to make fewer movements away from the egg mass,
suggesting some ability to orient to cues associated with the egg
mass (chemical or visual).

Parent female — offspring interaction — First instars associated
with 5 parent females averaged 51.8 (range 37 to 60) individuals with

138

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nymphs forming tight clusters occupying 2 to 5 linear cm of plant
surface. Egg hatch observed for 3 egg masses on the underside of the
leaf midrib was completed within a 24-hour period. In 1 egg mass,
15 nymphs after egg hatch lined up along the leaf midrib between
egg mass and leaf petiole facing the female next to the egg mass.
When the leaf was turned over, all nymphs moved toward the
female and clustered together under the egg mass until the leaf was
returned to its normal position. During the subsequent 24 hrs, the
remaining eggs hatched and all 60 nymphs with the parent female
had moved 1 foot from the depleted egg mass on the leaf to the main
woody branch. The behavior of nymphs and females on 2 other
leaves was similar. One group moved 27 inches to another large
branch within 24 hrs of egg hatch. Females which deposited eggs on
branches moved with their nymphs away from the old egg mass to
the apex of the branch, where they became associated with new
leaves.

Relocation movements of parent females and nymphs is not only
restricted to the 24 hr period after egg hatch. Four aggregations
during a 4 to 7-day observation period relocated naturally 1 to 2
times. Relocation is not simply a matter of moving to an adjacent
leaf or new shoot, but involves distances up to 3 feet in a 24-hr
period. For example, one female and apparently all her offspring
were observed to move down a main branch to a fork, then up the
2nd branch to the tip of a lateral twig.

The escape response of parent females and nymphs is different
from other treehoppers. Six marked females and their offspring on
isolated plants were disturbed by either moving the branch or
probing the female with a pencil. Each of 5 females was tested 2 to 5
times and 1 female was tested once for a total of 18 trials. Only 1
trial for a female was done each day, but some females were tested
on consecutive days. The amount of “violence” necessary to pro-
voke flight by the female varied from trial to trial. My approaching
the branch or touching it was sometimes effective, while in others,
females had to be pushed with a pencil 5 to 30 times before taking
flight. Distances females flew varied from 1 to 15 feet. Females
relocated nymphs in 9 trials within 24 hrs; in 2 trials in 1 .5 to 4.5 hrs,
but in 7 trials, they returned within 50 min. (range of 12 to 50 min.).
No one female consistently returned faster than others. Females
sometimes simply dropped down into the tree, flew off making a

1978] Wood — Parental Care in Guayaquila compressa

139

Figure 2. Female of G. compressa on egg mass [Original drawing by Sarah
Landry],

circular flight back, or flew to adjacent trees with a series of short
flights back to the host. One undisturbed female was observed to
desert and relocate offspring, but whether this was triggered by a
predator could not be determined.

Nymphs dispersed almost immediately in 12 of 18 trials after the
female was disturbed. Nymphs moved distances of up to several feet
and often reaggregated on new branches. In one trial, when the
female was probed once, she fanned her wings which produced an
audible clicking while nymphs dispersed up and down the branch.
This female did not fly off until all nymphs had dispersed. Careful
hand removal of 2 other females triggered immediate nymphal
dispersal and reaggregation within a 40-min period. When the
female was removed from 3 aggregations, nymphs remained to-
gether for 3 days.

When an injured or crushed nymph was presented, both siblings
and female responded. In 1 case, as 7 of 56 1st instars moved past
the female, she responded by a rapid twisting motion and followed
the nymphs for 6 inches. When the female started to move, the

140

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remaining nymphs dispersed within a minute and walked past her.
Lead nymphs stopped at a petiole 12 inches from the original site.
After nymphs dispersed from the petiole, the female followed them
up the branch, but then returned to the original site where 2 nymphs
remained. After a total of 10 minutes, all nymphs were on 9 leaves
on 3 different shoots. During the next 8 minutes, the female walked
back and forth at the original site before moving to the main
branch. Nymphs were still on several leaves but several groups were
increasing to the point where 1 leaf had 29 of the 56 nymphs. In the
next 70 minutes, the female repeatedly twisted laterally or walked
up and down the branch and nymphs continued to consolidate into
larger groups. Three groups were formed, one at the tip of shoot,
one at its base, and one on the next shoot up the branch. The female
after this period positioned herself below the group of nymphs at the
base of the shoot. These nymphs then moved to the tip of the shoot,
followed by the female. Within 2 min, all but 2 nymphs from the
shoot above joined the aggregation. This aggregation then remained
in the same place for 24 hours.

Single females on eggs or with offspring were typically the only
conspecifics on most host plants. However, 1 host plant about 6 feet
high had 2 females with offspring which were observed daily for 9
consecutive days. Initially, the eggs of 1 female had just hatched and
nymphs had lined up along the midrib of a mature leaf, while the
2nd female with 50 1st instars was located on the same trunk, but 12
inches below the leaf with the 1st female. During the next 24 hr,
nymphs with the 2nd female moved 2 feet up the trunk to new leaves
on the apical shoot. The 1st female and her 60 nymphs moved off
the leaf to the trunk while 10 nymphs remained on the leaf petiole.
On the 3rd day of observation, both females and their broods had
merged together on the terminal growth tip where they remained for
the next 5 days. On 2 separate days during this 5-day period, leaves
were touched with a pencil, triggering immediate movement of
several nymphs down the petiole toward the females. One or both
females responded to these nymphs by lateral twisting, back and
forth walking, or walking backwards. In both trials, some nymphs
went past females, but others which followed were stopped by
tapping movements made by a female’s prothoracic legs. In neither
trial was there massive dispersal and within 5 min., all nymphs had
reaggregated at the original site.

1978] Wood — Parental Care in Guayaquila compressa

141

On the 8th day of observation, both females were probed with a
pencil until they flew off and nymphs immediately dispersed from
the petiole to the main trunk of the plant. Within 23 min., all
nymphs had moved down the plant 2{/i feet and reaggregated along
the midrib of 3 mature leaves. Consolidation into 1 aggregation
took place during the next 17 minutes without the presence of either
parent female. Both females made a series of short flights back to
the host and at the end of 1 hr, one female had located nymphs while
the other was rapidly walking up and down branches off the main
trunk. This last female, during a 6-min. period, walked to 5 different
branches within 12 in. of nymphs. After 24 hrs, both females were
with nymphs, but had relocated 18 inches away from the previous
observation.

Adult aggregations — One aggregation of 34 teneral adults was
observed for 10 days before dispersal. I could not determine if the
parent female was present since coloration of teneral adults was
similar to that of other parent females. The number of adults
decreased during the first 5 days to 28, then to 15 during the next 4
days, with all individuals gone on the 10th day of observation. In the
first 5 days, 2 females deposited eggs, one on the same host, the
other on a host 15 feet away. I failed to locate other females on eggs
in the surrounding area during a 7-day period after complete
dispersal.

This adult aggregation was more sensitive to disturbance than
females on eggs or with nymphs. On the first day of observation, all
adults were together on the petiole of the leaf. When I accidentally
moved the tree trunk, there was an explosive, almost synchronous
dispersal with 3 individuals observed 10 feet away. In the following
60 min., individuals moved back to the host and began to form 2
aggregations near the original site. Once on the host, individuals
walked up and down branches or flew short distances until coming
to the one of the 2 groups where they stopped. At the end of 1 hr, 30
insects were in the 2 groups. In the next 48 hrs, 1 aggregation
attracted all but 5 insects from the other. When this aggregation was
disturbed again, dispersal occurred with all 28 adults back together
at a new site on the same plant within 24 hrs. Without further
disruptions, this aggregation remained in the same place for 3 days.

142

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Discussion

Female parental investment is well developed in the subfamily
Membracinae (classification of Dietz 1975), particularly the tribes
Hoplophorionini ( U . crassicornis and P. vittata) and Aconophorini
( G . compressa). Although the behavior of G. compressa is similar in
most respects to species in the Hoplophorionini, certain aspects of
nymphal behavior are similar to E. bactriana in the subfamily
Smiliinae (tribe: Polyglyptini).

Placement of eggs by female G. compressa on top of plant tissue
and held together by an accessory secretion is typical of the
Aconophorini (Hinton 1977), while in the Hoplophorionini or
Polyglyptini, eggs are inserted into plant tissue. Although all
females sit on egg masses, the insertion of eggs into, as opposed to
on top of, plant tissue may offer more protection from desiccation,
parasites or egg predators due to less exposed egg surface area.
Female G. compressa as with some Pentatomids with similar
ovipositional habits, may not be able to protect peripheral eggs
from parasites (Eberhard 1975).

Female G. compressa on egg masses are extremely sensitive to
disturbances while females of U. crassicornis and P. vittata usually
do not desert egg masses, even when given more violent treatment.
Physically displaced females of the latter 2 species can relocate egg
masses (Wood 1976b, Wood in preparation), but female U. crassi-
cornis do not recognize individual egg masses (Wood in prepara-
tion). Whether individual female P. vittata or E. bactriana recognize
their own egg masses has not been tested. Female G. compressa
return to egg masses, but this may be an artifact of this species’
patchy distribution and low population densities. A dislodged
female, which flew 10 or so feet, may encounter an egg mass which
has a high probability of being her own through random flights or
walking, but whether females recognize their own egg masses must
wait until choice tests can be made. Activity which results in egg
mass relocation by female G. compressa is adaptive since females
appear to be necessary in preventing mold growth, protection from
egg parasites or predators, and maintaining offspring aggregations.

Protective or defensive adaptations of female U. crassicornis on
eggs are cryptic coloration and lack of movement, but also involve
the shape and hardness of the pronotum (Wood 1975, 1977a).
Although mature female P. vittata are not physically protected by

1978] Wood — Parental Care in Guayaquila compressa

143

the pronotum, their cryptic coloration and lack of movement make
them difficult to find on woody branches of the host plant (Wood
1976b). The pronotum of female G. compressa is similar in shape
and hardness to that of mature P. vittata, but the black females
provide contrast to the green leaf background. Thus, rapid female
dispersal producing a startle response in a predator such as an
arboreal anole would be a viable alternative for an otherwise
unprotected female.

Herding of offspring by parent females within a host plant is
unique to G. compressa. Female U. crassicornis and P. vittata
remain with their offspring on the woody branches where eggs were
deposited until nymphs reach maturity. Eggs of E. bactriana are
deposited in leaves on herbaceous plants but nymphs move from
these leaves and reaggregate on new ones after being deserted by
females (Wood 1977a). Herding in G. compressa may permit
enhanced exploitation of the host plant by reducing localized
feeding damage.

Alarm or escape behavior actively involves both female G.
compressa and offspring, differing significantly from that of U.
crassicornis or P. vittata. Alarm displays by females of the latter
species are produced in response to injured nymphs or predators,
but nymphs do not disperse from the feeding site, nor are they
deserted by parent females (Wood 1976a, b). E. bactriana females
place themselves between predator and offspring. Nymphs remain
with females initially, but upon prolonged exposure to alarm
pheromones or injured females, nymphs disperse from the leaf.
Escape behavior of nymphs is also modified by the behavior of
various ant species which provide predator protection (Wood
1977a). Thus, the startle response produced by female G. compressa
provides time to permit nymphal dispersal. Dispersed nymphs
reaggregate with or without parent females and maintain an effec-
tive nymphal escape response if the parent female is captured or dies
from other causes.

Females in a number of membracid species deposit eggs close to
each other on a branch even when there appears to be an abundance
of oviposition space, suggesting cooperative brood care (Wood,
unpublished). G. compressa provides the first evidence to suggest an
hypothesis for the adaptive nature of this cooperation. Normal
herding behavior on a small host plant means 2 aggregations have a

144

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[March

high probability of contacting each other. Even if the females were
not related, it could be adaptive to merge aggregations to increase
the effectiveness of the startle response since 2 females dispersing
together or slightly out of phase may provide more time for
nymphal dispersal. Even if 1 female is captured, the remaining
female can facilitate reaggregation and maintenance of the aggrega-
tion. The interaction of 2 or more females may also facilitate normal
maintenance of nymphal aggregations, promoting increased feeding
efficiency and maturation.

Adult G. compressa maintain stable aggregates as do U. crassi-
cornis and P. vittata. In both of the Hoplophorionini species,
aggregations are stable for 15 to 20 or more days. During this
period, individuals within the aggregation become progressively
more sensitive to disturbances. Individual teneral adult U. crassi-
cornis are unpalatable to Anolis lizards, but in addition, employ
both an individual and collective cataleptic behavior to reduce
predation (Wood 1975, 1977b). Older aggregations disperse explo-
sively about the time of sexual maturity, but do not appear to
reaggregate. The explosive dispersal and subsequent reaggregation
of adult G. compressa is an extension of nymphal behavior and
appears to be an effective response to predators such as anoles
which are often seen walking or sunning themselves on branches
similar to those where treehoppers are found.

Acknowledgments

My thanks to Dr. Glenn Morris and Frank Hale for their help
with the field work, and Dr. Gary Hartshorn for the host plant
identifications. The faithful and useful criticisms by Dr. David Horn
on the manuscript were, as usual, very helpful. Mrs. Carole Kenney
typed several manuscript drafts and, as usual, kept her sense of
humor. Research funding was provided by NSF grants BNS
74-19764 and BNS 74-19764 A01 . I thank Ms Sarah Landry for the
excellent drawing in Figure 2.

Literature Cited

Bequaert, J.

1935. Presocial behavior among the Hemiptera. Bull. Brooklyn Entomol. Soc.
30: 177-191.

1978] Wood — Parental Care in Guayaquila compressa

145

Brown, R. L.

1976. Behavioral observations on Aethalion reticulatum (Hem., Aethalioni-
dae) and associated ants. Insect. Soc. 23(2): 99-107.

Deitz, L. L.

1975. Classification of the higher categories of the New World treehoppers
(Homoptera: Membracidae). Tech. Bui. No. 225, N. Carolina Agr. Exp.
Station, pp. 1-177.

Eberhard, W. G.

1975. The ecology and behavior of a subsocial pentatomid bug and two
scelionid wasps: Strategy and counterstrategy in a host and its parasites.
Smithsonian Contributions to Zoology, No. 205. pp. 1-39.

Haviland, M. D.

1925. The Membracidae of Kartabo. Zoologica. 6: 229-290.

Hinton, H. E.

1977. Subsocial behavior and biology of some Mexican membracid bugs.
Ecological Entomology 2: 61-79.

Melber, A., and G. H. Schmidt.

1975. Sozialverhatten zweier Elasmucha-arten (Heteroptera: Insecta). Z. Tier-
psychol. 39: 403-14.

1977. Sozialphanomene bei Heteropteren. Sonderdruck aus Zoologica. 127:
19-53.

Nault, L. R., T. K. Wood, and A. M. Goff.

1974. Tree hopper (Membracidae) Alarm Pheromones. Nature (London). 149
(5444): 387-388.

Ralston, J. S.

1977. Egg guarding by male assassin bugs of the genus Zelus (Hemiptera:
Reduviidae). Psyche. 84: 103-107.

Smith, R. L.

1976. Male brooding behavior of the water bug Abedus herberti (Hemiptera:
Belostomatidae). Ann. Entomol. Soc. Amer. 69: 740-747.

Wood, T. K.

1974. Aggregating behavior of Umbonia crassicornis (Homoptera: Membraci-
dae). Can. Ent. 106: 169-173.

1975. Defense in two presocial membracids (Homoptera: Membracidae). Can.
Ent. 107: 1227-1231.

1976a. Alarm behavior of brooding female Umbonia crassicornis (Membraci-
dae: Homoptera). Ann. Entomol. Soc. Amer. 69: 340-344.

1976b. Biology and presocial behavior of Platycotis vittata F. (Homoptera:
Membracidae). Ann. Entomol. Soc. Amer. 69: 807-811.

1977a. Role of parent females and attendant ants in the maturation of the tree-
hopper, Entylia bactriana (Homoptera: Membracidae). Sociobiology 2
(4): 257-272.

1977b. Defense in Umbonia crassicornis: The role of the pronotum and adult
aggregations (Homoptera: Membracidae) Ann. Entomol. Soc. Amer.
70: 524-528.

The illustration on the front cover of this issue of Psyche is a
reproduction of the published figure of the minute diapriid, Solen-
opsia americana (1.3 mm. long), described by C. T. Brues in Psyche
(1936, vol. 43, p. 17). The insect was taken in the nest of an ant,
Paratrechina parvula, in eastern Tennessee.

CAMBRIDGE ENTOMOLOGICAL CLUB

A regular meeting of the Club is held on the second Tuesday of
each month October through May at 7:30 p.m. in Room 154,
Biological Laboratories, Divinity Ave., Cambridge. Entomolo-
gists visiting the vicinity are cordially invited to attend.

BACK VOLUMES OF PSYCHE

Requests for information about back volumes of Psyche should
be sent directly to the editor.

F. M. Carpenter
Editorial Office, Psyche
16 Divinity Avenue
Cambridge, Mass. 02138

ISSN 0033 2615

QU

nei

PVH

Ent

A JOURNAL OF ENTOMOLOGY

founded in 1874 by the Cambridge Entomological Club
Vol. 85 June-September, 1978 No. 2-3

PSYCHE

t

CONTENTS

Geographical Distribution and Biological Observations of Cyphoderris (Or-
thoptera: Haglidae) with a Description of a New Species. Glenn K. Morris

and Darryl T. G wynne 147

Description of the Ergatoid Queen of Pogonomyrmex mayri with Notes on

the Worker and Male (Hym., Formicidae). Charles Kugler 169

Notes on Bryothinusa with Description of the Larva of B. catalinae Casey

(Coleoptera: Staphylinidae). Ian Moore and R. E. Orth 183

Grooming Behavior in Diplura (Insecta: Apterygota). Barry D. Valentine and

Michael J. Glorioso 191

The Dacetine Ant Genus Pentastruma (Hymenoptera: Formicidae). William L.

Brown, Jr., and Ronald G. Boisvert 201

Observations on a Population of Sialis itasca Ross in West Virginia (Meg-

aloptera: Sialidae). C. K. Lilly, D. L. Ashley, and D. C. Tarter 209

Structure and Relationships of the Upper Carboniferous Insect, Prochoroptera
calopteryx (Diaphanopterodea, Prochoropteridae). Frank M. Carpenter and

Eugene S. Richardson, Jr. 219

Division of Labor within the Worker Caste of Formica peripilosa Wheeler

(Hymenoptera: Formicidae). Carlos Roberto F. Brandao 229

Culture Techniques for Acanthops falcata, a Neotropical Mantid Suitable for
Biological Studies (with Notes on Raising Web Building Spiders.) Michael

H. Robinson and Barbara Robinson 239

Searching Behavior of Hippodamia convergens Larvae (Coccinellidae: Coleop-
tera). Kenneth W. Hunter, Jr 249

Further Studies of the Myrmicine Sting Apparatus: Eutetramorium, Oxyo-
pomyrmex, and Terataner (Hymenoptera, Formicidae). Charles Kugler . . 255

An Unusual Ascalaphid Larva (Neuroptera: Ascalaphidae) from Southern
Africa, with Comments on Larval Evolution within the Myrmeleontoidea.

Charles S. Henry 265

The Evolutionary Significance of Redundancy and Variability in Phenotypic-
Induction Mechanisms of Pierid Butterflies (Lepidoptera). Arthur M.
Shapiro 275

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1978-1979

President John A. Shetterly

Vice-President Barbara Thorne

Secretary Norman Woodley

Treasurer Frank M. Carpenter

Executive Committee Jo B. Winter

Margaret Thayer

EDITORIAL BOARD OF PSYCHE

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

ALFRED F. Newton, Jr., Curatorial Associate in Entomology, 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. E. SlLBERGLIED, Assistant Professor of Biology, Harvard University
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: $9.50, domestic and foreign. Single copies, $3.50.

Checks and remittances should be addressed to Treasurer, Cambridge Entomo-
logical 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 $24.50 per
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The March, 1978, Psyche (Vol. 85, No. 1) was mailed January 26, 1979

The Lexington Press, Inc., Lexington, Massachusetts

PSYCHE

Vol. 85 June-Septernber, 1978 No. 2-3

GEOGRAPHICAL DISTRIBUTION AND BIOLOGICAL
OBSERVATIONS OF CYPHODERRIS
(ORTHOPTERA: HAGLIDAE)

WITH A DESCRIPTION OF A NEW SPECIES

By Glenn K. Morris1 and Darryl T. Gwynne2
Introduction

With the exception of Prophalangopsis obscura (F. Walker) from
India, Cyphoderris are sole survivors of a primitive orthopteran
family, the Haglidae, abundant in the Triassic and ancestral to
modern Ensifera (Zeuner, 1939; Ander, 1939; Ragge, 1955; Sharov,
1968).

There are presently two recognized species of Cyphoderris: C.
monstrosa Uhler and C. buckelli Hebard. Their most dramatic
distinguishing feature is the presence in C. monstrosa, and the
absence in C. buckelli, of a prominent ventrally-directed sternal
process, shaped like the claw of a hammer and located on the IXth
sternum (Hebard, 1934). Specimens of both species have been
extensively collected from mountainous areas of the North Ameri-
can northwest.

When Uhler established Cyphoderris in 1864 he had before him
two adult male specimens. He published body measurements for
both of these and there is a substantial size difference e.g. body
length 22 mm for one specimen and only 16 mm for the other. These
specimens are in the Museum of Comparative Zoology, Harvard
University and we have examined them. The larger has a prominent

‘Erindale College and Department of Zoology, University of Toronto, Missis-
sauga, Ontario, L5L 1C6, Canada.

department of Zoology and Entomology, Colorado State University, Fort
Collins, Colorado, 80523, U.S.A.

Manuscript received by the editor November 15, 1978.

147

148

Psyche

[June-September

sternal process and is C. monstrosa of present usage; the smaller
specimen lacks the process and is C. buckelli of Hebard. It is the C.
buckelli specimen which bears a red ‘Type 792’ label. Uhler’s
description concentrates on the larger specimen. Thus monstrosa is
said to have a “pointed keel-like elevation, projected backwards
upon the segment, grooved and emarginated at tip”, i.e. a sternal
process. He does not indicate that such a structure is absent from
the other specimen.

Caudell (1904) described a variety of C. monstrosa which he
called Cyphoderris monstrosa piperi. His types, which we have seen,
are a male and a female from Mt. Rainier, Washington, housed in
the U.S. National Museum. The male has a sternal process identical
with that of Uhler’s larger specimen and the female has Ander’s
organs (see below).

In 1922 Fulton collected a series of males in Oregon about 30 mi.
southwest of Crater Lake (Fulton, 1930). He compared these with
specimens furnished him by E. R. Buckell from southern British
Columbia and found that Buckell’s specimens lacked a genitalic
process. Drawings were sent to Nathan Banks and to Caudell who
compared them with the types of monstrosa and piperi.

It might now have become apparent that Uhler’s types differed in
their genitalia and that only the larger was of the same species as
piperi. Since the published description applied substantially to the
larger specimen one would then have expected it to be designated as
monstrosa. But for some reason piperi was given specific status by
Fulton and applied to the taxon with the sternal process while
Uhler’s name was conferred upon the smaller of Uhler’s two species.
Probably it was at this point that the red type label was appended at
Harvard.

Hebard (1934), responding to Uhler’s published description,
recognized piperi as a synonym of monstrosa and gave the name
buckelli to the species without the sternal process. It is clear that he
did not examine Uhler’s types and was unaware that one of these
was his new species. Uhler did not designate a holotype and so in
accordance with Article 74 of the code and in the interest of
taxonomic stability, we here designate as lectotype the larger of the
two specimens in his type series, that possessing the sternal process.
This ensures that application of the name monstrosa continues in
conformity with present custom.

1978]

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149

A third species of Cyphoderris, C. strepitans, is described here. Its
distribution lies southeast of both monstrosa and buckelli (Figure
5), populations of strepitans being originally considered as southern
range extensions of monstrosa (Alexander, 1935; Willey and Willey,
1963). C. strepitans appears to be most similar in morphology and
calling song to buckelli but is readily distinguished from the latter
species by the structure of the male terininalia.

We have made use of the following abbreviations: ROM/ Royal
Ontario Museum, Toronto, Canada; UMMZ/ University of Michi-
gan, Museum of Zoology, Ann Arbor; ANSP/ Academy of Natural
Sciences, Philadelphia; CNC/ Canadian National Collection, Ot-
tawa, Canada; MCZ / Museum of Comparative Zoology, Harvard
University; USNM/ National Museum of Natural History, Smith-
sonian, Washington, D.C.

Cyphoderris strepitans new species
Figures 1 and 2

The specific name refers to the calling song: strepitans (Latin)
‘making a great noise’.

SYNONOMY:

Cyphoderris monstrosus, Thomas (not Uhler), 1876. Proc. Davenp. Acad. Nat. Sci.
I, p. 263. Wind River, Wyo.

Cyphoderris monstrosa, Hebard (in part not Uhler), 1934. Trans. Am. ent. Soc. 59,
p. 374. Pearl, Col.

Cyphoderris monstrosa, G. Alexander (not Uhler) 1935. Ent. News 46, p. 30. Park
Range (Pearl) Col.; ibid 1941. Univ. Col. Studies 1, p. 136.

Cyphoderris monstrosa, Willey & Willey (not Uhler) 1963. Ent. News 74, p. 200. Los
Pinos Pass, Col.

Cyphoderris monstrosa, Evans (not Uhler) 1970. Bull. Mus. Comp. Zool. Harv. 140,
p. 484. Jackson Hole, Wyo.

Holotype. Adult o. Park Range 24.7 mi. west of Cowdrey via
Pearl, nr Big Creek Lakes, Jackson Co., Col., U.S.A.; 19 June 1976;
Coll. G. K. Morris & D. T. Gwynne (Deposited ROM).

Description of male type. Body length (fastigium to para-
procts in dorsal view) 18.6 mm; pronotum inid-line length 7.2 mm;
caudal pronotum width 7.8 mm; maximum exposed tegminal length
in dorsal view 8.2 mm; length in lateral view of left metathoracic
femur 8.3 mm.

150

Psyche

[June-September

Figure 2. Adult female C. strepitans (both antennae broken).

1978]

Morris & Gwynne — Cyphoderris

151

Allotype. Adult o. Los Pinos Pass, Saguache Co., Col., U.S.A.;
17 June 1977; Coll. D. T. Gwynne (Deposited ROM).

Description of female allotype. Body length (fastigium to
extremity of epiproct in dorsal view) 21.5 min; pronotum mid-line
length 5.9 mm; caudal pronotum width 5.4 mm; length of ovipositor
2.0 mm; length in lateral view of left metathoracic femur 9.6 mm.

Diagnosis. Adult males of strepitans are similar in size and
coloration to buckelli, but readily distinguished by the presence of
the sternal process (Figures 3 & 4). Males of both strepitans and
buckelli are generally smaller than males of monstrosa. In life they
usually lack the vivid pink coloration of monstrosa" s venter, their
venters being instead cream white. The styli of the IXth sternum are
strongly depressed in monstrosa; viewed from above each stylus is
sublanceolate and broadest at its base; they are inserted on the
lamellate dorsal projection of the IXth sternum at a distance slightly
greater than the stylus length. By contrast the styli of strepitans are
distally dilated and broadly rounded (mitten-like), gently arcuate
and tapering slightly to the base; they are inserted close together
immediately adjacent to the inid-line with less than one stylus length
between their bases.

The sternal process of monstrosa, viewed in lateral outline,
follows a broadly concave arc beyond the base of the styli to where
it turns abruptly downward; in strepitans this arc is shorter and
much shallower. From the end of the arc the process of monstrosa is
more strongly reflexed than in strepitans and is often bent sharply
forward at its extremity.

We are unable at present to distinguish between females of
buckelli and strepitans but both of these species may be separated
from monstrosa by their lack of the ‘stridulatory’ organs of Ander
(1938). In C. monstrosa these structures are located dorsolaterally at
the junction of abdomen and thorax (Ander, 1938; Kevan, 1954;
Duinortier, 1963). Each organ consists of a row of robust posteri-
orly-directed recurved teeth on the slightly swollen posterolateral
edge of the metanotum. The teeth contact a patch of transverse
ridges on the first abdominal tergite during telescoping of the
abdominal segment. Ander’s organs are present in both sexes and
are readily seen in later stadia of immatures. While buckelli
sometimes possesses weak thoracic teeth, it never exhibits the ridged

152

Psyche

[June-Septernber

Figure 3. Posterior views of terminalia. A) C. strepitans, B) C. monstrosa, and C) C. buckelli. Symbols: VIIIT, 8th tergum;
IXT, 9th tergum; XT, 10th tergum; Cer, cercus; Eppt, epiproct; Papt, paraproct; Sty, stylus; IXS, 9th sternum; sp, sternal process.

1978]

Morris & Gwynne — Cyphoderris

153

Figure 4. Lateral view of IXth sternum, A) C. strepitans, B) C. monstrosa, and
C) C. buckelli.

patch. No stridulatory function has been established for these
structures.

Paratypes. 9 adult same data as holotype (ROM, MCZ, &
UMMZ); 1 adult $, same data as holotype (ROM); 4 adult
Cowdrey, 8.8 mi. west on road to Pearl, 4 June 1978 (G. K. Morris
& D. T. Gwynne) (ROM & Colorado State Univ., Ft. Collins); 1
adult (J, Park Range nr Pearl, Col., 17 Aug. 1932 (G. Alexander)
(ANSP); 1 adult $, Pearl, Col., 17 Aug. 1932 (G. Alexander)
(Museum of University of Colorado, Boulder, Col.); 7 adult
Los Pinos Pass, Col., 17 June 1977 (D. T. Gwynne) (USNM &
CNC); 1 irmn. Los Pinos Pass, Col., 17 June 1977 (D. T.
Gwynne) (ROM); 1 adult $, Dunraven Pass, Yellowstone Nat. Pk,
Wyo., 25 June 1930 (E. C. Van Dyke) (UMMZ); 1 adult $, Jackson
Hole Res. Stn, Grand Teton Nat. Pk, Wyo., Aug. 1967 (H. E. & M.
A. Evans) (USNM); 1 irmn. $, Jackson Hole Res. Stn, Grand Teton
Nat. Pk, Wyo. (H. E. & M. A. Evans) (USNM); 1 adult 9, Stratton
Exp. Watershed, nr Saratoga, Wyo. (J. M. Schmid) (USNM).

154

Psyche

[June-September

Key to Cyphoderris Species
(Males only)

1. Subgenital plate (IXth sternum) with prominent ventrally-

directed process (Figures 4a, b) 2

Sternal process absent (Figure 4c) C. buckelli Hebard

2. IXth sternum strongly produced posteriorly; sternal process
with an angularly forward-bent tip, often appearing terminally
cleft and toothed; styli of IXth sternum depressed, sublanceo-
late (Figure 3b); Ander’s organ ridge-patch present and thoracic

teeth robust; fastigiurn often weakly rugose

C. monstrosa Uhler

Posterior of sternal process not angular but rounded, never
terminally cleft and toothed (Figure 3a); Ander’s organ absent
or if present, only as weak thoracic teeth; styli of IXth sternum
mitten-like; fastigiurn smooth ... C. strepitans new species

Geographical Distribution

C. strepitans, as presently known, is confined to the mountains of
Colorado and Wyoming (Figure 5). Its distribution is disjunct from
that of buckelli and monstrosa. The broad valley of the Snake River
isolates it on the northwest from the most southerly populations of
Idaho monstrosa; if overlap occurs it must be north of Yellowstone
in southern Montana.

C. monstrosa is found from the Canadian Rockies in the south-
west corner of Alberta, west through southern British Columbia. It
reaches much farther north than buckelli, to Quesnel and to
Sinithers B.C. (This latter record exceeds the northern extent of our
map and could not be plotted; Sinithers is about 700 miles north of
the Canada/ U.S. border.) A western arm of monstrosa extends
down the Cascades, reaching almost to northern California. A less
documented eastern arm crosses western Montana to a cluster of
localities in the Salmon River Mts of central Idaho.

C. buckelli has a more restricted range. It lies between these arms,
overlapping broadly with monstrosa in southern B.C. and extending
south through northern Idaho. There are interesting isolated rec-
ords from Columbia Falls, Montana and from near Seneca in east
central Oregon. Though their distributions overlap substantially we
have not found monstrosa and buckelli together at the demic level.

1978]

Morris & Gwynne — Cyphoderris

155

Figure 5.

Geographical distribution of Cyphoderris.

156

Psyche

[June-September

However Buckell (1924) states that both species have been seen at
Nicola, B.C. “in large numbers during late May feeding together
upon flowers of Amelanchier . . .”.

We have examined about 300 specimens of monstrosa and 200 of
buckelli. In addition to the type, allotype and paratypes of strepi-
tans, we have seen 20 or so alcoholic specimens supplied by Dr. R.
Willey of the Univ. of Chicago. Dr. Willey’s material is from Los
Pinos Pass, Col., a locality which he discovered in 1962 (Willey &
Willey, 1963).

All plotted localities are based upon actual examination of
specimens excepting Wind River, Wyo. This record is taken from
Thomas (1876) on the strength of his illustration of an adult male.
Though the drawing is small the shape of the sternal process is
apparent and marks the specimen as strepitans. A listing of localities
is given below for each species.

C. strepitans: WYOMING: Dunraven Pass, Yellowstone N. Pk;
Jackson Hole, Grand Teton N. Pk.; Wind River, Fremont Co.;
Stratton Exp. Watershed, nr Saratoga, Carbon Co. COLORADO:
Park Range, nr Big Creek Lks, Jackson Co.; Cowdrey, 8.8 mi. west,
Jackson Co.; Los Pinos Pass, Saguache Co.

C. monstrosa: BRITISH COLUMBIA: Smithers; Quesnel; Chilcotin
nr Williams Lk; Lac La Hache; Clinton; Lillooet; Whistler Mt.,
Garibaldi Prov. Pk; Salmon Arm; Field, Yoho N. Pk; Glacier N.
Pk; Monck Prov. Pk; Merritt; Lumby; Peachland; Fish Lk, nr
Surmnerland; Manning Prov. Pk; Hedley. ALBERTA: Jasper N. Pk;
Mt. Eisenhower Cpgrd, Banff N. Pk; Sulphur Mt., Banff N. Pk;
Bragg Creek, w. of Calgary; Barrier Lk, Kananaskis Valley; Ka-
nanaskis Lks, Kananaskis Valley. Montana: Belton, Flathead Co.;
2 mi. s. Elliston, Powell Co. IDAHO: McCall, Valley Co.; Challis,
Custer Co.; Beach Cr., nr Bull Trout Lk, Custer Co.; Red Fish Lk,
Custer Co.; Centerville, Boise Co.; 22 mi. ne. Idaho City, n. fork
Boise R., Boise Co.; Camas Co.; Arco, Butte Co. WASHINGTON: Lk.
Wenatchee St. Pk, Chelan Co.; Entiat R. Trail, Chelan Co.; 2 mi. se.
Easton, Kittitas Co.; Stampede, King Co.; Paradise Valley, Mt.
Rainier N. Pk.; Berkeley Park, Mt. Rainier N. Pk; Gooseprairie,
Yakima Co.; Trout Lk Cpgrd, Klickitat Co. OREGON: Mt. Hood,
Hood R. Co.; Hat Point, Wallowa Co.; Middle Sister, Lane Co. &
Deschutes Co.; McKenzie Pass, Lane Co. & Deschutes Co.; Salt
Creek Falls, Lane Co.; Waldo Lk, Lane Co.; Lost Lk, Willamette
Nat. For., Linn Co.; North Santiam R., Linn Co.; Pinehurst,

1978]

Morris & Gwynne — Cyphoderris

157

Jackson Co.; Union Creek, Jackson Co.; Crater Lk N. Pk, Klamath
Co.; Douglas Co.; Olallie Lk, Marion Co.; Mt. Jefferson, Jefferson
Co.

C. buckelli: BRITISH COLUMBIA: Chilcotin; Paul Lk Prov. Pk;
Squilax; Salmon Arm; Nicola Lk; 2 mi. s. Merritt; Aspen Grove;
Vernon; Lumby; Kelowna, nr airport; 3 mi. se. Rutland; Okanagan
Falls; Rock Creek; Yahk; Ainsworth; Rosebery Prov. Pk; Cran-
brook; Boswell, Kootenay Lk. IDAHO: Reeder Bay, Priest Lk,
Bonner Co.; Sandpoint, Bonner Co.; 10 mi. s. Coeur d’Alene,
Kootenai Co.; Moscow Mt., Latah Co.; Moscow, Latah Co.;
Karniah, Lewis Co. WASHINGTON: Newport, Pend Oreille Co.;
Palouse, Whitman Co.; Pullman, Whitman Co. MONTANA: Colum-
bia Falls, Flathead Co. OREGON: 3 mi. e. Seneca, Grant Co.

Habitat and Feeding Behavior

The distribution of Cyphoderris corresponds roughly with the
Cordilleran forest province (Gleason & Cronquist, 1964). In south-
ern British Columbia C. buckelli occurs in the Dry Forest biotic
area (Cowan & Guiguet, 1965) characterized by yellow pine ( Pinus
ponderosa) and at higher elevations, by interior Douglas fir ( Pseu -
dotsuga menziesii). Amelanchier (serviceberry), Balsamorhiza (ar-
row-leaf balsam-root) and Berberis (tall Oregon grape) are common
understory plants in this association. In spring the nymphs and
adult females of C. buckelli feed upon the flowers of these plants;
night collecting at blooms is a good way to obtain specimens.

C. buckelli is also found in the Columbia Forest biotic area, the
so-called interior wet belt of British Columbia. In 1977 we located
large populations adjacent to Kootenay Lake near Boswell and at
Rosebery on Slocan Lake.

C. monstrosa does occur in Dry Forest e.g. at Monck Prov. Park
in southern British Columbia, but it is typically encountered in Sub-
alpine Forest. Lodgepole pine {Pinus contorta) and Englemann
spruce ( Picea engelmannii ) are characteristic of the Sub-alpine
biotic area. In the Kananaskis Valley of southern Alberta we
observed nymphs and adults to feed upon staininate cones of
lodgepole pine (this before the cones reach a ‘loose pollen’ stage).
Consumption was established by identifying cone bracts in the feces
of field-caught specimens. Also caged insects were given cones and
in most cases overnight they ate large portions.

158

Psyche

[June-September

It is presumably to feed upon staminate cones that C. monstrosa
nymphs and adult females are observed at dusk climbing high into
the trees. A useful method of collection is to search tree trunks with
a flashlight just after sunset. The insects are always discovered
oriented head upward. More often than not they occur in groups of
2 to 4 on the same trunk, which suggests that they may aggregate
during their daytime stay in the leaf litter of the forest floor.

David Lightfoot of Oregon State University has studied C.
monstrosa at Three Sisters in the central Oregon Cascades. He
found this species abundant there in drier, more open stands of
lodgepole pine and mountain hemlock ( Tsuga mertensiana) above
5000' elevation. In contrast to our observations of ascent as the
evening progresses Lightfoot notes that the singers begin high in the
trees (about 6 in) and gradually occupy lower and lower perches
until singing at ground level.

C. strepitans is found in both subalpine forest and high altitude
sagebrush prarie. The holotypic site near Big Creek Lakes is an open
forest of subalpine fir (Abies concolor ) and lodgepole pine at an
altitude of 8800'. At Los Pinos Pass, Colorado, strepitans is found
in aspen woods adjacent to open areas of prairie. The mature aspen
(Populus tremuloides ) has an understory of subalpine fir and
englemann spruce (altitude 10,200'). Two predominant ground
cover plants at both sites are kinnikinik (Arctostaphylos spp.) and a
shrubby juniper (Juniperus communis). In the high altitude (8400')
sagebrush (Artemesia tridentata) prairie of North Park, Colorado,
the density of singing males appeared to be much greater than in the
nearby pine forest of the holotypic site to the east. C. strepitans is
also very numerous in the sagebrush areas (altitude 6700') of Grand
Teton National Park, Wyoming. In late June, 1978, aggregations of
singing males were easily heard while we drove along park roads at
night. Thus, C. strepitans may be considered a predominantly
sagebrush species although occurring in open forest habitats in the
vicinity of sagebrush prairie.

Acoustic Behavior

Males of all three species produce a succession of short musical
trills, beginning in late evening and continuing well past midnight if
weather permits. C. buckelli invariably sing near the ground from
low shrubs (knee-height), the bases of tree trunks or on the forest

1978]

Morris & Gwynne — Cyphoderris

159

floor itself. The same is true of C. strepitans. Only C. monstrosa
climb high into the trees as the night’s signalling progresses. At
Monck Park singing heights in excess of 5 m were common and an
hour after sunset collection without climbing trees becomes im-
possible.

The calling songs are generated by tegminal stridulation. As in
Gryllidae the tegmina are morphological mirror-images, both left
and right bearing a functional file and scraper. Unlike gryllids
however, which maintain a characteristic ‘right above’ forewing
overlap, the overlap of a Cyphoderris male may change during his
lifetime and both files take part in his stridulation.

Certain Tettigoniidae also have mirror-image tegmina and two
functional files: Megatympanon speculatum Piza (Listroscelinae)
(Riek, 1976), Neduba macneilli Rentz & Birchim, Neduba sierranus
Rehn & Hebard (Decticinae) (Morris et al., 1975). Most tettigoniids
have structurally distinct left and right forewings and overlap them
‘left above’. In the Neduba species some individuals show left above,
some right above. Unlike Cyphoderris they appear to maintain their
particular overlap as individuals through life. Both overlaps were
represented by Riek’s two (pinned) specimens of M. speculatum.

Spooner (1973) analysed the calling song of C. monstrosa and
describes it as a trill of grylloid (sinusoidal) pulses at a carrier
frequency of 13 kHz. He noted substantial variation in the intensity
and frequency of pulses and suggested that these changes “reflect
irregular switching of tegmina from top to bottom position”. He
refers to this habit as “switch-wing singing” and regards it as
occurring several to many times in the course of a single trill.

Overlap at rest (i.e. between singing bouts) is very infrequently
changed in C. buckelli. The overlap of 16 individually-caged males
was monitored by examining them once a day during almost 2
weeks. Of 141 checks, only 4 reversals from the immediately
previous overlap were observed; the incidence of resting overlap
reversal was less than 3%. Thirteen of these males never showed an
overlap reversal.

Four C. monstrosa males checked over 5 days, gave similar
results: two were never found with reversed overlap (checked
respectively 5 and 6 times), one was reversed once in 5 checks and
one twice in 6 checks. If Cyphoderris alter overlap several times
within a single trill, it is strange that individuals end up so
consistently at the same overlap with which they began.

160

Psyche

[June-September

We recorded the calling song of a C. monstrosa specimen (Figure
6, 75-6) before and after damaging with a scalpel, several teeth in
the central region of his right tegmen file. In oscillograms of post-
mutilation recorded song, his use of the damaged file (i.e. right
above overlap) was apparent as a drastic mid-pulse drop in ampli-
tude. In one oscillogram, a portion of which makes up Figure 6
(second trace from bottom), 20 pulses in succession were ‘right
above’.

Switch-wing singing as suggested by distinctive pulse envelopes
within the same trill was only evident in our records on one
occasion. A male of C. monstrosa had been released in the
immediate vicinity (i.e. within antennal range) of a mature female
on the observer’s hand. He began to sing while walking about on the
hand and directing his attention toward the female. His song was
recorded and on analysis found to be a trill in which every other
pulse was identical in envelope and distinctly different from the
intervening pulse i.e. there were two pulse types occurring in
alternation without break in the sequence of the trill (Figure 6,
bottom trace). This was apparently a courtship song.

It is clear that pulse envelopes are highly variable in the genus,
though usually quite consistent for a particular recording session of
a particular individual. Switch-wing stridulation is probably not an
everyday feature of C. monstrosa calling song but it may occur
under special circumstances such as courtship.

Oscillograms of normal calling songs are given in Figure 6. The
pulses of C. strepitans and C. buckelli are apparently indistinguish-
able. They are usually wedge-shaped: each begins with a steep rise to
maximum amplitude, then falls steadily to the pulse’s end. The
pulses of C. monstrosa also have a steep onset but are usually of
longer duration. They are drawn out in an uneven envelope near
their maximum amplitude before dropping away to silence.

Carrier frequency spectra of all three species are highly similar.
Specimens were analysed ‘live’ (i.e. without tape-recording) by
directing the output of a Bruel & Kjaer sound level meter (2204)
fitted with a !4" microphone (4135) into a Tektronix 3L5 spectrum
analyser. This system will detect ultrasonic frequencies up to 100
kHz. No substantial sound energy exists in the ultrasonic range for
any of the Cyphoderris species. The sinusoidal nature of the
waveform is apparent in the narrowness of the dominant frequency

1978]

Morris & Gwynne — Cyphoderris

161

Figure 6. Oscillograms of Cyphoderris calling song. From top trace downward:
C. strepitans 76-8, 17.8° C; C. buckelli 76-14, 25.0° C; C. monstrosa 75-6, 24.5°C,
intact file; C. monstrosa 76-2, 8.7° C, field-recorded, Monck Prov. Pk B.C.; C.
monstrosa 75-6, 24.0° C, file teeth of right forewing damaged; C. monstrosa 75-10,
23.1°C, male recorded singing in immediate presence of female.

162

Psyche

[June-September

peak, suggesting the operation of a sharply-tuned (high Q) tegminal
resonator (Sales & Pye, 1974).

In the figured C. strepitans male (Figure 7), the dominant peak
centers on 12.7 kHz and there are very weak second and third
harmonics near 25 and 38 respectively. The C. buckelli specimen has
its principal peak near 13.3 kHz and a lesser peak occupies the range
28-30 kHz. Like Spooner (1973) we obtained 13 kHz as the
dominant carrier frequency of C. monstrosa.

Sound level measurements were obtained with the ‘/^'microphone
and the 2204 meter, the latter on ‘linear, fast’ setting. At 5 cm dorsal
aspect, the sound level of C. strepitans (76-7) was between 100.5 and
101.0 dB. A specimen of C. buckelli (76-3) was 102 ± 2 dB at 6.5 cm
dorsal.

Pulse rate varies linearly with temperature (Figure 8) as in other
acoustic Ensifera (Walker 1962, 1975). Both field and laboratory
recordings of calling song contributed to the regression lines. One C.
monstrosa plotted point is from Spooner (1973) (S in Figure 8); 5

Figure 7. Calling song frequency spectrograms traced from photographs.

1978]

Morris & G wynne — Cyphoderris

163

different males provide the other 6 points. C. bucked? s regression is
based on 12 different individuals, 3 at two different temperatures
each. Each of the 13 C. strepitans pulse rates derives from a different
individual; all those at temperatures of 8°C and below are field
recordings. Pulse rates were calculated from an oscilloscope display
in which a single beam sweep embraced 3-13 pulses. Successive,
single-sweep samples (3-6), were averaged to obtain each plotted
value. The coefficients of determination indicate a very good fit to
the calculated regression lines. Although the C. monstrosa regres-
sion line is different from the strepitans and buckelli lines the slopes
and Y intercepts of the latter two species are not significantly
different.

C. strepitans males stridulate at very low temperatures. Previous
reports cite minimal singing temperatures for acoustic Orthoptera
of about 7°C [e.g. Fulton (1925) for the tree cricket Oecanthus
fultoni (under the name of O. niveus) and Frings and Frings (1957)
for the katydid Neoconocephalus ensiger ]. On May 17, 1977, at the
holotypic site, one of us (D.T.G.) heard three of four males singing
from branches and logs near the ground when the air temperature at
waist level was —0.5° C. On June 4 and 5, 1978, tape recordings were
made of males singing at temperatures as low as 2° C (see Figure 8).
Following the recording the thermometer bulb was placed close to
the singing male’s perch. There is a suggestion in the plotted rates in
Figure 8 of a departure from linearity at very low temperatures.

In conclusion, the song of C. monstrosa differs from the other
two species in both the shape of the pulse amplitude envelope and in
pulse rate, both these parameters being useful diagnostic features.
C. buckelli and C. strepitans, however, have virtually identical
calling songs: song intensities, carrier frequencies, and pulse ampli-
tude envelopes provide no basis for human discrimination; the pulse
rates, especially, are indistinguishable at any given temperature.

It is interesting to note that Alexander (1969) has questioned the
traditional interpretation that reproductive isolating mechanisms
evolved to prevent “mating mistakes” between species. He suggested
(citing evidence from acoustical insects) that species differences
have most likely arisen as a result of the different selection pressures
operating on populations while they are in allopatry. He reasoned
that if this is so, among other things, we should rarely find identical
pair forming signals among allopatric or allochronic species. C.
strepitans and C. buckelli are allopatric (Figure 5) and by the above

164

Psyche

[June-September

1“ 1 1 1 1 T"

5 10 15 20 25 30

TEMPERATURE °C

Figure 8. Relation of pulse rate and temperature in Cyphoderris calling song
(S is from Spooner, 1973).

1978]

Morris & Gwynne — Cyphoderris

165

reasoning their songs should have diverged yet this is not the case.
Any difference between these two species (including the above
mentioned habitat differences) apparently have not affected their
pair forming signals.

Acknowledgements

These studies were supported by operating grant 4946 from the
National Research Council of Canada. Appreciation is extended to
the many persons who provided us with access to specimens: R. D.
Alexander (Univ. of Michigan), W. F. Barr (Univ. of Idaho), D. R.
Davis & A. B. Gurney (Smithsonian, Washington), K. Goeden
(Oregon Dept, of Agriculture), J. D. Lattin & D. C. Lightfoot
(Oregon State Univ.), J. E. H. Martin (Biosystematics, Ottawa), D.
Otte (Acad. Nat. Sciences Philadelphia), G. G. E. Scudder (Univ. of
B.C.), Margaret Thayer (Harvard), W. J. Turner (Washington State
Univ.), Bob Willey (Univ. of Chicago), S. M. Ulagaraj and V. R.
Vickery (Macdonald College) and G. B. Wiggins (Royal Ontario
Museum). We are grateful for the help of the staff of the Kananaskis
Environmental Centre in Alberta (Univ. of Calgary). Jim Fullard,
Darlene Morris and E. J. Morris helped with field studies. T. J.
Walker (Univ. of Florida) and T. Alloway (Univ. of Toronto)
transported specimens. Dr. Howard E. Evans criticized the manu-
script. Beyond all others we acknowledge the help of Ron Aiken of
the Univ. of Toronto.

References

Alexander, R. D.

1969. Comparative animal behavior and systematics. Syst. Biol. Proc. Internat.
Conf. Nat. Acad. Sciences, pp. 494-517.

Alexander, G.

1935. Orthoptera new to Colorado. Entomol. News 46: 30.

Ander, K.

1938. Ein abdominales Stridulationsorgan bei Cyphoderris (Prophalangopsi-
dae) und iiber die systematische Einteilung der Ensiferen (Saltatoria).
Opuscula Entomol. 3: 32-38.

1939. Vergleichend-anatomische und phylogenetische Studien iiber die Ensi-
fera (Saltatoria). Opuscula Entomol. Suppl. II, 306 pp. (Unpublished
translation by T. H. Hubbell.)

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Buckell, E. R.

1924. Additions and corrections to the list of British Columbia Orthoptera.
Proc. Entornol. Soc. British Columbia 21: 7-12.

Caudell, A. N.

1904. The genus Cyphoderris. J.N.Y. Entornol. Soc. 12: 47-53.

Cowan, E McT. and C. J. Guiguet.

1965. The Mammals of British Columbia. Handbook No. 11, British Colum-
bia Provincial Museum, 3rd ed., 414 pp.

Dumortier, B.

1963. Morphology of sound emission apparatus in Arthropoda. In Busnel,
R.-G., ed.. Acoustic Behaviour of Animals. Elsevier, Amsterdam, pp.
277-345.

Evans, H. E.

1970. Ecological-behavioral studies of the wasps of Jackson Hole, Wyoming.
Bull. Mus. Comp. Zool. 140: 451-51 1.

Frings, H. and M. Frings.

1957. The effects of temperature on chirp-rate of male cone-headed grasshop-
pers, Neoconocephalus ensiger. J. Exp. Zool. 134: 411-425.

Fulton, B. B.

1925. Physiological variation in the snowy-tree cricket, Oecanthus niveus
DeGeer. Ann. Entornol. Soc. Arner. 18: 363-383.

1930. Notes on Oregon Orthoptera with descriptions of new species and races.
Ann. Entornol. Soc. Arner. 23: 611-641.

Gleason, H. A. and A. Cronquist.

1964. The Natural Geography of Plants. Columbia Univ. Press, New York,
420 pp.

Hebard, M.

1934. Cyphoderris, a genus of katydid of southwestern Canada and the north-
western United States. Trans. Am. Entornol. Soc. 59: 371-375.
Kevan, D. K. McE.

1954. Methodes inhabituelles de production de son chez les Orthopteres. In
Busnel, R.-G., ed., L’Acoustique des Orthopteres. Ann Epiphyt., Fasc.
Special, pp. 103-141.

Morris, G. K., R. B. Aiken and G. E. Kerr.

1975. Calling songs of Neduba macneilli and N. sierranus (Orthoptera: Tet-
tigoniidae: Decticinae). J.N.Y. Entornol. Soc. 83: 229-234.

Ragge, D. R.

1955. The Wing-venation of the Orthoptera Saltatoria with Notes On Dicty-
opteran Wing-venation. British Museum Natural History, London,
159 pp.

Riek, E. F.

1976. The Australian genus Tympanophora White (Orthoptera: Tettigoniidae:
Tyrnpanophorinae). J. Aust. Entornol. Soc. 15: 161-171.

Sales, G. and D. Pye.

1974. Ultrasonic Communication by Animals. Chapman and Hall, London,

281 pp.

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Sharov, A. G.

1968. Phylogeny of the Orthopteroidea. Trans. Paleontol. Inst. Acad. Sci.
U.S.S.R. 118: 1-216. (English ed., Israel Prog. Sci. Transl. 1971,
pp. 1-251.)

Spooner, J. D.

1973. Sound production in Cyphoderris monstrosa (Orthoptera: Prophalan-
gopsidae). Ann. Entornol. Soc. Arner. 66: 4-5.

Thomas, C.

1876. A list of Orthoptera, collected by J. Duncan Putnam, of Davenport,
Iowa, during the summers of 1872-3-4 & 5, chiefly in Colorado, Utah
and Wyoming Territories. Proc. Davenport Acad. Nat. Sci. 1: 249-264,
pi. 36.

Uhler, P. R.

1864. Orthopterological contributions, Gryllodea. Proc. Entornol. Soc. Phil.
2: 543-555.

Walker, T. J.

1962. Factors responsible for intraspecific variation in the calling songs of
crickets. Evolution 16: 407-428.

1975. Effects of temperature on rates in poikilotherm nervous systems:
evidence from the calling songs of meadow katydids (Orthoptera: Tet-
tigoniidae: Orchelimum ) and re-analysis of published data. J. Comp.
Physiol. 101: 57-69.

Willey, R. B. and R. L. Willey.

1963. Range extension of Colorado Cyphoderris (Orthoptera: Prophalangop-
sidae). Entornol. News 74: 200-201.

Zeuner, F. E.

1939. Fossil Orthoptera Ensifera. British Museum Natural History, London,
321 pp.

DESCRIPTION OF THE ERGATOID QUEEN OF
POGONOMYRMEX MAYRI WITH NOTES ON THE
WORKER AND MALE (HYM., FORMICIDAE)*

By Charles Kugler
Department of Entomology
Cornell University
Ithaca, New York 14853

Introduction

During a recent stay in Santa Marta on the north coast of
Colombia, I had the opportunity to study Pogonomyrmex mayri,
the sole member of the subgenus Forelomyrmex, whose entire range
is the desert and dry deciduous forest below 200 m. on the
northwestern, western, and possibly southern skirts of the Sierra
Nevada de Santa Marta.

This ant was described by Forel (1899: 61-62, footnote) from
worker(s) and male(s) he collected. Neither he nor subsequent
entomologists, including P. J. Darlington, found females. The
reason females were unknown became clear as I worked in the area
and later began to look at the biology of P. mayri more closely.
Though I collected males from vegetation nearly year around (3
Sept, to 30 June), no winged females were seen in two years.
Furthermore, only after thoroughly excavating 10 nests were any
females found at all, one in each of 2 nests dug by my coworker,
Maria del Carmen Hincapie, and her assistant. Both were ergatoid
nest queens.

This paper presents a formal description of those queens, notes on
the worker and male supplementing Forel’s original description,
and a discussion of the taxonomic status of Pogonomyrmex mayri.
Notes on the biology of P. mayri will be reported later.

*A report of research of the Cornell University Agricultural Experiment Station,
New York State College of Agriculture and Life Sciences at Cornell University,
Ithaca, New York 14853.

Manuscript received by the editor October 20, 1978.

169

170

Psyche

[June-September

Queen

(Figs. 1-4, Table 1)

Two specimens (Museum of Comparative Zoology at Harvard
University) taken by Maria del Carmen Hincapie from nests in soil
behind the beach at Bahia Gairaca, about 20 km. by road east of
Santa Marta, Colombia, in Parque Nacional Tayrona, on 2 dif-
ferent days, April 22, 1978 (#780422-6) and April 26, 1978
(#780426-9). The following description is based primarily on the
first specimen, with occasional remarks on the second if they differ
in some way. For each measurement, that of the first specimen is
followed by that of the second.

Mandibles subtriangular; basal and masticatory borders meet at
obtuse angle; outer border gently convex, slightly flattened mid-
length; masticatory border with 5 teeth increasing in size apically,
the second tooth with a denticle on its distal edge (second specimen
with denticle reduced on one mandible, absent on the other). Palpal
formula 3,3 as determined from undissected specimens; neither palp
reaching posterior margin of buccal cavity. Labrum with rounded
lobes on either side of a sharp median emargination. Head (fig. 4)
about as wide as long; in full face dorsal view occiput broadly V-
shaped, with distinct occipital lobes; sides of head gently convex on
temples to small bulge behind the eye, then indented to flat cheeks.
Middle of clypeus weakly convex overall, both transversely and
longitudinally; sculpture stops a weak median carina short of
reaching apron; apron narrow, leading edge convex with median
tooth bracketed by low rounded lobes; lateral arms form low
narrowly rounded ridges in front of antennal fossae. Frontal
triangle reduced to a narrow, deep Y-shaped sulcus separating
clypeus from posterior % of frontal lobes. Frontal lobes short, well
separated; carinae convex. Eyes small, convex, ringed by a round-
bottom sculptured groove.

Antennae 12-merous. Scape short and rather thick, not reaching
vertex of head; bent at a nearly right angle within basal !4 of length
and largely flat beyond (extensor surface viewed from side). First 2
segments of flagellum longer than wide, L 0.21, 0.20; 0.18, 0.18;
segments 3-7 wider than long; segments 8-1 1 about as long as wide.
Apical segment much longer than any other, L 0.34, 0.37; apex
narrowly rounded. Last 4 segments constitute a weakly defined

1978]

Kugler — Pogonomyrmex mayri

171

Figs. 1-3. Queen of Pogonomyrmex mayri. Fig. 1, dorsal view. Fig. 2, lateral
view. Fig. 3, lateral view of trunk and waist (waviness of some hairs is an artifact of
charging).

172

Psyche

[June-September

antennal club. Width of flagellar segments increases uniformly from
first (0.13, 0.12) to last (0.18, 0.20).

Trunk in full dorsal view (fig. 1) with complete, fine, immobile,
promesonotal suture; mesonotum bisected at about 2/3 of length
by a faint shallow sulcus (probable remnant of scutoscutellar suture)
which drops out at lateral margins; metanoturn reduced to a sharp
groove between mesonotum and propodeuin. Propodeal spines
robust, acute, their bases widely separated by a broad concavity.
Propodeal spiracles prominent; their orifices circular, facing pos-
terolaterad.

From the side (figs. 2, 3) dorsal profile convex; mesonotum set off
by grooves, its outline higher and more convex than pronotum and
propodeum. Distinct sutures separate pronotum and mesonotum
from mesopleura; metapleura delimited by weak sulci scarcely
distinguishable from the sculpture, expecially in areas anterior to
the spiracle. Metapleural gland bulla well developed. Inferior pro-
podeal plates broad, blunt. Dorsal face of propodeum curves
broadly into declivous face.

Petiole robust (figs. 1-3), with broadly conical anterior peduncle
and weakly differentiated posterior peduncle. Venter seen from
below with 3 longitudinal ridges running from short acute tooth to
posterior peduncle. Node in side view with flat anterior face (except
for 2 low undulations) joining anterior peduncle at about a 135°
angle, meeting dorsal face of node at an acute angle; dorsal face flat,
broadly curving into short posterior face. Outline of node from
above ovoid, slightly wider than long, widest just caudad of
midlength. Anterior edge sharp (with low upturned burr in second
specimen), gradually softening to broadly rounded along sides.

Postpetiole large, inflated; connection with gaster broad. In
lateral view, dorsum strongly and evenly convex; venter much
shorter than dorsum, profile undulate, anterior lip broadly rounded.
From directly above (not as in fig. 1) node subtrapezoidal; posterior
edge a broad convex arc somewhat flattened mesad, posterior
corners broadly rounded, sides flat, tapered to corners of weakly
concave anterior edge.

Gaster greatly enlarged, subglobose, heavily sclerotized, slightly
longer than wide and slightly wider than deep; widest at midlength.
Sclerites not fused, but rigidly articulated. Measured from side
along axis, first tergum covers 90% of length of gaster; first sternum
53%.

1978]

Kugler — Pogonomyrmex mayri

173

Legs neither long and slender nor very robust. Femur lengths
front to back 1.48, 1.50; 1.45, 1.45; 1.64, 1.65 rnm., not incrassate
(W/L 18, 18; 17, 18; 16, 16%). Hind tibia length 1.33, 1.48 min.
Middle and hind tarsi each with one short simple apical spur. Hind
metatarsus 1.16, 1.29 mm.; tarsal claws simple.

Most of body with fine dense punctulate striae overlain by a
coarse broken wavy sculpture (figs. 1-4): clypeus, top, sides and
back of head, trunk (except those areas listed below), petiolar node
and postpetiole. In other places the overlying sculpture is much
reduced or absent, leaving largely flat surfaces of fine, punctulate
striae: gula (some weakly undulate sculpture present), anterior face
of pronotum, prosternum, forecoxae, dorsomedial surfaces of meta-
pleural gland bullae, declivous face of propodeum (striae coarser,
less dense), most of first gastric segment (striae weaken caudad).
Striation much reduced, leaving predominately fine, densely punc-
tate surfaces on: antennae (striae present but finer than elsewhere
and not predominate), legs except first coxae, and end of first and
most of succeeding segments of gaster (narrow coriaceous margins
on each). Mandibles finely longtitudinally striate, without punc-
tures. Inner surface of mandibles, peduncle and venter of petiole
smooth and shining; the latter with scattered punctures. Striation
more or less longitudinal on mandibles, antennae, clypeus, dorsum
and sides of head, mesonotum, sides of trunk (though much
confused in parts), coxae, anterior face and sides of petiolar node,
sides and venter of postpetiole, and gaster. Striation essentially
transverse on gula (bisected by median longitudinal ridge), pro-
notum, basal and declivous faces of propodeum (striae converge on
apices of spines from all sides), dorsal surface of petiolar node,
anterior and dorsal faces of postpetiole (transversely arcuate).

Whole body, except peduncle of petiole, covered with short, stiff,
erect, acute golden hairs, interspersed with shorter, more flexuous
recurved hairs. Apron of clypeus with longer flexuous hairs; hair on
mandibles more decumbent. No psainmophore of any sort. Color
uniformly dark ferruginous brown, except for brown to yellow
apical antennomere.

Worker
(Figs. 6-8)

The worker is most strikingly different from the queen in its jet
black color, its larger head, and smaller waist and gaster. It appears

174

Psyche

[June-Septernber

Figs. 4-5. Fig. 4, Head of queen of P. mayri, nearly full dorsal view. Fig. 5, side
of head and trunk of P. mayri male.

1978]

Kugler — Pogonomyrmex mayri

175

as if there is a longitudinal gradient of allornetric growth in the
queen such that the anterior half grows slower than, and the
posterior half faster than in the worker. A detailed description of
how the worker differs from the queen follows (see also table 1).

Head much larger, more elongate (fig. 6). Mandibles longer, with
6 distinct teeth and a low basal angle. Papal formula 4, 3. Median
carina complete to marginal tooth, but low; seen best from postero-
dorsal view. Eye smaller relative to size of head. Trunk more
completely fused; profile evenly convex, with poorly differentiated

Figs. 6-8. Worker of P. mayri. Fig. 6, head in full dorsal view. Fig. 7, trunk and
petiole in lateral view. Fig. 8, dorsal view of petiole.

176

Psyche

[June-September

neck and declivous propodeal face (fig. 7). Propodeal spiracle not
prominent; orifice oval, facing caudad. Superior propodeal spines
longer, spiniform; their bases narrowly separated. Inferior pro-
podeal plates short, spiniform. Petiole (figs. 7, 8) low, narrow;
anterior peduncle very slender in dorsal view, dorsoventrally cunei-
form; no posterior peduncle; venter with 2 longitudinal ridges.
Anterior ridge of petiolar node drawn out into an acute, dorso-
ventrally flattened, somewhat upturned tooth; sides of node seen
from above parallel, flat in anterior half, weakly convex in posterior
half. Postpetiole smaller than in female, especially in width and
height; more conical in dorsal view. Gaster much smaller in all
dimensions. Legs longer, more slender; middle and hind tibial spurs
finely pectinate. Sculpture on dorsum of trunk not broken by
sutures; transverse striae on anterior pronotum become longitudinal
on mesonotum, then transverse again on propodeum. Striation on
first tergum of gaster fades at about midlength; much of caudal half
smooth and shining. Color uniformly dull black in mature workers;
callows dark ferruginous brown. Petiole, postpetiole and gaster less
densely hairy than in female; color of erect hairs can vary from
black to golden on any one individual.

The sting apparatus of the worker is most like those of P.
( Ephebomyrmex ) naegeli and P. ( E .) imberbiculus (Kugler, 1978):
anterior apodeme of spiracular plate wide and of uniform width
along entire length of plate; lancets with 2 distinct barbs and no
dorsal ridge; triangular plate without dorsal and medial tubercles;
other parts as shown in Kugler (1978) figs. 1.8, 19, 21, 22, and 26.

Male
(Fig. 5)

Measurements (ranges from 5 individuals, including the largest
and smallest available from 7 collections) TL 7.29-8.10; HL 1.56-
1.74; HW just behind eyes 1.10-1.25 (Cl 70-73); eye L 0.31-0.34;
scape L 0.34-0.42; combined L of 2nd and 3rd flagellar segments
0.81-0.90; ML 0.12-0.19; WL 2.17-2.40; front wing L 4.00-4.10;
hind femur L 1.86-2.00; petiole L 0.80-0.94; postpetiole L 0.72-
0.93; gaster L 1.74-2.10.

Head remarkably elongate and flattened behind (fig. 5). Clypeus
rather long, broadly convex in both dimensions, middle subsiding
evenly to sides; free margin broadly arcuate except for slight median

1978]

Kugler — Pogonomyrmex mayri

177

flattening or ernargination, which possesses in some specimens a
very small acute or rounded tooth that is a continuation of a weak
median carina running the length of the clypeus. Frontal triangle
triangular, with a broad, shallow, V-shaped impression. Frontal
carinae reduced to rims encircling the sockets of the antennae.
Palpal formula 4, 3. Scape much shorter than combined length of

Table 1. Measurements and indices of Pogonomyrmex mayri females and
workers. Data for the first female specimen are listed first, followed by those of the
second. Worker data are ranges from 6 individuals selected to include the largest and
smallest available from 7 collections. Measurements are expressed in millimeters;
indices in percentages, and both follow the standard definitions (see Brown, 1953:
11-14; 1975: 3-4). Head width was measured just behind the eyes. Postpetiolar and
gaster measurements were taken separately, and from dorsal view.

Table 1

Measurements

TL

HL

HW

ML

Eye L

Scape L

WL

Petiole L
Petiole W
Postpetiole L
Postpetiole W
Gaster L
Gaster W
Fore femur L
Fore femur W
Hind femur L
Hind femur W
Indices
Cl
MI
SI

Scape W/L
Fore femur W/L
Hind femur W/L
Petiole W/L
Postpetiole W/L
Gaster W/L

Queens
8.21, 8.49
1.57, 1.58
1.45, 1.52
0.25, 0.27
0.25, 0.25
1.12, 1.20
2.02, 2.00
0.90, 0.90
0.72, 0.68
0.94, 0.94
1.25, 1.20
2.53, 2.80
2.18, 2.14
1.48, 1.50
0.27, 0.27
1.64, 1.65
0.27, 0.26

92, 96
16, 17
73, 77

15, 13
18, 18

16, 16
80, 75
133, 128
108, 107

Workers

7.85-9.04

2.00- 2.24
1.73-1.96
0.26-0.35
0.26-0.30
1.67-1.82
2.20-2.54
0.90-1.00
0.36-0.43
0.68-0.80
0.59-0.70
1.80-2.24
1.38-1.67

2.00- 2.20
0.30-0.35
2.32-2.62
0.30-0.34

85-88
13-16
90-96
11-12
15-16
12-13
39 54
79-88
70-78

178

Psyche

[June-September

second and third flagellar segments. Scape and flagellum fairly
densely covered with short erect hairs; segments 2-12 of flagellum
also very densely endowed with fine appressed pilosity.

Trunk uniquely shaped and proportioned, as if the propodeum
has grown forward, compressing the mesoscutum and pronotum,
and rotating the neck and head to a more ventral position (fig. 5).
Pronotum very constricted mesad. Mesonotum very short (meso-
scutal L/WL 20-23%);* seen from the side evenly and rather
strongly convex, from above nearly equilaterally triangular; with 2
short black lines that indicate underlying apophyses of the notauli.
Middle of scutellum and metanoturn raised into a prominent
subcircular bulge. Propodeum elongate (propodeal L/WL 49-52%),*
dorsal face curving insensibly into declivous face; unarmed. Petiole
long, cylindrical, nodeless. Legs long and slender (hind femur
L/ WL 83-86%); front coxae long, compressed front to back (fig. 5).

Wings evenly covered with fine hairs. Venation of fore wing
variable. Of 154 wings (77 individuals) from 7 collections, the most
common venation had a small closed discoidal cell with subequal
sides, a hexagonal cubital cell with a narrow opening to the costal
cell and sinuate lower edge (Rs), radial cell open distad, and veins
Rs and M unjoined by a cross vein beyond the discoidal cell (121
wings). In 1 1 wings the discoidal cell was open distad, but otherwise
the same. In 20 wings the discoidal cell was closed, but a cross vein
(r-m) connected the Rs and M veins, creating a second cubital cell.
That cross vein joined the Rs well proximad of the end of the first
cubital cell, except in one wing where it was almost even with the
end of the cell. Two wings had both open discoidal cells and the r-m
cross vein. Wings of different venation commonly appear on the
same individual.

Hypopygium in ventral view subtriangular; apex broadly rounded,
proximal corners square with slender truncate lateral projections,
middle of base with a long slender truncate anterior process.
Gonostyli (=parameres) of genital capsule tapered in side view, but
with apices broadly rounded; setae occur only around apices.
Digitus long, slender, blunt, strongly down-curved; not reaching to
apex of gonostylus. Cuspis short, pollicate when seen from the side.

'Mesoscutal and propodeal lengths measured from lateral view by taking their
maximum length along lines parallel with Weber’s length.

1978]

Kugler — Pogonomyrmex mayri

179

Aedeagus fairly slender, with serrate ventral margin; serrations
decrease in size to apex. Apex narrow, blunt; half smooth, half
finely serrate. Inner dorsal margins of gonocoxites form a long
narrow, gently convex V.

Sculpture like that of worker and female, with the following
exceptions: Striation on head largely transversely arcuate caudad of
antennae, clypeus without coarse undulations. Trunk and petiole
without overlying broken wavy sculpture; all but pronotum longi-
tudinally striate. Striation on postpetiole gives way to purely
punctate sculpture in caudal half. Gaster, first and second anten-
nomeres, and legs smooth and shining, except for weakly striate fore
coxae.

Discussion

Pogonomyrmex mayri clearly belongs to the genus Pogono-
myrmex as presently constituted; and is most closely related to
members of the subgenus Ephebomyrmex. It is most like Ephebo-
myrmex in 15 of the 22 characteristics used by Cole (1968) to
distinguish males and workers of the subgenera Ephebomyrmex and
Pogonomyrmex, the sting apparatus most resembles that of the
Ephebomyrmex species I have examined (Kugler, 1978), and some
of its most unusual characters, such as the elongate head of the
male, the Y-shaped frontal triangle of the queen and worker, and
the circumocular groove, may be seen as the extreme development
of characteristics of Ephebomyrmex species. Nevertheless, it is
remarkably different from any known Pogonomyrmex, with a
number of novel characters, and consequently has been placed in its
own subgenus ( Forelomyrmex Wheeler) since its description. The
following shows how other Pogonomyrmex species compare with
mayri9 s most distinctive features.2

The clypeus in most Pogonomyrmex has a concave leading edge
except in angustus, darlingtoni, odoratus, schmitti and townsendi
(all Ephebomyrmex ), none of which has a median tooth. The frontal
triangle is usually broadly triangulate, but is somewhat elongate,
laterally compressed, and except for a median carina, depressed

2Except where indicated, based on the MCZ collection containing 51 of the
estimated 67 presently standing species, subspecies and varieties of the subgenera
Ephebomyrmex and Pogonomyrmex.

180

Psyche

[June-Septernber

below the level of the clypeus only in darlingtoni, saucius and
schmitti. In none of these, however, is the front so narrow as in
mayri, nor is it at all Y-shaped. The back of the head in full dorsal
view is broadly and shallowly concave in most Ephebomyrmex
species, but is only excavated to such a degree that it has definite
occipital lobes in the majors of some subgenus Pogonomyrmex
species, e.g., badius. The cephalic index approaches that of mayri
only in cunicularius (84), an undetermined species near cunicularius 3 4
(84, 87), odoratus (85-87), and angustus (86-89). Only in the latter is
the occiput at all concave. In some species the sculpture is flattened
at the edge of the eye, but only in darlingtoni does it become at all
impressed, and then it is shallow, only weakly defined, and limited
to the dorsal half of the eye. All species examined have much larger
eyes, relative to the size of the head, than mayri. The petiolar node
in most Pogonomyrmex species is well rounded on top, sides and
apex, and has a distinct posterior peduncle. A few species have a
broad subacute to acute apex, but only in the sp. near cunicularius
does the node even superficially resemble that of mayri. On closer
examination, it also is quite distinct. Sculpture in the subgenus
Ephebomyrmex tends to be “coarsely rugo-reticulate” (Cole 1968:
35), but no species examined has the overlying broken, undulate
pattern of mayri.

Ergatogyny has occasionally been reported in Pogonomyrmex,
but most specimens are only occasional aberrations in species with
normal queens, e.g., comanche, maricopa, subnitidus, calif ornicus
(Cole 1968: 175), and pima (MCZ). Only one female has been
reported for cunicularius, and it is ergatoid (Santschi, 1931), but the
description indicates nothing more remarkable about it than a more
or less distinct scutellum. Ergatogyny seems to be the rule in
laticeps. Kusnezov (1951: 273-275) describes the range of ergatoid
forms, but makes no mention of enlarged waists or gasters, or of
reduced heads. Those characters are evidently unique to mayri.

The bizarre form of the P. mayri male is also apparently
unequalled in this genus (Cole, 1968; Creighton, 1952; Gallardo,
1932; Kusnezov, 1949, 1951). Some males of Aphaenogaster species
have elongate heads constricted behind. In the other Pogonomyrmex

3Two specimens collected by W. L. Brown in Argentina: Catarnarca, Cat. (airport),

4 Feb. 1967; Prov. Tucurnan, Krn 1333, Rte. 9, N. of Tapia, 25 Jan. 1967, bosque
chaqueno.

1978]

Kugler — Pogonomyrmex mayri

181

species I was able to examine directly, the head is at best only
slightly longer than wide in some species (Cl range of 4 Ephebo-
myrmex species 89-97). The mesoscutum is long (mesoscutal L/WL
33-50%), propodeum short (propodeal L/WL 26-37%) and the
petiole has a distinct node. The legs are shorter than in mayri (hind
femur L/WL 61-76%), and the front wing usually has a r-rn
crossvein that joins the Rs at or distal to the end of the first cubital
cell (see Cole, 1968: 25-26, plate 1 fig. 1). If the r-m vein is absent,
the first cuboidal cell is open.

Pogonomyrmex mayri may in fact deserve full generic status, but
for the present it seems prudent not to create a monotypic genus
before Pogonomyrmex is completely revised. The most recent
revisions (Kusnezov, 1951; Cole, 1968) have been regional in scope
and thus inadequate to address the question of whether the sub-
genera Pogonomyrmex and Ephebomyrmex are really two distinct
genera. Should such a split occur, mayri would no doubt be placed
in the separate genus Forelomyrmex.

Acknowledgements

I thank Norman F. Johnson for taking the SEM photographs,
Janet M. Hahn for helping prepare various stages of the manuscript,
W. L. Brown, Jr. for his advice and constant support, and E. O.
Wilson for the loan of specimens from the MCZ. This research was
supported by the NSF grant DEB-22427 (W. L. Brown, Jr.,
principal investigator).

References

Brown, W. L.

1953. Revisionary studies in the ant tribe Dacetini. Am. Midi. Nat. 50: 1-137.

1975. Contributions toward a reclassification of the Formicidae. V. Ponerinae,
tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichini,
and Aenictogitini. Search, Agrculture, Cornell Univ. 5(1): 1-116.

Cole, A. C.

1968. Pogonomyrmex Harvester Ants, A Study of the Genus in North
America. Univ. of Tennessee Press, Knoxville. 222 pp.

Creighton, W. S.

1952. Studies on Arizona ants (3), the habits of Pogonomyrmex huachucanus
Wheeler and a description of the sexual castes. Psyche 59: 71-81.

Forel, A.

1899. Biologia Cent. -Am., Insecta, Hym. 3 (Formicidae), pp. 1-169 + 4 pi.

182

Psyche

[June-September

Gallardo, A.

1932. Las Hormigas de la Republica Argentina, subfarnilia Mirmicinas,
segunda seccion Eurnyrmicinae, Genero Pogonomvrmex Mayr. An.
Mus. Argent. Hist. Nat. 37: 89-170.

Kugler, C.

1978. A comparative study of the myrmicine sting apparatus (Hymenoptera,
Formicidae). Stud. Entornol. 20: 413-548.

Kusnezov, N.

1949. Pogonomyrmex del grupo Ephebomyrmex en la fauna de la Patagonia.
Acta Zool. Lilloana 8: 291-307.

1951. El genero Pogonomyrmex Mayr. (Hym., Formicidae). Acta Zool.
Lilloana 11: 227-333.

Santschi, F.

1931. L’etude des fourmis de l’Argentine. An. Soc. Cient. Argent. 112: 273-282.

NOTES ON BRYOTHINUSA WITH DESCRIPTION OF
THE LARVA OF B. CATALINAE CASEY
(COLEOPTERA: STAPH YLINIDAE)*

By Ian Moore and R. E. Orth
Division of Biological Control, University of California
Riverside, 92521

A number of insects are restricted to special marine habitats along
the California seashore. At least one of these habitats is intertidal in
the sense that its fauna is regularly submerged by the tides. This is
the fauna of the reefs. Several species of staphylinids are known
from this region. Members of the staphylinid genera Liparocephalus,
Amblopusa and Diaulota are found on the rocky shores of northern
California; while in southern California only members of the genera
Diaulota and Bryothinusa are known. Several studies have dealt
with these insects including Chamberlain and Ferris (1929) and
Moore (1956a, 1956b). Larvae of some of the species have been
described. The larva of Bryothinusa catalinae Casey is described
and illustrated for the first time in this paper.

Bryothinusa catalinae (Fig. 1), the type species of the genus, was
described by Casey in 1904. Sawada (1955, 1971) described four
species from Japan under the name Halaesthenus. Several species
were described from the harbor at Hong Kong by Moore and
Legner (1971) and Moore, Legner and Chan (1973). Finally,
another species was made known by Moore and Legner (1975) from
the Gulf of California, bringing the total number of known species
to eleven. All but one of these is restricted to an intertidal habitat;
the exception being B. fluenta Moore et al. which was found by Tai-
Din Chan in a strictly fresh water habitat in a stream emptying into
Hong Kong harbor.

Bryothinusa catalinae is not common in collections probably
because of its restricted habitat. Derham Giuliani collected ten adult
specimens of this species incidental to other work in October, 1976,
near White Point, San Pedro, California, and presented the speci-
mens to us. We made two trips to the area in November and
December, 1977, where we found both adults and larvae fairly
common. This special habitat may be described as follows:

* Manuscript received by the editor September 15, 1978.

183

184

Psyche

[June-September

1

2

Figures 1-2. Bryothinusa catalinae Casey. Fig. 1, Habitus of imago; California,
Los Angeles Co., White Point, Dec. 6, 1977, R. E. Orth. Fig. 2, Habitus of larva;
same locality data as imago.

1978]

Moore <& Orth — Bryothinusa

185

White Point City Park, San Pedro, Los Angeles, California (Fig.
9, photograph), occupies approximately a mile of a narrow strip of
cliff and rocky beach along the Pacific Ocean just a few miles north
of Point Firmin. Access to the beach is by a steep road down the
cliff which joins a road along the beach. The beach area consists of
four rocky embayrnents between five short points. The second
embayment from the south is somewhat protected from the open
ocean by a 6 foot to 10 foot high ledge along the seaward side of
which is the ruin of a concrete wall. Between the concrete wall and
the road at the base of the cliff is a shallow reef which is exposed at
low-water and under water at high tide. The area is largely a field of
boulders two or three feet across with smaller stones and gravel in
sand. The staphylinids and their larvae were found beneath and on
the stones in an association with dense worm tubes, chitons, limpets,
small abalones, flat worms, small crabs and brittle stars. They were
most readily collected by a flotation method in which stones were
agitated in a bucket of seawater. The insects floated to the surface
where they were lifted with a camel’s hair brush and transferred to a
vial of alcohol. Collecting in the intertidal zone is best done in the
fall and winter when low tides occur during daylight.

The larva of Bryothinusa catalinae Casey
* (Fig. 2)

Color. Semitransparent-white except the apices of the mouth-
parts and the larger setae which are nebulously brown, and black
eye-spots on the head.

Head round, widest just behind the eye-spots, neck absent. Ocelli
apparently absent. With an ovoid heavily pigmented eye spot on
each side near the base of the antenna. Clypeal margin (Fig. 6) with
four small teeth. Antenna three-segmented (Fig. 5); first segment
wider than long; second segment about as wide as first, almost twice
as long as wide, apex with two articulated processes, each about
one-third as wide as apex of second segment; one process an “acorn-
like seta”, bears no seta on its surface, is almost twice as long as the
other and is somewhat sinuate; the other process which is the actual
third antennal segment, bears several ordinary setae and is about
twice as long as wide. Maxilla (Fig. 7) with the stipes longer than the
palpus, outer half of inner margin with closely placed short teeth.
Maxillary palpus three-segmented; first segment about as long as

186

Psyche

[June-September

Figures 3-8 Larva of Bryothinusa catalinae Casey. Fig. 3, left mandible; Fig. 4,
labium; Fig. 5, right antenna; Fig. 6, anterior margin of clypeus; Fig. 7, right maxilla;
Fig. 8, urogomphi and pseudopod.

1978]

Moore & Orth — Bryothinusa

187

wide; second segment somewhat narrower and almost as long as
first; third segment narrower and longer than second, tapered to
bluntly pointed apex. Mandible (Fig. 3) curved, pointed, with a
large internal apical tooth followed by four blunt denticles in the
apical half. Ligula (Fig. 4) rounded at apex, about as long as wide,
about as long as first segment of labial palpus. Labial palpus (Fig. 4)
two-segmented, segments longer than wide, second segment a little
narrower than first, tapered to bluntly pointed apex. Gular sutures
most approximate in the middle, widely divergent ahead and
behind.

Thorax. — Pronotum almost twice as wide as long, apical margin
straight, sides gently arcuate, base straight, angles narrowly rounded,
with a few scattered setae. Mesonotum and metanotum very similar
to pronotum.

Abdomen with the first six segments nearly parallel sided, seventh
segment narrower than sixth, eighth segment narrower than seventh.
First six segments with two discal setae, a row of about eight setae
along the basal margin and scattered setae at the sides. Eighth
segment with distinct raised opercula for an osineteriuin, the area

Figure 9. Low tide at White Point City Park, Los Angeles County, California,
habitat of Brythinusa catalinae Casey.

188

Psyche

[June-September

not pigmented. Urogornphus (Fig. 8) two-segmented, first segment
broadly triangular, second segment slender, nearly cylindrical,
about as long as the first segment. Membrane of apex of pseudopod
(Fig. 8) with four small dark hooks, each with a bifid base.

Length 3.0 mm.

Material examined. Two specimens 9 November 1977 and three
specimens 6 December 1977, White Point, San Pedro, Los Angeles,
California, associated with numerous adults under stones in coarse
sand and gravel on the seashore at about the 1 or 2 foot tide level
collected by R. E. Orth. In company with these specimens were
adults of Diaulota harteri Moore, the only other staphylinid genus
known from this habitat.

Notes. The larvae of Bryothinusa can be distinguished from
those of Diaulota , the only other staphylinid genus known from this
habitat in southern California, by the presence of four small
chitinized pigmented hooks with a bifid base imbedded in the apical
membrane of the pseudopod. Specimens of larvae examined by us
associated with adults of B. sawadai Moore et al., B. Hongkongen-
sis Moore et al., B. sinensis Moore et al. and B. chani Moore and
Legner all have pseudopodal hooks. Similar pseudopodal hooks
have been reported in the larvae of Alianta incana Erickson by
Paulian (1941). Paulian’s illustrations indicate they may be present
in other genera.

Literature Cited

Casey, T. L.

1904. On some new Coleoptera including four new genera. Can. Ent. 36:
312-324.

Chamberlain, J. C. and G. F. Ferris

1929. On Liparocephalus and allied genera (Coleoptera: Staphylinidae). Pan-
Pac. Ent. 5: 137-143, 153-162, illus.

Moore, Ian

1956a. A revision of the Pacific Coast Phytosi wirh a review of the foreign
genera (Coloptera: Staphylinidae) Trans. San Diego Soc. Nat. Hist. 12:
103-152, illus.

1956b. Notes on some intertidal Coleoptera with descriptions of the early stages
(Carabidae, Staphylinidae, Malachiidae). Trans. San Diego Soc. Nat.
Hist. 12: 207-230, illus.

Moore, Ian and E. F. Legner

1971. Bryothinusa chani, a new species of marine beetle from Hong Kong
(Coleoptera: Staphylinidae). Coleopt. Bull. 25: 107-108.

1978]

Moore & Orth — Bryothinusa

189

1975. A study of Bryothinusa (Coleoptera: Staphylinidae), comparing a
tabular key and a dichotomous key to the species. Bull. So. Cal. Acad.
Sci. 74: 109-112, illus.

Moore, Ian, E. F. Legner and T. Chan

1973. A review of the genus Bryothinusa with descriptions of three new species
(Coleoptera: Staphylinidae). Ent. News 84: 73-81, illus.

Paulian, Renaud

1941. Les premier etats des Staphylinoidea. Etudes de morphologie comparee.
Mem. Mus. Nat. Hist. Natur. Paris, n. ser. 15: 1-361, illus.

Sawada, K.

1955. Marine insects of the Tokara Islands. VIII. Family Staphylinidae
(Coleoptera). Publ. Seto Marine Biol. Lab. 5: 81-87, illus.

1971. Aleocharinae (Staphylinidae, Coleoptera) from the intertidal zone of
Japan. Publ. Seto Marine Biol. Lab. 19: 81-109.

GROOMING BEHAVIOR IN DIPLURA
(INSECTA: APTERYGOTA)

By Barry D. Valentine and Michael J. Glorioso
Departments of Zoology & Entomology respectively,

The Ohio State University, Columbus, Ohio 43210

Insect grooming studies are adding an important new dimension
to knowledge of comparative behavior and evolution. Recent
advances include an overview of a few selected movements of insects
and myriopods (Jander, 1966), studies of the functional morphology
of grooming structures (Hlavac, 1975), extensive reports about
individual orders (Coleoptera: Valentine, 1973; Hymenoptera: Far-
ish, 1972), quantitative studies at species levels (Chironomidae:
Stoffer, in preparation; Drosophila: Lipps, 1973), and many less
inclusive works. All such studies have difficulties which include the
inability to know when an observed sequence is complete, the
enormous number of potential taxa, the problem of generalizing
about families and orders from small samples of individuals or
species, and the absence of data from primitive or odd groups which
may be critical for interpreting evolutionary sequences. The first
three difficulties can be partially solved by increasing sample sizes
and combining observations; however, the fourth can be solved only
by availability. Grooming in the apterygote order Diplura is a good
example because we can find only incomplete reports on one
species. Recently, we have studied ten live specimens representing
two families and three species; the data obtained provide an
important picture of grooming behavior in one of the most primitive
surviving orders of insects. Our observations greatly extend the
limited discussion of grooming in the European japygid Dipljapyx
humberti (Grassi, 1886) reported by Pages (1951, 1967). Data on
Dipljapyx are incorporated here, but have not been verified by us.

Initially we asked two questions: The first concerned whether a
very primitive insect would enable us to observe a primitive
grooming repertory; what we actually observed were primitive
insects with grooming behavior beautifully tuned to a special and
restricted environment. The second question concerned the effects

*Manuscript received by the editor September 26, 1978.

191

192

Psyche

[June-September

of endognathous mouthparts on grooming. The invaginated, non-
condylar mandibles and maxillae of Diplura might reduce their
effectiveness in oral cleaning, and result in an increased importance
of leg rubbing movements. In fact, leg rubbing was seldom observed.
The rarity of rubbing has two possible explanations: either en-
dognathy does not significantly modify grooming or else most leg
rubbing movements have not yet evolved in Diplura.

Material Examined

Campodeidae (seven specimens and seven hours of recorded
observations plus about five additional hours of non recorded
observation which add no new data) Ohio, Franklin Co., Columbus,
Upper Arlington, 20 September, 1975, B. D. Valentine family, in
soil in back yard (1 specimen). Same data except 5 November, 1977,
in soil under boards and logs in back yard (6 specimens). Many
additional specimens were seen and collected by breaking up clods
of dirt in a garden.

Japygidae (three specimens and nine hours of recorded observa-
tion plus about four more hours which duplicate previous data).
Alabama, Butler Co., 2 mi. N.W. McKenzie on U.S. rte. 31, 7
December, 1975, B. D. Valentine, R. L. Stoffer, A. J. Penniman, in
rich humus under leaf litter (1 specimen). Ohio, Franklin Co.,
Columbus, 23 October, 1977, M. J. Glorioso, under large flat rock
at base of overgrown hill (1 specimen). Same data except 24
October, 1977 (1 specimen).

The campodeids key to the genus Campodea subgenus Campodea
Westwood, 1842, in Paclt (1957). Silvestri (1933a) and Pack (1957)
list two species of this subgenus occurring east of the Mississippi
River, Campodea (C.) fragilis Meinert, 1865, and Campodea (C.)
plusiochaeta Silvestri, 1912. Both are illustrated and described by
Silvestri (1912). Our specimens more closely match C. plusiochaeta
because the cereal setae are fairly long on all segments, as opposed
to the long basal and shorter distal cereal setae of C. fragilis, and
because there are bifurcate antennal setae, as opposed to the serrate
or plumose setae of C. fragilis. Nevertheless, the determination is
not firm and the specimens should be listed as Campodea ( Campo-
dea) ? plusiochaeta Silvestri, 1912. The Ohio japygids key in Pack
(1957) to the genus Metajapyx Silvestri, 1933. Using Smith and
Bolton (1964) they key to Metajapyx subterraneus (Packard, 1874)

1978]

Valentine & Glorioso — Grooming in Diplura

193

which is recorded from Ohio, Pennsylvania, Kentucky, Virginia,
and District of Columbia. It is the only species recorded from Ohio;
our Franklin County specimens constitute a new northern-most
record in the state, and are one of the very few American records of
the genus in glaciated territory. The Alabama japygid keys (in Paclt,
1957, and Smith and Bolton, 1964) directly to Metajapyx steevesi
Smith and Bolton, 1964, known from Mississippi, Alabama, Geor-
gia, South Carolina, North Carolina, Tennessee, and Virginia. Our
record is especially noteworthy because it marks the southernmost
limits of both the species and the genus in North America.

Results

CLEANING. Involves grooming with the mouthparts.

Antenna Clean. Passage of the antenna through the mouth is
accomplished in two major modes: unassisted and assisted. In
unassisted, which is the usual mode in Diplura, the antenna deflects
into the mouth due to its intrinsic musculature, and the legs are not
involved. In Campodea this movement is vertical to the substrate,
the antenna is curled ventrally under the head and is chewed by the
mouthparts; in japygids the movement is rarely vertical, the antenna
usually is curled along a more horizontal plane from an initial
position lateral of the head, and is usually drawn rapidly through
the open mouthparts; less frequently it is chewed by the maxillae. In
the much rarer assisted mode, the ipsilateral foreleg pulls the
antenna into the mouth and in both families is either returned to the
substrate or held in mid-air; in addition, the japygids were occasion-
ally observed using the ipsilateral foreleg to help hold the antenna in
the mouth by placing the leg crosswise in front of the mouthparts.
Pages (1967) points out that in Dipljapyx the foreleg holds the
antenna during chewing by the maxillae, but is not used when the
antenna is drawn through the maxillae without chewing movements.

Palp Clean. A maxillary palp is passed unassisted through the
mouthparts in the anterior mode in which the palp tip projects
posteriad and is drawn anteriad out of the mouth. This was
observed clearly in Metajapyx. (In Campodea, maxillary palpi are
one segmented and the labial palpi are vestigial.)

Foreleg Clean. A foreleg is raised and extended forward while
the head turns to the side to reach it; the leg is essentially in a ventro-
lateral position during cleaning, and is drawn posteriorly through

194

Psyche

[June-September

the mouth, tarsal claws last. This occurs in both families and all
three genera.

Midleg Clean. A midleg is brought forward alongside the body
and the head turns and dips to reach it, the limb moving posteriorly
through the mouth, tarsal claws last. There are three modes: under
L\, in which the foreleg is raised out of the way, in both families; L\
pull, in which the raised foreleg is used to pull the midleg into
the mouth, seen rarely in Campodea and reported in Dipljapyx by
Pages (1967); and over L\, where the foreleg remains on the
substrate and the midleg crosses above is, seen in Metajapyx.

Hindleg Clean. A hindleg is brought forward alongside the
laterally arched body and the head turns and dips to reach it, the
limb moving posteriorly through the mouth, tarsal claws last.
There are three modes in Diplura: under L\+2, where fore and mid
legs are raised out of the way, in both families; under L\, over L2,
which is self-explanatory and occurs in both families (in this mode
both families usually raise and partly extend Li, and japygids some-
times flex Li and position it under the body); and L\ pull, in which
the foreleg helps pull the hindleg to the mouth, in Campodea and
Dipljapyx.

Fore-Midleg Clean. Ipsilateral fore and midlegs are passed
simultaneously through the mouth in anterior-posterior sequence.
This infrequent action occurs in both Campodea and Metajapyx.
Sometimes both tarsi are involved, but usually the fore tarsus and
mid tibia are the parts cleaned.

Fore-Hindleg Clean. As above, the ipsilateral limbs moving
posteriorly through the mouthparts, observed rarely in Campodea.

Mid-Hindleg Clean. As above, except that the movement seems
to be a rare continuation of Hindleg Clean, under L\+2, where the
midleg becomes involved; in no case was the movement initiated
independently of Hindleg Clean. This movement was observed
rarely in Metajapyx.

Body Clean. Both families can bend double and use their mouth-
parts to groom body surfaces from the thorax to the cerci. These
movements are less frequent than other grooming, so it is not
known if the differences between the two families are real or
sampling error. Watching these animals, the observer rapidly gets
the impression that they can probably reach any body part they wish
except the pronotum. At present, the campodeids have been seen
cleaning all three coxae with the head directed ventro-posteriad;

1978]

Valentine & Glorioso — Grooming in Diplura

195

they also clean the lateral edge of the body, the styli, and the cerci
with the body curled laterally. Cereal grooming techniques appear
to be very diverse and are more controlled by position and substrate
irregularities than by a stereotyped program. For example, the cerci
can be held by Li, or by Li+i, or by Li+2, in each case the remaining
ipsilateral legs are under the cercus; other variants involve L3 raised
out of the way, L2 raised out of the way, and the cercus positioned
over all three ipsilateral legs. Body cleaning in japygids extends at
least from the mesonotum or mesosternum to the cerci, including
dorsal, lateral, and ventral surfaces; during cereal grooming, the
mouth can work the outer margin of a forceps from base to apex,
around the tip, then the inner margin to and across the anal area,
and out the inner margin of the contralateral forceps to its tip; the
far outer margin is not groomed until the insect straightens and
bends to the opposite side. In Dipljapyx Pages (1967) reports that
the thoracic legs hold the abdomen when the body is tightly curved
to clean from the mesothorax to the fifth abdominal segment.

rubbing. Involves progressive contact of body parts with each
other or with the substrate. In Diplura, all rubbing is of low
frequency.

Antenna- Foreleg Rub. The fore tarsus or tibia is used to rub the
dorsal surface of the ipsilateral antenna. This occurs in Campodea
where the movement is confined to the basal antennal segments, and
is sometimes combined with and precedes Antenna Clean, assisted.

Head-Foreleg Rub. In Campodea, the fore tarsus is used to rub
the venter of the head and the mouthparts; in Dipljapyx, Pages
describes head capsule rubs but does not indicate the areas involved.

Head-Midleg Rub. Also in Campodea, a midleg is used to rub
the venter of the head.

Head-Substrate Rub. Dipljapyx was observed rubbing the labial
region of the head on the substrate with a sideways motion.

Body-Midleg Rub. In Metajapyx, the midleg is used to rub the
dorsal and lateral surfaces of the thorax.

Body- Midleg- Midleg Rub. Also in Metajapyx, this is the bilat-
eral version of the previous movement, both midlegs rubbing
different thoracic regions simultaneously.

Body-Hindleg Rub. In Metajapyx, the hindleg is occasionally
used to rub the dorsal or lateral surfaces of the thorax. Body Rubs
can be combined, for on one occasion the thorax was rubbed

196

Psyche

[June-September

simultaneously by a mid and hind leg from opposite sides.

Body-Substrate Rub. Pages (1967) reports that Dipljapyx rubs
the thoracic sternum and abdominal base energetically on the
substrate. He recognizes that this may be territorial marking, but
believes that grooming is more probable.

Discussion

The grooming patterns of Campodeidae and Japygidae are
basically similar with one major exception. In Antenna Clean,
campodeids chew the antenna with the maxillae during passage
through the mouth, while japygids usually open the mandibles and
then scrape the antenna rapidly through the open maxillae without
chewing motions. Japygids can also chew the antenna but do so less
frequently. The differences in grooming suggest different maxillary
structures. Dissection of Metajapyx reveals an extraordinarily
complex lacinia with five pectinate lamellae along the mesal face.
Since we believe that these lamellae are the structures with which the
antennae are cleaned; their distribution and function should be
considered in future dipluran studies. Illustrations of these struc-
tures can be seen in the following works.

Japygidae: (note that the last genus is sometimes listed in a separate
family).

Indjapyx crivellari (Silvestri) as Parindjapyx (Silvestri, 1932, fig.
XXXI, 4).

Burmjapyx major (Grassi) as Japyx (Silvestri, 1922, fig. IV).

Metajapyx confectus Silvestri (Silvestri, 1947, fig. 2).

Monojapyx simplex profusa Silvestri as Japyx (Silvestri, 1932,
fig. XXI, 2).

Catajapyx confusus (Silvestri) as Japyx (Silvestri, 1929, fig. 2-5).

Heterojapyx gallardi Tillyard (Snodgrass, 1935, fig. 79).

Evalljapyx hubbardi (Cook) as E. sonoranus (Silvestri, 1947,
fig. 3).

Parajapyx isabellae (Grassi) (Paclt, 1957, fig. 37).
Anajapygidae:

Anajapyx vesciculosus Silvestri (Silvestri, 1905, fig. 4).

Anajapyx hermosus Smith (Smith, 1960, fig. 9).

Projapygidae:

Symphylurinus stangei Smith (Smith, 1960, fig. 7).

1978]

Valentine & Glorioso — Grooming in Diplura

197

The general localities of these twelve species are, in sequence: Is.
Rhodes; Mediterranean; Washington, D.C.; Greece; Greece; Aus-
tralia; se. Arizona; semi-cosmopolitan; Italy; California; Mexico;
Mexico.

The remaining two families of Diplura: Procampodeidae and
Campodeidae are described by Paclt (1957, p. 5) as having tongue-
like processes (“languettes”) on the lacinia, but lacking pectinate
lamellae, while Smith (1960) simply states that the two campodei-
form families are without “pectens”. In Procampodea the lacinial
apex has a mesal row of 4 projections, one of which is bifid; these
structures are illustrated by Silvestri (1905b: pi. XII, fig. 21). The
functional morphology of dipluran mouthparts is further compli-
cated by another feature: the presence of an antebasal serrate
prostheca (sometimes called a “lacinia mobilis”) on the mandibles of
campodeids (see Paclt, 1957, fig. 7), anajapygids and projapygids
(see Smith, 1960, figs. 10, 13 respectively), and anteapically on
procampodeids (see Silvestri, 1905b, pi. XII, fig. 19, 20). The
functions of these maxillary and mandibular structures have not
been demonstrated, but based on our observations, grooming is one
of the most probable uses of the pectinate lamellae.

Jander (1966-842) states that grooming “. . . the antennae and all
of the legs with the mouthparts ... is ... to be regarded as the
primordial mode of grooming . . .” in tracheate arthropods. It is true
that oral cleaning movements predominate in diversity and fre-
quency in primitive taxa, but it is also true that virtually all
primitive arthropods have rubbing movements too. In most cases it
is impossible to decide objectively which came first.

Many factors affect grooming, and all act on both primitive and
derivative taxa. For example, grooming movements have con-
straints imposed by body flexibility and degree of leg movement.
The configuration of a coxa and its cavity can be primitive or
derivative, but superimposed on this basic structure are the results
of selection for plane of leg movement, rotation, strength, speed,
body height, and grooming requirements. The resolution of these
diverse pressures must result in a morphological compromise which
affects grooming capability, but has little to do with primitiveness.
Additional examples are numerous. An elongate, flexible, soft-
bodied organism has different grooming patterns from a fatter,
more rigid, sclerotized organism; one with easily abraded scales will

198

Psyche

[June-September

be different from one with firm setae; and an interstitial inhabitant
will be different from a subcortical or leaf-litter inhabitant. The
point is that all of these kinds of organisms occur in Apterygota and
all are among the most primitive known hexapods.

In Diplura, grooming of the antennae, mid- and hindlegs involves
at least fourteen cleaning positions, all of which appear to be
satisfactory. This diversity is quite remarkable and is unequaled in
other insects (Valentine, unpubl.). The grooming of dipluran fore-
legs involves only one mode. The stereotypy of foreleg grooming
contrasts sharply with the diversity of antennal, mid- and hindleg
grooming. The logical explanation is that the single foreleg tech-
nique works in most or all situations, while no one technique works
for the other appendages. Environmental constraints appear to
require that the insect reach and groom its antennae, mid- and
hindlegs in several alternate ways. Diplura are basically interstitial
organisms. Almost all specimens were found in the soil under
undisturbed stones or boards, or in soil clods in gardens. A standard
technique for finding campodeids was to break up the damp clods in
a freshly plowed field or while digging potatoes. The very fine
tunnels and cracks in this unyielding substrate are inhabited prin-
cipally by small myriapods, Collembola, and Diplura. Since cam-
podeids do not burrow and japygids do so very weakly (Pages,
1967), they primarily use the interstices already present. In such a
habitat body configurations are subject to an infinite diversity of
living spaces. A grooming behavior possible in one crack may be
impossible in another; however, a modification may work. We
believe that the unequal grooming diversity in Diplura is a response
to the problems of an interstitial life style. Foreleg grooming, where
the leg is simply raised to the mouth, does not require any special
bending or movement, so one technique does the job. Antennal,
mid- and hindleg grooming require unusual movements of the
appendage or of the body. Such movements may be limited by the
varied configurations of the crawl space, and must accommodate to
those configurations; thus, a variety of alternate positions appears
to be a necessity.

It is important to contrast the remarkable freedom of grooming
positions of Diplura, with the very high degree of stereotypy in such
orders as Diptera and Hymenoptera. The point is that a discussion
of insect grooming based on Diptera or Hymenoptera is as biased
towards stereotypy as a discussion of Diplura is biased towards lack

1978]

Valentine & Glorioso — Grooming in Diplura

199

of stereotypy. Present literature emphasizes the stereotyped aspects
of grooming, but it should be obvious that generalizations based on
highly derivative or primitive orders are not valid for the entire class
and may be skewed in opposite directions. The order Thysanura
would be a heuristic study because of the diversity of surface
textures. There are scaly lepismatids, campodeid-like nicoletiids,
and sclerotized, non-scaly lepidotrichids. Grooming in these three
families may further clarify why the degree of stereotypy varies from
taxon to taxon.

Literature Cited

Farish, D. J.

1972. The evolutionary implications of qualitative variation in the grooming
behavior of the Hymenoptera (Insecta). Anim. Behav., 20: 662-676, fig.
1-2, tab. I-III.

Hlavac, T. F.

1975. Grooming systems of insects: structure, mechanics. Ann. Ent. Soc.
Arner., 68(5): 823-826, fig. 1-2.

Jander, Ursula.

1966. Untersuchungen zur Stammesgeschichte von Putzbewegungen von
Tracheaten. Zeitschr. Tierpsychol., 23(7): 799-844, fig. 1-21, tab. 1-4.
Lipps, K. L.

1973. Comparative cleaning behavior in Drosophila. Ph.D. Dissertation,
University of California, Davis.

Paclt, Jiri.

1957. Diplura. In Wytsman, Genera Insectorum, fasc. 212: 1-23, fig. 1-37,
tab. 1.

Pages, Jean.

1951. Contribution a la connaissance des Diploures, Bull. Sci. Bourgogne, 13:
Suppl. rnecan. 9: 1-97, fig. 1-149.

1967. Donnees sur la Biologie de Dipljapyx humberti (Grassi). Rev. Ecol. Biol.
Sol, 4(2): 187-281, fig. 1-26, tab. 1-15.

Silvestri, Filippo.

1905a. Nuova contribuzione alia conoscenza dell 'Anajapyx vesciculosus Silv.
(Thysanura). Annali della R. Scuola Superiore d’Agricultura in Portici,
6: 1-15, fig. 1-12.

1905b. Materiali per lo studio dei Tisanuri VII. Dexcrizione di un nuovo genere
di Campodeidae delTtalia meridionale. Redia, 2: 115-119, 120, pi. XII.
1912. Contribuzione alia conoscenza dei Campodeidae (Thysanura) d’Europa.
Bollettino del Laboratorio di Zoolegia Generale e Agraria della R.
Scuola Superiore d’Agricultura in Portici, 6: 110-147, fig. I-XXXI.
1929. Zoologische Forschungsreise nach den Jonischen Inseln und dem
Peloponnes von Max Beier, Wien III. Teil Japygidae (Thysanura).
Sitzungsberichte d. mathem-naturw. Klasse Acad. Wissensch. Wien,
Abt. I, 138(9-10): 457-461, fig. 1-4.

200

Psyche

[June-Septernber

1932. Nuovi contributi alia conoscenza della fauna delle isole Italiane dell’-
Egeo II. Thysanura-Entotropha (Insecta). Bollettino del Laboratorio di
Zoologia Generale e Agraria del R. Instituto Superiore Agrario in
Portici, 27: 61-111, fig. I-XLIV.

1933a. Quarto contributo alia conoscenza dei Carnpodeidae (Thysanura) del
Nord America. Ibid 27: 156-204, fig. I-XXX1I.

1933b. Sulle appendici del Capo degli “Japygidae” (Thysanura Ehtotropha) e
rispettivo confronto con quelle dei Chilopodi, dei Diplopodi e dei
Crostacei. Ve Congres Internatl. Entorn., Paris, 1932: 329-343, fig.
I-VII.

1947. On some Japygidae in the Museum of Comparative Zoology (Dicellura).
Psyche, 54(4): 209-229, pi. 17-19, text-fig. 1-6.

Smith, L. M.

1960. The family Projapygidae and Anajapygidae (Diplura) in North America.
Ann. Ent. Soc. Arner., 53: 575-583, fig. 1-25, tab. 1.

Smith, L. M. and C. L. Bolton.

1964. Japygidae of North America 9. The genus Metajapvx. J. Kansas Ent.
Soc., 37(2): 126-138, fig. 1-10.

Snodgrass, R. E.

1935. Principles of Insect Morphology. McGraw-Hill Book Company, Inc.,
New York and London, ix + 667, fig. 1-319.

Valentine, B. D.

1973. Grooming behavior in Coleoptera. The Coleopterists Bulletin, 27(2):
63-73.

$

THE DACETINE ANT GENUS
PENTASTRUMA (HYMENOPTERA: FORMICIDAE)1

By William L. Brown, Jr. and Ronald G. Boisvert
Department of Entomology
Cornell University, Ithaca, New York 14853

The genus Pentastruma was established by Forel ( loc . cit. infra)
on the basis of a single Taiwanese worker specimen that he
described (P. sauteri ) as having 5 antennal segments, a very unusual
number even for a member of the Dacetini, to which tribe he
indicated that it belonged. In several ways, the description read as
though based on a depilated species of Smithistruma, and when,
several years ago, Dr. Masao Kubota sent specimens of a nearly
hairless short-mandibulate dacetine from Japan, WLB suspected
that it might be close to Pentastruma sauteri, despite the fact that its
antennae displayed the 6-merous condition usual in strumigenite
dacetines.

Now we have finally discovered the location of the Hans Sauter
Collection of Taiwanese ants, in the Institiit fur Pflanzenschutz-
forschung (BZA) der Akademie der Landwirtschaftswissenschaften
der Deutsche Demokratik Republic in Eberswalde. Through the
kind offices of Dr. G. Morge we have been able to borrow some
critical formicid types from the Sauter material, among them the
unique specimen of Pentastruma sauteri. This worker proves to be
close to the Japanese species received from Dr. Kubota, but it is
specifically distinct. It does also have 6 antennoineres, with the
normal strumigenite proportions, and not 5 as stated by Forel. In its
general form, P. sauteri is a rather typical Smithistruma, except for
its complete lack of standing or other conspicuous hairs on head,
trunk and petiole, and the new Japanese species matches it in these
respects.

It seems logical that Pentastruma should eventually be merged
with Smithistruma, but the latter genus is itself not stable at this

*A report of Research from the Cornell University Agricultural Experiment
Station, New York State College of Agriculture and Life Sciences. The research was
supported by National Science Foundation Grant GB-31662.

Manuscript received by the editor September 15, 1978.

201

202

Psyche

[June-Septernber

time, because new species have been discovered (mostly unpub-
lished) that seem likely to link it with such senior genera as
Trichoscapa and Codiomyrmex.

Until more of these new species have been formally described and
properly analyzed, no firm classification of the short-mandibulate
strumigenites is practicable. It seems best to retain some of the
available generic names for now, if only to avoid excessive combina-
torial changes as the classification develops.

Accordingly, we retain the name Pentastruma for the time being.
We figure for the first time the type species, P. sauteri, and
supplement its original description, and we describe a second
species, P. canina, from Japan, based on all 3 castes.

Measurements and proportions, and their abbreviations, are
those standard in papers on Dacetini, e.g., Brown, 1953, Arner.
Midi. Nat. 50: 7 ff., and 1973, Pacific Insects 15: 259.

Pentastruma

> Pentastruma Forel, 1912, Ent. Mitt. 1: 50. Type species: Pentastruma sauteri Forel,
monobasic.

Worker: Like Smithistruma in size, and form of head, mandibles
and remainder of body, including the 6-merous antennae; small
funicular segments II and II separate and distinct. Clypeus with
median tumulus and broadly extended anterolateral apron; anterior
margin concave in outline. Mandibles depressed, porrect, with
rounded basal lamella and no diastema, up to 15 acute teeth and
denticles of varying length, including small apical tooth. Labrum
with 2 long, tapered lobes, as in Smithistruma.

Body densely reticulate-punctulate and opaque (feebly shining in
some views), but postpetiolar disc and gaster smooth and shining,
except for basigastric costulae. Head, trunk, petiole and appendages
without erect hairs, and even the pubescence reduced to a virtually
invisible (at 50X) dilute vestiture of minute, appressed to decumbent
hairs. The under-mouthparts have some small standing hairs.
Postpetiole and gaster with a few short, blunt-tipped or remiforrn,
standing hairs, mostly arranged symmetrically. Color testaceous to
light ferruginous.

Queen: Like the Smithistruma queen, but with differences cor-
responding to those of the worker. Thoracic dorsum with a few

1978]

Brown & Boisvert — Ant Genus Pentastruma

203

short, slender but stiff, erect hairs. Pronotum with a flat, C-shaped,
marginate dorsal platform; scutum rising abruptly above this.

Male: As in Smithistruma.

Distribution as known: Japan (Honshu and southward); Taiwan.
Almost certainly occurs on the Asian mainland, but not yet
collected there.

Pentastruma sauteri
Fig. 1

Pentastruma sauteri Forel, 1912:51, worker. Type loc.: Pilarn, Taiwan. Holotype
worker: TL 2.2, HL 0.62, HW 0.43, ML 0.06, WL 0.55 rnm; Cl 69, MI 10.

The figure shows well the full-face outline view of head and
mandibles. The head is shorter, with vertex more convex, than in P.
canina, and the mandibles are shorter and more “set into” the
anterior clypeal concavity. Although we have been unable to view
the mandibular dentition directly and in detail in the lone holotype
specimen, it seems that a rounded basal lamella is present, and that
a series of sharp teeth follows without a diastema. The number of
teeth (12?) may be slightly smaller than in P. canina, and the sizes of
the teeth seem to be more evenly graded.

Viewed from the side, P. sauteri is much as shown in the side view
of P. canina (Fig. 4), except that in sauteri, the upper vertex is more
prominent, so that the head is thicker at this point dorsoventrally,
and the transition from frontal to occipital faces of vertex more
abrupt. The truncal dorsum is more strongly sinuate in side view;
the propodeal teeth are also shorter and less acute than in canina,
and the propodeal lamella wider. The erect pilosity of the gastric
dorsum and apex is even more reduced, in hair number and size,
than in canina. In the sauteri type, the middle of the mesopleura is
weakly shining, though still sculptured, and the dorsolateral mar-
gins of the trunk are completely lacking.

P. sauteri still remains known only from this single specimen from
Taiwan.

Pentastruma canina new species
Figs. 2, 3, 4

Holotype worker: TL 2.9, HL 0.77, HW 0.55, ML 0. 16, WL 0.72,
petiole L 0.30, eye L 0.05, scape L 0.32, hind femur L 0.46, hind tibia
L 0.35 mm; Cl 71, MI 21.

204

Psyche

[June-September

Figures 1-4. Pentastruma spp., workers. Fig. 1, P. sauteri holotype, head in full-
face dorsal view. Fig. 2, P. canina new species, paratype from type nest series, head in
full-face dorsal view. Fig. 3, same, side view. Fig. 4, same, another paranidotype,
mandible greatly enlarged. Figs. 1-3 to same scale; scale line = 0.2 mm. Drawings by
Ronald G. Boisvert.

1978]

Brown & Boisvert — Ant Genus Pentastruma

205

Habitus as shown for the paratype in Figs. 2 and 4. Note the
depressed, flat mandibles and the 6-inerous antennae, with segments
proportioned as in Smithistruma, and also the broadly extended,
sharp-edged, lamelliform, free lateral margins of the clypeus, trans-
lucent in bright light. Mandibular armament shown in detail in Fig.
3. After the broadly rounded basal lamella there follow without a
diastema 15 teeth, of which the first, fifth, and ninth are the longest.
Between these, 2 groups of 3 smaller teeth each, in each such triplet,
the middle tooth a little longer than the 2 flanking it. A similar
triplet follows the ninth tooth, and after this 2 small subapical teeth
and a robust apical tooth. Labral lobes at rest extending beyond the
midlength of the mandibles. Mandibular surface very finely sculp-
tured, weakly sericeous-shining.

Pronotum with rounded, strongly marginate anterior margin
(excluding cervix), faintly indicated but rounded humeri, feebly
marginate or submarginate dorsolateral margins, widest (W about
0.31 mm) behind midlength, tapering caudad into subparallel-sided
posterior half of trunk (W about 0.16 mm across propodeal
dorsum), which even widens again very slightly caudad. Meta-
notal groove obsolete or nearly so as viewed from above; pro-
podeal teeth approximately parallel. Faint margins extend the
length of the dorsolateral borders of the trunk, but these are visible
only in certain views and lights.

Petiolar node subquadrate (rounded in front), slightly wider (W
about 0.15 mm) than long, its dorsal surface with sculpture partly
effaced, weakly shining. Postpetiolar disc transversely elliptical,
nearly twice as wide as long (L 0.15, W 0.27 mm), smooth and
shining, with a widely spaced pair of inclined hairs near posterior
border. Gaster with weak basidorsal costulae, effaced mesad, the
longest extending about a quarter of the length of the first terguin.
Erect, feebly enlarged hairs: 4 near base of first gastric tergite, a pair
near midlength, and a pair near posterior border of first tergite;
remaining segments with 2 or 4 hairs each on tergites, and a few fine
ventral, erect hairs also on apical half of gaster. The fine, short,
extremely dilute, appressed and decumbent pubescence is invisible
except at high magnifications (over 50X) and in special, strong
lights, and is best developed on mandibles, antennae, vertex, legs,
and gaster, though nowhere evident without a special effort to find
it.

Color medium testaceous; spongiform apendages sordid whitish-
testaceous.

206

Psyche

[June-September

Type deposited in the Museum of Comparative Zoology, Har-
vard University.

Queen (Based on 7 specimens from 4 localities, including type
nest series): TL 3.0-3. 3, HL 0.77-0.80, HW 0.57-0.59, ML 0.17-
0.18, WL 0.80-0.84, forewing L (3 specimens) 2. 4-2. 6 mm; Cl
73-75, MI 22-23. Largest specimen (from Kiyosumiyama, HL 0.80
mm) with scape L 0.33, eye L 0.13 mm.

With the usual differences from the worker. Pronotum with flat
dorsal pronotal platform, as seen from above, markedly constricted
before it joins the inesothorax. Scutellum rounded and bulging
caudad. Petiolar node broader than in worker, and tending to be
medially sulcate in front. Scutum irregularly reticulate-punctate, its
surface weakly shining in some lights, opaque in others. Color
testaceous to medium ferruginous, usually slightly darker than
worker.

Male (3 specimens from Manazuru and Shirahaina): TL 2.7-2. 8,
HL 0.56-0.60, HW including eyes 0.45-0.47, eye L 0.20-0.22, WL
0.82-0.83, forewing L 2. 3-2. 4 mm.

Color blackish-brown, gaster dark reddish-brown, legs and an-
tennae sordid, pale, dull brown; wings hyaline. Mandibles slender,
each with a weak tooth-like angle apicad of midlength, only slightly
curved, probably not opposable, tapered to a very acute apex.
Labrurn broad, the 2 lobes short and separately rounded.

Antennal scape broader and longer than pedicel, and about as
broad as the apical antennal segment; segments III through XII
slender and cylindrical, all longer than scape or pedicel. Pronotum
forming a flattened, C-shaped platform, something like that of
female. Mesonotum large and bulging; notauli present but short,
and not meeting behind to form a V or Y; parapsidal lines present.
Scutellum bulging, rounded caudad. Propodeal teeth low, subacute,
with narrow, concave infradental lamellae. Mesokatepisternum and
a patch on the side of the propodeuin with sculpture effaced, nearly
or quite smooth and shining.

Petiole claviform, with low, rounded, scarcely differentiated node
that is mainly smooth and shining above; spongiform appendages
reduced to a narrow mid-ventral strip and a fine, cariniform
posterodorsal collar. Postpetiole broader than long, rounded,
smooth and shining, with a narrow, posterior collar of transparent
lamella and an anteroventral spur trimmed narrowly with trans-
parent lamella. Gaster unadorned at base, smooth and shining, with
an extremely sparse, inconspicuous sprinkling of tiny, appressed

1978]

Brown & Boisvert — Ant Genus Pentastruma

207

pubescence hairs, and a very few obliquely erect, short, fine hairs on
terguin of basal segment, plus some more of these nearer the gastric
apex. Legs with dilute, fine, inconspicuous, decumbent and ap-
pressed pubescence. Head, trunk, antennae, legs and sides of petiole
predominantly finely reticulate-punctulate, opaque, with the excep-
tions noted above.

Paratypes: 38 workers, 8 queens (alate and dealate) and 2 males,
all from JAPAN. HONSHU: Manazuru, Kanagawa Pref. (type local-
ity), 3 nest series; 4 April 1968 (with males), 20 Oct. 1968, 5 Jan.
1973 (with winged queens), all leg. M. Kubota. Hamaoka, Shizuoka
Pref., 15 Nov. 1976, leg. R. Egawa. Kiyosumiyama, Chiba Pref., 17
Aug. 1976, leg. T. Kannari. Shirahama, Wakayama Pref., 6 Jan.
1971 (with males), leg. M. Kubota. KYUSHU: Miyazaki-jingu,
Miyazaki Pref., 18 July 1971, leg. M. Shindo. Deposited in the
collection of Dr. Masao Kubota, at Odawara, Kanagawa Pref.,
Japan; in the Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge, Massachusetts, U.S.A.; and in the Department of
Entomology, British Museum (Natural History), London.

The variation of worker paratypes is most marked in body size,
relative head width, acuteness and width of propodeal teeth and
trailing lamellae, and in depth of ferruginous coloration, often
varying to faded straw color (in callows?). TL 2. 5-2. 9, HL 0.68-
0.78, HW 0.48-0.56, ML 0.14-0.16, Eye L 0.04-0.06, WL 0.62-0.72
mm; Cl 66-74 (mean 71 for n = 16), MI 19-22.

P. canina, widespread in central and southern Japan, is readily
distinguishable from P. sauteri by the form of the head and
mandibles (Figs. 1 and 2). From the known Smithistruma and
Trichoscapa species of eastern Asia, the canina worker may be
separated by its head shape and by the total lack of standing pilosity
on head, scapes and trunk.

Dr. Masao Kubota, of Odawara, Kanagawa Prefecture, Japan,
deserves thanks, not only for the opportunity to study the many
excellently prepared specimens of P. canina, but also for notes on the
biology of the species summarized below.

P. canina is an uncommon species, found in the Kanto District and
southward. It inhabits the floor of coastal evergreen broadleafed
forest, which is generally subtropical. Nests are found in small pieces
of rotten wood, rotten fallen branches, under moist leaf litter, or at a
slight depth in the humus. The largest colony censused contained one
queen and 57 workers.

OBSERVATIONS ON A POPULATION OF
SIALIS ITASCA ROSS IN WEST VIRGINIA
(MEGALOPTERA: SIALIDAE)

By C. K. Lilly1, D. L. Ashley2, and D. C. Tarter2

Observations on the ecology of each species in an aquatic
community are necessary for the total understanding of community
dynamics. Several authors, including Davis (1903), Ross (1937),
Townsend (1939), Flint (1964), Azam and Anderson (1967), Wood-
ruin and Tarter (1973), Pritchard and Leischner (1973), Tarter and
Woodrum (1973), Tarter (1973), Tarter et al. (1976), and Tarter et
al. (1978) have reported on the taxonomy, distribution, life history,
and ecology of several Sialis spp. Other authors, including Roback
and Richardson (1969), Warner (1971), Nichols and Bulow(1973),
Tarter and Woodrum (1972) and Woodrum and Tarter (1973), have
noted the extreme tolerance of Sialis to acid mine drainage.

The primary objectives of this investigation were: (1) to make
observations on the life history and ecology of the alderfly S. itasca
in a small farm pond and (2) to determine the pH tolerance of this
population under laboratory conditions.

Materials and Methods

The population of S. itasca inhabits a small farm pond, 0.1
hectare, near Shoals, West Virginia which is 8 km south of
Huntington, West Virginia. This pond is located in the north-central
region of Wayne County. It is located at 82°29'40"W longitude and
38°21'50"N latitude.

This investigation was initiated in May 1975 and continued until
April 1976. Monthly samples were taken by a small seine (0.25 inch
mesh). The seine was placed in the water vertically and the mud and
debris on the bottom of the pond between the seine and the bank
were disturbed. The seine was then moved toward the bank while
dragging the bottom of the net on the bottom of the pond collecting

■Present address: 4074 - 40th Street, Nitro, WV 25143

2Dept. of Biol. Sci., Marshall Univ., Huntington, WV 25701
Manuscript received by the editor August 1, 1978.

209

210

Psyche

[June-September

mud and debris in the net. The larval alderflies were collected and
preserved in 70 percent ethanol.

Temperatures were taken with a Taylor maximum-minimum
thermometer placed on the pond bottom 0.5 meter below the water
surface. They were recorded in degrees Celsius once per month at
the time benthic samples were taken. Water chemistry tests were
performed in the field with a Hach chemical kit, Model AL-36-
WR. All tests were completed within one hour. Hydrogen-ion
concentration (pH) was measured colorimetrically. Dissolved oxy-
gen, carbon dioxide, hardness (magnesium and calcium), phenol-
phthalein and methyl orange alkalinity, free acidity, and total
acidity were measured and recorded in mg/1.

Size classes were determined by length frequency distributions
arranged in 1 mm length groups. Total length (exclusive of the
caudal filament) was measured under 7X magnification with calipers
and a plastic ruler (nearest 0.5 mm). Head width was measured with
an ocular micrometer between the inner edges of the eyes (nearest
0.01 mm). Differences in head width of 124 larvae were determined
to show the mean, range and standard deviation.

A total of 84 foreguts were examined to determine food habits.
The head was removed and the abdomen was split open to remove
the intestine. The contents of the intestine were removed and
examined under a dissecting microscope and a compound micro-
scope to identify their contents. The percent frequency of occur-
rence of each item was determined, and the monthly and seasonal
averages and various sizes were compared.

For the pH tolerance test, forty mature larvae collected from the
pond in April were taken to the laboratory for acclimatization over
a 24 hr period. The larvae were placed in groups of 10 in 4 finger
bowls. One bowl was filled one-third of the way with pond water. A
one molar solution of potassium dihydrogen phosphate (KH2P04)
was diluted approximately into the other three bowls, to set pH
values at 5.5, 4.0, and 2.5. The pH value of the control was 7.0. A
Model 5 Corning Scientific pH meter was used to determine pH
values. Oxygen was constantly supplied with air stones. The temper-
ature did not change significantly during the experiment and
averaged 12 C. The 96 hour TLm (median tolerance limit) test
(APH A, 1965) was used to measure the effect of low pH. The pH
value at which 50 percent of the alderfly larvae died after 96 hours
was determined by straight-line graphical interpolation.

1978] Lilly, Ashley, & Tarter — Population of Sialis itasca 211

Fifteen larvae were collected in April and returned to the
laboratory for rearing in vials containing pond water. These vials
were connected to vials containing sand by a short section of rubber
tubing (Pritchard and Leischner, 1973). Wire mesh and strips of
foam rubber were placed in the vials and rubber tubing to enable the
larvae (one per vial) to move freely between the vials. The vials were
kept at room temperature which was approximately 70 F.

Fecundity was determined by a direct count of ovarian eggs under
a compound microscope. The ovaries of 3 adults were removed and
a total of 1616 eggs were counted. The diameters of 90 eggs were
measured to the nearest 0.01 mm with an ocular micrometer using a
Bausch and Lomb compound microscope.

Results and Discussion
Pond Environment

Temperature. — The average annual temperature for the pond
was 18.1 C. The extreme monthly temperatures were 0.5 C in
February and 35.0 C in August and September.

Water Chemistry. — The average annual pH was 8.4 (7.5 to 9.0).
The average annual dissolved oxygen was 8.5 mg/ 1 (6.5 to 10.0).
The average annual total hardness was 139.6 mg/ 1 (102.6 to 153.9).
The average annual carbon dioxide concentration was 9.6 mg/ 1 (5.0
to 25.0). The average annual alkalinity was 110.5 mg/ 1 (68.4 to
145.4). The average annual total acidity was 12.4 mg / 1 (5.7 to 28.5).

Larval Stage

Development. — Length-frequency histograms indicated that the
population of S. itasca contained one size class (Fig. 1). Hatching
occurred at the end of May. There were 18 egg masses located in the
field at the end of May. The egg masses were found on the leaves of
hornbeam and buckeye trees. Three egg masses which were returned
to the laboratory hatched within 2 days. In July all the egg masses
were empty but no larvae were located. The earliest and smallest
larvae were collected on August 2, 1975. Their average length was
6.45 mm, and their average head width was 1.02 mm. The last and
largest larvae were collected on April 6, 1976. Their average length
was 13.67 mm, and average head width was 1.59 mm.

Head width was used to show the monthly variation in growth
rate and the percent increase in growth rate (Fig. 2). Due to the

212

Psyche

[June-Septernber

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Figure 1 . Length-frequencies at monthly intervals of S. itasca larvae from a farm
pond near Shoals, W. Va. The number of larvae is given for each month.

1978] Lilly, Ashley, & Tarter — Population of Sialis itasca 213

small sample size (4) in August, no statement can be made
concerning the growth rate from August to September. An eighteen
percent increase in growth rate was recorded from October to
November. There was a decrease in growth rate in December. Small
sample sizes in January (2) and February (6) prevented any growth
rate information. There was a six percent growth rate from March
to April. The largest mean head width was measured in April at 1.59
mm and ranged between 0.95 and 1.86 mm September and Novem-
ber, respectively. No larvae were located in May, June and July.

Woodrum and Tarter (1973) also found a decrease in growth
during the winter months in S. aequalis. Azam and Anderson (1969)
found a decrease in the growth rate for S. rotunda and S. californica
during the winter months.

0

1

M

5

Figure 2. Monthly variation of the head width in S. itasca larvae. Vertical lines =
ranges, horizontal lines = means, open rectangle = one standard deviation, numbers
= sample sizes, and dotted lines = temperature (C).

214

Psyche

[June-September

Food Habits. — This population of S. itasca was found to feed
almost exclusively on the ostracod, Cyclocypris sp. The only other
food items were 3 midges, Chironomus sp., which were found on 3
different occasions. Of the 84 foreguts analyzed, 21 percent were
empty and 79 percent contained food. Ostrocods were found in 71
percent of the foreguts, and midges were found in 4 percent of the
foreguts.

Excluding the month of August when a very small sample size (4)
was used, the largest number of empty foreguts occurred in January
(50%). The percent of empty foreguts increased again in April
supporting the findings of Woodrum and Tarter (1973) in S.
aequalis that the larvae probably do not feed just before pupation.

Azam and Anderson (1969) reported S. rotunda and S. Cali-
fornia to be indiscriminate feeders and reported cannibalism to be
frequent. Woodrum and Tarter (1973) found S. aequalis to be more
restricted in its feeding due to the limited choices of organisms
found in the acid mine stream in which they were located. They also
reported cannibalism to occur to a lesser extent. The S. itasca in this
investigation were found to be more restrictive feeders preying
almost exclusively on ostracods while having an abundant supply of
other organisms upon which they could feed. Cannibalism was
observed in the laboratory when larvae were confined for three days
without food.

Predation. — The stomachs of 18 odonates and 10 sunfish were
examined. No alderfly remains were found in the sunfish and only
one alderfly head was found in the odonates. Schwiebert (1973)
noted that the hellgrammite and trout are predators of the alderfly.

pH Tolerance. — The 96 hour TL™ value for S. itasca was found
to be 3.1. All ten larvae survived the 96-hour period at pH values of
7.0 and 5.5, 70 percent survived in a pH of 4.0, and 30 percent
survived at a pH of 2.5. Tarter and Woodrum (1972) found S.
aequalis from an acid mine stream to have a TLm value of 2.1. These
values would indicate that S. itasca and S. aequalis are quite
tolerant of low pH. Sialis spp. have been noted to be tolerant to low
pH conditions in western Pennsylvania streams (Roback and
Richardson, 1969), Roaring Creek in eastern West Virginia (Warner,
1971), and in the East Fork of the Obey River in Tennessee (Nichols
and Bulow, 1973).

1978] Lilly, Ashley, & Tarter — Population of Sialis itasca 215

Pupal Stage

Larvae placed in the laboratory rearing chambers moved to the
sand for pupation within 2 to 4 days. The pupal stage lasted for
approximately 2 weeks. The adults emerged during the night. No
pupae could be located in the bank of the pond.

Azam and Anderson (1969) reported S. rotunda and S. calif ornica
to pupate during April, May and June. Pritchard and Leischner
(1973) reported that S. cornuta pupated from May to mid-June.
Woodrum and Tarter (1973) found S. aequalis to crawl 1.5 to 5 m
out of the water onto a moist sandbank when the water temperature
reached 11 to 13 C and pupated in an earthen cell 1 to 7 cm below
the surface. They found the pupae to respond to a disturbance but
otherwise remained rather dormant.

Adult Stage

Number and Size of Eggs. — Fecundity of 3 adult alderflies
showed a range of 454 to 587 eggs per female; the average was 539
eggs. The eggs were cylindrical, rounded on the ends, had a curved
micropylar tubercle on one end, and averaged 0.31 mm in length by
0.14 mm in width. The number of eggs found in S. itasca was similar
to those of S. aequalis (657) (Woodrum and Tarter, 1973), S.
rotunda (300-500) and S. californica (400-700) (Azam and Ander-
son, 1969), and S. cornuta (615) (Pritchard and Leischner, 1973).
Egg masses of S. itasca were found on the underside of hornbeam
and buckeye leaves 0.5 to 3 m above the water surface. The eggs are
laid in rows in a nearly vertical position much like those of S.
rotunda (Azam and Anderson, 1969).

Mating. — Although copulation was not observed, a courtship
behavior was observed in the laboratory much like that described by
Azam and Anderson (1969) for S. rotunda and by Woodrum and
Tarter (1973) for S. aequalis.

Longevity. — Adults in the laboratory lived for 4 to 6 days. Only
one adult was captured in the field on 3 May 1975. Sialis rotunda
was first seen in mid-April and reached their peak in May (Azam
and Anderson, 1969) while S. californica was seen in May but did
not peak until mid-June (Azam and Anderson, 1969). Sialis aequalis
was observed in the field between April 21 and May 4 (Woodrum

216

Psyche

[June-September

and Tarter, 1973). Since the adults serve only to reproduce the
species, it is apparent that they have a brief life span to perform this
function. Azam and Anderson (1969) observed the female of S.
rotunda laying eggs within a day after emerging.

References

American Public Health Association, Inc.

1965. Standard Methods for the Examination of Water and Wastewater (12th
Ed.). New York, N. Y. 744 pp.

Azam, K. M., and J. H. Anderson.

1969. Life history and habits of Sialis rotunda and Sialis calif ornica in western
Oregon. Ann. Ent. Soc. Arner. 62: 549 558.

Davis, K. C.

1903. Sialididae of North and South America. Aquatic insects of New York
State. No. 7. N. Y. State Mus. Bull. 68: 442-486.

Flint, O. S., Jr.

1964. New species and new state records of Sialis (Neuroptera: Sialidae). Ent.
News 25: 9-13.

Nichols, L. E., Jr., and F. J. Bulow.

1973. Effects of acid mine drainage on the stream ecosystem of the East Fork
of the Obey River, Tennessee. J. Tenn. Acad. Sci. 48(1): 30-39.
Pritchard, G., and T. G. Leischner.

1973. The life history and feeding habits of Sialis cornuta Ross in a series of
abandoned beaver ponds (Insecta: Megaloptera). Can. J. Zool. 5:
121-131.

Roback, S. S., and J. W. Richardson.

1969. The effects of acid mine drainage on aquatic insects. Proc. Acad. Nat.
Sci. Phila. 121: 81 107.

Ross, H. H.

1937. Studies of nearctic aquatic insects. I. Nearctic alderflies of the genus
Sialis (Megaloptera: Sialidae). 111. Nat. Hist. Surv. Bull. 21: 57-78.
SCH WEIBERT, E.

1973. Nymphs. Winchester Press. 339 pp.

Tarter, D. C., and J. E. Woodrum.

1972. Low pH tolerance of the larvae of the alderfly, Sialis aequalis Banks,
under controlled conditions. Proc. W. Va. Acad. Sci. 44: 85-88.

1973. First record of the alderfly, Sialis joppa Ross (Megaloptera: Sialidae), in
West Virginia. Proc. W. Va. Acad. Sci. 45: 163-164.

1973. Distribution and new record of the alderfly Sialis (Megaloptera: Siali-
dae) in West Virginia. Ent. News 84: 147-148.

Tarter, D. C., D. L. Ashley, and C. K. Lilly.

1976. New record of the alderfly Sialis itasca Ross for West Virginia (Meg-
aloptera: Sialidae). Ent. News 87: 32.

Tarter, D. C., W. D. Watkins, D. L. Ashley, and J. T. Goodwin.

1978. New state records and seasonal emergence patterns of alderflies east of
the Rocky Mountains (Megaloptera: Sialidae). Ent. News 89: 231-234.

1978] Lilly, Ashley, & Tarter — Population of Sialis itasca 217

Townsend, L. H.

1939. A new species of Sialis (Megaloptera: Sialidae) from Kentucky. Proc.
Ent. Soc. Wash. 41: 224-226.

Warner, R. W.

1971. Distribution of biota in a stream polluted by acid mine drainage. Ohio J.
Sci. 71: 202-216.

WOODRUM, J. E., AND D. C. TARTER.

1973. The life history of the alderfly, Sialis aequalis Banks, in an acid mine
stream. Amer. Midi. Nat. 89: 360-368.

STRUCTURE AND RELATIONSHIPS
OF THE UPPER CARBONIFEROUS INSECT,
PROCHOROPTERA CALOPTERYX
(DIAPHANOPTERODEA, PROCHOROPTERIDAE)

By Frank M. Carpenter1 and Eugene S. Richardson, Jr.2

Prochoroptera calopteryx was described by Handlirsch (1911)
from a single specimen in a concretion from the Francis Creek Shale
(Pennsylvanian) in northeastern Illinois. Although poorly preserved,
the fossil showed several unusual features. Because of these and its
very incomplete preservation, the insect’s relationships have been
decidedly controversial. Fortunately, after a lapse of nearly seventy
years following the publication of Handlirsch’s account, three more
specimens have recently been found in concretions from the same
deposit. These additional fossils, which have been turned over to us
for study, provide considerably more information about the insect
than the type and enable more reliable conclusions about its
relations with other Paleozoic insects, even though much of its body
structure still remains unknown. It clearly belongs to the extinct
order Diaphanopterodea and is only the second species of that order
known from Pennsylvanian strata in North America. Descriptive
accounts of all four specimens of calopteryx are included below,
with a discussion of the relationships of the family Prochorop-
teridae.

For the opportunity of studying these new specimens we are
indebted to Helen and Ted Piecko of Chicago, and Mr. J. J. Fagan
of Burbank, Illinois. Handlirsch’s type of calopteryx has been
placed at our disposal by Jean S. Lawless, Division of Invertebrate
Paleontology, Peabody Museum, Yale University. Partial support
of this research is gratefully acknowledged to the National Science
Foundation, Grant No. DEB 78-09947, F. M. Carpenter, Principal
Investigator.

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

2Field Museum of Natural History, Chicago, Illinois 60605
Manuscript received by the editor, November 25, 1978.

219

220

Psyche

[June-September

Order Diaphanopterodea

Diaphanopteroidea Handlirsch, 1919, p. 575.

Diaphanopterodea Rohdendorf, 1962, p. 69; Carpenter, 1963, p. 253.

[= Palaeohyrnenoptera Haupt, 1941, p. 99.]

Family Prochoropteridae
Prochoropteridae Handlirsch, 1911, p. 375.

Fore wing: similar to that of the Elmoidae but with SC extending
much farther beyond the origin of RS than in Elmoa; stem of R
without a prominent bend at the point of divergence of CUA; cross
veins weakly developed, without branching. Hind wing: similar to
the fore, so far as known, but relatively broader and with its hind
margin more curved; coalescence of RS with MS apparently less
than in the fore wing. Thorax large, but prothorax small and
conical; head small (from above).

Type-genus: Prochoroptera Handlirsch. A second genus, Eu-
choroptera Carpenter (1940), from the Pennsylvanian of Kansas
(Stanton Formation), also appears to belong to this family; little is
known of its wing venation, but the body structures are like those of
other Diaphanopterodea.

Genus Prochoroptera Handlirsch
Prochoroptera Handlirsch, 1911, p. 376.

Since this genus is monospecific, its characteristics can only be
suggested. In all probability the coalescence of MA with RS and the
distal forking of MP are in this category. Prochoroptera differs
from Euchoroptera in having the branching of RS pectinate, instead
of dichotomous.

Type-Species: Prochoroptera calopteryx Handlirsch.

Prochoroptera calopteryx Handlirsch
Figures 1-3

Prochoroptera calopteryx Handlirsch, 1911, p. 376; 1919, p. 586; 1920, p. 172.

Haupt, 1941, p. 99. Laurentiaux, 1953, p. 444. Carpenter, 1940, p. 638; 1947, p. 45;

1963, p. 248.

The fore wing is known to a length of 20 mm; estimate of
complete length (based on Euchoroptera ), 36 mm; maximum width,
6 mm. Hind wing: known to a length of 20 mm; maximum width, 6

1978] Carpenter & Richardson — Prochoroptera calopteryx 221

■' 'L: 1 1

Figure 1. Prochoroptera calopteryx. Photograph of reverse of specimen HTP
12523 (X 3.8).

222

Psyche

[June-Septernber

rnrn. The details of venation are shown in figure 2. Body length,
from front of mesothorax to end of abdomen, 25 mm; length of
abdomen, 18-19 mm; width, 4-5 mm.

Holotype: no. YPM67, Peabody Museum, Yale University (ob-
verse and reverse).

As shown in Handlirsch’s figure, this fossil includes the thorax,
abdomen, and parts of both pairs of wings, which rest obliquely
backward. His figure of the wings is essentially accurate, although
his attempted restoration of the complete wing turns out to be
incorrect, at least to the extent that we now know it. The body is
also satisfactorily represented in his figure, although the prothorax
is too large. The abdominal segments of this specimen possess
paired lateral expansions, apparently strongly sclerotized and simi-
lar to those present in at least some Palaeodictyoptera. Handlirsch’s
figure depicts these as membranous areas bordering the terga, but
there is nothing in the fossil to support that interpretation. The 9th
abdominal segment bears a pair of longitudinal ridges, converging
distally, as shown in Handlirsch’s figure. These are almost certainly
the basal portions of the valves forming the ovipositor. Cerci are not
preserved.3 The dimensions of this specimen are as follows: fore
wing, length (as preserved), 20 mm; width uncertain; hind wing,
length (as preserved), 20 mm. Body length (from front of meso-
thorax to end of abdomen), 25 mm; pterothorax, length, 6.3 mm;
width, maximum, 6 mm; abdomen, length, 18 mm; width, maxi-
mum (3rd segment), 4.5 mm; prothorax, length, 3.5 mm; head
absent.

The three recently discovered specimens of calopteryx are as
follows:

1. No. JJF-IL1 1-007, collected by J. J. Fagan in Pit 11 (strip
mine in Will and Kankakee Counties, 3 to 5 miles south of
Braidwood, Illinois).4 This specimen is preserved in almost the same
position as the type. The prothorax and the wing veins are clearer
than in the type. Its dimensions are as follows: fore wing, length (as

3Most of the “whole” specimens of insects preserved in these concretions lack the
distal parts of the wings and the distal anterior and posterior appendages, such as
antennae and cerci. The insect’s body is ordinarily in the center of the concretion,
which is usually too small to include the distal portions of these appendages.

4Mr. Fagan has now deposited this specimen in the Field Museum of Natural
History, where it bears the number PE 25667.

1978] Carpenter & Richardson — Proehoroptera calopteryx 223

Figure 2. Proehoroptera calopteryx. Drawing of fore and hind wings, as known;
based mainly on specimen HTP 12523, with some details added from the holotype
and specimen JJF 1 LI 1-007. SC, subcosta; Rl, radius; RS, radial sector; MA, an-
terior media; MP, posterior media; CUA, anterior cubitus; CUP, posterior cubitus;
1A, first anal vein; +, convex vein; — , concave vein.

preserved), 18 inm; width, 6 mm; hind wing, length (as preserved),
18 mm; width, 6 mm. Body length (from front of inesothorax to end
of abdomen), 25 mm; pterothorax, length, 6.3 mm; width, 4.8 mm;
abdomen, length 19 mm; width, 4.8 mm; prothorax, length, 2.5 mm;
width at posterior edge, 3 mm; head absent.

2. No. HTP 12523, Helen and Ted Piecko collection. This is the
best specimen known. The obverse half represents a ventral view of
the insect and the reverse (figure 1), a dorsal view.5 The wings,

5In our terminology, the obverse is the counterpart that shows the topography of
the veins as seen on the dorsal surface of the living insect (i.e., SC, concave; Rl,
convex, etc.); the reverse shows the veins in the opposite topography. Generally
speaking, the reverse shows the dorsal surface of the insect, and the obverse shows
the ventral surface.

224

Psyche

[June-Septernber

pterothorax, and abdomen are especially clear, but the prothorax
and head are not preserved. The wings on one side, although lacking
the apical portions, are very clearly preserved; those on the other
side are twisted and folded together. Widely spaced pits in the
impressions of the veins and cross veins indicate the original
presence of trichia or small setae. The segmentation of the abdomen
is especially clear and the lateral extensions of the segments,
mentioned above in the type, are faintly preserved in the obverse
half of the fossil. Also, in that counterpart, which shows the ventral
view of the abdomen, the bases of the ovipositor valves can be seen
in the 8th and 9th segments. The ovipositor appears to be about 5
mm long before disappearing at the edge of the concretion. The
cerci are not preserved. The following are the dimensions of this
fossil: fore wing (as preserved), 15 mm; width, uncertain; hind wing,
length (as preserved), 15 mm; width, 6 mm. Body, length (from front
of mesothorax to end of abdomen), 25 mm; pterothorax, length, 5
mm; width, 4.8 mm; abdomen, length, 18.5 mm; width, maximum,
4.6 mm.

3. No. HTP 415, Helen and Ted Piecko collection. This is a
poorly preserved specimen, resting in the same position as the
previous one. The venation is clear enough for identification of the
species, and the prothorax and even the head, which is not shown in
other specimens, are discernible. Neither the ovipositor nor the cerci
are visible. The dimensions of this specimen are as follows: fore
wing (as preserved), 20 mm; width, uncertain; dimensions of hind
wing uncertain. Body length (from front of mesothorax to end of
abdomen), 25 mm; pterothorax, length, 6 mm; width, maximum, 4
mm; abdomen, length, 20 mm; maximum width, 4.5 mm; pro-
thorax, length, 3.3 mm; posterior width, 2.5 mm; anterior width, 1.5
mm; head, length, 3 mm; width 3 mm. The head as preserved shows
somewhat protuberant eyes; the head is obviously seen in dorsal
view, with no indication of the beak.

Discussion of Structure

The wings of calopteryx, as far as they are known at present, are
shown in figure 2, which is based mainly on specimen HTP 12523,
with some details added from the other specimens. The figure shows
more of the basal parts of both wings than was known to Hand-
lirsch. The homologies of the veins are obvious, the convexities and
concavities being clearly preserved. Handlirsch (191 1) confused MA

1978] Carpenter & Richardson — Prochoroptera calopteryx 225

with a branch of the media, but his interpretations were the
conventional ones at the time. In all four specimens the amount of
coalescence between MA and RS is distinctly less in the hind wing
than in the fore wing, but in other respects the venational patterns
are nearly identical, so far as they are known. The shape of the hind
wing, as preserved, suggests that it may have been slightly shorter
than the fore wing, the hind margin having a greater curvature. The
fore wing was apparently somewhat more elongate than indicated in
Handlirsch’s figure.

Figure 3, showing all of the known parts of the insect, is based
mainly on HTP 12523, with the prothorax and head as preserved in
HTP 415. The wings have been added to the figure to show their size
relative to the body, which appears to be surprisingly robust
compared to the wings (See figure 1). The prothorax, which is
preserved in HTP 415 and the holotype, is clearly conical in shape
and unusually small with respect to the rest of the thorax. The

Figure 3. Prochoroptera calopteryx. Drawing of wings and body, as known;
based mainly on specimen HTP 12523 (reverse) with prothorax and head added from
specimen HTP 415. TH2, mesothorax; TH3 metathorax; Al, first abdominal
segment.

226

Psyche

[June-September

outline of the head is faintly indicated in specimen HTP 415.
Handlirsch’s drawing (1911) of the type includes a suggestion of the
head or part of it, but we have been unable to discern the structure
that he has drawn. However, the head as shown in specimen HTP
415 is very small with respect to the rest of the insect, although of
course the head as preserved in the fossil is seen from above and
does not show its full length. The meso- and metathorax, on the
other hand, appear large relative to the rest of the insect. The
abdominal segments are virtually homonomous, with only a slight
reduction in the width posteriorly.

Affinities of the Prochoropteridae

Opinions of the affinities of calopteryx, based on the type, have
been diverse. Handlirsch (1911, 1919, 1920) considered it to belong
to the Megasecoptera. Carpenter (1940), following the discovery of
another and apparently related Pennsylvanian insect ( Euchoroptera
longipennis), concluded that the family Prochoropteridae had close
affinities with the Permian family Asthenohymenidae, then con-
sidered to be the most highly specialized of the Megasecoptera.
However, Haupt (1941), basing his conclusions on Handlirsch’s
brief account of Prochoroptera, designated a new order, Palaeo-
hymenoptera, for the family Prochoropteridae, assigning it to his
“Legion Hymenopteroidea”, which also included the Hymenoptera.
Unaware of that publication6 Carpenter (1947, 1954) proposed that
the families Prochoropteridae and Diaphanopteridae (Cormnentry,
France), along with two other Permian families, be placed in a
separate suborder of the Megasecoptera. However, Rohdendorf in
1962 adopted the ordinal name Diaphanopterodea for that series of
families, since the taxon had been named by Handlirsch (1919) for
the Diaphanopteridae. That term has subsequently been generally
accepted.

The order Diaphanopterodea is now recognized as belonging to
the Palaeoptera and as allied to the Palaeodictyoptera and Mega-
secoptera, all of their members possessing haustellate mouthparts
and long cerci. Alone in this series, however, the Diaphanopterodea
had the ability to “fold” their wings at rest along the abdomen. The

6The journal containing Haupt’s 1941 paper (Zeitschrift fur Naturwissenschaften,
Halle) was not received by the library at Harvard University until January, 1958.

1978] Carpenter & Richardson — Prochoroptera calopteryx 227

most distinctive feature of their venation is the coalescence of CUA
and M basally, and, except in the Diaphanopteridae, the coalescence
of CUA and M further basally with R. In all the families of the
order, M appears to arise at the point of divergence of R and CUA.

Although the body structure of Prochoroptera is very incom-
pletely known, the wing venation clearly shows the diagnostic
features of the order Diaphanopterodea, and of course, as noted
above, in all known specimens the wings are folded along the
abdomen. However, the position of the Prochoropteridae within the
order is still uncertain. As far as known, the most obvious evolu-
tionary changes within the Diaphanopterodea are the narrowing of
the wing base and the consequent increased coalescence of M A with
RS and ultimately with R itself (See Carpenter, 1963, plate 10, figs.
1-6). However, it now seems probable that these changes occurred
independently within several lines of the order (Kukalova-Peck,
1974). As a result, diagnoses of some families are very difficult to
make, especially of those in which the wings are not completely
known, such as the Prochoropteridae. However, it seems advisable
to retain the family Prochoropteridae as valid and as distinct from
such Permian families as Elmoidae and Martynoviidae on the basis
of its venational features as presently known.

References

Carpenter, F. M.

1940. Carboniferous insects from the Stanton Formation, Kansas. AMER J
SCI 238: 636-642.

1947. Lower Permian insects from Oklahoma. Part 1. Introduction and the
orders Megasecoptera, Protodonata, and Odonata. PROC AMER
ACAD ARTS SCI 76: 25-54.

1954. Extinct families of insects, in Classification of Insects (C. T. Brues, A. L.
Melander, and F. M. Carpenter). BULL MUS COMP ZOOL 108:
777-827.

1963. Studies on Carboniferous insects from Commentry, France. Part V. The
genus Diaphanoptera and the order Diaphanopterodea. PSYCHE 70:
240-256.

Handlirsch, A.

1911. New Paleozoic insects from the vicinity of Mazon Creek. AMER J SCI
31: 297-377.

1919. Revision der Palaozoischen Insekten. DENKSCHR ACAD WISS 96:
1-82.

1920. Palaeontologie, in Handbuch der Entomologie (ed. C. Schroder) (3) 7:
117-306.

228

Psyche

[June-Septernber

Haupt, H.

1941. Die altesten geflugelten Insekten und ihre Beziehungen zur Fauna der
Jetztzeit. ZEIT NATURWISS HALLE 94:60-121.

Kukalova-Peck, J.

1974. Wing-folding in the Paleozoic insects of the order Diaphanopterodea
(Palaeoptera), with a description of new representatives of the Elmoidae.
PSYCHE 81: 315-333.

1974b. Pteralia of the Paleozoic insect order Palaeodictyoptera, Megasecop-
tera, and Diaphanopterodea (Palaeoptera). PSYCHE 81: 416-430.
Laurentiaux, D.

1953. Classe des insectes, in Traite de Paleontologie (ed. J. Piveteau) 3:
397-527.

Rohdendorf, B. B., et al.

1962. Osnovi paleontologii [Principles of Paleontology], Moscow (ed. B. B.
Rohdendorf), Arthropoda: Tracheata and Chelicerata. Pp. 1-560.

DIVISION OF LABOR WITHIN THE WORKER CASTE
OF FORMICA PERPILOSA WHEELER
(HYMENOPTERA: FORMICIDAE)*

By Carlos Roberto F. Brandao

Museu de Zoologia, Universidade de Sao Paulo
Sao Paulo, Brasil

Introduction

Polymorphism, in the study of social insects, is defined as the
existence within an individual colony of two or more phases or
castes belonging to the same sex, without particular regard to their
genetic or environmental origin (Wilson, 1953). The adaptive result
of the development of female polymorphism is the division of labor
within the colony. In most ants this division is clearly seen between
reproductive and non-reproductive caste, but less evident within the
worker caste (Oster & Wilson, 1978). The present article utilizes the
Fagen & Goldman (1977) method for estimating the total repertory
size of behavioral categories of each worker subcaste listed on an
ethogram or behavioral catalog.

Ethograms are the essential first step of the comparative study of
behavior (Wilson, 1974). A behavioral catalog of Formica perpilosa
Wheeler, a weakly polymorphic species of the neogagates group
(Buren, 1968), was constructed in order to investigate behavioral
differences between the major size groups, defined here arbitrarily
as three worker subcastes. Formica perpilosa is a common ant in the
southern United States and northern Mexico (Gregg, 1963). It feeds
mainly on plant exudates and tends membracids of the genus
Pubilia (LaBerge, 1952). Its physiology has been relatively well
studied by Schumacher & Whitford (1974), Kay & Whitford (1975),
Whitford, Kay & Schumacher (1975) and Schumacher & Whitford
(1976). The genus Formica is of unusual interest because its species
are either monomorphic or weakly polymorphic and thus span the
early stage of caste evolution. Yet close studies of the polymorphic
species have not been undertaken.

* Manuscript received by the editor October 10, 1978.

229

230

Psyche

[June-September

Material and Methods

Founding queens were collected near Portal, Arizona, in May,
1975, by B. Holldobler and individually placed in test tubes, 14.8 cm
long by 23 mm inner diameter, kept moist by compact cotton plugs
that trap water at the bottom of the tubes. As the selected colony
grew the tube was moved to a plexiglass box 28 cm X 45 cm and
15 cm deep, the sides of which were coated with Fluon GP-1
(Northeast Chemicals Co., Woonsockett, R.I.) to prevent escape,
and four similar tubes added. The colony has since been maintained
on an artificial diet for ants (Bhatkar & Whitcomb, 1970) and honey
water three times a week and fed freshly killed cockroaches
( Nauphoeta cinerea ) once a week.

Before the first set of observations the colony was moved to a
glass nest made of two square plates 12.5 cm on a side, held apart by
small pieces of non-toxic plasticin and taped on three sides. This
simple nest permits close observation of the behavior of the entire
colony. The assemblage was put on the floor of the original
container next to the water tubes. The remainder of the floor served
as foraging space and was kept clear for observation.

The container was small enough to be placed under a swinging
arm dissecting microscope. During a period of 4 weeks, a total of 18
hours were dedicated to cataloging behavior; 2809 separate be-
havioral acts were recorded. The observation hours ranged ran-
domly from 9:00 A.M. to 11:00 P.M.; no differences in level or
pattern of activity were noted, related to time of day.

This species is polymorphic in the sense of Wilson (1935); for the
purpose of this investigation the workers were classified in 3 groups,
minors, medias and majors. Samples of 15 specimens of each group
were selected later for head width measurements in order to check
the adequacy of the classification. Specimens that could not be
readily placed in one of the size classes were not included in the
ethogram.

Results

The behavioral catalog of Formica perpilosa is presented on
Table 1. Twenty-eight behavioral categories were observed in the
minor category, 34 in the media, and 1 1 in the major. By fitting the
frequency data to a lognormal Poisson distribution (Fagen &
Goldman, 1977), the total numbers of categories, including those

1978]

Brartdao — Worker Caste of Formica perpilosa

231

Table 1. Relative frequencies of behavioral acts by the three worker categories of
a single colony of Formica perpilosa. (N, total number of behavioral acts recorded
for each caste). Approximate population of the nest: one nest queen, 150 minor
workers, 270 media workers, 30 major workers, 50 eggs, 7 larvae and 5 pupae.

Behavioral Acts
A — Grooming

1 — Autogrooming head, 1st pair of legs

2 — Autogrooming 2nd pair of legs

3 — Autogrooming abdomen, 3rd pair
of legs

4 — Allogrooming minor workers

5 — Allogrooming media workers

6 — Allogrooming major workers

7 — Allogrooming nest queen

B — Brood Care

8 — Standing at the brood pile

9 — Carry egg (or eggs in succession)

10 — Lick egg (or eggs in succession)

1 1 — Carry larva (or larvae in succession)

12 — Feed larva

13 — Lick larva (or larva in succession)

14 — Carry pupa

15 — Lick pupa

16 — Assist eclosion to adult

C — Regurgitation Behavior

17 — Regurgitation with minors
18 — Regurgitation with medias

19— Regurgitation with majors

20 — Donating to the Queen

21 — Lay trophic egg

22 — Feed queen trophic egg

23 — Feed larva trophic egg

24 — Carry infrabucal pellet

25 — Feed on infrabucal pellet

D — Working

26 — Foraging outside the nest

27 — Feed on diet

28 — Feed on honey

29 — Feed on cockroaches

30 — Carry live nestmate

31 — Carry dead nestmate

32 — Drag the nest Queen

33— Handle nest material

34 — Carry debris

35 — Lick nest wall

36 — Excavating

E — Other Behaviors

37 — Antennal tipping

38 — Jittering

Total

no. behavioral categories observed
per caste

minor

media

major

workers

workers

workers

N = 996

N = 1679

N = 134

.167

.131

.224

.060

.062

.052

.023

.022

0

.054

.019

0

.043

.095

.052

.007

.012

.030

.011

.049

.022

.066

.077

.134

.033

.055

0

.016

.053

0

.009

.024

0

0

.004

0

.011

.031

0

0

0

0

0

0

0

.002

.024

0

.070

.024

.052

.040

.165

.291

.007

.021

.045

0

.009

.008

0

0

0

0

.001

0

0

.001

0

.003

.009

.090

.007

.011

0

.082

.021

0

.022

.004

0

.029

.025

0

.020

.007

0

0

.001

0

0

.003

0

0

.001

0

.018

.005

0

.033

.003

0

.059

.034

0

.092

.012

0

.001

.002

0

.016

0

0

1.0

1.0

1.0

28

34

11

232

Psyche

[June-Septernber

not seen, were estimated to be respectively 29, 36 and 1 1, with 95 per
cent confidence intervals of (25, 33) (25, 47) and (5, 19).

Comments on Table 1:

Some behavioral categories deserve special mention:

1 . Carry pupa or lick pupa: Care of pupae consisted exclusively of assisting eclosion
to adult. The pupae remained in the brood chamber and only when the colony was
disturbed (which did not happen during the drawing of the catalog) media workers
carried them to the new nest.

2. Lay trophic egg: The actual laying of trophic eggs was not seen, but 3 times
media workers were seen offering the larvae and the queen small, round, shiny
objects, looking like eggs, but different from the normal ones laid by the nest queen.
It is not impossible that the laying of trophic eggs is a real but rare event.

3. Recruitment behavior: In order to determine whether the workers utilize any
kind of food recruitment, the colony was deprived of food for one week and the nest
connected to an arena by a bridge of round sticks. Honey water was presented on the
arena floor. Two periods of one hour observation were recorded. Initially all ants
that reached the food source were collected and not allowed to return by the bridge.
Between the two experiments the honey source, the bridge and the arena floor were
changed to avoid recognition by the ants. In the first period 35 ants reached the food
source, in the second 262. This experiment clearly shows that this species uses food
recruitment. Returning ants were observed to rub the tips of the abdomen on the
bridge sticks (probably laying a scent trail) and to display no nestmates. During the
construction of the catalog, however, no recruitment behavior was noted, probably
because the colony was kept fed to saturation.

4. Jittering and antennal tipping: These behavioral categories were described by
Wilson (1976), but nothing is known about their meaning.

5. Defensive behavior: Our perpilosa colony was not stressed to ellicit defensive
behavior. However, a stray individual of the ant Novomessor cockerelli was found on
the nest floor being attacked by media and minor workers of Formica perpilosa: not
even in this situation the majors were observed outside the nest. This of course does
not mean that majors cannot play a role in defending the nest against predators or
raids by other ants, but it is apparent defense is not their characteristic behavior. A
more detailed account of the defensive behavior of this species should be useful.

After the behavioral catalog was drawn, 50 specimens had their
head widths and lengths measured and plotted. The linear ana-
morphosis afforded a linear regression of head length on head
width, where the coefficient of determination is r2 = .964. When
logarithmically plotted, in order to compare with Wilson’s model

1978]

Brandao — Worker Caste of Formica perpilosa

233

(1953), the log-log anamorphosis afforded a linear regression of
head length on head width. The equation is

log L = .2795 - .8602 log W

or, alternatively,

L = 1.9033 W"8602

The coefficient of determination, r2 is also .964, showing of course
an excellent fit.

I have chosen the following criteria for the definition of the
categories of workers: minors, head width 1 mm or less; medias, 1 to
1.4 mm; majors, larger than 1.4 mm.

The majors represent only 6.6% of the nest population and their
behavioral catalog comprises merely 1 1 behavioral categories. Their
behavior, as seen in Table 2, is mostly directed toward grooming
and regurgitation with nestmates.

Discussion and Conclusion

The behavioral catalog and the head shape curve of our Formica
perpilosa colony show that the worker caste is strongly polyethic, in
spite of being only slightly polymorphic. The media category is
responsible for most of the behavioral categories. The majors are
specialized in regurgitation with nestmates and may act as a trophic
subcaste as in Camponotus (Colobopsis)fraxinicola (Wilson, 1974).
F. perpilosa majors also possess a relatively large abdomen and

Table 2. Relative frequencies of groups of behavioral acts listed on Table 1 by
the three worker subcastes in a single colony of Formica perpilosa. (N. total
number of behavioral acts recorded for each subcaste.)

Groups of
behavioral acts
(from Table 1)

A — Grooming
B — Brood Care
C — Regurgitation
D — “Working”

E — Jittering & Antennal tipping

minor

media

major

workers

workers

workers

N = 996

N = 1679

N = 134

.365

.390

.380

.137

.248

.134

.127

.241

.486

.355

.116

0

.017

.002

0

1.0

1.0

1.0

234

Psyche

[June-September

were never observed performing any defensive or “working” be-
havior. They were recorded as using almost half of their time doing
trophallaxis or offering infrabucal pellets to minors and medias.

King and Walters (1950) have found a very similar situation in
“Formica rufa melanotica” (=F. obscuripes) in natural nests. They
were able to prove a correlation between polymorphism and
polyethism, showing that minor workers were specialized in attend-
ing aphids. The major workers again remained inside the nest most
of the time.

On Table 2 one can notice that all the categories of F. perpilosa
spend the same amount of time performing grooming behavior. The
minors are almost exclusively responsible for several tasks grouped
as “working” and the medias spend a more uniform amount of time
performing each group of behavioral acts.

Wilson (1953) proposed a model for the origin and evolution of
polymorphism in ants. In 1971 he reviewed the ant caste system and
said that five steps can be recognized in the evolution within the
worker caste: monomorphism, monophasic allometry, diphasic
allometry, triphasic allometry and complete dimorphism.

Age polyethism is the responsible phenomenon for caste structure
in monomorphic species, and the polyethic classes are sometimes
referred to as physiological classes. Traniello (1978) reported an
apparent lack of temporal division of labor in the primitive ponerine
ant Amblyopone pallipes, which appears to have the most primitive
caste system yet documented in ants.

Monophasic allometry is the commonest manifestation of non-
isometric growth. Studies on the division of labor in weakly
polymorphic species showed a “by preponderance” division of
labor, i.e., any given worker should be capable of performing any
given normal task.

In Formica polyctena size variation is weakly correlated with
division of labor. The behavioral variation observed by Otto (1958)
consisted mainly of age polyethism and individual peculiarities.
Although the F. perpilosa subcastes were not divided using age
parameters, I believe that age does not account for a large fraction
of the total behavioral variation.

Majors of advanced polymorphic ant species, especially com-
pletely dimorphic species, where intermediates no longer exist and
the two remaining classes are remarkably different in morphology,

1978]

Brandao — Worker Caste of Formica perpilosa 235

usually act as soldiers. This is not true in the present case, where
another kind of specialization was revealed.

In the complete dimorphic species there is no behavioral overlap
between the castes. Each category can be easily distinguished from
the others morphologically and behaviorally.

Majors of Zacryptocerus varians (members of one of the nine
genera in which complete dimorphism is easily recognized) function
primarily in defense (Wilson, 1976) and deserve the title of soldiers.
Yet they sometimes wash and manipulate larvae and pupae. But
during a high intensity attack, majors block the nest entrance with
their saucer-shaped heads and are more persistent and effective than
minors in forcing the enemy back to the entrance and finally out of
the nest.

Dr. Edward O. Wilson, reviewing the manuscript, was kind
enough to suggest that F. perpilosa majors represent an early stage
in the evolution toward repletes, as seen in Proformica and Myr-
mecocystus (Wilson, 1971). The specialization shown by perpilosa
majors (see Table 2) agrees with this view.

F. perpilosa seems to occupy, from the viewpoint of evolution of
division of labor, an intermediary position between monomorphic
and completely dimorphic species.

Acknowledgments

I am indebted to the “Fundagao de Amparo a Pesquisa do Estado
de Sao Paulo” for a scholarship (Biologicas 77/1208). I wish to
express my appreciation to my parents for supporting my visit to
Harvard University; and to Prof, and Mrs. B. Patterson, Dr. and
Mrs. W. L. Brown, Jr., and Dr. E. E. Williams for their kind
hospitality.

I am specially grateful to Dr. E. O. Wilson for the opportunity to
learn ant rearing techniques, for the suggestion of the problem,
assistance and uses of facilities during the research and critically
reviewing the manuscript. I thank Dr. and Mrs. B. Holldobler,
Gary Albert, Hiltrud Engels and specially Katherine Horton and
James Traniello for advice and help with the ethogram method, and
also the latter for showing me his unpublished data on Amplyopone
behavior; Mr. O. Schmidt for conducting the repertory estimations;
Dr. Francisca C. do Val for critically reading the manuscript.

236

Psyche

[June-September

Finally, I am grateful to Dr. P. E. Vanzolini for the helpful
suggestions during the elaboration of this paper.

References

Bhatkar, A., and W. H. Whitcomb.

1970. Artificial diet for rearing various species of ants. Florida Entornol. 53:
229-232.

Buren, W.F.

1968. Some fundamental taxonomic problems in Formica (Hymenoptera:
Formicidae). J. Georgia Entornol. Soc., 3(2): 25-40.

Fagen, R. M., and R. N. Goldman.

1977. Behavioral Catalogue Analysis Method. Anim. Behav., 25(2): 261-274.

Gregg, R. E.

1963. The ants of Colorado. University of Colorado Press, Boulder, Colorado,
xvi + 792 pp.

Kay, C. A., and W. G. Whitford.

1975. Influences of temperature and humidity on oxygen consumption of five
Chihuahuan desert ants. Comp. Biochem. Psysiol. A., 52: 281-286.

King, R. L., and F. Walters.

1950. Population of a colony of Formica rufa melanotica Emery. Proc. Iowa
Acad. Sci., 57: 469-473.

LaBerge, W. E.

1952. Locality records of two ants found in Kansas. J. Kansas Entornol. Soc.
25: 59.

Oster, G. E., and E. O. Wilson.

1978. Cast and Ecology in the Social Insects. Princeton University Press,
Princeton, New Jersey (in press).

Otto, D.

1958. Uber die Arbeitsteilung im Staate von Formica rufapratensis minor
Gosswald und ihre. Verhaltensphysiologischen Grundlagen: Ein Beitrag
zur Biologie der Roten Waldarneise. Wiss. Abh. Deutsche Akad.
Landw. — Wiss Berlin 30: 1-169.

Schumacher, A. M., and W. G. Whitford.

1974. The foraging ecology of two species of Chihuahuan desert ants: Formica
perpilosa and Trachymyrmex neomexicanus (Hymenoptera: Formici-
dae). Ins. Soc. 21: 317-330.

1976. Spatial and temporal variation in Chihuahuan desert ant faunas. South-
west Natur. 21: 1-8.

Traniello, J. F. A.

1978. Caste in a primitive ant: Absence of age polyethism in Amblyopone.
Unpublished manuscript.

Whitford, W. G., C. A. and A. M Schumacher.

1975. Water loss in Chihuahuan desert ants. Physiol. Zool. 48: 390-397.

1978]

Brandao — Worker Caste of Formica perpilosa 237

Wilson, E. O.

1953. The origin and evolution of polymorphism in ants. Quat. Rev. Biol. 28:
136-156.

1971. The Insect Societies. Belknap Press of the Harvard University Press,
Cambridge, Massachusetts, ix + 548 pp.

1974. The soldier of the ant Camponotus (Colobopsis) fraxinicola as a trophic
caste. Psyche 81(1): 182-188.

1976. A social ethogram of the neotropical arboreal ant Zacryptocerus various
(Fr. Smith). Anirn. Behav. 24(2): 354-363.

CULTURE TECHNIQUES FOR ACANTHOPS FALCATA,
A NEOTROPICAL MANTID SUITABLE FOR
BIOLOGICAL STUDIES

(WITH NOTES ON RAISING WEB BUILDING SPIDERS)*

By Michael H. Robinson and Barbara Robinson

Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa,
Canal Zone (Panama).

Introduction

“It is both expensive and difficult to maintain a year-round
colony of mantids and a continuous supply of living insects upon
which to feed them. Because of their cannibalism, mantids must be
raised in individual containers, and the smaller males are invariably
in short supply.” Roeder, 1963, p. 141.

We think that we have found the solutions to these problems. We
have devised a simple and inexpensive culture regime and found a
species that is easy to manage in captivity. This species, Acanthops
falcata Stol, is small enough to raise in large numbers in a modest
amount of space and large enough to be convenient for a wide
number of biological investigations. Females that are sexually
receptive can be triggered to mate by a dark/ light transition and
males also become sexually active following such a transition.
Matings are thus readily manipulate by the experimenter. In
addition, the species is fecund and hardy.

Our interest in solving the problems of raising predatory arthro-
pods began in 1971, when we needed naive predators in order to
investigate instinctive behavior. We used two species of araneid
spiders, Argiope argentata and A. aemula, feeding them on dead
drosophiloid flies (Robinson & Robinson, 1976a). Since then we
have raised several generations of A. argentata and successfully
hand reared two other species of web building spiders from egg
cocoons, without the restriction of using dead prey, which makes
the problem simpler.

* Manuscript received by the editor September 8, 1978.

239

240

Psyche

[June-September

Later we became interested in the developmental biology of
sexual size dimorphism and intrasexual size poymorphism (Robin-
son & Robinson, 1976b, Robinson, B. & M. H. Robinson, 1978).
However, orb-weaving spiders are not easy subjects for this type of
study, and we turned our attention to mantids, in particular to the
dead leaf mantid, Acanthops falcata. A. falcata proved to be an
ideal subject and could become a most useful laboratory'animal. It
is an extremely efficient predator, attacking, at all instars, any live
prey from drosophiloid flies up to insects of almost its own size. In
captivity it becomes very ‘tame’ and will wait for food to drop when
the cage lid is raised, and will often take food directly from the
forceps. Although the species has to be raised in separate cages, the
containers we used were small, inexpensive, readily available and
easily manageable.

We think that A. falcata has great potentialities as an experi-
mental animal. It has an extensive behavioral repertory, with an
efficient predatory strike and a complex startle display (first de-
scribed by Crane, 1952). We have demonstrated (Robinson &
Robinson, in prep.) that receptive females secrete an attractant to
which males fly. The pheromone remains to be isolated. The female
has a relatively large head capsule and brain and the large eyes have
excellent movement detection and depth perception. It should be an
excellent subject for neurophysiological studies. Our success rate of
rearing nearly 1,000 individuals over several generations has been
about 95%, and out methods should prove of interest to ethologists,
neurophysiologists, biochemists and developmental biologists wish-
ing to work with this mantid.

Culture Methods

THE INSECT.

Acanthops falcata is a small neotropical mantid of the family
Hymenopodidae. For a mantid it has an unusual degree of sexual
dimorphism. The flightless female (Figure 1) resembles a curled
dead leaf and weighs 400-500mg. The male, which flies well,
resembles a flat dead leaf, and weighs under 200mg.

A brief summary of the life history is given here. Details of instar
duration, sizes and weights at each instar, variability of number of
instars and mating behavior are to be published separately (Robin-
son & Robinson, in prep.).

1978]

Robinson & Robinson — Acanthops falcata

241

Figure 1 . A female Acanthops falcata hanging from a twig in its cryptic posture.
Body length ca 45mm.

In captivity the females have a life expectancy of about 6 months.
They lay long slender oothecae which may contain as many as 60
eggs, but more usually, 25-35. Eggs can be produced every 8 days
and hatch in 16-19 days. We have records of females, that after one
mating, laid 16 fertile oothecae. In Panama, at ambient temperature
(about 25°C-30°C), development from egg to adult is usually
accomplished in 7 molts, including the molt from pronymph to first
instar, that takes place at eclosion. Postembrionic development
takes about 2 months, depending on food supply. We have kept
males alive for over a month and for most of this time they remain
sexually active. Even old males can be effectively mated. Unmated
females secrete a pheromone for a limited period each day until they
are mated. The period of secretion coincides with the brief daily
period when males fly. In the wild, this period is immediately after
dawn, but in the laboratory will occur after a dark/ light transition
(Robinson & Robinson, in prep.).

242

Psyche

[June-September

Cannibalism does not appear to be part of the mating behavior,
as it is in many other species of mantids (Roeder 1935, 1963).
Although males fall within the prey range of females, they seem to
escape readily unless conditions are overcrowded. Copulations last
from 20 minutes to 1 hour. After mating, the male drops from the
female and usually survives to inseminate several females.

THE CAGES.

To raise large numbers of mantids in individual cages we made
use of materials that are readily available and inexpensive. Trans-
parent plastic party “glasses” need only a loose fitting cover to make
a single plastic container into a cage. We found that the covers of
plastic (disposable) petri dishes would fit one brand of 6oz shot
“glasses” while the bottoms would fit another. These lids can be
drilled to take a cotton ball insert that can be wetted to provide a
high humidity within the cages. (In Panama’s 80-100% relative
humidity, this is not necessary.) The plastic cages can be stored on
wooden trays, and stacked so that several hundred mantids occupy
relatively little space. Fiber-tipped pens, sold as freezer markers, are
excellent for marking index numbers and other data on the plastic
cages. In natural conditions, A. falcata rests hanging from a thin
twig. To provide a perch, we placed a piece of bamboo in each cage,
resting diagonally from the top at one side to the bottom of the
opposite side (Figure 2).

6oz shot “glasses” are large enough for an adult female, but
occasionally prove too small for an adult female to emerge success-
fully from the final molt. Ecdysis is diurnal, quick and usually
without complications, taking about 30 minutes for all but the final
molt, which takes about twice as long. As the majority of the insects
ecdyse between 10am and 3pm, we solved the cage size problem by
watching individuals that were due to molt and removing them from
the cage. Once molting had started, the stick with the molting
mantid on it was placed in a piece of florists’ clay or Play-Doh. Here
the mantid ecdysed, turned and extended its wings. After a success-
ful molt we returned the mantid, on its stick, to the cage.

We used several different cages in which to introduce males and
females for mating. Out of doors we used a 2m X linX lm screened
cage which was placed over several shrubs. Pheromone secretion by
females, male flights and mating occurred only in the first 15
minutes after dawn. We also used a twelve gallon glass aquarium as

1978]

Robinson & Robinson — Acanthops falcata

243

Figure 2. Left. Rearing cage for Acanthops falcata. (See text) A is a plastic petri
dish and B a 6oz shot “glass”. The female is shown in her resting posture on a piece of
bamboo (C).

Right. A larger cage, suitable for spiders and large mantids. (See text) D is a cylinder
of transparent acetate sheeting, lOcrn high, joined with cellulose tape. E shows the
double lid made from a plastic petri dish and cover. The inner section has a hole of 5
cm diameter cut from it. The outer section, which is plugged with a moist cotton ball,
can be raised to introduce food, without disturbing the animal. The positions of the
cylindrical toothpicks which provide web attachment points and footholds are
indicated.

a mating arena. This was fitted with a screen top and held two or
three potted plants. In the enclosed space males seemed to have
difficulty locating a secreting female and if males and females were
left together in such a container outside the mating period, males
were eaten. Several males can be left together, however, and females
can be introduced when they are secreting. The aquarium had the
advantage that the mantids could be kept in it in the dark, and we
could initiate mating behavior without disturbing them, by bringing
them into a lighted area. Later we developed a lin X lm X .5m
screened cage as a mating arena. Mantids from the dark phase of a
controlled light regime can be brought into the light and released
into such a cage, where they immediately become sexually active.
Results are better if the light intensity is low. Since secreting females
maintain a specialized posture (Robinson & Robinson, in prep.) it is
easy to determine when they are receptive.

244

Psyche

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Females produce perfectly normal oothecae in the rearing cages.
These are long and they hang down from the substrate to which they
are attached by a long stalk. We removed oothecae from rearing
cages, dated them and put them in 8oz clear plastic tunblers to
hatch. To allow the pronymphs to emerge normally and complete
their first molt on the outside, the ootheca must be attached to the
lid of the hatching cage so that it can hang downwards, surrounded
by free space. We tried various methods of attaching the ootheca
stalk to the lid. This can be done with adhesive tape, but the small,
fragile nymphs may stick to the exposed edges of the tape. It is more
satisfactory to pass the stalk through a small hole and tape it to the
outside of the lid. No additional moisture was necessary for most of
the year in Panama, but during the dry season eggs were sprayed
towards the end of the developmental period.

The pronymphs develop within the ootheca and at eclosion
ecdyse to the first instar. They stand on the ootheca for several
hours before dispersal and are easier to transfer to individual
rearing cages at this stage. During the first instar they are not
cannibalistic and can be kept communally but should be separated
soon after the next molt.

CULTURE TECHNIQUES.

Normally mantids will not strike at motionless prey. The first
three instars were fed exclusively on live drosophiloid flies. Some-
time during the 4th instar the diet was changed to include live
domestic crickets, Acheta domestica, supplied by Fluckers Cricket
Farm, and by the 5th instar, only live crickets were fed. The
Drosophila sp. were caught over a massive outdoor fruit culture in a
fine meshed net, which was then placed in a freezer. By trial and
error we determined the length of time necessary to immobilize the
flies without killing them, and how long it took for them to become
active again. The fastest way of introducing the flies into the cages
was also the simplest. We transferred a quantity of immobile flies
from the net onto a sheet of paper, and lifting each lid in turn, we
shook the required number of prey into each cage. At the first instar
this number was only two or three; towards the end of the fourth
instar it was about ten. We quickly learned by experience how many
mantids we could feed before the flies recovered on the paper, and,
by looking at the cage debris, could tell if the mantids were under or
overfed. The stage at which we changed the diet to crickets

1978]

Robinson & Robinson — Acanthops falcata

245

depended on the size of the crickets which were available. A. falcata
will reach maturity entirely on a diet of Drosophila, but the
developmental period is longer, and the adults that we raised on this
diet were abnormally small (Robinson, B. & M. H. Robinson, in
prep.). To make the crickets easier to handle, they too were placed
in a freezer for a short time, or stored in a refrigerator before being
dropped into the cages from forceps. Uneaten crickets left in cages
will eat ecdysing mantids or oothecae. The former is not a serious
problem with A. falcata which molts during the day, as it is with
species which molt at night, or whose molting time is longer. We
have raised two other species of mantid, Phyllovates chlorophaea
(see below) and Chaerododis rhombicollis (see Robinson 1969).
Both these species were vulnerable to cricket attack for about two
hours during each molt, which takes place at night. We had to
remove live crickets from their cages at the end of each day.

In the humid tropics drinking water does not seem to be necessary
for mantids that are fed daily. However, when they were transferred
to the mating cages containing wet plants, they were observed to
drink, males more than females. In air-conditioned or heated
laboratories, it may prove necessary to supply moisture. As de-
scribed above, cotton balls can be inserted into holes in the lids of
the cages. These can be sprayed to keep them moist. Care should be
taken that water does not form in droplets in the cages of first instar
nymphs, as they will not be able to escape from the surface tension.
We always use rainwater rather than tap water for spraying insects
or providing moisture in cages.

CAGES AND CULTURE TECHNIQUES SUITABLE FOR
OTHER ARTHROPODS.

We have also raised several other species of mantids successfully,
using the above methods for small species. For larger species we
used these techniques for early instars adapting them as necessary as
the mantids grew too large for the cages. Phyllovates chlorophaea is
a large, long-legged mantid, which we have raised successfully. For
this large insect 6oz shot glasses were suitable until the 4th-5th
instar. This species hangs below a leaf rather than from a twig. In
order to give it a foothold on the lid of the cage, we attached a piece
of masking tape, approx. 3cm - 5cm to the undersurface. The cages
that we used for later instars and adults are adaptations of
lepidoptera rearing cages, designed by R. Silberglied and A. Aiello.

246

Psyche

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These consisted of a cylinder of metallic hardware cloth, 10cm high
and with a diameter very slightly larger than that of a plastic petri
dish. The petri dish fitted snugly inside the cylinder as a base, and a
cover was placed over this, enclosing the bottom of the cylinder
between the two. The top was bound with masking tape, and a
second petri dish cover was used as a lid. Screened cages cannot, of
course, be used until the mantid has transferred to a diet of crickets.
For the final molt, when the mantid hangs from its exoskeleton until
the legs harden, a 20cm high cage was needed. Rather than build
new cages for this very short period, two cages were joined together,
by removing the base of the upper one and attaching it to the top of
the lower one with masking tape. After ecdysis, the double cage was
disassembled and the adult mantid was returned to its original cage.

Feeding procedures were exactly the same for P. chlorophaea as
those described for A. falcata (above). However, either P. chloro-
phaea attacks smaller prey, or domestic crickets are not preferred
prey. It was necessary to feed several small crickets daily, and to
remove any uneaten prey at the end of each day. We raised P.
chlorophaea to study its life history. It needs more care, more space
and more complex cages than A. falcata. In addition, its mating
behavior is similar to that of Mantis religiosa (Roeder, 1935), where
males frequently do not survive to inseminate more than one female.

The lepidoptera rearing cage (above) can be modified so that live
Drosophila can be fed, if the cylinder is made of transparent acetate
sheeting. With the following modifications we found that they made
excellent cages for raising web-building spiders, particularly those
with horizontal webs. (See Figure 2.) The acetate cylinder was made
to fit tightly into a plastic petri dish base. An inner lid was made by
cutting a circle of 5cm diameter out of a second petri dish, using an
electric soldering iron. A petri dish cover was used as an outer lid.
The outer lid, which was plugged with a cotton ball to keep the cage
humid, could be raised for feeding the spiders, without disturbing
web foundation lines. To give the spiders web attachment points,
four cylindrical toothpicks were pushed through the acetate until
they almost met in a cross, 2cm from the top and 2cm from the
bottom of the cage.

We initially kept web building spiders in glass vials with snap caps
(Robinson & Robinson, 1976a). These are not ideal, as the web is
often attached to the cap, and the spider cannot be fed without the
web being damaged. Later we discovered the 1 inch diameter
transparent plastic storage tubes, supplied by Forestry Supplies Inc.

1978]

Robinson & Robinson — Acanthops falcata

247

These come in 18 inch lengths, with separate plugs. They can be cut
to the required size and fitted with a removable plug at each end.
One cap was fitted with a moist cotton ball and either cap could be
used to introduce food. Webs were rarely attached to both caps.

FEEDING SPIDERS.

Drosophila, the most readily available food, are probably larger
than the natural prey of 2nd instar spiderlings of most species and
when used live usually break the fragile web and escape. Dead
Drosophila placed in a spiderling’s web are usually found and eaten
soon after being introduced. If they are ignored at this stage, the
spider feeds on them when the web is taken down. For later instars,
prey of an appropriate size, immobilized in the freezer, can be
placed or dropped in the webs. Spiders drink water from their webs,
and even in the humid tropics, once the spider has been transferred
from a vial to a cage we spray the webs with rain water daily.

We are (September 1978) presently undertaking a long research
trip to Papua New Guinea and have given our culture of Acanthops
falcata to the Insect Zoo, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, who hope to
maintain it.

References

Crane, J.

1952. A comparative study of innate defensive behavior in Trinidad mantids
(Orthoptera, Mantoidea), Zoologica, N.Y., 37: 259-263.

Robinson, B. & M. H. Robinson.

1978. Developmental studies of Argiope argentata (Fabricius) and Argiope
aemula (Walckenaer). Symp. Zool. Soc. Lond., 42: 31-40.

Robinson, M. H.

1969. The defensive behaviour of some orthopteroid insects from Panama.
Trans. R. ent. Soc. Lond. 121: 281-303.

Robinson, M. H. & B. Robinson.

1975. Techniques in the field study of spiders. Bull. Brit. Arach. Soc., 3: 160-165.

1976a. Discrimination between prey types: an innate component of spider
predatory behaviour. Zeit. f. Tierpsychol. 41: 266-276.

1976b. The ecology and behavior of Nephila maculata: a supplement. Smith-
sonian Contrib. Zool. 218: 1-22.

Roeder, K. D.

1935. An experimental analysis of the sexual behavior of the praying mantis
( Mantis religiosa L.), Biol. Bull. 69: 164-184.

1963. Nerve Cells and Insect Behavior, Harvard Univ. Press.

SEARCHING BEHAVIOR OF
HIPPODA MIA CONVERGENS LARVAE
(COCCINELLIDAE: COLEOPTERA)*

By Kenneth W. Hunter, Jr.

Uniformed Services University of the Health Sciences
Bethesda, Maryland 20014

Introduction

Survival and development of predaceous Coccinellidae depend in
large part on their ability to find food (Hodek, 1973). Coccinellid
larvae exhibit different searching patterns before and after finding
prey; the path of a larva just after consuming a prey item is generally
more tortuous than before the encounter (Banks, 1954; Banks, 1957;
Kaddow 1959). When the prey are gregarious, this type of altered
searching behavior is thought to increase the chance of finding
additional prey (Banks, 1957). In the present study I describe the
searching behavior of larvae of the convergent lady beetle, Hippo-
damia convergens (Guerin), before and after feeding on the spotted
alfalfa aphid, Therioaphis maculata (Buckton).

Materials and Methods

Adult H. convergens were collected from a field of alfalfa located
at the Arizona State University Experimental Farm, Tempe, AZ.
Copulating pairs were isolated for 48 hrs, then the females were
removed and placed in six-dram plastic vials lined with paper
toweling. The toweling was moistened periodically. Isolated females
were supplied daily with thirty fourth-instar or adult T. maculata
(Nielson and Currie, 1960) collected from the alfalfa field by sweep
netting. The vials were incubated at 32° C for 4 or 5 days during
which time the fertilized females deposited clusters of eggs on the
toweling; then the female beetles were removed and the brood
chambers incubated for another two days. After hatching, the first
instar larvae remain clustered around the egg shell, but because of
cannibalism it was necessary to immediately separate the newly
hatched larvae. Individual larvae were transferred to new vials, by

* Manuscript received by the editor October 17, 1978.

249

250

Psyche

[June-September

the use of a fine camel-hair brush, together with five immature
aphids. After moulting to the second instar and thereafter the larvae
were provided thirty aphids per day. Following the third molt the
new fourth instar larvae were used for experimentation.

To study searching behavior before and after feeding, an artificial
searching arena was constructed similar in design to the arenas of
Fleschner (1950) and Banks (1957). A twelve inch square plywood
board was covered with buff-colored art paper and surrounded by
an electrically heated wire. The arena was uniformly illuminated by
a fluorescent light suspended two feet above the center, and all tests
were performed at room temperature. Tracks of the larvae were
recorded by tracing lightly with a pencil, and 30 sec intervals were
noted on the tracks. Track diagrams were enlarged with an over-
head projector. The number of degrees of each turn, whether to the
right or left, was measured with a protractor. The data were
expressed as the total number of degrees turned, which was a
function of the number and type of turns the larvae made. Statistical
analysis was done with Student’s t-test.

Results

Sixteen fourth instar H. convergens larvae were starved for 4 hrs
prior to testing. Each larva was placed in the center of the arena
under a vial, and timing commenced when the vial was removed.
The larva was followed by a pencil tracing and 30 sec intervals were
recorded on the track for a total test period of 5 min. At this time an
adult T. maculata was placed directly in front of the larva; the aphid
was seized and rapidly consumed. The track was again traced when
the larva began to move after feeding, and 30 sec intervals were
recorded for 5 min. This test procedure was repeated for each of
sixteen larvae.

The movements of larvae before feeding were less tortuous and a
much larger portion of the arena was searched; after feeding the
larvae concentrated their search in the vicinity where the aphid was
discovered. The track consisted of numerous turns and frequently
areas previously searched were revisited several times. Analysis of
the tracks confirmed these observations (Fig. 1). For three minutes
following the consumption of an aphid the larvae made more turns
than they did prior to feeding, but in the final two minutes of
observation the fed larvae appeared to revert to their pre-fed

1978]

Hunter — Hippodamia convergens Larvae

251

MINUTE INTERVALS

Figure 1. Searching movements of fourth instar H. convergens larvae during a
five minute interval before and after consuming one adult T. maculata. The bar
represents the cumulative mean ± one standard deviation of the numbers of degrees
turned by larvae for five 1 -minute-intervals prior to feeding.

252

Psyche

[June-September

searching pattern. The total number of degrees turned was 6200.3 ±
992.0 before feeding and 12058.8 ± 1798.8 after feeding(p< 0.001).

Discussion

The searching movements of fourth instar H. convergens larvae
before finding prey generally consist of wide sweeping turns. After
consuming one adult T. maculata the searching movements are
modified; initially movements consist of many small turns in the
immediate vicinity of the previously consumed prey. With time the
movements become more characteristic of the pre-fed state. Banks
(1957) noted a similar searching behavior in a study of Adalia
bipunctata (L.) and the aphid Myzus persicae (Sulz.), and Kaddow
(1959) found the same searching movements in larvae of Hip-
podamia quin que si gnat a (Kirby) fed pea aphids, Macrosiphum pisi
(Kaltenbach). Modification of turning movements after finding the
first host is common to other entomophagous and parasitic insects
(Fleschner, 1950; Laing, 1937).

It appears that coccinellid larvae lack sophisticated sensory
apparatus and do not discover their prey until actual contact occurs
(Hodek, 1973). Furthermore, this undirected searching is very
inefficient in that much time and energy are wasted revisiting areas
previously searched (Banks, 1957). If one assumes that the searching
behavior of coccinellid larvae is indeed undirected, then the number
of encounters between predator and prey would be merely a
function of their respective population densities. However, this
assumes that the chance of finding a second prey is the same as that
for the first. On the other hand, if after the first prey encounter the
predator alters its pattern of search in such a way as to increase the
likelihood of capturing additional prey, then the predator has
become more efficient even though its searching is still undirected
(in terms of sensory capability). T. maculata are colonial and not
uniformly dispersed throughout their habitat. With this character-
istic in mind it would seem advantageous for H. convergens larvae,
having encountered a single T. maculata, to modify their searching
behavior in order to exploit the gregarious nature of their prey.
Since survival and reproduction of predators like H. convergens is
dependent upon finding adequate food (Hodek, 1973), a strong
selection pressure probably exists for behaviors which improve food
finding capabilities. Therefore, it is suggested that the change of

1978]

Hunter — Hippodamia convergens Larvae

253

searching pattern by H. convergens larvae after finding one prey
item is adaptive.

Acknowledgements

I would like to thank Drs. John Alcock and Gordon Bender of
the Department of Zoology, Arizona State University, for their
encouragement of this study. Publication costs were covered by
funds from the Uniformed Services University of the Health
Sciences, Department of Defense.

References

Banks, C. J.

1954. The searching behavior of Coccinellid larvae. Br. J. Anim. Behav. 2:
37-38.

1957. The behavior of individual Coccinellid larvae on plants. Br. J. Anim.
Behav. 5: 12-24.

Dixon, A. F. G.

1959. An experimental study of the searching behavior of the predatory
Coccinellid beetle Adalia decempunetata (L.) J. Anim. Ecol. 28: 259-
281.

Fleschner, C. A.

1950. Studies on the searching capacity of the larvae of three predators of the
citrus red mite. Hilgardia. 20: 233-265.

Hodek, I.

1973. Biology of Coccinellidae. Academia Publishing House of the Czechoslo-
vak Academy of Sciences, Prague, 1973, 260 pp.

Kaddow, I.

1959. The feeding behavior of Hippodamia quinque signal a (Kirby) larvae.
Univ. Calif. Pubs. Entomol. 16: 181-228.

Laing, J.

1937. Host finding by insect parasites. I. Observations on the finding of hosts
by Alysia manducator, Mormoniella vitripennis, and Trichogramma
evanescens. J. Anim. Ecol. 6: 298-317.

Nielson, M. W., and W. E. Currie

1960. Biology of the convergent lady bird beetle when fed a spotted alfalfa
aphid diet. J. Econ. Entomol. 53: 257-259.

FURTHER STUDIES OF THE MYRMICINE STING

APPARATUS: EUTETRAMOR1UM, OXYOPOMYRMEX,

AND TERATANER (HYMENOPTERA, FORMICIDAE)*

By Charles Kugler
Department of Entomology
Cornell University
Ithaca, New York 14853

Introduction

In an earlier investigation (Kugler, 1978, 1979) I described the
sting apparatus of representatives of 63 genera of myrmicine ants. In
so doing, it was shown that this complex structure has clear
potential for defining myrmicine genera and perhaps generic group-
ings. Furthermore, its morphology may have played an important
role in the evolution of some genera.

Here I present descriptions of members of 3 genera that could not
be included in that work. Structural affinities with the sting appa-
ratus of other genera are discussed in order to assist those reclassify-
ing this taxonomically problematical subfamily.

Methods

The materials and methods used are described in detail in Kugler
(1978). For the sake of brevity, figures from that earlier study are
cited here in italics; new figures of this paper are cited in Roman
type. Voucher specimens are deposited in the Harvard Museum of
Comparative Zoology, labeled “Kugler study 1978.” Scale lines in
the figures are in millimeters.

Eutetra morium

Species examined: E. mocquerysi, 2 workers.

Spiracular plate: (Fig. 1) Body subtriangular, only slightly longer
than wide. Anterior apodeme wide along nearly whole length, end-
ing abruptly near dorsal margin of plate. Ventral and posterior

*A report of research of the Cornell Agricultural Experiment Station, New York
State College of Agriculture and Life Sciences at Cornell University, Ithaca, New
York 14853.

Manuscript received by the editor September 15, 1978.

255

256

Psyche

[June-Septernber

Figures 1-4. Sclerites of the sting apparatus of Eutetramorium mocquerysi. Fig.
1, spiracular plate, side view. Fig. 2, apical !4 of lancet, side view. Fig. 3, sting and
furcula, side view. Fig. 4, sting and furcula, ventral view. Abbreviations: AAP,
anterior apodeme; ALP, anterolateral process of sting base; BD, body; BR, basal
ridge; DA, dorsal arm of furcula; DR, dorsal ridge; HE, hernocoel; IAP, internal
apodeme; LA, lateral arm of furcula; PAP, posterior apodeme; SBLB, sting bulb;
SS, sting shaft; VC, valve chamber; VR, ventral ridge.

apodemes present, well sclerotized. No dorsal notch or posterodor-
sal lobe. Spiracle relatively small, located caudad of center.

Quadrate plate: Body and apodeme both subrectangular, equal in
width; body extends well ventrad of apodeme. Dorsal edge of plate
convex, with moderately wide medial and lateral lobes terminating
in an acute anterodorsal corner.

Anal plate: Very poorly sclerotized, ill-defined; no sensilla.

1978]

Kugler — Myrmicine Sting Apparatus

251

Oblong plate: Posterior arm with narrow, well sclerotized apo-
derne, body widens abruptly just caudad of articulation with fuicral
arm, a lightly sclerotized ventral ridge arcs from intervalvifer
articulation to opposite fuicral arm. Anterior apodeme wide, promi-
nent, bluntly rounded. Ventral arm wide, broadly spindle-shaped
fuicral arm meets posterior arm perpendicularly, postincision deep.

Gonostylus: Two-segmented as indicated by bimodal sensilla
pattern and ill-defined membranous region; evenly tapered to end,
neither long and slender nor short and wide, not strongly dorso-
ventrally compressed. Distal segment with 7 long setae and isolated
dorsoterminal chaeta; proximal segment with 10 setae of varied
lengths. Distal segment grades evenly into terminal membrane. No
distal notch or basiconic sensilla.

Triangular plate: Body wide, basal portion abruptly tapered;
ventroapical process a distinct extension from body, narrowly
rounded apically. Dorsal and medial tubercles present.

Lancet: (Fig. 2) Straight, acute, well sclerotized, with distinct
ventral ridge and, distally, a dorsal ridge. Each lancet with 2 valves
of moderate size; the caudal one smaller, less sclerotized. No barbs
present.

Sting: (Figs. 3, 4) Long, slender, well sclerotized; sting bulb, valve
chamber and sting shaft regions distinguishable in profile. Sting
base not arched, anterolateral processes present. Valve chamber
internally much lower than sting bulb. Sting shaft with distinct
hemocoel throughout length, height of hemocoel greater apically
than subapically. No terminal flange. Campaniform sensilla present
on sting shaft and distal 2/3 of valve chamber. Index of reduction
26.8.

Furcula: (Figs. 3, 4) T-shaped in anterior view, lateral arms wrap
around sting base, dorsal arm well developed.

Discussion

The sting apparatus of Eutetramorium mocquerysi seems most
closely related to those of the Tetramoriini and the Leptothorax
genus group (see Kugler, 1978). Some common characters are: the
wide anterior apodeme and overall shape of the spiracular plate
(Fig. 57); the medial and lateral lobes, and shape of the anterodorsal
corner of the quadrate plate (Fig. 58); the shape of the anterior
apodeme, ventral ridge and body of the posterior arm of the oblong

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Psyche

[June-September

plate (Figs. 52, 59); much of the shape of the sting (Figs. 55, 56). The
gonostyli are much like those of Leptothorax (Fig. 53), but not as
long and narrow, and they lack the companion seta. The sting base
and furcula have characteristics of those of Liomyrmex cf. aurianus
(Figs. 69, 70).

In spite of its similarities with the Tetramoriini, E. mocquerysi
lacks the dorsal flange on the end of the sting and the character-
istically stubby gonostyli of that tribe. On the basis of the flange
character and other external characters Bolton (1976) removed
Eutetramorium from its traditional association with the Tetra-
moriini. He prefers to place it in the Myrmicini, but as an alternative
hypothesis, based admittedly on one character system, I would
suggest a position nearer Leptothorax and related genera.

OXYOPOMYRMEX

Species examined: O. tuneticus, 1 worker; O. tuneticus var. thoraci-
cus, 1 worker.

Spiracular plate: Body subrectangular, with straight anterior
edge, slightly concave posterior edge and convex dorsal and ventral
edges. Anterior apodeme forms a square anteroventral corner, then
a thin margin along anterior edge to dorsad of body of plate; medial
connection membranous. Large V-shaped dorsal notch present; no
distinct posteroventral tubercle or posterodorsal lobe.

Quadrate plate: Subtriangular, both body and apodeme much
narrowed at base. Dorsal edge straight, anterodorsal corner broadly
rounded. Anterior edge poorly sclerotized, expecially dorsad.

Anal plate: Weakly sclerotized, longer than wide, with 4 long
setae on terminal edge in O. tuneticus, 2 setae in O. t. var.
thoracicus.

Oblong plate: Dorsal ridge and body of posterior arm long and
narrow, uniform in width, without ventral ridge. Anterior apodeme
very long and slender. Ventral arm tapered and subtruncate distally;
fulcral arm slender, weak, diffuse dorsad, forming acute angle with
posterior arm.

Gonostylus: In O. t. thoracicus, narrow from side view, with no
indication of segmentation; 17-20 setae scattered more or less
evenly along distal 2/3 of ventrolateral surface. Dorsoterminal
chaeta, companion seta and terminal membrane present. No basi-
conic sensilla. Gonostyli of O. tuneticus lost in preparation.

1978]

Kugler — Myrmicine Sting Apparatus

259

Figures 5-6. Sting and furcula of Oxyopomyrmex tuneticus. Fig. 5, O. tuneti-
cus, side view. Fig. 6, O. tuneticus var. thoracicus, ventral view.

Triangular plate: In O. tuneticus body and ventroapical process
very slender and distinct; in O. t. var. thoracicus body wider dorsad
and merging more with ventroapical process. In both specimens,
body evenly tapered to ramus; neither dorsal nor medial tubercle
present.

Lancet: Long, weak, and spatulate distally, with broadly rounded
apex; about twice as deep as end of sting shaft. One large lancet
valve per lancet.

Sting: (Figs. 5, 6) Sting bulb large with weak basal ridge and
distinct anterolateral processes. Valve chamber well developed,
clearly distinguishable from sting bulb and sting shaft in profile and
in ventral view; internal apophysis long. Sting shaft short, very
slender and weak; no dorsal flange. Index of reduction 21.9 for O.
tuneticus, 20.4 for O. t. var. thoracicus.

Furcula: (Figs. 5, 6) Slender arch, uniform in diameter; no dorsal
arm.

Discussion

The sting apparatus of Oxyopomyrmex tuneticus and O. t. var.
thoracicus are very similar to those of Messor and Aphaenogaster.
The large dorsal notch and thin anterior apodeme of the spiracular

260

Psyche

[June-Septernber

plate (Fig. 130)', the subtriangular quadrate plate (Fig. 131)', the long
slender posterior arm and anterior apodeme and the inclined fulcral
arm of the oblong plate (Figs. 132, 133)', and the shape of the sting
bulb, valve chamber and furcula (Figs. 138, 139) are common
characteristics of the species so far examined in these 3 genera. The
Oxyopomyrmex species, however, are distinct in two important
respects: the sting shaft has no dorsal flange (Fig. 138), and the
lancets are spatulate and broadly rounded apically, rather than
acute (Fig. 137).

The close affinities of Oxyopomyrmex, Messor and Aphaeno-
gaster as suggested by the sting apparatus are consistent with the
views advanced by Emery (1922) and Wheeler (1910: 139-140, 268;
1922).

Terataner

Species examined: T. alluaudi, 2 workers.

Spiracular plate: (Fig. 7) Body subrectangular. Anterior apodeme
narrow, well sclerotized, continues dorsad to make completely
sclerotized connection with opposite side. Dorsal notch and ventral
and posterior apodemes present.

Quadrate plate: (Fig. 8) Apodeme subtriangular, no medial or
lateral lobes, anterodorsal corner rounded. Body subrectangular,
extending below apodeme.

Anal plate: Wider than long, clear suture between plate and anal
arc; moderately well sclerotized proximally, posterior and lateral
borders unsclerotized, undefined. Thirty-one long setae cover plate;
no sensilla basiconica.

Oblong plate: (Fig. 9) Posterior arm short and straight, dorsal
ridge strong, ventral ridge weak, ill-defined. Ventral arm wide
apically with narrow fulcral arm perpendicular to posterior arm;
postincision deep. Anterior apodeme rather short, acute.

Gonostylus: One-segmented. Dorsal surface free of sensilla,
broadly triangular. Lateral surface narrower, uniformly and densely
covered with long thin setae along distal Vi of length, longest setae
proximad; small dorsoterminal chaeta embedded among setae; no
obvious companion seta; no basiconic sensilla. Short terminal
membrane present.

Lancet: Long, slender, weak and flagelliform. Two intermediate-
sized valves per lancet, caudal valve smaller.

Figures 7-11. Sclerites of the sting apparatus of Terataner alluaudi. Fig. 7,
spiracular plate, side view. Fig. 8, quadrate plate, side view. Fig. 9, oblong plate, side
view. Fig. 10, sting and furcula, side view. Fig. 11, sting and furcula, ventral view.
Abbreviations: AAP, anterior apoderne; ADC, anterodorsal corner; AP, apoderne;
BD, body; DN, dorsal notch; FA, fulcral arm; IVA, intervalvifer articulation; PA,
posterior arm; PI, postincision; VA, ventral arm; VR, ventral ridge.

Sting: (Figs. 10, 11) Sting bulb wide, with heavy basal ridge.
Valve chamber small, indistinguishable from sting shaft in profile;
topped by heavy internal ridge; 2 large ventrolaterally projecting
prongs seem to originate on the internal apophysis. Sting shaft long,
slender, tapering to weak apex; hemocoel clearly visible in basal
2/3. Index of reduction 20.1.

Furcula: (Figs. 10, 11) A simple low arch, no dorsal arm;
extremities dilated at articulation with base of sting.

262

Psyche

[June-September

Discussion

The sting apparatus of Terataner alluaudi bears most resemblance
to several genera normally considered unrelated by most modern
inyrmecologists. It has a variety of derived characters in common
with Atopomyrmex mocquerysi, such as: shape of the anal plate;
form of the gonostyli; long, flagelliform lancets (Fig. 215 ); form of
the bulb, valve chamber and shaft of the sting, including 2 prongs
extending into the sting bulb from the internal apophysis (Figs. 216,
217). These species, however, differ markedly in the shapes of the
spiracular and oblong plates, and in the lack of a furcula and anal
setae in A. mocquerysi. With the Cephalotini, T. alluaudi shares the
following characters: complete medial connection of the anterior
apodeme of the spiracular plate (may not be derived); anal plate
wider than long and with numerous dorsal sensilla; long, well-
defined fulcral arm of the oblong plate; gonostylus shape and
setation (Fig. 192); shape of the lancets (Fig. 194); sting bulb shape,
low valve chamber and flagellate sting shaft as in Procryptocerus
scabriusculus (Figs. 196, 197). The main differences are the shape of
the spiracular plate (Fig. 198); and in T. alluaudi the lack of the long
pollicate anterodorsal process of the quadrate plate (Figs. 189, 198),
the furcula not appressed to the sting base, the more elongate sting
shaft containing a hernocoel, and the prongs in the sting bulb (cf.
Figs. 190, 191, 196, 197). With Cataulacus tardus, T. alluaudi shares
the medial connection of the spiracular plate anterior apodernes; the
shape and setation of the gonostyli; flagelliform lancets (though
much shorter in C. tardus); and prongs in the sting bulb (the number
of which is different, Figs. 203, 204). Such prongs are known only
from Terataner, Atopomyrmex and Cataulacus.

Both Emery (1922) and Wheeler (1922) placed Terataner in the
tribe Myrinecinini with Podomyrma, Lordomyrma, Atopomyrmex,
Myrmecina, Pristomyrmex and Acanthomyrmex, along with other
genera, the stings of which have not yet been examined. Emery more
specifically placed it with the first 3 genera in the subtribe Podo-
myrmiti after earlier creating the genus Terataner from some of the
species then in Atopomyrmex (Emery, 1912). The sting apparatus
does not support the groupings of the above genera into tribe
Myrmecinini (see also discussions in Kugler, 1978), but the view of
Terataner and Atopomyrmex as distinct, related genera is seen here
as likely. Sting apparatus morphology also suggests a relationship

1978]

Kugler — Myrmicine Sting Apparatus

263

between Terataner, the Cephalotini and possibly Cataulacus that
should be considered in future classifications of the Myrmicinae.

Acknowledgements

I thank W. L. Brown for supplying the specimens for dissection,
and for commenting on the manuscript. Research was supported by
the NSF grant DEB-22427 (W. L. Brown, Jr., principal investiga-
tor).

References

Bolton, B.

1976. The ant tribe Tetrarnoriini (Hymenoptera: Formicidae), constituent
genera, review of smaller genera and revision of Triglyphothrix Forel.
Bull. Br. Mus. (Nat. Hist.) Entornol., 34(5): 283-378.

Emery, C.

1912. Etudes sur les Myrmicinae. Ann. Soc. Entornol. Belg., 56: 94-105.
1922. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. Gen. Insect.
174: 1-397, 7 pi. (1921-1922).

Kugler, C.

1978. A comparative study of the myrmicine sting apparatus (Hymenoptera,
Formicidae). Stud. Entornol., 20: 413-548.

1979. Evolution of the myrmicine sting apparatus. Evolution, in press.
Wheeler, W. M.

1910. Ants, Their Structure, Development and Behavior. Columbia Univ.
Press, New York, xxv + 663 pp.

1922. Keys to the genera and subgenera of ants. Bull. Am. Mus. Nat. Hist. 45:
631-710.

AN UNUSUAL ASCALAPHID LARVA (NEUROPTERA:
ASCALAPHIDAE) FROM SOUTHERN AFRICA,

WITH COMMENTS ON LARVAL EVOLUTION
WITHIN THE MYRMELEONTOIDEA*

By Charles S. Henry

Biological Sciences Group, University of Connecticut, Storrs 06268

Ascalaphidae is a fairly large family of planipennian Neuroptera
that has received little attention from taxonomists since Weele’s
1908 monograph. Life history and behavioral studies of the group
have suffered even greater neglect; what is known has been sum-
marized in a previous work (Henry, 1977). As an order, Neuroptera
is taxonomically intractable, largely due to difficulty in interpreting
wing venation: extreme convergence is common in distantly related
families, yet divergence often occurs within a single subfamily or
genus. For this reason, the immature stages have proven to be more
reliable indicators of relationship than the adults, and our present
understanding of evolution within the order is based more on larval
than adult features (Withycombe, 1925; MacLeod, 1970). Since
such considerations apply as strongly to Ascalaphidae as they do to
other Neuroptera, all larval or life-history data are of paramount
importance to phylogenetic studies of the family.

Described here is a peculiar ascalaphid larva from Mkuze Game
Park, Natal, collected in November of 1967 by J. A. Slater and T.
Schuh. Although it shows all the diagnostic features of the family,
certain details of its setal morphology and of the form and
distribution of its thoracic and abdominal extensions (scoli) are
unique and of great phylogenetic importance. As the larva is
(necessarily) unassociated with an adult, I will describe its major
features informally and compare them critically with those of larvae
of known taxa in an attempt to deduce the general systematic
position of the insect. In addition, I will summarize what is known
of larval morphology in the superfamily Myrmeleontoidea, so that
conclusions concerning the Mkuse specimen can be placed in
perspective.

* Manuscript received by the editor December 12, 1978.

265

266

Psyche

[June-September

Methods and Materials

The larva was preserved in 70 percent ethanol. Wild™ M5 and
M5A stereoscopic dissecting microscopes were used to observe the
specimen, while drawings were made by means of integral camera
lucida attachments for these instruments.

Description of Larva

The African specimen is a typical ascalaphid immature in most
respects (Fig. 1), displaying robust jaws with three mandibular
teeth, large squarish head with pronounced occipital angles, prom-
inent ocular tubercles each bearing six dorsal and one ventral
sternrnata, and compact, lightly sclerotized body fringed by numer-
ous finger-like, setigerous extensions. It measures about 7 mm from
labral margin to anal spinneret and is probably a partially grown
second instar. Previous studies of ascalaphid immatures (Henry,
1976) indicate that important features of the mature larva that may
be absent or distorted in the first instar are expressed quite clearly
by the second stadium; for this reason, the following description can
be compared with existing descriptions of third instar larvae.

The head capsule is of the generalized ascalaphid type in shape,
more square than cordate, without extreme dorso-ventral flattening.
Ocular tubercles (Fig. 2-A) are cylindroid, and the ventral sternma
on each is but slightly reduced compared with the dorsal ones. The
antennal tubercles are poorly developed. Jaws are straight-shanked,
each tapering smoothly from proximal tooth to tip; the distal tooth
is markedly smaller than the others. Ventrally, sclerites of the
mouthpart bases and the pieces of the labium (Fig. 2-B) have a
generalized form and relationship to one another (Henry, 1976).
The manner of ventral articulation of the mandibles to the head
capsule is also relatively unspecialized: each condyle bears against a
median lobe at the end of the subgenal ridge rather than being
retained more positively by a U-shaped socket.

The body of the specimen also exhibits several apparently
primitive traits. Twelve pairs of long primary scoli fringe the body
from mesothorax to eighth abdominal segment (Fig. 1); in addition,
a pair of equally prominent ventro-lateral secondary scoli occurs on
abdominal segments I-VII (Fig. 3). All scoli are slightly flattened
dorso-ventrally and possess a border of specialized setae (see
below). The eight pairs of abdominal spiracles are situated laterally

1978]

Henry — Ascalaphid Larva

267

Figure 1 . Mkuze ascalaphid larva, probably second instar. ScMs = mesothoracic
scoli, ScMt = metathoracic scoli, ScAbdi = scoli of first abdominal segment, Dol =
dolichasterine seta.

268

Psyche

[June-September

in a linear arrangement between the primary and secondary series of
scoli; the eighth abdominal segment, lacking ventral scoli, bears its
spiracles beneath the primary scoli.

Nearly all setae are highly modified (Figs. 1-3). Those clothing
most of the dorsal and ventral surfaces of the head, body, and scoli
are very short and flattened into round discs or scales. Large, dorso-
ventrally flattened dolichasters project from the labral margin and
fringe the ocular tubercle and each primary and secondary scolus.
Spoon-shaped dolichasters occur dorsally in a double row down the
mid-line of the body and in a triple series on each side of the head
capsule. Long setae of conventional shape are present in small
numbers on the tips of the first and third scoli of the thorax and
singly on the tip of each scolus of the ventral abdominal series; the
last primary scolus also bears a few terminal setae of this type.

The pigmentation pattern of the specimen is due primarily to the
scale-like setae that cover nearly all parts of the body. In general the
insect has a mottled appearance, as though adapted for crypsis in a
relatively exposed or open microhabitat; however, its true colors are
of course unknown. The ocular tubercles are conspicuously darker
and the tips of the scoli lighter than other parts of the larva;
otherwise, mottling is quite uniform.

Discussion

The Ascalaphidae is one of six families in the superfamily
Myrmeleontoidea, a complex defined by a common larval type
exhibiting an array of cephalic traits that apparently evolved
together to provide structural support for the large muscles and
condyles of the jaw “trap” mechanism (MacLeod, 1964, 1970). In
contrast to the hemerobiiform larva of other Planipennia, that
of Myrmeleontoidea (=Infraorder Myrmeleontiformia) displays
1) maxillary blade lance-like, never as broad as mandible; 2) robust,
sickle-shaped mandibles; 3) ventral surface of large quadrate head
convex and heavily sclerotized, with sclerites of maxillae and labium
confined to medial anterior region; 4) pronounced anteriad migra-
tion and torsion of the tentorium so that it assumes a dorso-ventral
orientation connecting the anterior tentorial pits above to the
)OSterior ones directly below; 5) relatively short antenna with thick
.cape but filiform distal portion; and 6) two- to four-segmented
labial palps each arising from the tip of a large, palpimere-like
structure that actually represents half of the divided prelabium.

269

1978] Henry — Ascalaphid Larva

Figure 2. Mkuze larva, details of head capsule. A = anterior dorsal aspect, B =
anterior ventral aspect. Cd = maxillary cardo, Dol = dolichaster, ER = epistomal
ridge, Gu = gular area, Gul = gular line, LmM = labral margin, MdCV = ventral
mandibular condyle, OT = ocular tubercle, Plb = postlabium, Prlb = prelabial lobe,
Pip = labial palp, Scl = scale-like seta, SgR = subgenal ridge, St = maxillary stipes,
TAP = anterior tentorial pit, TPP = posterior tentorial pit.

270

Psyche

[June-September

Psychopsidae appears to be the most generalized of myrmeleon-
toid families with respect to these larval features (Fig. 4). Basic
specializations that have originated within the assemblage include
the development of mandibular teeth (all families except Psychopsi-
dae), elaboration of setigerous tubercles or scoli on the sides of the
body (all except Psychopsidae and Nemopteridae), increase in the
number of pairs of stemmata to seven (all except Psychopsidae and
most Nymphidae), appearance of distinct ocular tubercles (Asca-
laphidae, Stilbopterygidae, and Myrmeleontidae), fusion of tarsus
with tibia in the metathoracic leg (same three families), and great
enlargement of hind tarsal claws (Stilbopterygidae and Myrmeleon-
tidae). Available evidence suggests a sister-group relationship be-
tween Nymphidae and all other families except Psychopsidae
(MacLeod, 1970); Ascalaphidae in turn is probably the sister-group
to Myrmeleontidae (Riek, 1976), while the stilbopterygids — at
least, those from Australia — may prove to be nothing more than
specialized ant-lions (manuscript in preparation).

As discussed in an earlier paper (Henry, 1976), several larval
specializations appear within the Ascalaphidae (Fig. 4). For ex-
ample, New World ascalaphine (split-eyed) forms of the genus
Ululodes Currie and Colobopterus Rambur manifest extensively
modified cordate heads and a complex of mouthpart specializa-
tions, all relating to an extreme 270° “trap” position of the jaws;
these larvae also possess fewer and longer body scoli (ten pairs) than
other taxa, show no trace of a ventral scolus series, and bear al

« — 1 mi" i SpAbda

Figure 3. Mkuze larva, lateral view. SpAbds = spiracle of fifth abdominal
segment.

1978]

Henry — Ascalaphid Larva

271

■ MYRMELEONTIFORMI A -

no scoli
on abd.
seg.3Znr

uj w
< <
o a

o pi

2 O

■ASCALAPHIDAE

NEUROPTVNGINAE

.dorso-ventral flattening;
all scoli co-planar;
loss of dorsal abd.
scoli I+H;
abd. spiracles I+H
dorsally located

r

ASCALAPHINAEn

greatly
enlarged
hind claws

head specialization for
270° jaw trap position;
loss of all abd. scoli
of ventral series-,
loss of I pair of scoli each
on meso- and metathorax

^'reduction of ventral scolus series on abdomen;
littering of dorsum

^-fJcute occipital angles on head capsule

ocular tubercle; 7 stemmata, tibio- tarsal fusion in meta- leg
complete dorsal and ventral series of abdominal scoli
6 or 7 stemmata; mandibular teeth

Jr-*

distinct

\' o or r siemmaia; manaiDuiar reein

N head and jaw specializations; 5 stemmata /side-, dolichasterine setae

suctorial mouthparts

Figure 4. Cladogram of myrmeleontoid families of planipennian Neuroptera,
based upon larval features.

abdominal spiracles ventrally. In contrast, New World neuroptyn-
gine (entire-eyed) forms like Ascaloptnyx Banks and Byas Rambur
(Henry, 1978) show extreme dorso-ventral flattening and a much
larger number of scoli — twelve primary and six secondary (smaller)
pairs, the latter placed just anteriad of the former on abdominal
segments III-VIII. In addition, abdominal spiracles I and II are
dorsally located in these neuroptyngines, and specialized scale-like
dolichasters predominate on their body surfaces. Known Old World
ascalaphines, on the other hand, resemble Nymphidae in possessing
both a dorsal and ventral series of scoli on the abdomen and
laterally located abdominal spiracles; however, at least in Ascala-
phus Fabricius,1 the ventral series is largely vestigial and the first

'Tjeder (1972) points out that Ascalaphus as used here and as previously
understood should be replaced for nomenclatural reasons by Libelloides Schaffer;
Ascalaphus Fabricius then replaces Helicomitus McLachlan.

272

Psyche

[June-September

two pairs of abdominal spiracles show signs of dorsal migration
(Henry, 1976, Fig. 10). Finally, the Oligocene Neadelphus protae
MacLeod displays ventral spiracle location and is devoid of ventral
scoli, vestigial or otherwise, but those it possesses number twelve
rather than the ten of Ululodes and its relatives; it also shows no
setal modification (MacLeod, 1970; Henry, 1976, Fig. 9).

MacLeod (1970) interprets the secondary abdominal scoli of New
World neuroptyngines as having migrated from the ventral series of
a non-flattened nyinphid-like ascalaphid ancestor. My own view,
based on analysis of changes in spiracle location and comparison of
scolus shapes, is that the primary rather than secondary scoli of
these neuroptyngines have been derived from the ventral series, and
that the primary abdominal scoli of Neuroptynginae and Ascala-
phinae are therefore not homologous (Henry, 1976).

The Mkuze specimen described here shares its general head
capsule morphology and its twelve pairs of primary body scoli with
Neadelphus and known extant Neuroptynginae and Old World
Ascalaphinae; in both respects, it is a primitive or generalized
ascalaphid. Its most remarkable feature, however, is the double row
of scoli on each side of its abdomen with spiracles placed laterally
between the rows: although Ascalaphus shows traces of the ventral
series and New World neuroptyngines bear the latter in the same
plane as the dorsal series, no larva possesses such a fully developed
double series nor so closely approaches MacLeod’s hypothetical
nymphid-like ancestor as this specimen.

The setal patterns on the scoli of the specimen may help to
homologize these protuberances in this and other ascalaphid larvae.
For example, it is not known which (if either) of the two pairs of
scoli on the mesothorax or on the metathorax represent the dorsal
series, since spiracles are not present on either segment and all
known taxa showing the condition bear both pairs in the same
plane. The Mkuze larva possesses sharp-tipped setae on the tips of
the first and third pairs of thoracic scoli and on all ventral scoli,
suggesting a ventral origin for the more anteriorly placed pair of
scoli on each thoracic segment. Actually, the same relationship is
also present in Ascaloptynx: the first and third thoracic scoli
resemble in shape those deduced to belong to the ventral abdominal
series (Henry, 1976, Fig. 5). It then follows that, in neuroptyngine
larvae bearing all acoli in a common plane, the anterior scoli on
each thoracic segment did not originate from the same series as did

1978]

Henry — Ascalaphid Larva

273

the anterior ones on each abdominal segment. This conclusion
should be tested further.

Although the African specimen is primitive or generalized in most
respects, its scale-like setae and flattened dolichasters are a signifi-
cant specialization shared only with New World Neuroptynginae.
Based upon this single apomorphy, the larva could be classified with
the Neuroptynginae, for which no larvae are known from the Old
World; the dorsal location of abdominal spiracles I and II of New
World forms could then be interpreted as an additional specializa-
tion within a subgroup of the subfamily, associated with scolus
migration in response to dorso-ventral flattening and exposed living
habits.

Acknowledgements

This work was made possible by grants to the author from the
Research Foundation of The University of Connecticut (Storrs) and
from the National Science Foundation (DEB77-1 2443). I thank Dr.
James A. Slater (University of Connecticut) for loan of the Mkuze
specimen from his personal collection and for his constructive
comments on the manuscript. Ms. Marian Gergler kindly typed and
edited the manuscript, while Ms. Mary Jane Spring prepared the
cladogram reproduced in Fig. 4.

References

Henry, C. S.

1976. Some aspects of the external morphology of larval owlflies (Neuroptera:
Ascalaphidae), with particular reference to Ululodes and Ascaloptynx.
Psyche 83(1): 1-31.

1977. The behavior and life histories of two North American ascalaphids.
Ann. Entomol. Soc. Am. 70(2): 179-195.

1978. The egg, repagulum, and larva of Byas albistigma (Neuroptera: Asca-
laphidae): morphology, behaviour and phylogenetic significance. Syst.
Entomol. 3: 9-18.

MacLeod, E. G.

1964. A comparative morphological study of the head capsule and cervix of
larval Neuroptera (Insecta). Ph.D. thesis, Harvard University, Cam-
bridge, Mass.

1970. The Neuroptera of the Baltic Amber. I. Ascalaphidae, Nymphidae, and
Psychopsidae. Psyche 77: 147-180.

Riek, E. F.

1976. The family Stilbopterygidae (Neuroptera) in Australia. J. Aust. Entomol.
Soc. 15: 297-302.

274

Psyche

[June-September

Tjeder, B.

1972. Two necessary alterations in long-established genus nomenclature in
Ascalaphidae (Neuroptera). Entomol. Scand. 3: 153-155.

Weele, H. W. van der.

1908. Ascalaphiden: Monographisch Bearbeitet. Coll. Zool. Selys-Long-
champs fasc. 8: 1-326.

WlTHYCOMBE, C. L.

1925. Some aspects of the biology and morphology of the Neuroptera. With
special reference to the immature stages and their possible phylogenetic
significance. Trans. R. Entomol. Soc. London. 72: 303-411.

THE EVOLUTIONARY SIGNIFICANCE OF
REDUNDANCY AND VARIABILITY IN
PHENOTYPIC-INDUCTION MECHANISMS OF
PIERID BUTTERFLIES (LEPIDOPTERA)*

By Arthur M. Shapiro
Department of Zoology
University of California
Davis, California 95616, U.S.A.

One of the major discoveries in the study of seasonal polyphenism
in butterflies was the role of larval photoperiodic exposure (Dani-
levskii, 1948). Following this discovery, experiments on two Pierid
species — Colias eury theme Bdv. (Watt, 1969; Hoffman, 1974) and
Pieris protodice Bdv. & LeC. (Shapiro, 1968) — appeared to
establish that temperature played no role in the seasonal polyphen-
isms of that family. This was clearly not the case in at least one
Nymphalid, Araschnia levana (L.): Siiffert (1924) had shown a
temperature effect, and later work by Danilevskii (1948), Muller
(1955, 1956, 1960), Reinhardt (1969, 1971), and Muller and Rein-
hardt (1969) showed that photoperiod and temperature interact in a
characteristic way. Long-day larvae or young pupae, normally
destined to produce the summer form prorsa, if experimentally
cooled will produce either the winter form levana or an intermediate
form {porima). Short-day larvae give rise to diapause pupae which
always give levana, regardless of temperature. Thus photoperiod,
acting on 4th- and 5th-instar larvae, can irreversibly determine the
vernal phenotype, but not the estival one, which can be overridden
by temperature acting on the young pupa.

Shapiro (1977) established that pupal diapause and adult pheno-
type, normally tightly coupled in the Pieris napi (L.) group of
Pieridae, could be decoupled in P. n. venosa Scudder. In napi
generally, diapausing pupae give rise to vernal phenotypes and non-
diapausing ones to estival phenotypes. Populations consist of a
mixture of obligate diapausers, apparently determined genetically,
and facultative ones responsive to daylength. In some but not all
populations, inducing photoperiods can be overridden by high

* Manuscript received by the editor December 30, 1978.

275

276

Psyche

[June-September

developmental temperatures, but low temperatures cannot induce
diapause in long-day animals. (This system was first demonstrated
for P. rapae (L.) by Barker, Mayer, & Cohen (1963).) When non-
diapause, long-day pupae of P. n. venosa are chilled, they produce
vernal phenotypes; thus diapause is not necessary to produce them.
However, it is not known whether all diapause individuals are
irreversibly determined phenotypically, because they have a manda-
tory chilling requirement and break diapause in mid-winter, thus
assuring that every individual will receive some post-diapause
chilling. The entire system is summarized in Fig. 1.

Although the necessary experiments to clarify this point have not
been completed with P. n. venosa, they have been in the literature
for 50 years in a very important paper which has been universally
overlooked by English-speaking workers (Lorkovic, 1929). Lorkovic
worked with P. rapae and, to a lesser extent, Pontia daplidice (L.).
It is worth quoting at some length from a translation of the
summary (“homodynamic” pupae are non-diapausers; “heterody-
namic” are diapausers):

“Not only the homodynamic but also the heterodynamic
pupae of P. rapae are strongly influenceable in respect to
the butterfly’s markings. If the homodynamic pupae are
put in heat (25-30° C) during the sensitive period (which at
25° sets in about 12 hours after pupation), they produce
strongly black-spotted butterflies, while cold (5° C) brings
about a disappearance of these spots as well as a densely
dark powdering of the hindwing underside. Naturally
there are formed at corresponding temperatures also all
intermediates between the two extremes. The hetero-
dynamic pupae produce as a whole intermediate forms . . .
but, contrary to the results of Siiffert (on A. levana —
A.M.S.), the heat form can also be produced by a high
temperature of 32° C; but the black marking of the fore-
wing always approaches the heat form more than the hind-
wing underside, which can never attain the extreme grade
of the heat form. At lower temperatures the spotless form
is occasioned. The influencing of the heterodynamic pupae
by temperature is successful only during the last section of
development; pupae which for 3 months were kept at
lower temperatures (—5° to +10°) produced heat forms
after 6-8 days of exposure to higher temperature (32°).

, . diapause post-diapause chilling?

(genetic — ** ► vernal adult

pupa

diapauser

1978]

Shapiro — Pierid Butterflies

277

1. Schematic representation of developmental and phenotypic options available to Pieris napi. Not all options
in all populations, but all except those queried have been demonstrated in the laboratory.

278

Psyche

[June-September

(Lorkovic used pupal weight to estimate time of breaking
of diapause. — A.M.S.) It must therefore be taken that the
“heterodynamic sensitive period” sets in only after the
completed latency. (Compare Papilio zelicaon Lucas,
Shapiro, 1976. — A.M.S.) There are species in which the
duration of the pupal dormancy shows great lack of uni-
formity, varying from 1-8 years, without the difference
being reflected at all in the markings of the butterflies . . .”
Indications of the same phenomena are apparent in temperature-
manipulation experiments with the aforementioned P. protodice
and C. eurytheme, which had been thought to employ photoperi-
odic cues alone. In both species vernal phenotype is irreversibly
determined by short days (actually long nights), regardless of
temperature and with no linkage to diapause (there are no obligate
diapausers in P. protodice, and C. eurytheme has no diapause at
all). However, as in both A. levana and P. n. venosa, long-day
animals can be induced to develop the full vernal phenotype by
pupal chilling. Examples are shown in Fig. 2. That these responses
have been missed in the past is not surprising; they are difficult to
work with. In a given group of sibs a few individuals will respond
strongly to a given treatment, while others respond slightly or not at
all. It is difficult to keep track of precise pupation times for large
numbers of individuals and to obtain statistically meaningful num-
bers of even-aged pupae for treatment, and the precise characteriza-
tion of the responses will take several years, just as it did for the
Nymphalid Nymphalis urticae (L.) (history reviewed in Shapiro,
1976). However, it is already clear that there is considerable
intrapopulational variability in the timing of the “sensitive period,”
that it is quite short (there is no statistical difference in the
distribution of phenotypes in batches of P. protodice chilled at the
same age and held for 1, 2, 3, or 4 weeks), and that the mean
responses among geographic conspecific populations differ.

Precisely the same phenomena emerge in previous studies of
phenotypic plasticity which is not involved in regular seasonal
polyphenism: Aricia spp. (Hoegh-Guldberg, 1974a, b; Jarvis, 1974)
(Lycaenidae); various moths (Kettlewell, 1963); and especially the
“elymi” series of aberrations in Vanessa spp., and similar aberra-
tions in Nymphalis spp. (all Nymphalidae; see Shapiro, 1976). These
temperature-induced variations are “morphoses” (Schmalhausen,

1978]

Shapiro — Pierid Butterflies

279

Fig. 2. Examples of redundant mechanisms in phenotypic induction. Ventral
surfaces of males of Pieris protodice (left) and Colias eury theme (right). Top: estival
phenotypes, 24 L:0 D, 25° C, pupa unchilled. Center: vernal, same conditions, young
pupa chilled. Bottom: vernal, 10L: 14D, 25° C, pupa unchilled.

280

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[June-September

1949), nonadaptive developmental reactions. But the reactions of
Pierids are the same as those regularly produced by photoperiod,
and are presumably adaptive.

Probably most or all individuals of P. protodice and C. eury theme
can be switched from an estival to a vernal phenotype by some
degree of chilling at some time in the first 24 hours after pupation. If
this is true, then temperature can reinforce photoperiod in pro-
ducing seasonally appropriate adult phenotypes in multivoltine
species. For diapaused individuals of P. protodice and the closely
related P. occidentalis Reak., the irreversibly determined vernal
phenotype may be reinforced by post-diapause chilling, producing
the characteristically greater phenotypic expression than is seen in
lab-reared animals. The same is perhaps true for the (non-diapaus-
ing) late winter pupae of C. eurytheme. The period of post-diapause
or late-winter chilling is so prolonged in the wild that virtually all
overwintered animals will be affected. This process will tend to
smooth over, or conceal, not only the aforementioned genetic
variance in “sensitive period” but that which characteristically
occurs for chilling requirement (strength of diapause), as well. Yet
all of this variance is potentially adaptive in fluctuating environ-
ments, particularly when the suitability of spring weather for
butterflies is highly uncertain, and we should expect selection to
match norms of reaction to environmental uncertainty in popula-
tions with sufficient genetic integrity. Does it?

Hoffman (1978) reports that Rocky Mountain Colias philodice
eriphyle Edw. differs from both C. eurytheme and C. p. philodice
Godart in having adult phenotype decoupled from photoperiodic
control (though it may be temperature-sensitive). He attributes this
to high environmental uncertainty which renders photoperiod a
poor seasonal predictor. Similar predictions about developmental
phenology were made by Bradshaw (1973) and Istock (1978).
Similarly, Shapiro (1973) found phenotype less reliably cued by
photoperiod in the montane Pieris occidentalis than in P. protodice,
a lowland species. Crosses of C. p. philodice X C. p. eriphyle would
be informative as to how simple or complex genetic control of
photoperiodic coupling can be. Right now everything which is
known points to a definite evolutionary sequence in the history of
insect polyphenism: a pre-existing phenotypic response to tempera-
ture which happens to be adaptive in some environments but not
others is subject to selection for modifiers which affect threshold of

1978]

Shapiro — Pierid Butterflies

281

expression and ultimately couple it to a reliable seasonal predictor,
normally a pre-existing photoperiodic control of diapause.

This postulated evolutionary sequence can be further rationalized
by noting that the pattern of interaction among genetics, photo-
period, and temperature is completely in accord with Darwinian
predictions. For multivoltine species, which in Pieridae are weedy
colonizers, photoperiodic determination of diapause can be over-
ridden by high temperature in some individuals, allowing them to
gamble on an extra generation in an unusually warm autumn. But
photoperiodic determination of vernal phenotype is absolute; this
makes sense insofar as autumn is colder than summer and any
direct-developing butterflies will be aided in their feeding and repro-
ductive activities by the thermoregulatory properties of the vernal
phenotype. Estival phenotypes are not irreversibly determined by
summer photoperiods. Cold can act on the young pupa just a few
days before hatch to produce more or less of the vernal phenotype,
giving rise to darker animals on short notice in unseasonably cold
conditions. Hoffmann’s Rocky Mountain C. p. eriphyle and Sha-
piro’s coastal P. n. venosa are commonly exposed to unseasonable
cold and switch readily without regard to photoperiod. But lowland
C. eurytheme and P. protodice, which rarely face such conditions,
are arcane enough in their “sensitive period” characteristics that
they were missed altogether in laboratory experiments for years.

Acknowledgements

I thank Sydney Bowden for bringing Lorkovic’s paper to my
attention and for sending me a translation of the German summary,
and Prof. Dr. Z. Lorkovic for lending me an original copy of the
paper, for amplifying on its contents, and for, in his words, feeling
“content to live so long to see that I was discovered after 49 years,
indeed!”

Literature Cited

Barker, R. J., A. Mayer, and C. F. Cohen.

1963. Photoperiod effects in Pieris rapae. Ann. Entomol. Soc. Amer. 56:
292-294.

Bowden, S. R.

1978. Seasonal polyphenism in Artogeia napi (L.) (Lep.: Pieridae). Ent. Rec.
90: 176-180.

282

Psyche

[June-September

Bradshaw, W.

1973. Homeostasis and polymorphism in vernal development of Chaoborus
americanus, Ecology 54: 1247-1259.

Danilevskii, A. S.

1948. Photoperiodic reactions of insects in conditions of artificial illumina-
tion. Compt. Rend. Acad. Sci. U.R.S.S., N.S. 60 (3): 481.

Hoegh-Guldberg, (Z).

1974a. Natural pattern variation and the effect of cold treatment in the genus
Aricia. Proc. Trans. Brit. Ent. Soc. 7: 37-44.

1974b. Polymorphism in Ariciae (Lep., Rhopalocera) in the field and labora-
tory. Natura Jutlandica 17: 99-120.

Hoffmann, R. J.

1974. Environmental control of seasonal variation in the butterfly Colias
eurytheme: effects of photoperiod and temperature on pteridine pig-
mentation. J. Ins. Physiol. 20: 1913-1924.

1978. Environmental uncertainty and evolution of physiological adaptation in
Colias butterflies. Amer. Nat. 112: 999-1015.

Istock, C. A.

1978. Fitness variation in a natural population, in H. Dingle, ed., Evolution of
Insect Migration and Diapause. Springer-Verlag, New York. pp. 171 —
190.

Jarvis, F. V. L.

1974. The biological relationship between two subspecies of Aricia artaxerxes
(F.) and temperature experiments in an F2 generation and on A.
artaxerxes ssp. salmacis. Natura Jutlandica 17: 121-129.

Kettlewell, H. B. D.

1963. The genetical and environmental factors which affect colour and pattern
in Lepidoptera, with special reference to migratory species. Entomolo-
gist 96: 127-130.

Lorkovi<5, Z.

1929. Razlike izmedu homodinamskoga i heterodinamskoga razvitka insekata.
Godisnjak Sveucilista kraljevine jugoslavije u Zagrebu. Za skolske
godine 1924/25-1928/29. pp. 283-297.

Muller, H. J.

1955. Die Saisonformenbildung von Araschnia levana, ein photoperiodisch
gesteuter Diapause-Effekt. Naturwiss. 42: 134-135.

1956. Die Wirkung verscheidener diurnaler Licht-Dunkel-Relationen auf die
Saisonformenbildung von Araschnia levana. Naturwiss. 43: 503-504.

1960. Die Bedeutung der Photoperiode im Lebenslauf der Insekten. Z. angew.
Entomol. 47: 7-24.

Muller, H. J. and R. Reinhardt.

1969. Die Bedeutung von Temperatur und Tageslange fur die Entwicklung der
saisonformen von Araschnia levana L. Entomol. Berichte 1969: 93-100.

Reinhardt, R.

1969. Uber den Einfluss der Temperatur auf den Saisondimorphismus von
Araschnia levana L. (Lepidoptera: Nymphalidae) nach photoperiodi-
scher Diapause-Induktion. Zool. Jb. Physiol. 75: 41-75.

1971. Modifizierung der photoperiodisch bedingten Saisonformen von Arasch-
nia levana L. durch Temperaturveranderungen. Limnologica 8: 538.

1978]

Shapiro — Pierid Butterflies

283

SCHMALHAUSEN, I. I.

1949. Factors of Evolution. Blakiston, Philadelphia.

Shapiro, A. M.

1968. Photoperiodic induction of vernal phenotype in Pieris protodice Bois-
duval & LeConte. Wasmann J. Biol. 26: 137-149.

1973. Photoperiodic control of seasonal polyphenism in Pieris Occident alis
Reakirt. Wasmann J. Biol. 31: 291-299.

1976. Seasonal polyphenism. Evol. Biol. 9: 259-333.

1977. Phenotypic induction in Pieris napi L.: role of temperature and photo-
period in a coastal California population. Ecol. Ent. 2: 219-224.

SUFFERT, F.

1924. Bestimmungsfaktoren des Zeichnungsmuster beim Saisondimorphismus
von Araschnia levana/ prorsa. Biol. Zentralb. 44: 173-188.

Watt, W. B.

1969. Adaptive significance of pigment polymorphisms in Colias butterflies.
II. Thermoregulation and photoperiodically controlled melanin varia-
tion in Colias eurytheme. Proc. Nat. Acad. Sci. USA 63: 767-774.

CAMBRIDGE ENTOMOLOGICAL CLUB

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ISSN 0033-2615

PSYCHE

A JOURNAL OF ENTOMOLOGY

founded in 1874 by the Cambridge Entomological Club
Vol. 85 December, 1978 No. 4

CONTENTS

Tergal and Sternal Glands in Ants. Bert Holldobler and Hiltrud Engel . . 285

Life History of Dynastor darius (Lepidoptera: Nymphalidae: Brassolinae) in
Panama. Annette Aiello and Robert E. Silberglied 331

A Close Relationship Between Two Spiders (Arachnida, Araneidae): Curimagua
bavano Synecious on a Diplura Species. Fritz Vollrath 347

Systematics and Evolution of Forest Litter Adelopsis in the Southern Appa-
lachians (Coleoptera: Leiodidae: Catopinae). Stewart B. Peck 355

External Sex Brand Morphology of Three Sulphur Butterflies (Lepidoptera:
Pieridae). Richard S. Vetter and Ronald L. Rutowski 383

Intrasexual Aggression in the Stick Insects, Diapheromera veliei and D.
covilleae, and Sexual Dimorphism in the Phasmatodea. John Sivinski . . . 395

Sexual Calling Behavior in Highly Primitive Ants. Caryl P. Haskins 407

A Restudy of Two Ants from the Sicilian Amber. William L. Brown, Jr. and
Frank M. Carpenter 417

Index to Authors and Subjects 425

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1978-1979

President John A. Shetterly

Vice-President Barbara Thorne

Secretary Norman Woodley

Treasurer Lrank M. Carpenter

Executive Committee Jo B. Winter

Margaret Thayer

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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
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R. E. SlLBERGLIED, Assistant Professor of Biology, Harvard University
E. O. WILSON, Baird Professor of Science, Harvard University

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The Lexington Press, Inc., Lexington, Massachusetts

PSYCHE

Vol. 85 December, 1978

No. 4

TERGAL AND STERNAL GLANDS IN ANTS*

By Bert Holldobler and Hiltrud Engel

Department of Biology
MCZ Laboratories, Harvard University
Cambridge, Massachusetts

Introduction

Chemical signals, or pheromones as they are generally called, play
a central role in the complex communication system of ant societies.
During the last 20 years a number of exocrine glands have been
identified as the anatomical sources for a diversity of pheromones
which mediate sexual and social behavior in ants (for reviews see
Wilson 1971, Blum 1977, Holldobler 1978).

In recent years, however, several hitherto unknown exocrine
glandular structures have been discovered in ants and the behavioral
functions of some of them have already been determined. In this
paper we will review these findings and will report the new results of
our comparative morphological study of tergal and sternal glands in
ants.

Material and Methods

For histological investigations live specimens were fixed in alco-
holic Bouin (Dubosqu Brasil) or Carnoy (Romeis 1948), embedded
in methyl methacrylate, and sectioned 8 /i thick with a Jung
Tetrander I microtome (Rathmayer 1962). The staining was Azan
(Heidenhain). The SEM pictures were taken with an AMR 1000 A
Scanning Electron Microscope. For some of the species which could
only be identified to the generic level, the respective number is given
of the voucher specimens, which are deposited in the ant collection
of the MCZ (Harvard University).

* Manuscript received by the editor May 3, 1979.

285

286

Psyche

[December

Results

Tergal glands
a. Pygidial gland

In a detailed anatomical study of Myrmica rubra Janet (1898)
described a pair of clusters of a few glandular cells, located under
the third gastric tergum. Each cell sends a duct through the
intersegmental membrane between the third and fourth gastral
terga. We discovered a similar, but considerably larger paired
glandular complex at the same anatomical position in Novomessor
cockerelli and N. albisetosus (Holldobler et al 1976). Kugler (1978)
recently investigated a number of myrmicine ants and in many of
them he found the gland, which had “distinct reservoirs, produced
by invagination of the intersegmental membrane between abdomi-
nal tergum 6 and tergum 7 (pygidium)”. Kugler suggested that these
glandular organs be called pygidial glands. We accept this terminol-
ogy, because it describes the anatomical designation of the organ
more precisely than the term “dorsal gland” or “tergal gland”,
originally suggested (Holldobler et al 1976, Holldobler and Haskins
1977). However, it has to be pointed out that the pygidium of the
ants (the last exposed tergum) is the 7th abdominal tergum and is not
homologous to the pygidium of the Coleoptera (8th abdominal
tergum). Hence, the pygidial glands of ants are not homologous to
the pygidial glands of Coleoptera.

In a previous study (Holldobler and Haskins 1977) we found
pygidial glands with large reservoirs in several ponerine and myr-
meciine ants ( Amblyopone , Paraponera, Ectatomma, Odontoma-
chus, Pachycondyla, Platythvrea, Rhytidoponera, Myrmecia ) and
we demonstrated that the virgin females of Rhytidoponera metallica
attract males by the release of a pheromone from these glands. In his
anatomical studies of Rhytidoponera metallica and R. convexa,
Whelden (1957, 1960) described a pair of cell clusters each com-
prising 8-15 glandular cells. Each cell sends a duct through the
membrane connecting the 6th and 7th abdominal segments. We are
now certain that Whelden already had discovered the pygidial gland
in Rhytidoponera; his histological methods, however, may not have
enabled him to detect the large reservoirs associated with the
glandular cell clusters. Similar paired glandular structures were
found by Whelden (1957) in the ponerine species Stigmatomma
(= Amblyopone) pallipes.

1978]

Holldobler & Engel — Glands in Ants

287

Finally, independently of our investigations, Maschwitz (pers.
communication) found a pygidial gland in Leptogenys chinensis,
which he called the “dorsal gland” (Maschwitz and Schonegge,
1977).

The new anatomical investigations presented in this paper reveal
that the pygidial glands are much more common in ants than
previously assumed. Usually the organ consists of a pair of lateral
clusters of glandular cells, each cell sending a duct through the
intersegmental membrane between the 6th and 7th abdominal terga.
Depending on the species, the intersegmental membrane can be
invaginated to different degrees, so that it can form a more or less
voluminous reservoir (Fig. 1, 2, 3, 4). If no reservoir is present, the
glandular structures can easily be missed during the dissection and
histological sectionings are therefore required to determine whether
or not the pygidial gland is present. As we have already indicated for
Novomessor and as confirmed by Kugler (1978) for several other
myrmicine species, the pygidial gland can be associated with a
special cuticular structure on the pygidium (7th tergum), (Fig. 5, 6,
7). Our histological studies demonstrated, however, that the absence
of such structures does not necessarily indicate the absence of

Figure 1. Schematic illustration of glandular cells that send ducts through the
intersegmental membrane. When the membrane is increasingly more invaginated, it
forms an increasingly larger reservoir (a to c).

288

Psyche

[December

pygidial glands. Thus, in several myrmicine species, in which we
(Holldobler et al 1976) and Kugler (1978) previously assumed the
pygidial gland to be absent, we find that we now detect this organ by
histological methods. Tables la and lb list the species of the major
subfamilies that we investigated histologically and indicate the type
of tergal glands found.

b. Postpygidial gland

Dorsal glandular structures which open posteriorly to the py-
gidial gland, between the 7th and 8th abdominal terga (spiracular
plate), we call postpygidial glands (Fig. 8). Whelden (1957, 1960)
described glands in the 5th gastral segment of Rhvtidoponera
convexa and R. me tallica. For R. convexa he writes: “Even the
largest of these is less than half the size of the fourth-segment glands
.... In the extreme case, there may be but a single gland cell on each
side. It is often difficult to distinguish such a unicellular gland from
an oenocyte, despite the usually distinct difference in size of the two
cell types. Only the identification of a duct certainly distinguishes
such a gland from the ductless oenocyte. In many individuals this
second pair of glands could not be found”. Whelden (1960) makes
similar statements for R. metaUica. Our results are somewhat
different. We found well developed postpygidial glands in the 4
species of Rhvtidoponera investigated (Table la). In all specimens
we found a pair of clusters of glandular cells. Each cluster contains
about 15-20 cells and each cell sends a duct through the interseg-
mental membrane close to the spiracular plate (Fig. 9, 10). In some
ant species the postpygidial gland consists of only a few glandular
cells, in others the postpygidial gland is associated with a well
developed reservoir (Table la, lb).

Sterna! glands

In several species we discovered intersegmental sternal glands
(Table 2). They can consist of a few glandular cells that send their
channels through the intersegmental membrane, or of large clusters
of glandular cells associated with voluminous reservoirs. These
reservoirs are formed by invaginations of the intersegmental mem-
branes (Fig. 8).

In several Leptogenys species (Fig. 12) we found two large sternal
glands with reservoirs between the 7th and 6th, and 6th and 5th sterna.
The latter glandular organ is usually associated with a special
cuticular structure on the 6th sternum (Fig. 12). In Paltothyreus

1978]

Holldobler & Engel — Glands in Ants

289

tarsatus we found well developed sternal glands between the 7th and
6th, 6th and 5th, and 5th and 4th sterna, but no reservoirs. Instead, the
duct openings are associated with filament-like protrusions of the
intersegmental membrane (Fig. 13, 14).

Other abdominal glands

The glandular venom apparatus of ants is composed of the
Dufour’s gland (alkine or accessory gland) and the poison gland.
Although the venom apparatus of ants is very well studied (see
reviews by Maschwitz and Kloft 1971, Blum and Hermann 1978 a,
b), other glands, such as Koschevnikov’s gland (sting gland),
Bordas’s gland and sting sheath glands, known from other Hymen-
optera, have not been firmly established in ants.

Koschevnikov (1899) found in honeybees and Vespa paired
clusters of glandular cells located laterally near the intersegmental
membrane between the quadrate plate and the spiracular plate.
Each individual cell sends channels into gathering ducts, which
connect with the intersegmental membrane. Altenkirch (1962) found
similar glands in most Apidae that she had studied.

There are indications that this gland might also be present in
some species of the primitive ant subfamilies Myrmeciinae and
Ponerinae. Whelden (1957) described a pair of clusters of gland
cells, located slightly dorsally on each side of the sting of Stigma-
tomma ( =Amblyopone ) pallipes. Each cell sends a “rather tortuous
duct . . . down and inward, to open through a membrane which is
above the sting”. Robertson (1968) found “sting glands” in Rhy-
tidoponera “toward the region of the triangular plate, where they
are attached to the intersegmental membrane”. She described
similar glands in Bothroponera sp. ( —Pachycondyla ), Leptogenys
sjostedji and Myrmecia gulosa. In the latter species the glands are
described as “two well formed masses of gland cells, each cell
attached to the intersegmental membrane in the region of the
triangular plate by a long, simple, cuticular duct”.

Table 3 (A) lists the species in which we found paired glandular
structures, closely resembling the “sting glands” described by
Robertson. In all cases the glandular cells are located near the
triangular plate, and from each cell a rather long duct leads
downwards and opens through a membrane near the base of the
sting (Fig. 15). Although we could not precisely locate the openings
of the ducts, we assume they opened in the sting chamber.

290

Psyche

[December

Altenkirch (1962) and Maschwitz (1964) discovered a so-called
sting sheath gland in several bee species. It consists of a palisade
epithelium located in the sheath valves. In some ant species we
found a distinct palisade epithelium in the sheath valves and/or
single glandular cells with long individual ducts (Table 3 (B), Fig.
16, 17). Janet ( 1 898) describes similar single gland cells, located near
the sheath valves, in Myrmica rubra.

Bordas’s glands, as they were described by Bordas (1895) in
Terebrantia and reexamined by Rathmayer (1962) in several sphecid
wasps, could not be identified in ants, although some of clusters of
the single gland cells which send their ducts through the membrane
of the sheath valves could be related to the Bordas’s glands. It is
obvious to us that the glandular structures, associated with the sting
apparatus of ants, need to be investigated in greater detail in future
studies.

In several ant species (Table 3 (C)) we found a highly developed
palisade epithelium in the 7th sternum (Fig. 8). It is especially
conspicuous in several Leptogenys species and in the army ants
Eciton and Neivamyrmex, but it is not strongly developed in
Dorylus. In the dolichoderine species and in Aneuretus this epithe-
lium seems to be closely associated with the sternal gland (Pavan’s
gland).

In the African weaver ant ( Oecophylla longinoda ) we discovered
a sternal gland under the 7th sternum, which is quite different from
the glandular epithelium described above (Holldobler and Wilson
1976, 1978). This structure consists of an array of single glandular
cells that send short channels into cuticular cups on the outer
surface of the sternite. In none of the other formicine species
investigated, listed in table 1 b, did we find this type of sternal gland.
But in Camponotus sericeus we detected different clusters of
glandular cells in the last sternum. Each cell sends a long channel
through the intersegmental membrane near the vagina into the
ventral part of the “sting chamber”. We discovered similar paired
glandular cell clusters in most myrmicine species we investigated.
The gland is especially distinct in Novomessor and Veromessor,
where the glandular cell channels penetrate the membrane near the
vagina (Fig. 18).

The function of the intersegmental glands:

The functions of most of the glandular structures described in this
paper are not yet known, but in a few species the function of the

1978]

Holldobler & Engel — Glands in Ants

291

pygidial gland has already been identified. In Novomessor cockerelli
and N. albisetosus the strongly smelling secretion of the pygidial
glands releases a “panic alarm” response in workers, apparently
specifically designed against army ant predation (Holldobler in
prep.). Kugler (in press) demonstrated that in Pheidole biconstricta
the pygidial glands produce an alarm-defense secretion. A quite
different function has been discovered in Rhytidoponera metallica.
Here the wingless virgin females attract males by the release of a
pheromone from the pygidial gland (Holldobler and Haskins 1977).
Since Rhytidoponera workers also have a well-developed pygidial
gland and are attracted to its secretions, we believe we have not yet
discovered the whole functional spectrum of this organ. In Lepto-
genys chinensis, Maschwitz and Schonegge (1977) demonstrated
that the pygidial gland secretions serve together with poison gland
substances as a recruitment trail pheromone. We obtained similar
results when we recently reexamined the anatomical source of the
trail pheromone of Paehycondvla ( =Termitopone ) laevigata. This
ant species conducts well organized predatory raids on termites.
During raiding the workers move in a single file, one closely behind
another, along a powerful trail pheromone laid down by leading
scout ants. Blum (1966) has identified the hindgut as the source of
this recruitment trail pheromone. We cannot confirm his findings.

In our experiments with artificial trails laid with extracts from
several abdominal glands, only the pygidial gland secretions re-
leased massive trail-following behavior in P. laevigata (Holldobler
and Traniello in prep.). A careful observational study of the trail-
laying behavior of P. laevigata workers revealed that not the anus
but rather the pygidial gland is dragged over the ground. Although
the pygidial gland of P. laevigata has no definite reservoir, it is very
well developed and is associated with an elaborate cuticular struc-
ture on the 7th tergum (Fig. 19, 20). The glandular secretion is
apparently stored in the many cavities of this structure. When
trailing, the ants rub this structure with its special applicator surface
over the ground and deposit thereby the trail pheromone. Traniello
(pers. communication) observed species of Odontomachus during
nest emigrations performing the same trail laying behavior. We
suspect that in Odontomachus also the pygidial gland secrets a trail
pheromone (Fig. 21).

In Bothroponera ( —Pachycondyla ) tesserinoda we previously
analyzed the signals involved in the tandem running recruitment
technique (Holldobler et al 1973, Maschwitz et al 1974). We

292

Psyche

[December

discovered that the cues responsible for “binding” the follower
behind the leader ant include both a surface pheromone and
mechanical stimuli. Although we could extract this surface phero-
mone, we were not able to identify its anatomical source; all
experiments with secretions from the known exocrine glands had
negative results. After the recent discovery of the pygidial gland in
Pachycondyla we have begun to conduct tandem running experi-
ments with Pachycondyla crassa* and P. harpax*, using dummies
contaminated with pygidial gland secretions. Our preliminary re-
sults strongly indicate that pygidial gland substance might be the
source of the tandem running pheromone in these species.

In the doryline army ants raiding and emigrations are conducted
along chemical trails deposited by workers. For Neivamyrmex,
Watkins (1964) and Watkins et al (1967), and for Eciton hamatum,
Blum and Portocarrero (1964), identified the hindgut as the source
of the trail pheromone. In addition, Chadab and Rettenmeyer
(1975) and Topoff and Mirenda (1975) demonstrated that besides
the relatively long-lasting hindgut trail-substance, other signals
(possibly more volatile secretions) are involved in the organization
of “mass recruitment” in Eciton and Neivamyrmex.

We believe that our morphological investigations provide new
possibilities for the analysis of chemical communication in army
ants. Both Neivamyrmex and Eciton have large pygidial glands with
distinct reservoirs (Fig. 22, 23). The postpygidial gland is smaller,
but still considerably larger than in most of the other investigated
speciesf . In both army ant species the 7th tergum is relatively small.
Therefore, the reservoirs of the pygidial gland and postpygidial
gland open directly above the anus at the abdominal tip (Fig. 23). In
workers (all castes) of Eciton the dorsal membrane near the exits of
the reservoir of the pygidial glands is conspicuously modified to a
brush-like structure (Fig. 24). These morphological features strongly

* P. crassa was observed tandem running by W. L. Brown, Jr. (pers. communica-
tion) at the western base of Ubombo Mts., Zululand, and by B. Holldobler in
Shimba Hills Reserve (Kenya). P. harpax was observed tandem running by S.
Levings (pers. communication) on Barro Colorado Island, Panama.

t Whelden (1963) described two glands at the extreme posterior end of the gaster of
Eciton burche/li workers. Although the description is not very accurate, from his
drawings we can conclude that he found the pygidial gland and postpygidial
glands.

1978]

Holldobler & Engel — Glands in Ants

293

suggest that the tergal glands might be involved in the chemical trail
communication of army ants. We have begun to test this hypothesis
with Eciton hamatum. The pygidial gland secretion of E. hamatum
has a strong, characteristic smell. The secretion is probably skatole
(Traniello pers. communication), the substance that gives army ants
their typical “fecal odor”. Recently Brown et al (1978) demonstrated
that skatole is an effective growth inhibitor for bacteria and fungi
and repels insectivorous snakes (Watkins et al 1969). Our first,
preliminary tests demonstrated that Eciton workers follow artificial
trails drawn with crushed pygidial glands. When we simultaneously
offered trails drawn with hindgut contents and pygidial gland
secretions, the latter were significantly preferred during the first
minute. When we used trails drawn with secretions of the poison
gland or Dufour’s gland as controls, the ants always followed the
pygidial gland trails. We have to stress, however, that these
experiments must be considered pilot tests. The preliminary results,
however, are striking enough to warrant a more detailed investiga-
tion in the future. It is interesting to note that the anatomy of the
pygidial gland in the African army ant, Dorylus molesta, is quite
different from that of Eciton and Neivamyrmex (Fig. 25). In this
species the 7th tergum is considerably larger than in species of the
latter genera, and the reservoirs of the pygidial glands do not open
at the abdominal tip. In Dorylus, however, we found single
glandular cells with channels opening directly at the anus, a feature
we have not detected in other ant species (Fig. 25, 26).

Finally, our morphological study of the pygidial gland of Ve-
romessor pergandei has led to results that are suggestive of the
function of this organ. In this species the 7th tergum is relatively
small, and as a result the large reservoirs of the pygidial glands open
at the tip of the gaster (Fig. 27). Veromessor forages in well-
organized columns (Went et al 1972; Wheeler and Rissing 1975;
Bernstein 1975). Several observations suggest that these foraging
columns are organized by a trail pheromone, though no trail
pheromone gland has yet been identified. Clearly, the large pygidial
gland has to be considered as a possible source for the trail
pheromone.

The function of most of the newly discovered sternal glands is
unknown. Only in Paltothyrcus tarsatus could we demonstrate
experimentally that foragers lay a recruitment trail with sternal
gland secretions (Holldobler in prep.).

294

Psyche

[December

Conclusions

Since we first found the pygidial gland widespread in the sub-
families Myrmeciinae and Ponerinae, we speculated that this gland
might be a primitive monophylogenetic trait in ants generally
(Holldobler and Haskins 1977). The results reported in the present
paper fully confirm this assumption. A well-developed pygidial
gland was found in the most primitive ant, Nothomyrmecia ma-
crops, and in representatives of all major subfamilies except in the
Formicinae (Table lb). We agree with Kugler (1978) that the “anal
glands” of the Dolichoderinae and Aneuretinae are homologous to
the pygidial glands of other ant subfamilies. Considering the
variation in the morphology of the pygidial glands, even within a
single subfamily, we think that the morphological variation of the
“anal glands” of dolichoderine and aneuretine species does not
warrant a separate terminology. In fact the term “anal gland” is
misleading, because the glands do not exit from the anal opening of
the gaster, as is sometimes inferred, but between the 6th and 7th
abdominal terga (Fig. 28). This was clearly demonstrated by Pavan
and Ronchetti (1955). It is our view and also Kugler’s (pers.
communication) that the “anal glands” should be called pygidial
glands.

Kugler (1978) concluded from his comparative studies of myrmi-
cine species that usually those species that have reduced or modified
stings also have well-developed pygidial glands. He assumes that the
pygidial gland replaces the sting apparatus as a chemical defense
device. Our finding that well-developed pygidial glands occur in
Pogonomyrmex, a genus with a very effective sting apparatus, and
in many stinging ponerine species, does not support Kugler’s
conclusions.

Acknowledgements

This paper would not have been possible without the help of
many people. We would like to thank all the collectors mentioned in
Table 1, including Donald W. Windsor, who helped finding the
acacias in the Canal Zone. Special thanks to Robert W. Taylor, who
sent us the precious Nothomyrmecia. Barry Bolton, William L.
Brown, Jr., William H. Gotwald, Jr. and Roy Snelling identified
many species for us. We are grateful to Ed Seling for his superb
assistance during the SEM work. Frank M. Carpenter’s many
suggestions improved the manuscript greatly. This work was sup-
ported by NSF grant BNS 77-03884.

1978] Holldobler & Engel — Glands in Ants 295

Table la

List of species of the poneroid complex (Taylor 1978) that were investigated histo-
logically, and the types of tergal glands found. When the histological series was
incomplete and we could not make a definite statement, the column is marked with
When the cuticular structure on the pygidium was only slightly sculptured, we
marked the column with

JC

3

O

JS

* E

W 3

Subfamily/ Species

Collector

and

% .3

■§ i

JS %

DO <D

I i

£ 8
DO <D

j l

-§ s £

^'5

c

-2

•2 f

.*2 j;

"c 1-

iS 'o
eo >

Lh

2 -
-o K

Locality

13 <u

11
CLh T3

13 <u

11
Cl, "O

.2 s. g

: 2 " 3

bo 3

>, .-3 V-

CX £ ts

>■, <D
CL

to -3
O .-3

Oh £

5b

>> 3

D, o

a? -3
O .13
CL £

Myrmeciinae

Myrmecia

R. J. Bartell

+

+

pilosula

R. W. Taylor
Brindabella
Ranges,
Australia

PONERINAE

Amblyopone

C. P. Haskins

+

australis

Manjimup,
W. Australia

Amblyopone

J. Traniello

+

-

-

-

pallipes

Carlisle, Mass.

Platythyrea

K. Horton

+

-

+

cribinoda

R. Silberglied
Shimba Hills,
Kenya

Rhytidoponera

C. P. Haskins

+

-

+

metallica

Blackwell

Range,

Queensland,

Australia

Rhytidoponera

C. P. Haskins

+

-

+

perthensis

Boddington,
W. Australia

Rhytidoponera

C. P. Haskins

+

-

+

purpurea

Black Mountain,

Kuranja,

Queensland

296

Psyche

[December

Collector

Subfamily/ Species and

Locality

3

O

-C

£ .!=

H I

iS %

DO U

■ - c

DO ^
u

Cl tj

-o

c

00

.2

■5

'do

>>

CL

E

3

"O

'do

o.

c

o

<L>

3

o

3

C/2

■a

c

3

00 !_

•— <D
DO c/2
>,
n.

o •-

CL £

Rhytidoponera

violacea

Paltothyreus

tarsatus

Pachycondyla

crassa

Pachycondyla

laevigata

( = Termitopone )

C. P. Haskins +

Kings Park,

W. Australia

B. Holldobler
Shimba Hills,

Kenya

B. Holldobler +

Shimba Hills,

Kenya

N. Franks
J. Traniello
BCI, Panama

+ +

+

+

+

? ?

Plectroctena

strigosa

Leptogenys

neutralis

B. Holldobler +

Shimba Hills,

Kenya

C. P. Haskins +

Manjimup,

W. Australia

+ +

+ +

Leptogenys

pavesii

B. Holldobler
Shimba Hills,
Kenya

+

Leptogenys

B. Holldobler

+

nitidia

Shimba Hills,
Kenya

Leptogenys

B. Holldobler

+

regis

Shimba Hills,
Kenya

Odontomachus

haematoda

C. P. Haskins
BCI, Panama

+

+ +

+ +

+ +

+ +

Table la Continued

Postpygidial gland
without reservoir

1978]

Holldobler & Engel — Glands in Ants

297

% --

'% •-

‘ob

Collector

■§ 1
ja «

1 l

a u

? 3 £

a — c

Subfamily/ Species

and

oo u

— c n

00 <L>

»•§ z

Locality

cd <u

|1
CL tj

75 <u

-s

00

^ <D

Cl

.2 CL 3

.-3 2 o

3

>% .- t-
CL £ tS

Dorylinae

Neivamyrmex

nigrescens

B. Holldobler
Arizona

+

Eciton

hamatum

A. Aiello
R. Silberglied
J. Traniello
BC1, Panama

+

Dorylus

molesta

B. Holldobler
Shimba Hills,
Kenya

+

+

PSEUDOMYRMECINAE

Pseudomyrmex

ferruginea

A. Aiello
R. Silberglied
Canal Zone,
Panama

+

Pseudomyrmex

pallidus

Tetraponera
spec. (78/26)

M. Moglich
Florida

B. Holldobler
Shimba Hills,
Kenya

+

+

Myrmecinae

Myrmica

americana

J. Traniello
Carlisle, Mass.

+

Tetramorium

caespitum

J. Traniello
Carlisle. Mass.

+

—

Pogonomyrmex

desertorum

B. Holldobler
Arizona

+

-(+)

Pogonomyrmex

californicus

B. Holldobler
Arizona

+

"(+)

-o

c

oo u.

• — <u

00 c/3

>' p

C_ *“

o

CL £

+

+

+

+

?

Table la Continued

Postpygidial gland
without reservoir

298 Psyche [December

sz

£ .S:

3

O

SZ

£ .i=

u E

CT3 3

3 i

■ ^ DO

73

C

aj

Collector

n !

JS 2o

1 i

■§ 3 &

cd "ti ^

"ob u.
15 ©

Subfamily/ Species

and

00 <D

— c/l

DO <D

?i °

• — >
.12 fc

Locality

15 4)

j§ 'E

CL TD

15 4)

ii 'E

00 IZ

£ -a

2 a 3

.■2 “ i

00 ~ 3

>>.-!_
Cl ^ to

8

a.

to

O •-
CL £

Pogonomyrmex

badius

B. Holldobler
Florida

+

-

-

Veromessor

pergandei

G. Alpert
Mexico

+

-

—

Novomessor

cockerelli

B. Holldobler
Arizona

+

+

-

Novomessor

albisetosus

B. Holldobler
Arizona

+

+

—

Aphaenogaster

rudis

J. Traniello
Concord, Mass.

+

-

-

Aphaenogaster

huachucana

B. Holldobler
Arizona

+

-

-

Pheidole militicida
(worker & soldier)

B. Holldobler
Arizona

+

-(+)

—

Pheidole

desertorum

B. Holldobler
Arizona

+

+

—

Atta sex dens
(several castes)

N. Weber
Timehri, Guyana

-

-

-

-

Table la Concluded

Postpygidial gland
without reservoir

1978] Holldobler & Engel — Glands in Ants 299

Table lb

List of species of the formicoid complex (Taylor 1978) that we investigated histo-
logically, and the types of tergal glands found.

G

o

^ £

'% .1=

-C

a p
3 TJ
3 'So

G

jd

Collector

"O >

-a >

TD g £

Subfamily/ Species

and

c C
£ %

1 §

G r-

cd o

.2 1

Locality

60 <u

GO

GO -r? °

- £ a

-a >-

.— (U

bO c«

73 <u

2 CU P

>.

3 • s
60 <-

>. <3

3 '£

GO

.3 “ o

60 S =S
»-

Cl, ^

to

O . -

CU -o

cu -o

CU £ to

CU £

Nothomyrmeciinae

Nothomyrmecia

R. W. Taylor

+

macrops

Eyre Peninsula
S. Australia

Aneuretinae

Aneuretus

E. O. Wilson

+

simoni

Ratnapura,
Sri Lanka

Dolichoderinae

Liometopum

B. Holldobler

+

_

apiculatum

Arizona

Conomyrma

B. Holldobler

+

-

-

bicolor

Arizona

Iridomyrmex

B. Holldobler

+

-

-

pruinosus

Arizona

Formicinae

Oecopylla

B. Holldobler

longinoda

Shimba Hills,
Kenya

Pachycondyla spec.

R. Silberglied
Nairobi
Arboretum,
Kenya

Myrmecocystus

B. Holldobler

-

-

-

-

mexicanus

Arizona

Myrmecocystus

"

-

-

-

-

mimicus

Myrmecocystus

mendax

—

—

—

Formica perpilosa

"

-

~

-

-

Camponotus

M. Moglich

-

-

-

sericeus

Sri Lanka

Postpygidial gland
without reservoir

300

Psyche

[December

Table 2

List of species of the poneroid complex in which we found intersegmental sternal
glands.

Species Location of glands between abdominal segments

7 and 6

6 and 5

5 and 4

reservoir

reservoir

reservoir

yes

no

yes no

yes n

Myrmecia pilosula

+

Rhvtidoponera purpurea

+

Paltothvreus tarsatus

+

+

Pachyeondyla crassa

+

+

Leptogenys neutralis

+

+

Leptogenys pavesii

+

+

Leptogenys nitida

+

+

Leptogenys regis

+

+

Tetraponera sp. (78/26)

+

+

sternum ( C) were found .

1978]

Holldobler & Engel — Glands in Ants

301

iS iS
3 3 1

T3 "O

e c

3

|S|

f * -a

> (U C

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0) 4-1

BO

£ ?>
.£ 4>

'« *5

’ex 3
<U O
t_. o
3 <L)

3 £

£ c
bo hj

g u

■Sf

03 £

~ O

ffl «

3" o

T3 c

e 3
3) ^

c §
■- 5b

c« O

3

< ts

+ .£
i £
'* 1

?!

03

C Oh

G &n

U T3
cfl u

■5 3

03 d

J* i

ca 3

bo e

3 T3
U3 <U
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3 3
'£. £

U <D
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3 =2

■o o
3 3
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3) _e

T3 u
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3)
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3 ^

<D -3
U< ^

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<u 5

^ c/3

Oh <l>

£ *o

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<D o

£ 3

-G 5

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60
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U </3

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(U c

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<X X x
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^ 3 Hj ^

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c a o o

£ ^ l>5 >>

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o. o Xi <X

£ x 2 £

o c o o

3 3 3 'C

+ + +

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^ 5 ■?

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3 o £

^ ^ ^

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52 O O O X

-2 Cx Q, £. S

K 2 O O o C

-o -c>

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q c:

•2 o

O sT' s £ -£ -2 3 <x

« ^ 2
^ § -2- 2
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£C a, <X

£. S3 C C

Q o o o
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'Oh G
oj T3

302

Psyche

[December

i

Figure 2. Above: Sagital section through pygidial glands of Amblyopone
australis. The gland has no distinct reservoir, but 10-15 glandular cells (GC) with
channels in each cluster. Below: Pygidial gland of Amblyopone pallipes with distinct
reservoir (R) and 30-40 glandular cells in each cluster.

1978]

Holldobler & Engel — Glands in Ants

303

Figure 3. Above: Sagital section through pygidial glands of Nothomyrmecia
macrops: Very large reservoir (R); each cluster consists of 50-80 cells; GC =
glandular cells; CH = channels of glandular cells. Below: Pygidial gland of Myrmica
americana: No distinct reservoir, 10-15 cells in each cluster.

304

Psyche

[December

Figure 4. Above: Sagital section through pygidial glands of Pogonomyrmex
calif ornieus: No definite reservoir, but large cell clusters, each containing 30-35
glandular cells (GC). Below: Pygidial gland of Pseudomyrmex ferruginea: No distinct
reservoir; intersegmental membrane forms a chitinous duct (D) into which the
individual glandular channels open.

1978]

Holldobler & Engel — Glands in Ants

305

Figure 5. Above: Sagital section through the pygidial gland of Leptogenys
pavesii (GC = glandular cells; CH = channels of GC; CS = cuticular structure on the
7th tergum). Below: Close-up view of sagital section through cuticular structure of 7th
tergum of Leptogenys neutralis.

306

Psyche

[December

Figure 6. Above: Sagital section through pygidial gland of Leptogenys regis.
Below: SEM photograph of cuticular structure of 7th tergum of L. regis. The tergum
is cut so that the well-structured cavities under the surface can be seen. (Lettering as
in Fig. 5).

1978]

Holldobler & Engel — Glands in Ants

307

Figure 7. SEM photograph of cuticular structure of 7th tergum of Plectroctena
strigosa. Above: View of the anterior part of the tergum. The cuticular grooves are
clearly visible. Below: Close-up view of cuticular structure; the tergum is cut, so that
the cavities beneath the grooves are visible.

308

Psyche

[December

Figure 8. Schematic illustration of tergal and sternal glands in Leptogenys.
(Lettering as in Fig. 5; sg = sternal gland; ge = glandular epithelium; ppg = post-
pygidial gland; pg = pygidial gland).

1978]

Holldobler & Engel — Glands in Ants

309

Figure 9. Above: Sagital section through pygidial gland and postpygidial gland,
with large reservoirs of Rhytidoponera perthensis. Below: Postpygidial gland only;
note the glandular cells with channels. (Lettering as in Fig. 8; R = reservoir).

310

Psyche

[December

Figure 10. Above: Sagital section through pygidial gland and postpygidial
gland of Plectroctena strigosa. Below: Pygidial gland. (Lettering as in Fig. 5; PPG =
postpygidial gland; PG = pygidial gland; R = reservoir).

1978]

Holldobler & Engel — Glands in Ants

311

50ju GC

Figure 11. Above: Sagital section through the pygidial gland of Platythyrea
cribinoda. The large reservoir is folded in this section. Each cell cluster contains
50-70 cells. Below: Section through postpygidial gland of P. cribinoda; each cluster
contains 15-20 cells. (Lettering as in Fig. 3).

312

Psyche

[December

Figure 12. Above: Sagital section through sternal gland between 5th and 6th
sterna of Leptogenys neutralis. Usually the 6th sternum has a distinct cuticular
structure (CS), very similar to that on the 7th tergum. Below: Section through both
sternal glands (Between 5th and 6th, and 6th and 7th sterna) of Leptogenys nitida.
(Lettering as in Fig. 5; R = reservoir).

1978]

Holldobler & Engel — Glands in Ants

313

Figure 13. SEM photograph of the sternal gland area between the 6th and 7th
sterna of Paltothyreus tarsatus. The picture below is an enlargement of the lighter
rectangle in the picture above. The openings of the glandular cell channels in the
intersegmental membrane are clearly visible.

314

Psyche

[December

Figure 14. Above: SEM photograph of the glandular cell openings of the sternal
gland of Paltothyreus tarsatus. Note the filament-like protrusions of the interseg-
mental membrane. Below: Sagital section through the same area. (Lettering as in
Fig. 5).

1978]

Holldobler & Engel — Glands in Ants

315

Figure 15. “Sting glands” of Myrmecia pilosula (above) and Amblyopone
pallipes (below). (Lettering as in Fig. 5).

Jefil

316

Psyche

[December

Figure 16. Glands associated with the sting sheath. Above: Sagital section
through sheath valve with glandular cluster and channels in Odontomachus haema-
toda. Below: Myrmecia pilosula. (Lettering as in Fig. 5).

1978]

Holldobler & Engel — Glands in Ants

317

Figure 17. Glands associated with sting sheath in Paltothryeus tarsatus. Above:
Sagital section through glandular cells with channels. Below: Sagital section through
glandular cell cluster with channels. (Lettering as in Fig. 5).

318

Psyche

[December

Figure 18. Glandular cell clusters in the T" abdominal sternum of Novomessor
cockerelli (above) and Veromessor pergandei (below). GE = glandular epithelium in
the 7th sternum. (Lettering as in Fig. 5).

1978]

Holldobler & Engel — Glands in Ants

319

Figure 19. SEM photograph of the 7th tergum of Pachycondyla (= Termitopone )
laevigata. Glandular applicator surface (GA) is shown in greater detail in the picture
below.

320

Psyche

[December

Figure 20. Above: Sagital section through the pygidial gland of Pachycondyla
( =Termitopone ) laevigata. Below: SEM photograph of the cuticular structure (GA in
Fig. 19) on 7th tergum. The cut open area shows the large cavities associated with the
structure. (Lettering as in Fig. 5 and Fig. 18).

1978]

Holldobler & Engel — Glands in Ants

321

Figure 21. Sagital section through the gaster of Odontomachus haematoda,
showing the well developed reservoir of the pygidial gland (PG).

322

Psyche

[December

Figure 22. Above: Sagital section through the gaster of Neivamyrmex nigrescens,
showing the pygidial gland (PG), postpygidial gland (PPG) and anus (A). Below: Close
up of sagital section through pygidial gland and postpygidial gland of N. nigrescens.
(Lettering as in Figs. 5 and 8; R = reservoir).

1978]

Holldobler & Engel — Glands in Ants

323

Figure 23. Above: SEM photograph of tip of the gaster of Eciton hamatum.
Below: Sagital section through the same segments. (Lettering as in Fig. 8; S = sting;
GE = glandular epithelium in 7th sternum).

324

Psyche

[December

Figure 24. Eciton hamatum: SEM photograph of the intersegmental membrane
between 6th and 7th terga, where the pygidial glands open. Below: Close up of the brush-
like structure.

1978]

Holldobler & Engel — Glands in Ants

325

pg

Figure 25. Schematic drawings of sagital sections through the gaster tips of
Dorylus molesta (left) and Eciton hamatum (right). (Lettering as in Fig. 8; a = anus;
ag = anus glands).

Figure 26. Sagital section through anus of Dorylus molesta worker; AG = anus
gland; A = anus.

Psyche

[December

Figure 27. Left- SEM photograph of abdominal tip of Veromessor pergandei. “PG” indicates opening of pygidial gland
between 6‘ and 7th terga; “A” indicates opening of anus. Right: Sagital section through pygidial gland of Veromessor, which has
a large reservoir (R) and each cell cluster contains 50-70 glandular cells (GC).

1978]

Holldobler & Engel — Glands in Ants

327

Figure 28. Above: SEM photograph of abdominal tip of Liometopum apicu-
latum. “PG” indicates where the pygidial gland opens between 6th and 7th terga.
“A” indicates opening of anus; SG = sternal gland (Pavan’s gland). Below: Sagital
section through gaster of L. apiculatum. PG = reservoir of pygidial gland; 7. = 7th
tergum.

328

Psyche

[December

References

Altenkirch, G.

1962. Untersuchungen liber die Morphologie der abdominalen Hautdriisen
einheimischer Apiden (Insecta, Hymenoptera) Zoologische Beitrage, 7,
161-238.

Bernstein, R. A.

1974. Seasonal food abundance and foraging activity in some desert ants. The
American Naturalist, 108, 490-498.

Blum, M. S.

1966. The source and specificity of trail pheromones in Termitopone, Mono-
morium and Huberia, and their relation to those of some other ants.
Proceedings of the Royal Entomological Society of London, 41, 155—
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1977. Behavioral responses of Hymenoptera to pheromones and allomones.
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Blum, M. S. and H. R. Hermann.

1978a. Venoms and venom apparatuses of the Formicidae: Myrmeciinae,
Ponerinae, Dorylinae, Pseudomyrmecinae, Myrmicinae and Formici-
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Blum, M. S. and H. R. Hermann.

1978b. Venoms and venom apparatuses of the Formicidae: Dolichoderinae and
Aneuretinae. In: Handbook of Experimental Pharmacology, Springer-
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Blum, M. S. and C. A. Portocarrero.

1964. Chemical releasers of social behavior. IV. The hindgut as the source of
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Bordas, L.

1895. Appareil glandulaire des Hymenopteres. Ann. Sci. Nat. Zool. 19, 289-
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Brown, C. A., J. F. Watkins II and D. W. Eldridge.

1979. Depression of bacteria and fungi by the army ant secretion: Skatole.
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Chadab, R. and C. Rettenmeyer.

1975. Mass recruitment by army ants. Science 188, 1124-1125.

Holldobler, B.

1978. Ethological aspects of chemical communication in ants. Advances in the
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Holldobler, B., M. Moglich and U. Maschwitz.

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Holldobler, B., R. Stanton and H. Engel.

1976. A new exocrine gland in Novomessor (Hymenoptera: Formicidae) and
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HGlldobler, B. and C. P. Haskins.

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Janet, Ch.

1898. Etudes sur les Fourmis, les Guepes et les Abeilles Note 17: Systeme
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1899. Zur Kenntnis der Hautdriisen der Apidae und Vespidae. Anat. Anz. 15,
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1964. Gefahrenalarmstoffe und Gefahrenalarmierung bei sozialen Hymenop-
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Maschwitz, U., B. Holldobler and M. Moglich.

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1962. Methylmetacrylat als Einbettungsmedium fUr Insekten Experientia
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Robertson, P. L.

1968. A morphological and functional study of the venom apparatus in repre-
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1978. Nothomyrmecia macrops: A living-fossil ant rediscovered. Science 201,
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1975. Trail-following by the army ant Neivamyrmex nigrescens: Responses by
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Watkins, J. F.

1964. Laboratory experiments on the trail following of army ants of the genus
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22-28.

Watkins, J. F., T. W. Cole and R. S. Baldridge.

1967. Laboratory studies on interspecific trail following and trail preference of
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1969. Attractant-repellent secretions in blind snakes ( Leptotyphlops dulcis )
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1972. Feeding and digestion in some ants ( Veromessor and Manica). Bio-
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1975. Natural history of Veromessor pergandei — II. Behavior. The Pan-
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1957. Notes on the anatomy of Rhytidoponera convexa Mayr (“violacea”
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1957. Notes on the anatomy of the Formicidae I. Stigmatomma pallipes
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1960. The anatomy of Rhytidoponera metallica F. Smith (Hymenoptera:
Formicidae). Ann. Entmol. Soc. Amer. 53, 793-808.

1963. The anatomy of adult queen and workers of the army ants Eciton
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1971. The insect societies. The Belknap Press of Harvard University Press,
Cambridge, Mass.

LIFE HISTORY OF DYNASTOR DARIUS
(LEPIDOPTERA: NYMPHALIDAE: BRASSOLINAE)
IN PANAMA

ByAnnette Aiello and Robert E. Silberglied*
Smithsonian Tropical Research Institute
P.O. Box 2072, Balboa, Panama

Dynastor darius (Fabr.) (Figure l)1 is a large crepuscular butterfly
found from Guatemala south into Brasil (Blandin, 1976; Blandin and
Descimon, 1977). The larva is known to feed on various bromeliads.
In the interests of amplifying the scattered and fragmentary descrip-
tions of the immature stages by previous workers, and commenting
upon certain discrepancies between their findings and our own, we
here present data on twelve individuals of Dynastor darius reared on
Barro Colorado Island (BCI), Panama in 1978.

On 21 June we captured a female of Dynastor darius, and over the
period 23-25 June obtained twelve eggs (Figure 2). These were
spherical, 2.2 mm in diameter, and cream colored with a pattern of
anastomosing striae oriented from pole to pole. The first instars
emerged eleven days after oviposition and we presented them with
leaves of a variety of monocot species: Musa paradisiaca (Musaceae),
Heliconia sp. (Heliconiaceae), Monster a sp. (Araceae), Cocos nuci-
fera (Palmae), Zingiber sp. (Zingiberaceae), Philodendron triparti-
tum (Araceae), Ananas comosus (Bromeliaceae), and an unidentified
grass (Gramineae). The larvae ate only the pineapple (Ananas
comosus).

First Instar Larvae

Newly hatched larvae were 7.6-8 mm in length excluding the
caudae, which were 2 mm long, orange on the basal half and black
apically. At the tip, each cauda bore a 1 mm long seta which was black
on the basal half and white apically, plus several appressed setae
toward the base. The tip of the body between the caudae was red. The

* Present address: Museum of Comparative Zoology, Harvard University, Cam-
bridge, Massachusetts, U.S.A. 02138.

Manuscript received by the editor February 26, 1979.

‘Figures 1 and 11 represent the individual in rearing Lot 78-75. Figures 2-10
represent the twelve individuals in rearing Lot 78-84.

331

332

Psyche

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Figure 1 . Dynastor darius, adult male. Left: dorsal; Right: ventral. Scale = 20 mm.

1978]

Aiello & Silberglied — Dynast or darius

333

head capsule (Figure 9, A and B) displayed a complex pattern of
beige, brown, and black, almost obscured by a dense covering of 1
mm long black, plumose setae. Four pairs of low bumps marked the
positions where horns appeared in the second instar.

Before taking their first meal, each larva (Figure 4) had a red-
brown dorsal stripe plus four yellow and three red-brown lateral
stripes, and an oval red dorsal spot (Figure 7A) between abdominal
segments 3 and 4. After feeding and expanding, the larval stripes
(Figure 5) became green and yellow, with several extra yellow stripes
appearing between old ones. The dorsal spot became black, flanked
on the sides with red and orange, and bordered on each end with
orange and white (Figure 7B). A second and much smaller dorsal
spot, white at the anterior end and black posteriorly, appeared
between abdominal segments 5 and 6. The positions of these spots
correspond to positions 6-7, and 8-9 of the system used by
Burmeister (1873) and Muller (1886) who numbered thoracic and
abdominal segments as one series.

A pinkish red eversible gland (Figure 3), located ventrally on the
prothorax between the legs and head, became particularly evident in
later instars. When disturbed, larvae reared up and waved from side
to side, emitting a pineapple-like odor that seemed to come from the
large everted gland.

Further Larval Development
Number of instars

Although larvae were reared separately and given food ad libitum,
the number of instars varied among the ten individuals surviving to
pupation. Three reached pupation after completing five instars, six
after six instars, and one after seven. Due to this variability in
development, larval size could not be used to identify instar number
(Figure 6).

Development time

For the eight individuals surviving to adulthood2, average devel-
opment time, from oviposition to eclosion, was 80.9 days (range:
73-88 days, s: 6.6 days, N = 8). The amount of time spent in each stage
by each individual is shown graphically in Figure 8. The average time

2Of the remaining four, one died as a first instar, one sixth instar was killed for
preservation, and two underwent poor pupations.

334

Psyche

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Figures 2 and 3. 2. Eggs. Scale = 1 mm. 3. Fifth instar larva (individual no. 4) displaying everted prothoracic gland. Scale — 2 mm.

Figures 4-6. 4. Early first instar larva (individual no. 1 ). 5. Late first instar larva (no. 1 ). 6. Fifth instar larvae (no. 1 (small)and

no. 4 (large)). Note that no, 4 has “final” instar head capsule. All scales = 3 mm.

336

Psyche

[December

H 9 reen

I black
o orange
p pink

pb pink-brown
rb red-brown
w white

Q-rb

A

Figure 7. Dorsal spot at junction of abdominal segments 3 and 4. A: early first
instar (individual no. 1); B: late first instar(no. 1); C: fourth instar(no. 8); D: early fifth
instar (no. 4); E: late fifth instar (no. 12). Scale = 2 mm.

spent in any given instar was 9.6 days (range 6-16 days, s: 2.4 days,
N= 63). Every larva spent four days molting from each instar to the
next: new cuticle formation occurred during the first two days,
ecdysis took place on the thrid day, and by the end of the fourth day
the new cuticle had hardened. No feeding took place during these
four-day periods. These data reveal that the larvae were engaged in
molting-related activities approximately 40% of the time, and feeding
and resting 60% of the time.

Head capsule

Beginning with the second instar, the head bore four pairs of horns
(Figure 9, C and D) radiating from it in a transverse plane
perpendicular to the body axis. The larger and most dorsal two pairs
were slightly expanded towards their rounded tips, while the smaller
and more lateral two pairs were conical. The fine white simple setae
which covered the head did not obscure pattern details. Setae on the
horns were longer and projected from their tips.

1978]

Aiello & Silberglied — Dynast or darius

337

bO

O Of

338

Psyche

[December

The complex beige, brown, and black head pattern of the second
instar persisted through all succeeding instars except the final one.
With the final instar, regardless of the number of previous instars, the
head capsule took on a new, less complex pattern (Figure 9, E and F)
of brown and black. All larvae which attained this new head pattern
pupated at their next molt.

Head capsule widths, measured for all instars of each individual,
indicate that size variation increased with each succeeding instar
(Figure 10). However, the final instar, regardless of number (5th, 6th,
7th), showed slightly less size variation than did the next to last instar,
suggesting that once a larva has reached a certain critical size it molts
to the final instar, and from there to the pupa.

Body

With growth, the body colors underwent gradual change and
additional stripes were added continually. The yellow and green
stripes passed through a pale and dark green stage, and finally
became dark brown and dark green. In mature larvae they anas-
tomosed slightly.

The large dorsal spot changed from the pattern described for the
first instar, to one with a greenish and black center, ringed with black,
followed by pink-brown, pink, green, black (narrow), and green
(Figure 1C). This pattern gave way to one with a black and green
reticulated center, ringed with black (wide), pink (narrow), black
(narrow), green (narrow), black (narrow), and green (medium)
(Figure 7D). In mature larvae, the central black and green reticulate
pattern was surrounded by a wide black ring, followed by rings of
green, black, green, black (narrow and irregular), and green (wide)
(Figure 7E).

The smaller dorsal spot also changed with growth. Gradually the
anterior white portion expanded and encircled the black, and the
center became reticujated black and white. By maturity the small spot
had become an exact miniature of the larger one.

As larvae reached their full size, four smaller spots appeared, at the
junctions of abdominal segments 1-2, 2-3, 4-5, and 6-7. Initially
these spots were red-brown, but later changed to black. In addition,
the nine pairs of spiracles showed as black spots along the sides of the
body (prothorax, abdominal segments 1 through 8).

Caudae

From the second instar on, the caudae lacked apical setae, and

1978] Aiello & Silberglied — Dynastor darius 339

Figure 9. Head capsules of larva no. 3. Lateral(A, C, E) and frontal(B, D, F) views
of instars one (A, B), two (C, D), and five (E, F). Scale = 2 mm.

were covered with short appressed setae. With growth the caudae
became shorter in proportion to the body, and widened basally,
approaching a conical shape in large larvae. Meanwhile, the black
portion decreased until only the tips of the caudae were dark. The
orange changed to red-brown and at last to dark brown, and the red
area disappeared from the tip of the abdomen.

Pupation

Pupation and the processes leading to it took four days. During the
first day, the larvae stopped eating, turned dark brown, emptied their
guts, and shortened their bodies. The second day was spent crawling
upon a support and spinning a silk stalk. During the third day, larvae
grasped the stalk with their anal prolegs and hung upside down, and

340 Psyche [December

Figure 10. Heights of larval head capsules of each instar. Small numbers indicate
frequencies greater than one.

on tne morning of the fourth day they shed the last larval skin. By the
evening of the fourth day, the pupae were hardened and darkened.

Pupa

Based on the eleven individuals available to us, the pupa (Figure
1 1) of Dynastor darius is about 4 cm long and bears a remarkable
resemblance to the head of a snake. Its brown, beige, and whitish

1978]

Aiello & Silberglied — Dynastor darius 341

Figure 1 1 . Pupa. Scales = 5 mm.

342

Psyche

[December

pattern, coupled with the constriction across the wings, the false
scales beneath, the snake-like eyes (located where the true eyes of the
adult butterfly will form), and large size give a very convincing
illusion of snakeness, despite the fact that the eyes are in the “wrong”
position for snake eyes. The question which arises when trying to
understand such resemblances, is, “what good does it do to mimic a
snake if, during those precious moments when your surprised
predator hesitates, you cannot carry through by turning and
running?” The answer may be that the predator itself turns and flees
after suddenly coming face-to-face with a realistic “snake” that waves
violently back and forth, as does the pupa of Dynastor darius when
disturbed.

Like many brassoline pupae, the pupa of Dynastor darius has two
lateral pairs of keels. The first begins on the outer margin of the wing
very near the tornus, and runs across the wing perpendicular to the
outer margin, ending at the edge of the discal cell. The second begins
near the midpoint of the inner margin and follows that margin to the
base of the wing.

The ground color of the pupa is whitish. The ventral side is
irregularly striped with fine brown longitudinal lines which become
more dense along a line following the spiracles. These lines fade into
pseudo-reptilian scaling toward the anterior end of the pupa.
Dorsally, the abdomen bears a complex brown pattern of fine lines
and spots, which ends abruptly at its interface with the darker, ventral
pattern.

The fore wings (concealing the hind wings) are covered with
reticulations, which are beige except at the tornal angle where they
are brown. The dark tornal angle lies between the keels and is marked
off sharply from the rest of the wing by a dark brown line which
begins at the midpoint of the inner margin (beginning of the second
keel), runs perpendicular to it until reaching the end of the first keel,
then turns sharply, following that keel to the outer margin where it
joins the dark line which follows the spiracles.

The eye is beige with tiny black spots and is surrounded by a fine
black line.

The pupal stage lasts an average of 15. 1 days (range: 13-17 days, s:
1 .8 days, N = 8). The day before eclosion, the pupa turns dark brown.
The dorsal pattern of the fore wings begins to show through the pupal
integument several hours before emergence of the adult butterfly.

1978]

Aiello & Silberglied — Dynastor darius

343

Discussion

The life cycle

Burmeister’s (1873) brief description and figure of a last instar
larva of Dynastor darius from Rio de Janeiro agrees with our
observations in all important respects, differing only slightly in
details of the dorsal spots and the color of the head. The dorsal spots
of Burmeister’s larva were described as “oval eyes, black, with a
yellowish pupil, surrounded by a white outer margin, bordered with
black.” It is not surprising that descriptions of the complex and ever
changing dorsal spots would vary. The black and green reticulated
center might be interpreted as dull yellow from a distance, the dark
green or brown might be seen as black, and the outermost green rings,
which blend with the pattern of the rest of the body could easily be
overlooked. The head of Burmeister’s specimen was “yellow-brown
with a large dark maroon spot on each side immediately in front of
the horns which are of the same color.” His observations may have
been made from a newly molted, partially darkened individual, or
Brasilian specimens might be paler than Panamanian ones.

During a long visit to southern Brasil, Muller (1886) reared
Dynastor darius from egg to adult, but did not describe the egg. His
larva passed through five instars, the period from hatching to adult
lasting 71 days. His detailed description notes changes in the dorsal
spots and in the exact number of stripes present on the body. Our
larvae showed considerable variation in number of body stripes
among individuals of the same size. Muller’s first instar was “red-
brown and white,” in contrast to ours which were red-brown and
yellow. As did Burmeister, Muller also described the dorsal spots as
having a yellow center. His pupa was “greenish white with very dense,
fine, irregular long brown stripes.” All eleven of our pupae were
brown. Perhaps the pupal color varies geographically, genetically, or
depends upon light conditions during or prior to pupation.

Rothschild (1916) briefly described and figured the last instar larva
and pupa of Dynastor darius from an unpublished sketch by E.
Hartgen, in the Tring Museum Library. The larva, as far as can be
determined, was essentially the same as ours. Rothschild’s figure of
the “deep grass-green” pupa is difficult to interpret; the wings appear
to cover the dorsal rather than the ventral side of the body. Being a
figure from a sketch, perhaps something was lost in translation.

344

Psyche

[December

None of these authors made reference to the snake-like appearance
of the pupa. The resemblance may not be so obvious in green pupae.

Ontogenetic variability

With respect to the number of instars of Dynastor darius, the
situation may be similar to that of Manduca sexta (Sphingidae). The
larvae of M. sexta initiate the sequence of events leading to pupation
at a critical weight, independent of the rate of growth (Nijhout and
Williams, 1974a, 1974b). Like Manduca, Dynastor exhibits variabil-
ity in the number of larval instars, corresponding with variability in
growth rate, and it appears that the larvae commit themselves to
pupation when a critical size is reached. In contrast to Manduca
however, commitment in Dynastor may take place during the next to
last instar, as evidenced by the fact that the final instar always has a
head capsule pattern different from that of previous instars. Another
possibility is that there may be two commitments: one to the final
head capsule pattern, and another to pupation.

Preserved material (head capsules, pupal skins, adults, and larvae
in alcohol) of Dynastor darius, upon which this study was based,
labeled as Lots 78-75 (an individual collected as a pupa) and 78-84
(reared from eggs), has been deposited in the Museum of Compara-
tive Zoology, Harvard University.

We would like to thank the Smithsonian Tropical Research
Institute, Panama, for use of their facilities, Mr. Gary Stump for
locating a wild pupa of Dynastor darius on BCI, Mr. Nigel Franks for
assisting with rearing, Mr. Gordon B. Small for identification of the
butterfly, and Mr. Norman Woodley for reading the manuscript.

References

Blandin, P.

1976. La distribution des Brassolinae (Lepidoptera-Satyridae). Faits et prob-
lemes. pp. 161-218. In H. Decimon, ed., Biogeographie et Evolution en
Amerique Tropicale, Publ. Lab. Zool. de l’Ecole Normale Superieure
No. 9.

Blandin, P. and Descimon, H.

1977. Contribution a la connaissance des Lepidopteres de Fequateur. Nouvelles
donnees sur les Brassolinae [Nymphalidae] de l’occidente. Annls. Soc.
ent. Fr„ (N.S.) 13: 75-88.

Burmeister, H.

1873. De Morphonides Bresiliens. Rev. Mag. Zool. Pureet Appl., (3)1: 17-47.

1978]

Aiello & Silberglied — Dynast or darius

345

Muller, W.

1886. Siidamerikanische Nymphalidenraupen Versuch eines natiirlichen Sys-
tems der Nymphaliden. Zoologische Jahrbucher. Zeitschrift fur Syst.
Geogr. Biol. Thiere, 1: 417-678.

Nijhout, H. F. and Williams, C. M.

1974a. Control of moulting and metamorphosis in the tobacco hornworm,
Manduca sexta (L.): growth of the last-instar larva and the decision to
pupate. J. Exp. Biol., 61: 481-491.

1974b. Control of moulting and metamorphosis in the tobacco hornworm,
Manduca sexta (L.): cessation of juvenile hormone secretion as a trigger
for pupation. J. Exp. Biol., 61: 493-501.

Rothschild, L.

1916. Notes on Amathusiidae, Brassolidae, Morphidae, etc., with descriptions
of new forms. Novit. Zool., 23: 299-318.

A CLOSE RELATIONSHIP BETWEEN TWO SPIDERS
(ARACHNIDA, ARANEIDAE):

CURIMAGUA BAYANO SYNECIOUS
ON A DIPLURA SPECIES*

By Fritz Vollrath
Smithsonian Tropical Research Institute
P.O. Box 2072, Balboa, Canal Zone, Panama

Spiders are known to be strictly carnivorous and, moreover,
often cannibalistic as well. Yet some animals have achieved a status
of close coexistence with several spiders, notably web spinners.
Web-building spiders do not change web sites frequently. They are
rather sedentary (Turnbull, 1964), which makes their webs a suitable
habitat for associates. Some wasps ( Eumenidae , for example) build
their mud nests within the webs of the “social” spider Anelosimus
eximius (Vollrath, in prep.) whose webs also harbor certain moth
larvae (Robinson, 1977a) and Hemiptera (Vollrath, in prep.). The
most striking examples of coexistence, however, are the kleptopara-
sitic spiders of the genus Argyrodes (Theridiidae) that inhabit the
snares of many web-building spiders (Brignoli, 1966; Robinson
and Robinson, 1973; Vollrath, 1977).

In this article I describe some aspects of the biology of another
spider that lives in close association with a web-building spider. It is
the only spider known so far that is unable to capture its food but
has to rely on its host to catch and, even further, to predigest the
prey.

CURIMAGUA AND DlPLURA

Curimagua bayano (Symphytognathidae; Forster and Platnick,
1977) has only been found riding on the cephalothorax and crawling
on the webs of Diplura sp. (undescribed species of Mygalomorphae,
specimen in the American Museum of Natural History) in Panama.
Although the diplurid host is not uncommon in different localities
in the monsoon forest at Pipeland Road (Canal Zone), only in the
Bayano limestone cliffs (the stream banks of the Rio Maje and Rio
Tigre) did their webs harbor Curimagua. On a recent trip to Iquitos,

* Manuscript received by the editor January 15, 1979.

347

348

Psyche

[December

Peru, I found a similar association between a Curimagua and a
Diplura in the “white sand forest” of Mishana.

Diplura sp. is a beautifully colored spider that measures up to 4
cm in length and builds sheet- or funnelwebs spreading up to 50 X 50
cm. The spider rests in the entrance of its retreat, a silken tube,
waiting for prey to fall onto or crawl over the sheet. The prey is
bitten in a sudden lunge and dragged into the retreat for consump-
tion. In a few minutes the prey (e.g., a cricket) will be a mushy and
liquified body, due to the combined action of the masticating
movements of the relatively large chelicerae and the regurgitated
digestive enzymes of the spider. Diplura s prey, as judged from the
remains, seems to consist mainly of beetles (Coleoptera), crickets
(Gryllidae), and ants (Hymenoptera). Millipedes (Diplopoda) and
tailless whipscorpions (Amblypygi) were also found, as was a partly
digested dendrobatid frog (Anura, Dendrobatas auratus).

In contrast to the diplurid host, Curimagua bayano is a tiny,
mitelike spider. The pale body of the adult female is globular and
measures not more than 1 .3 mm in length. The male is equally small;
for exact measurements, see Forester and Platnick (1977). Curi-
magua bayano was found in the retreats of several webs of Diplura.
The highest number per host was three females and one male, all in
the relatively small web (30 X 30 cm) of a male Diplura. An egg case
which was also found in the host’s retreat, and which was assumed
to be of curimaguan origin, consisted of four small eggs incorpo-
rated into a transparent and fluffy sphere (diameter 0.8 mm) of
loosely spun silk. Curimagua crawls about between the silken
threads of the host retreat, several centimeters behind the entrance,
near the place where the host feeds. Because of its minute size,
Curimagua has no difficulty in negotiating the fine mesh of silk.
Often the animals were observed riding on the diplurid, perched on
the center of its cephalothorax, apparently clinging to the fine hairs
that cover it. Even their host’s violent lunge for prey does not shake
them off. I never saw the host react to Curimagua crawling about in
the web or traversing the body of the host, even when it moved
across the host’s eyes (Figure 1). Diplura collected on Pipeline Road
(where Curimauga was absent in ten Diplura webs inspected) also
showed no hostile reaction when Curimagua climbed “aboard,”
suggesting that instead of being ignored the associate may not be
perceived by its host.

1978]

Vollrath — Relationship Between Two Spiders

349

Figure 1. The mygalomorph spider Diplura sp. in its silken retreat, feeding on a
grasshopper. Perched on its eyes and on its chelicerae is the symphytognathid asso-
ciate Curimagua bavano.

When I brushed Curimagua off the dipluran cephalothorax, they
crawled about in the web and, as soon as contact was made with an
extremity of the host, they started to climb it to regain their former
position. During an experiment, a Curimagua female was taken
from the back of a Diplura and confined to a Petri dish (diameter 5
cm) with another mygalomorph spider of approximately the same
size as the diplurid. The Curimagua did riot attempt to mount this
spider.

Food of Curimagua

Scanning photographs of a female Curimagua taken in frontal
view of the “head” region (frontal ridge of cephalothorax: clypeus)
show the distinct feature of this group, that is, the pedipalpal coxae
lack the palpae (Figure 2). The coxae and the clypeus form a
circular depression which is densely covered with fine hairs as well
as trichobothria. The chelicerae — which in spiders are normally

350

Psyche

[December

used to hold and kill prey — are tiny and fused at the bases (see also
Forester and Platnick, 1977). Moreover, the chelicerae are located
deep inside the mouthlike basin. Therefore, I consider it unlikely
that they could be used to hold or tear even the smallest prey, thus
restricting Curimagua to liquid food. I confined Curimagua to Petri
dishes (diameter 5 cm) which contained a variety of small arthro-
pods (Collembola, Coleoptera, Ricinulei) found in Diplura webs, as
well as dead food items, but never did I find the spiders feeding on
any of these offerings. Curimagua bayano shares fused chelicerae
with its congenerics (Forster and Platnick, 1977), some of which
construct very finely meshed orbicular webs with a diameter of only
several mm (Eberhard, pers. comm.). The prey of these free-living
congenerics are unknown. Their nets with a mesh size of approxi-
mately 125 /I (Patu saladito) may serve to filter for aerial plankton
such as pollen and fungus spores (Vollrath, in prep.).

When a Diplura that is carrying Curimagua bayano is fed an
insect, the Curimagua climbs forward from its resting position on
the central cephalothorax, over the eyes, and down the mighty
chelicerae of its host until it reaches the digestive fluids enveloping
the prey item. The abdomen of the pirating associate visibly swells
and only a few minutes pass before the Curimagua climbs back onto
the cephalothorax. Individuals moving about in the silk of the
retreat are also attracted to a feeding Diplura, perhaps alerted by
the vibrations generated by the masticating movements of the host.

Other Associates of Diplura

Curimagua is not the only associate of Diplura webs. Other
spiders ( Mysmenopsis ssp.; Platnick and Shadab, 1978) have been
found by Kraus (1955) in Guatemala, and three species of Mys-
menopsis (M. dipluroamigo, M. ischnamigo, and M. gamboa) were
observed by Kirkendall (in prep.) in Panama in the webs of three
differently sized diplurid spiders. These associates move about in the
sheet region of the diplurid web, approaching and catching small
insects that failed to attract the host. Some individuals might also
move toward a feeding host, but I rarely saw them sharing its prey.
Most often their approach is detected by the Diplura, which then
turns around in the retreat, spreading silk with its long spinnerets
and thus hindering the advance of the would-be pirates. I never
observed this behavior of Diplura directed toward Curimagua. The

1978]

Vollrath — Relationship Between Two Spiders

351

ability of Mysmenopsis dipluroamigo to survive without the
“proper” host is shown by the fact that Kirkendall and 1 found a
complex space web in a bush, resembling the web of the “social”
spider Anelosimus eximius (Simon, 1892). This web (34 X 37 X 27
cm) contained 9 females, 10 subadult females, 16 males, 2 subadult
males and 25 juvenile M. dipluroamigo, in addition to 2 Ischnothele
guianenesis (Walck.). Other associates of Diplura webs encompass
Coleoptera, dipteran larvae (Kirkendall, in prep.), lepidopteran
larvae (Robinson, 1977), and therediid spiders of the genus Argy-
rodes (Vollrath, 1977).

Figure 2. Scanning electron microscope photograph (350X) of the frontal cepha-
lothorax of Curimagua bayano showing the two frontal eyes (E), the coxae and
trochanter of legs 1, the frontal ridge of the cephalothorax (the clypeus = Cly), the
coxal processes (Cpx) of the pedipalpae with the reduced pedipalp (P), and, finally,
the chelicerae (Ch) with the cheliceral claws (CC).

352

Psyche

[December

Discussion

Curimagua bavano is an extreme case of specialization in spiders.
It is the first spider described which is not a predator and not even a
carnivore sensu strictu, but an ectoparasite or — in the terms of
Robinson and Robinson (1977) — a dipsoparasite, resembling, in
a way, representatives of another and most successful Arachnid
order, the mites (Acarina) which produced many parasite genera
(Vollrath, E., 1978). Curimagua bayano should not be called a
commensal (Gray, 1967) since the fluids imbibed are either the
host’s costly digestive enzymes (5% protein solution in Tegenaria
’trica; Mommsen, 1977) or liquefied prey tissue. In both cases,
Curimagua deprives its host of nutritive proteins, thus reducing its
fitness.

Acknowledgmemts

My thanks go to Drs. H. Levi and N. Platnick for identification
of the spiders, to Dr. R. Foelix for help with the scanning electron
microscopy, and Drs. M. Robinson, D. Morrison, and D. Windsor
for comments on the manuscript. Figure 1 was drawn by Arcadio
Rodaniche after photographs and a specimen. The work was
supported by a Noble Fellowship from the Smithsonian Tropical
Research Institute.

References

Brignoli, P. M.

1966. La societa eterotipiche degli Araneidi 1. Rend. Acad. Naz. 16: 219-246.
Forster, R. R. and Platnick, N. I.

1977. A review of the spider family Symphytognathidae. Am. Mus. Nov. 2619:
1-29.

Gray, P.

1967. Dictionary of the Biological Sciences. Reinhold, New York.

Kraus, O.

1955. Spinnen aus El Salvador. Abh. Senckenb. Naturf. Ges. 493: 31.
Mommsen, T. P.

1977. Zusammensetzung und Funktion extraintestinaler Verdauungsfluessig-
keit und Gift einer Spinne ( Tegenaria atrica Koch). Dissertation, Univ.
Freiburg.

Platnick, N. I. and Shadab, M. U.

1978. A review of the spider genus Mysmenopsis. Am. Mus. Nov. 2661: 1-22.

1978]

Vollrath — Relationship Between Two Spiders

353

Robinson, M. H.

1977. Symbioses between insects and spiders: an association between lepidop-
teran larvae and the social spider Anelosimus eximius. Psyche 84:
225-232.

Robinson, M. H. and Robinson, B.

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

1977. Associations between flies and spiders: bibiocommensalism and dipso-
parasitism? Psyche 84: 150-157.

Simon, E.

1892. Observations biologiques sur les Arachnides. 1. Araignees sociables.
Ann. Soc. Ent. France, 60: 5-14.

Turnbull, A. L.

1964. The search for prey by a web-building spider Achaearanea tepidariorum.
Can. Entomol. 96: 568-579.

Vollrath, E.

1978. Milben und ihre okologischen Beziehungen zu Insekten. Staatsex-
amensarbeit. Univ. Freiburg.

Vollrath, F.

1977. Zur Okologie und Biologie von kleptoparasitischen Argyrodes elevatus
und synoken Argyrodes- Arten. Dissertation, Univ. Freiburg.

SYSTEMATICS AND EVOLUTION OF
FOREST LITTER ADELOPSIS
IN THE SOUTHERN APPALACHIANS*
(COLEOPTERA: LEIODIDAE; CATOPINAE)

By Stewart B. Peck
Department of Biology

Carleton University, Ottawa, Ontario K1S, 5B6, Canada

The beetle genus Adelopsis was described by Portevin in 1907,
and was proposed for a microphthalmic soil-inhabiting species
from Bolivia. Since then, some 20 additional species have been
described, and the genus is known to be distributed through the
Neotropics from Mexico to southern Brazil and adjacent Argentina.
The species are generally large-eyed and winged, and are probably
all scavengers of decaying organic matter in mesic tropical lowland
and montane forests. They may occasionally occur in caves. The
beetles are seldom represented in collections, but they may be
frequently collected by sifting forest litter, or by using dung or
carrion-baited pitfall traps. My field program of such collecting in
the Neotropics since 1966 shows the genus to be far more diverse,
abundant, and widespread than indicated in the present literature
(Peck, 1977).

Adelopsis was first found to occur outside the Neotropics when I
(Peck, 1973) realized that Adelops mitchellensis (Hatch, 1933) from
Mt. Mitchell, North Carolina, actually belonged in the genus
Adelopsis . Some authors had placed the species in the genus
Ptomaphagus. At that time I noted that I also had material of other
species from North Carolina, Georgia, Tennessee, Alabama and
West Virginia. My indication that they were also in New Mexico
was in error (Peck, 1978). The purpose of this paper is to describe
these species which occur in forest litter and soil habitats in the
southeastern United States, and to consider their distributional and
evolutionary history.

Generic diagnosis. The characters are those oI/Leiodidae, Cato-
pinae, Ptomaphagini (as given in Peck, 1973). As such they are
small beetles with a loose antennal club, with segment 8 always
smaller than 7 and 9. They have transverse pronotal and oblique

* Manuscript received by the editor May 1, 1979.

355

356

Psyche

[December

elytral strigae, and the summits of their tibiae are armed with a
comb of many short and equal spines, as well as by two longer spurs
(see Peck, 1973, 1977 and their cited references).

Adelopsis is generally difficult to separate from its nearest
neighbor, the genus Ptomaphagus, using only external characters.
Tropical species of Adelopsis (but not those of the U.S.) usually
differ by being smaller (length 2.6 mm or under) and in having the
antennal club more loosely composed of gradually larger segments,
of which the last two are often conspicuously lighter in color. The
genera are more reliably separated by the chitinized internal repro-
ductive structures of the aedeagus and spermatheca. In Adelopsis
the spermatheca (as far as is known) is a more simple curved tube,
and the aedeagus has a tip which is more elaborately sculptured, or

Map 1. Distribution of known species of Adelopsis in the southeastern United
States. Heavy irregular line indicates maximum extent of Pleistocene glaciations.
A, A. mitche/lensis. B, A. steevesi. C, A. alleghenvensis. D, A. suteri. E, A. rich-
landensis. F, A. appalachiana. G, A. jonesi. H, A. bedfordensis. I, A. cumberlanda.
J, A. scottsboroensis. K, A. nashvillensis. L, A. fumosa. M, A. alta. N, A. joanna.
O, A. pisgaherisis. P, A. orichalcum.

1978] Peck — Adelopsis in the Southern Appalachians 357

is broader and more blunt. The orifice is dorsally sub-terminal and
cuts the left side of the aedeagus as in Ptomaphagus. The south-
eastern U.S. Adelopsis species are all forest litter and soil inhabi-
tants. They are all wingless and have eyes reduced to a collection of
about 20 pigmented facets. These structural features will help to
distinguish them from most Ptomaphagus, which are either winged
and with larger eyes, or are more subterraneanly adapted (for life in
caves) and have more-reduced eyes.

Description. The following applies only to the range of variation
known in the species in the southeastern U.S. Additional details on
Neotropical species may be found in Jeannel (1936) and Szymcza-
kowski (1964, 1968, 1969, 1975). Length 2.3 to 2.6 mm, width 1.2 to
1.3 mm. Form elongate oval, compact, convex (Figs. 1, 2). Color
light to dark reddish brown. Pubescent, with numerous short
recumbent hairs. Head finely punctured; eyes reduced to collection
of about 20 pigmented facets (Fig. 3); eye width 1/3 width of space
from antennal base to head margin across eye. Antennae short,
compact; club darker, somewhat flattened; reaching from middle to
hind margin of pronotum when laid back; segment 3 shorter than 2,
segment 6-10 wider than long; 8 over twice as wide as long
(segments usually longer and thinner in upper elevation deep-litter
species than in lower elevation litter-soil species). Last article of
maxillary palp slightly shorter than preceding; conical, thinner.
Pronotum widest 1/3 before base, 1.4 to 1.5 times as wide as long;
hind angles acute, hind margin straight; sides arcuate; covered with
seta-bearing punctures strongly to faintly organized into striae.
Elytra fused, sides gradually tapering to apex in both sexes; external
apical angles broadly rounded; sutural angles rounded; apex
oblique; strigae distinct, oblique, composed of seta-bearing punc-
tures. Metathoracic (flight) wings reduced to tiny scales. Meso-
sternal carina low, its notch distinct. Legs not noticeably short and
compact (seemingly adapted for running and not digging); protibiae
bowed-in, mesotibiae bowed-out, metatibiae straight; comb of
spines limited to tibial apex; protibial apex oblique and rounded in
both sexes; sexual dimorphism only in protarsi, males with first 4
protarsal segments expanded and spongy-pubescent beneath.
Aedeagus curved, stout, blunt, with orifice cutting to dorsal surface
through left side; internal sac with curled, short, terminal stylet with
surface ridges on one side of tip; about 6 sensory hairs on under

358

Psyche

[December

Figures 1-3. SEM photomicrographs of Adelopsis mitchellensis { Hatch). 1. Lat-
eral view, left proleg removed. 2. Dorsal view. 3. Lateral view of head showing eye
reduced to collection of about 20 poorly defined facets.

1978] Peck — Adelopsis in the Southern Appalachians 359

surface of each side of tip. Parameres fused to aedeagus at base;
with three terminal hairs. Spiculum gastrale short, thick, less than !4
its length projecting beyond anterior end of genital plates. Sperma-
theca thin and curved; anterior recurved and often with flattened
crest; posterior end often laterally curved-back on itself.

Bionomics. The known United States species are all inhabitants
of moist moss, litter, and soil of forests in the Appalachian
Mountain Chain, south of the limits of glaciation, from the
lowlands inside the Fall-line to the summits of the highest moun-
tains. They are occasionally found in soil-filled rock crevices or
under large rocks deeply embedded in forest soils, but are more
usually captured by sifting litter and moss and by extracting them
with Tulgren-Berlese funnels. The litter at the sides of rotting logs
seems to be a favored habitat. They may be locally abundant in
association with decomposing fleshy fungi or material richly im-
pregnated with fungal hyphae, but some may be taken at dung or
carrion baits in forests, or in caves. The frequency with which they
have been collected in caves reflects only that this is a way by which
a collector can gain easy access to the faunas of deep soils. In caves,
the beetles are found near cave entrances only and not in the deep
regions of caves. The beetles are not morphologically adapted for
caves as such. This is evident when they are compared to cave-
evolved species of Ptomaphagus (Peck, 1973), so they should be
termed edaphophiles (or endogeans or edaphobites), rather than
troglophiles, or troglobites. In litter, they seem to be more fre-
quently encountered in the springtime and early summer, probably
because they are more commonly present in the upper layers of
litter and soil which are cool and moist at these seasons. They
probably retreat downwards with the increasing warmth and dry-
ness of summer. Records do not indicate it, but I think they would
be active and accessible to the collector in the late fall and at certain
times of the winter as well.

Life cycle characteristics have been determined only from speci-
mens captured on several occasions in Morrison’s Cave, Dade
County, Georgia, and kept in laboratory culture at 15°C. The
techniques are those used in culturing Ptomaphagus beetles (Peck,
1973, 1975). Eggs are laid singly by the females on the soil surface of
the culture vessel. These hatch in a mean time of 12 days (range
9-15, n = 7). There seem to be three larval instars, and these feed for
about 20 days before constructing a mud igloo or pre-pupation cell,

360

Psyche

[December

in which the larvae spend a mean of 14 days (range 12-16, n = 7)
before pupating. The total larval duration then has a mean time of
34 days (range 23-40, n = 7). The pupal stage lasts for a mean of 20
days (range 14-24, n = 6), and pupal darkening is evident for the last
two days before eclosion. The newly emerged adult remains in the
pupal chamber for 5 days (n = 3) before emerging. The cycle is
similar to that of several litter and cave species of Ptomaphagus at
the same temperature (Peck, 1973, 1975, and unpubl.). Total
longevity of adults is unknown. Adults of unknown age collected in
April lived in culture for up to 10 months. Oviposition frequency
could be determined for only one paired female which laid 9 eggs in
30 days at intervals of from 1 to 10 days.

The larvae are very similar to those of Ptomaphagus and I am
unable to confidently distinguish the two. There is no strong
evidence for seasonality in reproduction. In caves (and deep soil?) it
may occur whenever moisture and food conditions are suitable and
this probably holds for forest litter situations with cool-moist
seasons being better than warm-dry ones.

Systematics

Methods and Materials. Specimens were borrowed from the
following collections and curators: American Museum of Natural
History (AMNH), Lee Herman; Field Museum of Natural History
(FMNH), Henry Dybas; Illinois Natural History Survey (INHS),
Milton W. Sanderson; United States National Museum (USNM),
John Kingsolver; Museum of Natural History, University of Ala-
bama (UANH), Herbert Boschung; Museum of Comparative Zool-
ogy (MCZ), John Lawrence and Alfred Newton. Specimens not
attributed to these collections are in that of the author. Types are
deposited in the Canadian National Collection (CNC) unless other-
wise indicated. Initials of collectors who frequently found material
are as follows: SBP, Stewart B. Peck, often helped by Alan Fiske,
James Peck, and Jarmila Peck; WBJ, the late Walter B. Jones, often
helped by A. Flannagan, J. M. Valentine, and others; HRS,
Harrison R. Steeves, Jr., often helped by J. Patrick. WRS, Walter
R. Suter. All HRS and WRS material is in the FMNH.

My collecting methods are described in sufficient detail elsewhere
(1973, 1977, and Newton and Peck, 1975) and need not be repeated.
This is also true for the methodology of specimen preparation, with
one major exception. A scanning electron microscope (SEM) was

1978] Peck — Adelopsis in the Southern Appalachians 361

found to be of great help in viewing and understanding the three-
dimensional structure of the tips of the minute aedeagi. With this
understanding, they can be more easily interpreted in the usual slide
or glycerine jelly mounts and preparations. SEM photomicrographs
were used to make the drawings of the aedeagal tips. A set of three
views is usually necessary to interpret and understand the geometry
of the aedeagal tip. Although a set of about 6 hairs occurs on each
side of the undersurface of the aedeagal tip in all species, these have
little value in helping to characterize the species. These hairs have
been shown on only a few drawings. Internal sacs were fully seen
only in polyvinyl-lactophenol slide mounts.

Variation. Little variation is evident in these species, and little is
known of variation within the other species of the genus. Variation
over the geographic range of a species is known and illustrated only
for A. simoni (Portevin), known from Brazil, Venezuela and
Mexico (Szymczakowski, 1968) and in A. brunneus Jeannel, from
cave and forest sites from Columbia through Venezuela to Trinidad
(Szymczakowski, 1975). Apparent variation in spermathecae is
partly due to the difficulty in preparing these fragile structures.
When material is limited and the normal spermathecal shape is not
known with certainty, several may be illustrated.

Individual diagnoses are not given in the following species
accounts. In all cases it would indicate that the species are character-
ized by the combination of characters of their geographic ranges,
and of the aedeagal tips, and possibly of the spermathecae.

The gender of the genus is treated as feminine, following the use
of its author.

The mitchellensis species group

This group is probably not a closely related assemblage, and is
used as a grouping of convenience. Each of the five species is clearly
defined, and each may be as phyletically old as the cluster of 1 1
species placed in the following appalachiana species group.

1. Adelopsis mitchellensis (Hatch)

Figs. 1-5, 50

Adelops mitchellensis Hatch, 1933: 208. Synonymy given in Peck 1973: 55.

Material seen. North Carolina. Yancey County. Mt. Mitchell,
no other data, holotype male, allotype female, and paratype male
50133, USNM. Mt. Mitchell 4500-6000' (=1475-1967 m); 17-24. VI.

362

Psyche

[December

1939, E. D. Quirsfeld, 1 male, MCZ. Mt. Mitchell, 6400' (2098 m),
3-10. IV. 1967, S. Peck, carrion trap 214 in summit hemlock and
moss forest, 1 female. Black Mountains, no other data, 1 female,
USNM; 1 female, FMNH. Black Mts., 8.VIII.1911, 2 females,
AMNH.

Description. Aedeagus (figs. 4-5) with dorsal section of tip
flattened and projecting posteriorly over orifice on upper left side.
Ventral section broad with central ogival point. Spermatheca
fishhook-like (fig. 50) with simple posterior portion, and prominent
crest on anterior end.

Distribution. The species is probably limited to the Black
Mountains of Yancey County, containing Mt. Mitchell, the highest
point in the eastern United States (2191 m). This is the only locality
known to have two species of Adelopsis, for here A. mitchellensis is
sympatric with A . alta. Some early collection records confused these
two.

Notes. The species is probably most common in the upper
elevation forests of spruce-fir or birch-maple. Adults have been
collected in April, July, August, and September.

2. Adelopsis steevesi n. sp.

Figs. 6, 7

Holotype male and allotype female in CNC. Type data: North
Carolina. Macon County. Norton, Coweeta Hydrological Labora-
tory, 24.X. 1965, 4,000' (13 11 m), log-litter, HRS. Paratypes: 17 with
same data; 12 with same data but 22. V. 1965, 4100' (1344 m) rot
wood debris.

Description. Dorsal section of aedeagus tip (figs. 6-7) incised,
upraised as knob on left, and as flattened vertical blade on right.
Ventral section deeply emarginate, nearly same length as dorsal
section. Spermatheca similar to that of A. joanna, A. alta, A.
fumosa, A. richlaYidensis, and A. suteri.

Etymology. Named for Mr. Harrison R. Steeves, Jr., of Bir-
mingham, Alabama, in recognition of his extensive field collecting
of litter beetles in the southeastern United States (collection de-
posited in FMNH).

Distribution. The species should be expected in other sites in the
vicinity of Highlands, North Carolina.

1978] Peck — Adelopsis in the Southern Appalachians 363

3. Adelopsis alleghenyensis n. sp.

Figs. 8-10, 51, 52

Holotype male and allotype female in CNC. Type data: West
Virginia. Pendleton County. 5 mi (8 km) S Witmer, 3000' (915 m),
16. VII. 1971, SBP, litter Ber. 217. Paratypes: 11 with same data;
Spruce Knob, 3500' (1 148 m), 8. VI. 1967, SBP. Ber. 58, 1 male and 1
female; Pocahontas County, Hills Creek Falls, 19. VI. 1971, W.
Shear, Berlese, 2 males and 2 females.

Description. Dorsal section of aedeagal tip low and simple (figs.
8-10), with recurved flange on left; ventral section broadly pointed,
with tip downcurved, extending far beyond dorsal section. Sperma-
theca (figs. 51-52) thick and gently curved, anterior crest large.

Etymology. The name refers to the northwestern flank of the
Appalachians called the Allegheny Mountains, lying along the
border of Virginia and West Virginia and extending into Pennsyl-
vania.

Distribution. The species probably has a wider range than the
known 60 mile (100 km) long NE-SW line along the Allegheny
Mountains of West Virginia.

4. Adelopsis suteri n. sp.

Figs. 11, 12, 53, 54

Holotype male and allotype female in CNC. Type data. Georgia.
Rabun County. Mountain City, Black Rock Mountain State Park,
15. VI. 1973, W. R. Suter, litter at log under Rhododendron. Para-
types: 1 with same data; Black Rock Mountain State Park, 3000'
(915 m), 21. VII. 1967, 2 males. North Carolina. Macon County. 5 mi
(8 km) NE Highlands, 27. X. 1969, W. Shear, Rhododendron litter, 1
male. Jackson County, 7 mi (1 1 km) S Cashiers, 1 1. VI. 1973, White-
water Falls, elm-maple pseudofork, WRS, 1 female.

Description. Dorsal section of aedeagal tip (figs. 11-12) inflated,
upturned, with broad and shallow depression in middle, equal in
length to uniformly emarginate ventral section. Spermatheca (figs.
53-54) with sharp bend in posterior end, which projects strongly
above plane of central curved section, anterior crest high.

Etymology. Named for Dr. Walter R. Suter of Carthage Col-
lege, Kenosha, Wisconsin, in recognition of his extensive collecting

364

Psyche

[December

1978] Peck — Adelopsis in the Southern Appalachians 365

of litter beetles in the southeastern United States (collections
deposited in FMNH).

Distribution. The species is to be expected in other localities in
the Coweea Mountains in the vicinity of Highlands, North Caro-
lina, which has been found to be a region rich in species of ground
dwelling beetles.

5. Adelopsis richlandensis n. sp.

Figs. 13, 14, 55, 56

Holotype male and allotype female in CNC. Type data. North
Carolina. Haywood County. Richland Balsam, 6000' (1967 m),
7-26. VIII. 1965, S. & J. Peck, carrion trap. Paratypes; three females
with same data; and 1 female, Haywood-Jackson Counties, Rine-
hart Knob, 6000' (1967 m), 1. VIII. 1960, T. C. Barr.

Description. Dorsal section of aedeagal tip (figs. 13-14) uni-
formly rounded, but with slight emargination; sloping downwards
from high crest on right to low and flat left side, ventral section
uniformly rounded and projecting beyond dorsal section. Sperma-
theca as in figs. 55-56, characteristic form not known with certainty.

Etymology. The name refers to Richland Balsam, the type
locality, and highest point on the Blue Ridge Parkway.

Distribution. There seems little reason to expect only this species
on Richland Balsam. Mt. Pisgah, with A. pisgahensis, is on the
same mountain ridge and there are no intervening lowland barriers
to dispersal of these flightless beetles. Soco Gap and Balsam Gap to
the northwest, each above 3000' (1000 m) separate Richland Balsam
from the Plott Balsams and Balsam Mountains, but these also seem
inadequate barriers to isolate Richland Balsam from species in-
habiting these other mountain regions. Intensive collecting should
resolve the questions of how these species are distributed with
regards to each other.

Figs. 4-18. Structures of Adelopsis. 4. Left lateral view of entire aedeagus of
A. mitchellensis, with internal sac (IS), ventral blade of tegmen (VBT), and dorsal
(D) and ventral (V) sections of aedeagal tip. 5. Dorsal view of aedeagal tip of
A. mitchellensis indicating right (R) and left (L) sides, and dorsal (D) and ventral
(V) sections of the tip. 6-7. Dorsal and lateral views, aedeagal tip, A. steevesi.
8-10. Dorsal, lateral, and posterior view, aedeagus, A. alleghenyensis. 11-12. Aedea-
gal tip. A. suteri. 13-14. A. richlandensis aedeagal tip. 15. Internal sac A. appalach-
iana. 16-18. A. appalachiana, note that parameres and hairs on ventral surface of
ventral section of aedeagus are not shown in most drawings.

366

Psyche

[December

The appalachiana species group

This group is seen as a naturally based and closely related phyletic
unit. It is based on A. appalachiana, the most widespread species in
both lowland and montane regions. Progressive modification of the
relatively simple aedeagus (assumed to be generalized and pleiso-
typic) in the nominate species could readily produce the other
aedeagal types known in the group.

6. Adelopsis appalachiana n. sp.

Figs. 15-18, 57-60

Holotype male and allotype female in CNC. Type data: Alabama.
DeKalb County. Fort Payne, Manitou Cave, 1937, W. B. Jones,
trap. Paratypes: 49 with same data; 8 with same data except 1939;
and the following: Alabama. DeKalb County. Kelly Girls Cave, 1
mi (1.6 km) SE Collinsville, 25. VIII. 1965, SBP, 2; Cook Cave (Ala
253), 13. VIII. 1958, WBJ, 2; Lykes Cave, 27.VIII.1965, SBP, fungus
on entrance talus, 22; Cherokee Cave, 4.5 mi (6.5 km) NE Ft. Payne,
15. VIII. 1967, SBP, 1. Blount County. Blount Springs, 19. III. 1966,
SBP, litter in limestone crevice, 1; 7 mi (1 1 km) S Cleveland, outside
Horseshoe-Crump Cave, 28.VI.1967, SBP, litter Ber. 70, 2; outside
Bangor Cave, 19. III. 1966, SBP, rotten tree roots, 4; outside Bangor
Cave, 5. IV. 1967, SBP, 1; inside Bangor Cave, 20.XII.1965, SBP, 1;
near Inland Lake, 28. IX. 1958, HRS, forest floor debris, 4. Shelby
County. Helena, 3.1.1959, HRS, treehole, 1. Jefferson County. Near
Purdy Cave #1, 13. IX. 1958, HRS, forest debris in rock crevice, 1.
Morgan County. 3lA mi (6 km) SE Fayette, 21. V. 1972, SBP, Ber
241, litter outside cave entrance, 14. Georgia. Dade County. Mor-
risons Cave 13. VII. 1967, SBP, 6; 16.IX.1968, SBP, 1; 16-23. VI.
1972, SBP, 3; 4 mi (6.5 km) NE Rising Fawn, 15.VII.1967, SBP,
forest litter Ber 78, 4; Johnson Crook Cave #2, 4 mi (6.5 km) NE
Rising Fawn, 14. VII. 1967, SBP, 2 in cave entrance; outside Johnson
Crook Cave #2, 14. VII. 1967, SBP, 6 in litter.

Tentatively associated material. Alabama. Etowah County.
Wright Cave, 11. VII. 1958, WBJ, 1 male, 1 female. Morgan County,
Inge Cave, 2. V. 1959, T. C. Barr and HRS, under debris, 2 males.
Cleburne County, Cheaha Mountain State Park, 330 m, 13. VI. 1967,
SBP, litter Ber 59, 2. Marshall County. Guntersville, near Griffith
Cave, 30. IV. 1960, HRS, 1 female; Kelly Ridge Cave, near Warren-
town, 23. VIII. 1958, HRS, 1 female. Georgia. Walker County.

1978] Peck — Adelopsis in the Southern Appalachians 367

McLemore Cove, 9 mi (15 km) SW LaFayette, 14. VII. 1967, forest
log litter Ber 79, SBP, 3. Dawson County. Mt. Oglethorpe, 2000'
(610 m), 20. VIII. 1967, SBP, litter Ber 80, 1 damaged male.

Description. Dorsal section of aedeagus tip (figs. 16-18) regu-
larly curved, unadorned, shortest on left side and longest on right;
ventral section regularly curved with downturned tip; posterior view
with dorsal section forming a uniformly arched genital orifice, with
a uniform border; setae on ventral surface lying in groove rather
than separated depressions. Internal sac thick, not narrowed into
curved tube at apex (fig. 15). Spermatheca (figs. 57-60) smoothly
curved, anterior part broadened and without crest, posterior part
recurved off structural plane.

Variation. The Morgan County, Alabama, sample has a lower
arch, with a flattened middle to the genital orifice when seen from
behind. This opening is most highly arched in the Cheaha Moun-
tain, Alabama, male which also has a differing left margin to the
ventral section.

Etymology. The name refers to the Appalachian Mountains,
around the southern end of which this species is distributed.

Distribution. This is the most widely distributed species, ranging
from north-central Alabama, south of the Tennessee River, east-
wards into northwestern Georgia.

Field Notes. This is the most frequently collected species because
it has a wide range, and occurs in a region which has received much
collecting attention. It has been found in forest litter and in the
entrance zone of caves, in the months of January, March, May,
June, July, August and September. It is probably active throughout
the year.

7. Adelopsis jonesi n. sp.

Figs. 19-21

Holotype male in CNC. Type data. Alabama. Colbert County.
Wolf Den Cave (Ala. 83), near Maude, 25. IX. 1940, W. B. Jones.

Tentatively associated material. Alabama. Colbert County.
McClusky Cave, near Maude, 15. IX. 1940, WBJ, 1 male missing
aedeagus; Gist Cave, 13.V.1940, WBJ, 2 females. Franklin County,
The Dismals, 30. VIII. 1958, HRS, forest floor debris, 1 female. Hale
County. Moundville State Park, 7. VII. 1967, SBP, log litter Ber 75,
3 females. Tuscaloosa County. No other locality, VI, Alabama
Museum Expedition, 1 female (UANH).

368

Psyche

[December

Figs. 19-33. Views of aedeagal tips of Adelopsis. 19-21. A. jonesi. 22-24. A.
bedfordensis. 25-27. A. cumberlanda. 28-30. A. scottsboroensis. 31-33. A. nashvil-
lensis. Scale of aedeagal tips approximately equal.

1978] Peck — Ade/opsis in the Southern Appalachians 369

Description. Dorsal section of aedeagal tip with highly raised
edge forming large arch over genital orifice (figs. 19-21); ventral
section smoothly rounded. Spermatheca of tentatively associated
females indistinguishable from those of A. appalaehiana.

Relationships. This species is easily derived in aedeagal type
from A. appalaehiana by an increased arching of the dorsal section
of the genital orifice. It probably represents an isolated and now
differentiated peripheral population of the latter.

Etymology. The species is named for the late Dr. Walter B.
Jones, of Huntsville and Tuscaloosa, Alabama, who, as former
State Geologist and Director of the Alabama State Museum of
Natural History, did so much to promote studies of the natural
history of the southeast.

8. Adelopsis bedfordensis n. sp.

Figs. 22-24

Holotype male in FMNH. Type data. Tennessee. Bedford County.
Campbell Cave, 17.1.1960, H. R. Steeves, Jr., under rock in dark
zone.

Description. Aedeagal tip (figs. 22-24) with uniformly arching
dorsal section without upturned flange in dorsal and lateral views.
Genital orifice moderately arched in posterior view and asym-
metrical. Internal sac long, thin, and curved.

The aedeagus is difficult to distinguish from that of A. appa-
lachiana, except for the striking differences in their internal sacs.

Etymology. The name refers to the county in which the type
was collected (and to which it may be approximately restricted in
distribution), located in the eastern Highland Rim of south-central
Tennessee.

9. Adelopsis cumberlanda n. sp.

Figs. 25-27, 61

Holotype male and allotype female in CNC. Type data: Alabama,
Jackson County. 5 mi (8 km) N Garth, 19. V. 1972, S. Peck, leaf litter
Ber 239. Paratypes: 65 with same data; 2 with same date but in dung
baited pit traps; and the following: Alabama. Jackson County.
Paint Rock, Nat Mountain, 15. III. 1966, SBP, 3 under log; Keel
Cave, Sharp Mt., 11. VI. 1940, WBJ, 1; Clemons Cave, Sharp Mt.,

370

Psyche

[December

9. VI. 1940, WBJ, 1; Rousseau Cave, July, 1968, R. Graham, 1;
Rousseau Cave, 12. VII. 1973, SBP, 4 baited near entrance; Paint
Rock, near Crossing (Stewart) Cave, 5. VII. 1958, HRS, 2 in rock
crevice debris; 5 mi (8 km) NW Princeton, 19. V. 1972, SBP, forest
litter Ber 240, 1. Madison County. Monte Sano State Park,
17. V. 1972, SBP, log litter Ber 238, 5; Monte Sano, Huntsville, 1936,
R. Hopping, 2 (U ANH). Monte Sano, VI. 1937, Ala. Mus. Exped., 1
(UANH); Tumbling Gap (NE ridge of Monte Sano), June 3-8,
1911, H. P. Loding, 2 (UANH); Sharps Cove, 5. VII.1965, SBP, leaf
Ber., 1. Marshall County. Grant, Guffey Cave, 25. V. 1958, 1 in floor
debris in entrance light zone, HRS; Grant, near River Cave,
25. V. 1958, 4 under debris at entrance light zone, HRS; Creek Cave,

25. V. 1958, HRS, 1 in cave debris; 1 mi N Guntersville Dam,

26. VI. 1967, litter Ber 68, SBP; Hampton Cave, Guntersville, 1 1 .VII.
1973, SBP, 1.

Description. Dorsal section of aedeagal tip short (figs. 25-27)
thickened and upcurved, especially at right side where small broad
tooth exists (best seen from right side); ventral section projecting,
tip truncate; posterior view with uniformly broad arch over orifice.
Internal sac thick and slightly curved. Spermatheca similar to that
in A. seottsboroensis (fig. 62) but with more bend in posterior part
when seen from right, and with a lower crest on the anterior part.

Etymology. The name refers to the Cumberland Plateau, flank-
ing the NW face of the Appalachian Mountains.

Distribution. The species range is in the forested slopes and
valleys of the southern end of the Cumberland Plateau, in north-
eastern Alabama, north and west of the Tennessee River. This river
is a natural boundary of the species range, separating it from that of
A. appalachiana south of the river.

Notes. The species has been found in moist forest litter and cave
entrances in the months of March, April, May, June, July and
August.

Relationships. The species is derivable from A. appalachiana by
an increased arching of the genital orifice, a thickening of the dorsal
orifice lip, and an increase in the length of the ventral section.

10. Adelopsis seottsboroensis n. sp.

Figs. 28-30, 62, 63

Holotype male and allotype female in CNC. Type data: Alabama.
Jackson County. 10 mi (16 km) N Scottsboro, 16. V. 1972, SBP,

1978] Peck — Adelopsis in the Southern Appalachians 371

forest litter Ber 237 (from limestone crevices near Tumbling Rock
Cave). Paratypes: 6 with same data.

Description. Dorsal section of aedeagal tip short (figs. 28-30)
and upturned, with right side free and reflexed; in posterior view
right margin of arch above orifice projecting and deflexed to right.
Spermatheca thin and uniformly curved as in figs. 62-63.

Etymology. Named for the city of Scottsboro, which sits at the
base of a spur of the Cumberland Plateau, near the margin of the
drainage basin probably containing the species range.

Distribution. The species should be expected to occur in addi-
tional sites below the Plateau escarpment in the headwaters of Mud
and Coon Creeks, Jackson County. Some beetles and other inverte-
brates are known to be endemic to this area, so some general
mechanism may have worked to isolate them in this section of the
Cumberland Plateau escarpment. How the species range overlaps or
intergrades with that of A. cumberlanda (occurring a short distance
to the west and south along the same general face of the Plateau,
and in the drainage of the Paint Rock River) is not known.

Relationships. The species can be considered a local isolate of A.
cumberlanda, from which the aedeagal tip is easily derived.

11. Adelopsis nashvillensis n. sp.

Figs. 31-33

Holotype male in USNM. Type data: Tennessee. Davidson
County near Nashville, 3 November 1937, S. F. Austin, “elm-maple,
562”. Tentatively associated material; one female, Tennessee, War-
ren County, Hubbard Cave at Irving College, 16. X. 1948, W. B.
Jones and J. M. Valentine.

Description. Aedeagus with dorsal section of tip (figs. 31-33)
upturned, and recurved on right side; in posterior view genital
orifice deflected up and to the right.

Etymology. The name refers to the city near where the species
was collected.

Relationships. The species seems to be closest to A. cumber-
landa, from which the aedeagal tip can be easily derived.

12. Adelopsis fumosa n. sp.

Figs. 34-37, 64, 65

Holotype male and allotype female in FMNH. Type data.
Tennessee. Sevier County. Great Smoky Mountains National Park,

372

Psyche

[December

Figs. 34-49. AedeagaFstructures of Aclelopsis. 34-36. A. fumosa. 37. Internal
sac, A. fumosa. 38 40. A. aha. 41- 43. A. Joanna. 44 46. A. pisgahensis. 47-49. A.
orichalcum.

1978] Peek — Adelopsis in the Southern Appalachians 373

16. IX. 1953, 2.5 road miles (4 km) above Chimney Campground,
3600 ft. alt. (1180 m), H. S. Dybas, leaf litter Berlese. Paratypes: 7
with same data; and the following from Great Smoky Mountains
National Park. Cherokee Orchard, 2-3000 ft (610-910 m), 15. IX.
1953, H. S. Dybas, 2 in Tilia treehole; 3 in forest floor litter; 4 in
flood debris (all in FMNH). State road to Newfound Gap, 3500 ft
(1147 m), 16.IX.1953, H. S. Dybas, litter, 8 (FMNH). Chimney
Campground, FIX. 1948, Ross and Stannard, 3, (INHS). Gatlin-
burg, 14.IX.1948, Quirsfeld, 5 (MCZ). Clingman’s Dome Summit,
6640 ft (2177 m), 6-26. VIII. 1965, SBP, carrion trap; 8 mi (13 km) S
Gatlinburg, 3100 ft, 17. V. 1972, A. F. Newton, 2 in carrion trap;
Buckeye Nature trail, 10. VIII. 1974, J. L. Bengtson, maple litter.
Georgia. Walker County. Bible Spring Cave, 2 mi (3 km) NE
Lookout, 20. VI. 1967, SBP), male.

Description. Dorsal section of aedeagal tip (figs. 34-36) uni-
formly rounded and arched; ventral section on right enlarged to
enclose right dorsal section. Internal sac as in fig. 37. Spermatheca
(figs. 64-65) with prominent crest.

Variation. The single male from the summit of Clingman’s
Dome seems to show differences that, with adequate material, could
prove to be specifically distinct. The aedeagal tip seems more
bilobed on the dorsal section and with a more prominent recurved
flange on the right. The internal sac seems more elongate and less
curved in its distal half.

Etymology. The name is derived from the Latin for smoke,
referring to the species range in the Great Smoky Mountains,
However, the Georgia population is a disjunct some 200 km to the
southwest.

Relationships. The aedeagal tip shows the closest similarity to,
and is most easily derived from that of A. appalachiana.

13. Adelopsis alta n. sp.

Figs. 38-40, 66

Holotype male in USNM. Type data: North Carolina. Yancey
County. Mt. Mitchell, 6500 ft (2131 m), 25. VI. 1937, E. Shoemaker.
Paratypes: Mt. Mitchell, 4-6000 ft. (1311-1967 m) 17-24. VI. 1939,
Quirsfeld, 4 (INHS); Mt. Mitchell, 12. VI. 1973, litter under Rho-
dendron, WRS, 1.

374

Psyche

[December

Description. Dorsal section of aedeagal tip (figs. 38-40) higher
and thicker on left, right side with downturned process curving into
genital orifice; ventral section thickened and upturned at base on
both right and left sides. Spermatheca (fig. 66) with central piece
curved inwards like grip of an archer’s bow.

Etymology. The name is the Latin adjective for “high” and refers
to the species’ high elevation habitats in the Black Mountains and
Mt. Mitchell.

Notes. This species is sympatric with A. mitchellensis, with
which it has been confused, but to which it is not at all closely
related judging from the dissimilarity of the aedeagal tips.

Relationships. The species is most easily derived from a gen-
eralized ancestor like A. appalachiana by modification of the
aedeagal tip through stages similar to A. alta, A. cumberlanda, and
A. nashvillensis, though these may not be direct ancestors.

14. Adelopsis joanna n. sp.

Figs. 41-43, 67, 68

Holotype male and allotype female in CNC. Type data: North
Carolina. Cherokee County. Joanna Bald, near Andrews, 4700 ft
(1540 m), 26. VII. 1967, S. B. Peck, litter Ber. 96. One paratype
female with same data.

Description. Dorsal section of aedeagal tip (figs. 41-43) not
smoothly curved, but more angular and forming offset arch over
genital orifice; ventral section upcurved on left but more so on right,
forming prominent notch between it and the dorsal section. Sper-
matheca thin, irregularly curved (figs. 67, 68).

Etymology. The name refers to the type locality, and is used as a
feminine noun in apposition.

Relationships. The species seems to be derived from a cumber-
landa- like ancestor through a form like that in A. alta, and may be
an isolate of the ancestor of this species.

Figs. 50-68. Spermathecae of Adelopsis. 50. A. mitchellensis indicating anterior
(A), posterior (P), right (R), and left (L) sides as oriented in the beetle. 51-52. A.
alleghenyensis. 53-54. A. suteri. 55-56. A. richlandensis. 57-58. A. appalachiana,
Morrison Cave, Georgia. 59-60. A. appalachiana, outside Bangor Cave, Alabama.
61. A. cumberlanda, paratype. 62-63. A. scottsboroensis. 64. A. fumosa, 8 mi S
Gatlinburg, Tennessee. 65. A. fumosa, Clingman’s Dome, Tennessee. 66. A. alta.
67-68. A. joanna. All drawn to same scale.

1978] Peek — Adelopsis in the Southern Appalachians 375

65

68

376

Psyche

[December

15. Adelopsis pisgahensis n. sp.

Figs. 44-46

Holotype male, allotype female, and paratype male in MCZ.
Type data: North Carolina. Haywood-Transylvania County line,
Mt. Pisgah, about 1475 m (on Blue Ridge Parkway), 1 1-12. IX. 1934,
Quirsfeld.

Description. Aedeagal tip (figs. 44-46) with dorsal section
broadly swollen on left and with sharp point on right; ventral
section low and regular on left but on right broadly expanded and
upraised along notch between it and dorsal section.

Etymology. The name refers to the type locality. Mt. Pisgah is a
section of the mountain ridge continuous with Richland Balsam,
inhabited by A. richlandensis. There is no evident barrier to
dispersal between the two and they must be assumed to be
parapatric or sympatric.

Relationship. The species has a highly derived type of aedeagal
tip and may be descendant from an ancestor similar to A. alta.

16. Adelopsis orichalcum n. sp.

Figs. 47-49

Holotype male in CNC. Type data: Georgia. Union County.
Brasstown Bald, el. 2415 ft (790 m), 24. X. 1966, H. R. Steeves, forest
debris.

Description. Dorsal section of aedeagal tip (figs. 47-49) uni-
formly thicker from left to right, ending in upcurved spur; ventral
section upcurved on left, depressed and emarginate in middle, and
more upcurved on right.

Etymology. The name is Latin for brass, is used as a noun in
apposition, and refers to Brasstown Bald, the highest point in
Georgia, in the vicinity of which the species is probably restricted.

Relationships. The species is derivable from a generalized stock
like A. appalaehiana through intermediates that were probably also
ancestral to A. pisgahensis, A. joanna, and A. alta.

Evolutionary and Distributional History

Jeannel (1936, 1964) considered the evolutionary center of the
genus (and the tribe) to be in Colombia and Venezuela. Species here
have an aedeagus with a copulatory orifice displaced onto a nearly

1978] Peek — Adelopsis in the Southern Appalachians 377

sagital plane. This condition was judged to be a form from which all
other aedeagal tips could be derived. He also saw this genus as
ancestral to Ptomaphagus, which developed a more “successful and
harmonious” and less variable aedeagal type, and then spread
through North America, and across the proto- Atlantic to Europe in
the Eocene. He also saw an earlier connection from South America
in the Jurassic to southeast Asia, where another derivative genus,
Ptomaphaginus, now occurs. Szymczakowski (1964, 1969) agrees
with the idea of an American evolutionary origin, but sees little
evidence to support arguments for specific despersal routes. The
survival of Proptomaphaginus in relictual species in the Caribbean
and Mexico supports the idea of an American origin for the stock
that became Ptomaphaginus in the Oriental Region.

I do not yet have a suitable overview of the genus as a whole to
elaborate on or offer alternative hypotheses to those of Jeannel and
Szymczakowski. This is best postponed until after my now massive
tropical American collections are studied. However, some ideas can
be proposed for the collection of species limited to the southeastern
United States.

The genus probably originated in tropical America in the late
Mesozoic. Whether this was in what we now know as South,
Central or North America probably cannot be known with certainty.
However, the greatest present generic diversity in the tribe is now
known to be in or adjacent to Mexico. This area (and Central
America) has been an important evolutionary center in its own
right. Savage (1966) suggested that broad, terrestrial, forested
connections united these three regions in the Paleocene and per-
mitted north-south faunal movements, so that mesic tropical cli-
mates and forests were continuous through South America up to
what is now the central United States. The patterns of distribution
of forest litter reptiles and amphibians may be informative and
related because they occupy environments similar to Catopinae,
though their dispersal abilities are probably very different. How-
ever, Savage (1974) has reconsidered the evidence and now finds less
substantial support for such broad connections, and now thinks
there was a significant water gap separating South America from
“nuclear Central America” and North America from the Cretaceous
up to the Pliocene. This is more consistent with other current ideas
of plate tectonic biogeography (Raven and Axelrod, 1975). This
gap, however, could have been crossed by island hopping and waif

378

Psyche

[December

dispersal, of which winged Adelopsis (and Ptomaphagus) should
have been capable.

These considerations then suggest an origin of Adelopsis in
“tropical” North or Central America perhaps in the late Cretaceous
or early Tertiary, and a movement through forests in the early
Tertiary into the Appalachian Region. The continuous distribution
of the genus was then broken with the developing xeric climates of
the Miocene, forming an effective non-forested barrier across what
is now northern Mexico (Axelrod, 1960, Graham, 1973). This
disjunction is observed in many groups (see Rosen, 1978). Since this
disjunction the assumed large-eyed and winged ancestral stocks
have vanished from what became the United States, leaving only the
small-eyed and wingless soil species of the Appalachians. Past
workers have suggested that the mesic forests of the Appalachians
and Mexico were rejoined in Pleistocene glacials, but most recent
analyses do not tend to support this (Graham, 1973; Martin and
Harrell, 1957; Rosen, 1978).

The comparative uniformity of overall morphology and aedeagal
types in the Appalachian species suggests that a single ancestral
species became an inhabitant of soil and deep litter. Subsequent
(late Tertiary?) isolations perhaps produced the set of less-similar
species in the mitchellensis species group. Differentiation of these
species may have been reinforced by Pleistocene climatic fluctua-
tions and distributional displacements. One of these species may
have become something like A. appalachiana and it experienced
speciation by range expansion, contraction, and population isola-
tion in response to Pleistocene climatic fluctuations. The more cool
and wet conditions accompanying glacials (Watts, 1970, 1975;
Whitehead, 1965) would allow their spread through the lowlands.
Warmer and drier conditions of interglacials would separate popu-
lations of these seemingly poorly mobile beetles as they retreated to
favorable and more isolated sites at higher elevations or in mesic
forest refugia of canyons, gorges, and protected spots below topo-
graphic escarpments. By this process, an A. appalachiana-\\ke
ancestor differentiated into the species of this more closely knit
group.

The production of 16 species from one ancestor seems only a
modest amount of differentiation if the southeastern Adelopsis have
truly been isolated from their Neotropical congeners for some 35-40
million years, especially when compared to some examples of

1978] Peck — Adelopsis in the Southern Appalachians 379

“explosive” species production on islands in 10 million years or less.
However, the fossil record shows that other groups with the same
Appalachian-Mexican disjunction have undergone very little dif-
ferentation in the same time, especially in certain tree genera
(Axelrod, 1960, p. 269; Rosen, 1978). It should also be noted that
these papers deal more with examples that seem to be northern in
origin and which have moved into Mexico and southwards rather
than the opposite (Graham, 1973).

It might at first seem puzzling that a Neotropical genus, charac-
teristic of lowland mesic tropical forests should have species isolated
on the highest peaks of the southern Appalachians. However, it is
not difficult to see the origin of the situation. When they were first
colonized the Appalachians seem to have had a tropical or sub-
tropical climate, and to have had a more gentle relief, sometimes
called the “Schooley Peneplain”. Their present relief is due to later
Tertiary uplift and erosional rejuvenation, a process to which the
beetles could undoubtedly adapt as their populations either were
slowly elevated, or slowly moved into higher sites.

The broad patterns of species distribution seem similar to those of
other flightless beetles inhabiting deep litter and/or soil in the
Appalachians (see Darlington, 1943). Close comparisons can be
made with the eyeless Anillinus carabids (Barr, 1969; pers. comm.),
Arianops pselaphids (Barr, 1974), and more distant ones with
Trechus carabids (Barr, 1962, 1969), and Catopocerus leiodids
(Peck, 1974). These show species to be generally confined to upland
areas or in deeper soil (collected in caves) or protected gorges in the
lowlands. Some of these groups have species which may be more
widespread in the northern end of the unglaciated Appalachians,
but most contain more species, but with not much more sympatry,
in the vicinity of the junction of North and South Carolina and
Georgia (the vicinity of Highlands, North Carolina). The mechanism
that acted to generate and then preserve the diversity near Highlands
might have worked in a relatively similar manner on these and many
other groups of forest litter-inhabiting insects and other arthropods.

Conclusion

The genus Adelopsis , otherwise Neotropical in distribution, is
found to have 16 species in the southern part of the Appalachian
region of the United States. All these species are small-eyed and

380

Psyche

[December

wingless, are inhabitants of forest soil and deep litter, and must have
limited dispersal abilities. Higher elevation Appalachian localities
seem generally to be occupied by distinct, more distantly related,
and very localized species. Lower elevation sites are more often
occupied by more similar species which are more often of a wider
distribution. This last generalization is reflected in the appalachiana
species group which may have resulted from a more recent series of
speciation events. There is not enough evidence to suggest how
ranges may have been affected and populations isolated with the
changed climatic conditions of the Pleistocene, but these are known
to have markedly lowered vegetational zones in the southern
Appalachians in the last glacial. Since most of the material reported
on in this paper was incidentally gathered by persons in persuit of
other beetles, many more species undoubtedly remain to be dis-
covered. These may possibly more than double those already
known, particularly in the poorly collected regions extending for
some 260 km (160 mi) along the Appalachian and Blue Ridge crests
between the NE corner of North Carolina, through Virginia, to
Spruce Knob, West Virginia. A more firmly based interpretation of
the evolutionary history of the genus in the southeastern United
States will be possible when these taxa and additional distributional
data become known. Because of inadequate material, this paper is
only a preliminary report. This is a reflection of the still inadequate
status of knowledge of the soil and litter faunas of North America in
general.

Acknowledgements

I am deeply appreciative of the work of Hana Kukal in making
the drawings. L. E. C. Ling, Carleton University, Department of
Biology, assisted with the SEM examination of specimens. The
museums from which I borrowed material, and their curators, as
well as private collectors, are thanked for their courtesies. Mrs.
Hazel B. Jones and the late Dr. Walter B. Jones offered many
months of hospitality in Huntsville, Alabama, in support of my field
work in Alabama and adjacent southeastern states. Field work from
1965 to 1970 was supported by NSF grants to the Evolutionary
Biology Committee, of Harvard University, and from 1971 to
present by Canadian National Research Council operating grants.
Drs. Alfred F. Newton, Henry F. Howden and John V. Matthews,
Jr. reviewed the manuscript.

1978] Peck — Adelopsis in the Southern Appalachians 381

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Axelrod, D. 1.

1960. The evolution of flowering plants, pp. 227-305, in S. Tax, (ed.), Evolu-
tion After Darwin, vol. 1, The Evolution of Life. Univ. Chicago Press.

Barr, T. C., Jr.

1962. The Genus Trechus (Coleoptera: Carabidae: Trechini) in the southern
Appalachians. Coleopterists Bulletin 16: 65-92.

1969 Evolution of the (Coleoptera) Carabidae in the southern Appalachians,
67-92, in P. Holt (ed.). The distributional history of the biota of the
southern Appalachians, part 1: Invertebrates. Research Div. Mono 1,
Virginia Polytech. Inst., Blacksburg, Va. 295 pp.

1974. The eyeless beetles of the genus Arianops Brendel (Coleoptera : Psela-
phidae). Amer. Mus. Nat. Hist. Bull., 154(1): 1-51.

Darlington, P. J., Jr.

1943. Carabidae of mountains and islands: data on the evolution of isolated
faunas, and on atrophy of wings. Ecol. Mono. 13: 37-61.

Graham, A.

1973. History of the arborescent temperate element in the northern Latin
American biota. Ch. 8, pp. 301-314. In Graham, A. (ed.). Vegetation
and vegetational history of northern Latin America. Elsevier Publ. Co.,
Amsterdam.

Hatch, M. H.

1933. Studies on the Leptodiridae (Catopidae) with descriptions of new
species. J. New York Ent. Soc. 41: 187-239.

Jeannel, R.

1936. Monographic des Catopidae. Mem. Mus. Nat. Hist. Natu., Paris, nouv.
ser. 1, 433 pp.

Martin, P. S. and B. E. Harrell.

1958. The Pleistocene history of temperate biotas in Mexico and eastern
United States. Ecology 38(3): 468-480.

Newton, Alfred F. and S. B. Peck.

1975. Baited pitfall traps for beetles. Coleop. Bull. 29: 45-46.

Peck, S. B.

1973. A systematic revision and the evolutionary biology of the Ptomaphagus
(Adelops) beetles of North America (Coleoptera : Leiodidae : Catopi-
nae), with emphasis on cave-inhabiting species. Bull. Mus. Comp. Zool.
Harvard Univ., 145(2): 29-162.

1974. The eyeless Catopocerus beetles (Leiodidae) of eastern North America.
Psyche SI: 377-394.

1975. The life cycle of a Kentucky cave beetle, Ptomaphagus hirtus (Coleop-
tera : Leiodidae : Catopinae). Int. J. Speleology 7: 7-18.

1977. The subterranean and epigean Catopinae of Mexico (Coleoptera :
Leiodidae). Ass. Mexican Cave Studies Bull. 6: 185-213.

1978. New montane Ptomaphagus beetles from New Mexico and zoogeog-
raphy of southwestern caves (Coleoptera : Leiodidae : Catopinae).
Southwest. Natur. 23: 227-238.

Portevin, G.

1907. Clavicornes nouveaux du groupe des necrophages. Ann. Soc. Ent. Fr.
74: 67-82.

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Raven, P. H. and D. I. Axelrod.

1975. History of the flora and fauna of Latin America. Amer. Sci. 63: 420-429.

Rosen, Donn E.

1978. Vicariant patterns and historical explanation in biogeography. Syst.
Zool. 27: 159-188.

Savage, Jay M.

1966. The origins and history of the Central American herpetofauna. Copeia
1966: 714-766.

1974. The Isthmian Link and the evolution of Neotropical mammals. Cont.
Sci. Los Angeles County Natur. Hist. Mus. 260. 51 pp.

SZYMCZAKOWSKI, W.

1964. Analyse systematique et zoogeographique des Catopidae (Coleoptera)
de la region orientale. Acta Zool. Cracoviensia 9(2): 55-289.

1968. Sur quelques Catopidae (Coleoptera) de la region neotropicale. Acta
Zool. Cracoviensia 13(2): 13-27.

1969. Decouverte d’un representant des Ptomaphaginini a Cuba (avec une
equisse de la systematique et la geonemie de cette tribu) (Coleoptera :
Catopidae). Acta. Zool. Cracoviensia 14(4): 87-97.

1975. Formes cavernicoles d'Adelopsis brunneus Jeann. du Venezuela et de
Hie de Trinidad (Coleoptera : Catopidae). Bol. Soc. Venezolana Espel.
6: 13-24.

Watts, W. A.

1970. The full-glacial vegetation of northwestern Georgia. Ecology 51: 17-33.

1975. Vegetation record for the last 20,000 years from a small marsh on Look-
out Mountain, northwestern Georgia. Geol. Soc. Amer. Bull. 86:
287-291.

Whitehead, D. R.

1965. Palynology and Pleistocene phytogeography of unglaciated eastern
North America, pp. 417-432. In H. E. Wright, Jr., and D. G. Frey,
(eds.). The Quaternary of the United States. Princeton Univ. Press,
Princeton, 922 pp.

EXTERNAL SEX BRAND MORPHOLOGY OF
THREE SULPHUR BUTTERFLIES
(LEPIDOPTERA: PIERID AE)*

By Richard S. Vetter
and

Ronald L. Rutowski
Department of Zoology
Arizona State University
Tempe, Arizona 85281

Introduction

Males of many pierid butterflies have specialized scales, in
patches or scattered over the wing surface, that are thought to be
involved in the dissemination of a chemical signal (Chapman, 1971).
Recent experiments with two sulphurs, Eurema lisa Boisduval and
Leconte and Colias philodice Latreille, have borne out this assump-
tion (Rutowski 1977, 1979). In these species, the male has a patch of
cells and scales in the friction area of either the ventral forewing {E.
lisa) or the dorsal hind wing (C. philodice ) that produces a chemical
signal used to seduce females during successful courtship. Females
respond to the chemical signal by extending the abdomen ventrally
out from between the hindwings, thereby permitting copulation.

In many sulphur butterflies the males’ sex brands are much more
distinctive than those found in E. lisa and C. philodice. It is the
purpose of this report to document the external morphology of the
male sex brands of three species of pierids, Nathalis iole Boisduval,
Eurema nicippe Cramer, and Colias cesonia Stoll, (Figure 1A).
These species were chosen for two reasons: 1) they appear to
represent a range of sex brand morphologies typical of that found in
the pierid tribe, Coliadini (sulphurs), and 2) they have close
affinities to species for which the function of the sex brand is well-
documented. A special effort will be made to understand the
possible adaptive significance of this variation in sex brand
structure.

* Manuscript received by the editor February 16, 1979.

383

384

Psyche

[December

Methods and Materials

All specimens were collected in Arizona between 17 July and 16
September 1978. The dainty sulphur, N. iole, was collected at Clint’s
Well, the sleepy orange, E. nicippe, near Rye, and the dog face, C.
cesonia, near Tortilla Flat. In addition, N. iole was reared through
several generations in a greenhouse on the Arizona State University
campus, Tempe, Arizona, on their natural foodplant, the tickseed
( Coreopsis cardaminefolia Torrey and Gray or C. atkinsonia Doug-
las). To assess the effect of age on sex brand color or condition in N.
iole and E. nicippe, individual males were assigned to one of three
age classes on the basis of wing condition and scale loss: 1) fresh —
no noticeable scale loss or wing tattering, 2) slightly worn — visible
scale loss and some tattering, especially along wing margins, and 3)
worn — substantial scale loss and/or wings with deep nicks and
tears. The categories used to describe sex brand color variation are
presented in the results. Sex brand condition in N. iole was also a
function of scale number and morphology. Scales in the sex brand
were scored for whether or not they were present and, if so, whether
they were flat, withered, or between these two extremes (half
withered). Lab-reared males were examined on the day they eclosed;
wild males were examined either alive or within 6 hours after they
had died.

Changes in sex brand color with time since death were docu-
mented for N. iole. The wings of 16 freezer-killed males were stored
in glassine envelopes at room temperature and the coloration of
their sex brands scored at weekly intervals until all attained the final
pale yellow color.

Wings of all three species were mounted on aluminium pegs with
DagR, coated with gold in a D. C. sputterer for 3 minutes, and
examined with an AMR-1000A scanning electron microscope at 20
KeV accelerating voltage. In some cases severe charging of the
specimens necessitated longer coating times or low accelerating
voltages (5 KeV). All sex brands viewed with the SEM were from
either freshly-killed males, live males, or males killed and stored at
— 5°C. To minimize the effects of desiccation at high vacuum some
N. iole wings were dehydrated in ethanol, and passed through
Freon-1 13 before critical point drying using Freon-13 as the transi-
tion fluid.

Scale densities were estimated for the sex brand and discal areas
of the wings from scanning electron micrographs of wings whose

1978] Vetter & Rutowski — Sulphur Butterflies

385

Figure 1. A, males of C. cesonia (left), E. nicippe (upper right), and N. iole (lower right); B, dorsal hindwing of male
N. iole (note sex brand under tip of arrow); scanning electron micrographs of N. iole sex brand without (C) and with (D)
scales. Scale lines: A, 3 cm; B, 2 mm; C, 300 D, 200 n .

386

Psyche

[December

Table 1. The relationship between sex brand color in N. iole and several variables.

% of observations

Variable

Dark Light

orange orange

Mottled

Pale

yellow

No. of
males

Days since death

3

88

0

6

6

16

10

38

50

6

6

16

17

6

56

25

13

16

24

6

44

31

19

16

31

0

6

31

62

16

38

0

0

X2 = 107.6,

0

p < 0.0001

100

16

Wing condition/ age

fresh

94

4

2

0

48

worn

71

29

0

0

17

very worn

57

21

X2 = 21.42,

11

p < 0.005

11

28

Scales in sex brand

present

82

13

3

1

76. 5a

absent

Condition of scales in sex

61

brand

18

X2 = 7.06,

9

p < 0.05

12

16. 5a

shrivelled

82

13

5

0

38

half-shrivelled

81

11

0

7

13.5a

flat

86

14

X2 = 6.85,

0

p > 0.3

0

25. 5a

fractional values indicates

a male whose sex brands differed with respect

to this

variable.

scales had been removed with a small brush. The area of the wing
represented in each micrograph was corrected for the fact that all
specimens were photographed from an angle of 45°. Scale attach-
ments intersecting the bottom and left hand borders of each
micrograph were tallied while those intersecting the top and right
hand borders were not. This procedure reduced the possibility of
inflated density estimates.

Results

The sex brand of N. iole

Males of N. iole have an orange, ovoid sex brand (about 0.5 X 1.5
mm) on each hindwing between the subcostal and the radius veins
near the wing base (Figure 1 B). The brand is surrounded by a bar of

1978]

Vetter & Rutowski — Sulphur Butterflies

387

melanic scales that runs along the leading edge of the hindwing from
the wing base to the outer margin. The most unique aspect of the sex
brand in N. iole is that it is frequently devoid of scales (Table 1,
Figure 1C). This condition is not a result of wear as it occurs in
many newly-emerged lab-reared males (Table 2). When scales are
present in the sex brand (Figure ID) they may cover up to 100% of
the brand and vary in form from a scale that is tan, flat, and similar
to other scales on the wing to one that is black, shrivelled, and
contorted (Figure 2A-C; Table 2). Within an individual male the
form of the sex brand scales and the extent to which they cover each
sex brand is usually uniform.

Scale attachments in the sex brand differ from those on other
areas of the wing in appearance and in density. Unlike those in
other areas, a typical sex brand scale attachment has fewer ridges
running up its side, is more rounded, and has a pronounced swelling
or bulge immediately behind the opening of the attachment (com-
pare Figure 2C and D). This bulge is distinctly collapsed in air dried
specimens compared with those that are critical point dried. The
density of scale attachments in the sex brand is almost twice that of
scale attachments in the immediately adjacent discal cell of the wing
(Table 3; t = 40.4, p < 0.001; df = 7).

Examination of descaled wings or wings whose scales had been
cleared in a dilute solution of sodium hypochlorite reveals that the
color of the sex brand in live males is due to an orange pigment in
the integument of the wing. This orange pigment typically extends
little if at all beyond the edges of the sex brand. The pigment is
visible under the light microscope in the space between the dorsal
and ventral surfaces of wings that were crudely sectioned with a
razor blade. Scanning electron microscopy of these same prepara-
tions reveals an ill-defined matrix of material, probably pigment, in
this space that is restricted to the sex brand (Figures 2E and F).

As pointed out by Clench (1976) the orange color changes to pale
yellow after the death of the butterfly. The complete transition takes
about 6 weeks (Table 1) and does not occur at a uniform rate
throughout the sex brand. Splotches of orange pigment remain in
some areas while others have changed to yellow (“mottled”). We
also made the unexpected observation that sex brand color changes
with male age (Table 1). All newly-emerged, lab-reared males (n =
52) have dark orange sex brands while older males, as evidenced by
wing condition, have significantly higher frequencies of mottled and
yellow sex brands (Table 1).

388

Psyche

[December

Table 2. Differences between lab-reared and wild males of N. iole with respect to
the presence and condition of scales in the sex brand.

Scales in sex brand

Lab-reared

Wild

%

No. observed

%

No. observed

present

67

82

52

95

absent

33

18

X2 = 3.81,

p > 0.05

withered

53

49

half-withered

23

35

18

77

flat

24

33

II

©

p > 0.4

The sex brand of E. nicippe

In E. nicippe, the sex brand appears as an orange, triangular
patch of scales about 3 mm across on the ventral surface of each
forewing between the second branch of the cubitus and the second
anal vein (Figure 3A). The scales in the brand are typically brilliant
orange in color in contrast with the white to light-orange scales that
surround the brand. Orange pigment extends through the base of
each scale in the wing integument where it can be seen as an
amorphous blob underlying each scale attachment in the sex brand.
Unlike that of N. iole, the sex brand of E. nicippe is always covered
with scales that occur at about twice the density of scales in the
discal cell of the ventral forewing (Table 3; t = 78.4, p < 0.001,
df = 7).

The scales in the sex brand and their attachments are quite
different in form from those found on other areas of the wings
(Figures 3B and C). The scales are shorter (60 ^um as compared to
80 /im for scales in discal cell) and they extend from the wing surface
at a greater angle. Other differences in scale shape are evident in
Figures 3B and C. The sex brand scale attachments are much larger
than those in the discal area of the wing and as in N. iole have a
pronounced bulge or swelling behind the opening.

The orange color of the sex brand is not constant in all specimens
(Table 4). Older specimens, as measured by wing wear, typically
have a bright white sex brand. In males in intermediate ages, one
often finds an intermediate coloration, with the upper half of the sex

1978]

Vetter & Rutowski — Sulphur Butterflies

389

brand white in color and the lower half a light orange. The sex
brand color does not change with death.

The sex brand of Colias cesonia

Colias cesonia males have a sex brand on the dorsal hindwing
between the subcosta and radius veins near the wing base (Figure
3D). The color of the brand ranges from lemon yellow to a light
yellow with no consistent changes with age or death that we have
observed. The color is solely attributible to the scales as the

Figure 2. Scanning electron micrographs of N. iole wings. A-C, variation in the
structure of sex brand scales; D, a scale from the discal cell; E, crude cross section
through the sex brand showing pigment between the dorsal and ventral wing
surfaces; F, crude cross section of the wings in the discal cell. See text for details.
Scale lines: A-D, 20 /jl\ E and F, 5/j,.

390

Psyche

[December

Figure 3. Eurema nicippe: A, male ventral forewing (note sex brand under tip of
arrow); scanning electron micrographs of male scales from the sex brand (B) and the
discal cell (C). Colias cesonia: D, male dorsal hindwing (note sex brand under tip of
arrow); scanning electron micrographs of male scales from the sex brand (E) and the
discal cell (F). Scale lines: A and C, 5mm; B, 1 0/x ; C and F, 20 E, 30yu.

integument of the wing under the scales is clear. The scales
surrounding the sex brand are of a color similar to that of the brand.
However the sex brand is still distinct from the rest of the wing in
that the density of scales is twice as high (Table 3; t = 42.9, p <
0.001 , df = 4). In fact, the scales in the brand are almost perpendicu-
lar to the wing surface.

The scales are also unique in form (Figures 3E and F). In
particular, the pedicel of each scale arises from the surface of the
scale against the wing rather than projecting from the base in the

1978]

Vetter & Rutowski — Sulphur Butterflies

391

same plane as the scale. The base of each scale in the sex brand
extends beyond the point where the pedicel emerges and partially
overlies the scale socket on the wing surface. The spacing of the
longitudinal and transverse ridges on these scales also appears more
variable than it is on other types of scales.

Discussion

Detailed information is now available about the sex brand
morphology of five species of sulphur butterflies in the pierid tribe,
Coliadini. They include Eurema lisa (Rutowski, 1977) and Colias
philodice (Rutowski, 1979) as well as the three species discussed in
this report. Several generalizations have appeared.

1) All male sex brands are located in the friction areas of the
wings, that is, the area of overlap between the forewings and
hind wings. Rutowski (1979) has suggested that this placement
of the sex brand helps to minimize evaporation from an
otherwise exposed and non-eversible scent producing struc-
ture.

2) The scales observed in the sex brand are not greatly differenti-
ated from those found in other areas of the wings. This
contrasts with the scent scales found in many eversible scent-
producing organs that are very hair-like (e.g. danaids (Pliske
and Salpeter, 1971), noctuids (Birch, 1970)) and with the
presumed scent scales on the wings of some pierids and
nymphalids which have fringed distal borders (Barth, 1950;
Bergstrom and Lundgren 1973; Tinbergen et al., 1942).

3) The scales are associated with cells in the integument of the
wing that reside in a swelling in the wing immediately behind
the scale socket. Presumably these cells are secretory trichogen
cells like those found in scent-producing structures that are
histologically better known, (e.g. danaids (Pliske and Salpeter,
1971)).

These similarities reflect not only the phylogenetic affinities of
these species but also the action of similar selection pressures. The
behavior of the male during courtship is similar in all five species.
The male buffets the female with his wings while she flies or perches
on vegetation (Rutowski, 1978; Silberglied and Taylor, 1978; pers.
obs.). Selection has not acted to favor any divergence from this
pattern among these species and hence there have been no major

392

Psyche

[December

Table 3. Scale attachment densities for the wings of three pierid butterflies.

Attachments/ mm2 Area/ sample (mm2) Sample

Species

Area

X

SD

X

SD

size

N. iole

sex brand

662.9

156.9

0.050

0.003

5

discal cell

393.1

22.4

0.144

0.106

4

E. nicippe

sex brand

974.3

142.2

0.048

0.003

5

discal cell

427.2

63.2

0.234

0.031

4

C. ce sonia

sex brand

411.3

40.5

0.231

0.012

3

discal cell

216.3

21.1

0.216

0.016

3

changes in the basic structure of the sex brands. Still, there are some
striking differences in their fine structure for which there is no ready
explanation. These differences include: 1) placement of sex brand
(forewing vs. hindwing), 2) scale density (some species have sex
brands in which there is no apparent increase in sex scale density
compared to surrounding areas), 3) presence or absence of scales in
the sex brand, and 4) changes in color with male age or death. The
biological rationale for these differences will have to await more
detailed information on the chemistry and cellular structure of these
sex brands.

This study was supported by funds from NSF Grant BNS
78-11211 and an Arizona State University Faculty Grant-in-Aid,
both to R. L. Rutowski. We thank Dr. D. J. Pinkava for identifica-
tion of plant specimens and Mr. L. Marshall for helpful comments
on the manuscript.

Table 4. The relationship between sex brand color and wing condition in
E. nicippe.

Sex brand color (no. of males)

Wing

Condition

Orange

Half orange,
Half white

White

Fresh

6

14

3

Worn

0

3

9

Very worn

0

0

X2 = 20.8, p < 0.0005

5

1978]

Vetter & Rutowski — Sulphur Butterflies

393

References

Barth, R. H.

1950. Vergleichend morphologische Studien iiber die Duftschuppen der Pieri-
den Pieris brassicae und Pieris rapae und der Satyrine Coenonympha
pamphilus. Zool. Jahrb. (Anatomie) 70: 397-426.

Bergstrom, G. and L. Lundgren.

1973. Androconial secretion of three species of butterflies of the genus Pieris
(Lep., Pieridae). ZOON Suppl. 1: 67-75.

Birch, M. C.

1970. Structure and function of the pheromone-producing brush-organs in
males of Phlogophora meticulosa (L.) (Lepidoptera: Noctuidae). Trans.
R. ent. Soc. Lond. 122: 277-292.

Chapman, R. F.

1971. The Insects: Structure and Function. American Elsevier, New York.

Clench, H. K.

1976. Fugitive color in the males of certain Pieridae. J. Lep. Soc. 30: 88-90.

Pliske, T. E. and M. M. Salpeter

1971. The structure and development of the hairpencil glands in males of the
queen butterfly, Danaus gilippus berenice. J. Morph. 134: 215-242.

Rutowski, R. L.

1977. Chemical communication in the courtship of the small sulphur butterfly,
Eurema lisa (Lepidoptera, Pieridae). J. comp. Physiol. 115: 75-85.

1978. The courtship behaviour of the small sulphur butterfly, Eurema lisa
(Lepidoptera: Pieridae). Anim. Behav. 26: 892-903.

1979. Male scent-producing structures in Colias butterflies: Function, localiza-
tion, and adaptive features. J. Chem. Ecol., in press.

SlLBERGLIED, R. E. AND O. R. TAYLOR, JR.

1978. Ultraviolet reflection and its behavioral role in the courtship of the
sulphur butterflies Colias eury theme and C. philodice (Lepidoptera,
Pieridae). Behav. Ecol. Sociobiol. 3: 203-243.

Tinbergen, N., B. J. D. Meeuse, L. K. Boersma, and W. W. Varossieau.

1942. Die Balz des Samtfalters, Eumenis (=Satyrus) semele (L.) Z. Tierpsy-
chol. 5: 182-226.

INTRASEXUAL AGGRESSION IN THE STICK INSECTS
DIAPHEROMERA VELIEI AND D. COVILLEAE AND
SEXUAL DIMORPHISM IN THE PHASMATODEA*

By John Sivinski

Department of Entomology and Nematology
University of Florida
Gainesville, Florida 32611

Introduction

Pairing1 in the Phasmatodea is notable for its duration. The
Indian stick insect Necroscia sparaxes may remain coupled for up to
79 days (a record for insects) and accounts of matings lasting days
or weeks are commonplace (LeFeuvre 1939; Korboot 1961; Gan-
grade 1963; Gustafson 1966; Clark 1974). Intromission may occur
only initially or intermittently. In either case, a substantial propor-
tion of male time-investment is not spent in ejaculate transfer. In
captivity, Diapheromera veliei and D. covilleae pair for 3 to 136
hours and the penis may be inserted and removed up to 9 times. The
genitalia are not in contact for ca. 40% of this period, and
attachment is maintained by a male clasping organ.

According to the best current explanation, males remain with
females when not actively engaged in insemination to guard against
the introduction of rival ejaculates and so avoid sperm competition
(the competition of sperm from 2 or more males for the fertilization
of an ovum; Parker 1971, 1974). Male aggression for possession of
mating females has been described in a variety of insects (Parker
1970). In the only account known to me of attempted mate theft by
a stick insect, an intruding male Orxines macklotti made several
futile efforts to engage his genitalia, climbed off and walked away
(Robinson 1965). Also indicative of a passive defense was a
congregation of Carausius alluaudi collected in the Seychelles
Islands consisting of 6 males clasping a female’s abdomen at various
points (a seventh male was nearby) (Bolivar and Ferriero 1912).
While sedentary guarding of the female might be typical, lack of any

* Manuscript received by the editor January 27, 1979.

'The vocabulary describing attachment between male and female is often inade-
quate for insects. In this paper mating, coupling, and pairing imply continuous
attachment and not constant intromission.

395

396

Psyche

[December

observed aggression may result from a paucity of observers, a
consequence of the Phasmatodea being . . less studied than any
other order of the Orthopteroidea . . (Bradley and Galil 1977).

This paper describes conflict between males in the walkingsticks
Diapheromera veliei and D. covilleae. Results of an experiment are
presented relating male size to ability to monopolize females. Data
on sex-related size-dimorphism in the 2 species are compared to
published values for other members of the order to reach tentative
correlations between population structure, mating strategies and the
relative size of males.

Materials

Adult Diapheromera veliei were obtained in Bernalillo Co., New
Mexico, from the legume Dalea scoparia, and specimens of D.
covilleae from creosote bush ( Larrea tridentata ) in Dona Ana Co.,
New Mexico. Observations were made on insects contained with the
appropriate host plant in 10-gallon aquaria. Densities within
aquaria varied from 3 to 10 adults. These densities, while high, were
not unnatural. Up to 7 adult males and a single female have been
found on a bush whose greatest dimension was 1.5 meters. Insects
are often unevenly distributed in nature, and it was not unusual to
see several adult males within 30 cm. of a mating pair.

The effect of size on the ability of male D. veliei to maintain
attachment to females was examined by keeping a small (x = 67
mm.) and a large (x = 87 mm.) male with a female in each of ten 10-
gallon aquaria. Tandem duration was recorded by checks at 2-hour
intervals between 8:00 A.M. and 8:00 P.M.

To determine the distribution of sexual size-dimorphism within
the Phasmatodea, body lengths of 155 species were obtained and a
male-over-female ratio calculated for each (data from material
deposited in the Florida State Collection of Arthropods, Gaines-
ville, Florida; random selections in Brunner Von Wattenwyl and
Redtenbacker 1908; the whole of the relevant material in Leigh
1909; Hebard 1923; Rehn and Rehn 1939; Gurney 1947; Salmon
1955; Korboot 1961; Bedford and Chinnick 1966; Gustafson 1966;
Stroheker 1966; Paine 1968; Moxey 1971; Clark 1974). Many of
the specimens were dried, and the possibility of differential shrink-
age of the sexes was examined with specimens of D. veliei.
Shrinkage in both sexes was 0 to 3% of live length after a drying
period of 2 weeks. The sexual dimorphism of D. veliei and D.

1978]

Sivinski — Stick Insects

397

covilleae was determined by measurement of freshly killed speci-
mens ( D . veliei — 48 males and 34 females; D. covilleae — 20 males
and 19 females).

Results and Discussion

Mating in both species is initiated by the male mounting the
female dorsally and gripping her body and legs with his tarsi.
Females seldom attempted to dislodge a suitor, and were never
successful. The male abdomen curves underneath the female and his
clasping organ attaches just anterior to the vulva (sometimes
attachment is initially well above the vulva). The resulting position
is typical of the order (Key 1970; Bedford 1978). Some male
copulatory postures may aid in avoiding takeovers. Alexander’s
(1964) phylogenetic scheme of orthopteroid mating positions initi-
ates with female above which then radiates to include a number of
male above or end to end arrangements. These derived postures are
conceivably more effective for male control of the area around the
mating pair. In Diapheromera the dorsal position blocks the length
of the female abdomen and facilitates additional means of prevent-
ing attachment by a second male.

At the approach of another walkingstick the consort almost
invariably bends the tip of the female abdomen down upon itself
with his clasping organ. The angle is sometimes sufficiently acute to
bring the ventral surfaces of the female abdomen on both sides of
the claspers into contact. A large part of the typical site of attach-
ment is thereby denied to competitors. Less frequently, mating
males strike out at approaching males with their forelegs.

Striking motions and manipulation of the abdomen are effective
defenses and most attempts of intruding males to attach to a mating
female were futile. Occasionally second males clasped the female
abdomen and sometimes succeeded in inserting their genitalia into
the vulva. This usually occurred when the consort was no longer in a
dorsal position but had moved to the side of the female to feed. For
periods of a few minutes to an hour, such “menages a trois” were
maintained with incident. In 6 of over a score of double couplings
and combats observed in D. veliei the entire sequence was recorded
from approach of the second male to resolution of the competition.
On one occasion the intruder left in less than an hour without
harassment. In all other cases, fights occurred which followed a
similar pattern.

398

Psyche

[December

The males lean backwards, pulling at each other and often
anchoring themselves by grasping foliage. Eventually, both become
freely suspended, held only by their clasping organs. They then
direct rapid sweeping blows against their opponent (Fig. 1). The
forelegs are used in a boxing manner. After a few seconds to several
minutes, one of the antagonists releases his grip and the victor
shortly regains the dorsal position. Once attachment of the second
male to the female is accomplished, takeover attempts are often
successful. In the 5 fights in which the original male was known, he
was displaced 3 times.

In D. veliei and D. covilleae, a spine is present on the mid femora
of both sexes. Those of the male are enlarged and hooked. In D.
covilleae combats, the opponent’s thorax is held in the joint of the
mid tibia and femur. By flexing the legs, the spine is brought against
the body, and it was once seen to puncture the integument, drawing
blood. Spines used in defense by stick insects are invariably more
highly developed in males (Lea 1916; Robinson 1968; Bedford 1975)
possibly because of their significance in male fighting.

Well-developed male clasping organs (as in Diapheromera spp.)
are not universally present in the order. In reviewing the Australian
Phasmatodea, Key (1970) found male cerci only occasionally modi-
fied into claspers. Perhaps varied cereal design is due to differing
probabilities of takeover.

The mean male over female length ratio of a sample of 155
phasmid species (approximately 8% of described species) is .727.
The average male D. veliei is .922 of the mean female length. This is
an unusually slight difference in body lengths (see Fig. 2), with 94%
of the sample having relatively smaller males (D. covilleae has a
similar male/ female ratio: .916). It might be generally expected that
males would be smaller than females when fecundity is dependent
on size. By spending less time in development or consuming less
food, males take fewer risks in reaching maturity. Even tiny males in
species with internal fertilization are capable of producing an
adequate ejaculate. Given that the niches of the sexes are similar, the
degree of sexual dimorphism is apt to result from a balance of
reproductive pressures acting on the male. These forces might
include maximizing female encounter rates (mobility, material
reserves affecting life span), the ability to invest materially in the
success of progeny, and maintaining copulations by aggression.

1978]

Sivinski — Stick Insects

399

Figure 1 . The combat of two male D. veliei both attached to a female by their
clasping organs (drawing from 35 mm prints of captive specimens).

400

Psyche

[December

MALE/ FEMALE LENGTH RATIO

Figure 2. Male length/ female length frequency distribution for the order Phas-
matodea (generated from citations in materials section). Dots along the curve
represent midpoints of categories spanning 5% on the male/ female length axis and
the number of species occupying these categories. The dashed vertical line represents
the position of D. veliei (.922), the solid vertical line the mean dimorphism of high
density species (.814, see text).

The positive relationship between copulatory success and size is
well established for some polygynous vertebrates (citations in
Trivers 1972). The great male bulk and armature of the dynastine
scarab beetles are agonistic adaptations (Beebe 1944; Eberhard
1977). Larger males of the bibionid fly Plecia nearctica are more
often found copulating than smaller males (Thornhill 1976). Potter
et al (1976) showed size to be a critical factor in the outcome of
combats between male mites.

In the 2 observed fights between male D. veliei of markedly
disparate size, the larger won. Two males from the extremes of the
size continuum were placed in 10 aquaria with single females and
924 hours of coupling were recorded. Large males accounted for 608

1978]

Sivinski — Stick Insects

401

hours (62%). The difference in the mating durations between the size
classes borders on significance (.10 > p > .05).

Density and sex ratio are major components of the sexual
environment and affect the extent of intrasexual competition
(Parker 1974). Unlike the majority of walkingsticks, which are
uncommon and widely dispersed (Key 1970; Craddock 1972), D.
veliei and D. covilleae are locally abundant (as many as 22
individuals in a 1.53 meter bush). Strongly male-biased adult sex
ratios (up to 4:1) persist until late in the season, and are perhaps due
to a combination of earlier male maturation and selective predation
by birds during the early summer nesting period (Sivinski 1977).

In 9 other phasmids known to exist consistently or occasionally at
high densities (numerous enough to cause defoliation of trees or at
least 5 individuals occupying a bush or refuge), the mean male
length/female length ratio is .814 (Wattenwyl and Redtenbacher
1908; Bedford 1975; Lea 1916; Gurney 1947; Wegner 1956; Stockard
1908; Hetrick 1949; Severin 1911; Key 1970; Paine 1968; Wilkins
and Breland 1951). Over % of the complete sample is more sexually
dimorphic (p = .2201) (see Fig. 2). Kentromorphic phases, morpho-
logically distinct forms existing at different densities, exist in several
stick insects. As in locusts with density phases, dimorphism usually
declines as populations reach their greatest concentrations (Key
1957; Uvarov 1966).

Individuals of Eurycantha spp. and Dryococelus australis are
found in the closest spatial proximity of all known stick insects.
Both congregate in tree hollows during the day (Gurney 1947).
Aggregation sex ratios vary enormously and as many as 68 phas-
mids have been found in a single cavity (Lea 1916; Bedford 1975).
Length dimorphism is minimal (.908) and males have massive hind
femora studded with spines of sufficient magnitude to have been
used as fishhooks (Balfour 1915).

Sexual dimorphism in some phasmids is exaggerated in compari-
son to other orthopteroid taxa. Species with below average di-
morphism engage in the longest matings recorded in the order
( Acrophylla tessellata, male/ female = .493, duration 11 days,
Korboot 1961; Anisomorpha buprestoides, .612, 3 weeks; Clark
1974; N. sparaxes, .700, 1-79 days, Gangrade 1963; Timema
calif ornica, .700, 5 weeks, Gustafson, 1966). Protracted couplings
might contribute to selection for male diminution. Disruption of
crypticity may be lessened and male maintenance cost kept to a

402

Psyche

[December

minimum. Since the mating female bears much of the male’s weight,
a dwarfed male may be more fit in two additional contexts:

1) By allowing the female greater activity, the rate at which
additional females are encountered is increased. A potentially
polygynous male can search for mates while in copulo (blocking the
female genitalia until the opportunity for another copulation arises).
Mating durations in D. veliei are shorter when unmated females are
present (Sivinski 1977). 2) By relieving the female’s copulatory
burden resources could be invested in ova the consort might
fertilize. The longer the pairing the more likely this circumstance.

Summary

Intrasexual combats, while common in the Orthopteroidea, have
apparently not been previously recorded in the Phasmatodea.
Fights between males in Diapheromera veliei and D. covilleae are
described. The minimal sexual size-dimorphism of the two species in
comparison to other walkingsticks may be due to a high level of
intrasexual competition brought about by atypical population
parameters (high density, male-biased sex ratios). It is suggested
that extreme dimorphism in the order relieves the burden an
attached male places upon female resources during lengthy copula-
tions.

Acknowledgements

I would like to thank Randy Thornhill, James Lloyd, Thomas
Walker, Bruce Woodward, and Pat Sivinski for their numerous
criticisms and suggestions which led invariably to improvements. I
am grateful to Joan Martin and Susan Wineriter for the illustra-
tions.

References

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1964. The evolution of mating behavior in arthropods. Insect Reproduction
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Balfour, H.

1915. Note on a new kind of fishhook from Goodenough Island d’entrecas-
teaux group New Guinea. Man. 15: 171.

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Bedford, G. O.

1975. Defensive behavior of the New Guiana stick insect Eurycantha (Phas-
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1978. Biology and ecology of the Phasmatodea. Ann. Rev. Entomol. 23:
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Bedford, G. O., and L. J. Chinnick.

1966. Conspicuous displays in two species of Australian stick insects. Anim.
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Beebe, W.

1944. The function of secondary sexual characteristics in two species of
Dynastinae (Coleoptera). Zoologica. 29(3): 53-58.

Bolivar, I., and C. Ferriero.

1912. No. XVII Orthoptera, Phasmidae of the Seychelles. Trans. Linn. Soc.
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Bradley, K. C., and B. S. Galil.

1977. The taxonomic arrangement of the Phasmatodea with keys to the sub-
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Brunner von Wattenwyl, K., and J. Redtenbacher.

1908. Die insektenfamilie der phasmiden. Leipzig.

Clark, J. T.

1974. Stick and leaf insects. Barry Shurlock and Co., Winchester.
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1972. Chromosomal diversity in the Australian Phasmatodea. Aust. J. Zool.
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Eberhard, W. G.

1977. Fighting behavior of male Golofa porteri beetles (Scarabeidae:Sp.
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Gangrade, A.

1963. A contribution to the biology of Necroscia sparaxes Westwood (Phas-
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Gurney, A. B.

1947. Notes on some remarkable Australian walkingsticks including a synop-
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Hebard, M.

1923. Studies in the Mantidae and Phasmidae of Panama (Orthoptera). Trans.
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Hetrick, L. A.

1949. Field notes on a color variant of the two-striped walkingstick Aniso-
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Key, K. H. L.

1957. Kentromorphic phases in three species of Phasmatodea. Aust. J. Zool.
5: 247-284.

Key, J. H. L.

1970. Phasmatodea. In CS1RO, The Insects of Australia. Melbourne Univ.
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Korboot, K.

1961. Observations on the life histories of Acrophvlla tessellata Gray and
Extatosoma tiaratum Macleay (Phasmida). Univ. of Queensland papers.
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Lea, F. E. S.

1916. Notes on the Lord Howe Island Phasma and on an associated longicorn
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Le Feuvre, W. P.

1939. A phasmid with spermatophore. Proc. Royal Ent. Soc. Lond. (A)14: 24.
Leigh, H. S.

1909. Preliminary account of the life history of the leaf insect Phyllium cruri-
folium Serville. Proc. Zool. Soc. Lond. 8: 103-113.

Moxey, C. F.

1971. Notes on the Phasmatodea of the West Indies: two new genera. Psyche.
78: 67-83.

Paine, R. W.

1968. Investigations for the biological control in Fiji of the coconut stick insect
Graeffea crouanii (Le Guillon). Bull. Entom. Res. 57(4): 567-604.
Parker, G. A.

1970. Sperm competition and its evolutionary consequences in the insects.
Biol. Rev. 45: 525-567.

1974. Courtship persistence and female guarding as male time investment
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Potter, D. A., D. L. Wrensch, and D. E. Johnston.

1976. Aggression and mating success in male spider mites. Science. 193:
160-161.

Rehn, J. A. G., and J. W. H. Rehn.

1939. The Orthoptera of the Philippine Islands. Part 1 — Phasmatidea:Sp.
Olriminae. Proc. Acad. Nat. Sci. Phil. 90: 389-487.

Robinson, M. H.

1965. The Javanese stick insect Orxines macklotti De Haan (Phasmatodea:
Phasmidae). Entomol. Mon. Mag. 100: 253-259.

1968. The defensive behavior of the stick insect Oncotophasma martini
(Griffin) (Orthoptera: Phasmatidae). Proc. Royal Ent. Soc. Lond.
( A)43( 10—12): 183-187.

Salmon, J. T.

1955. The genus Acanthoxyla (Phasmidae). Trans. Royal Soc. N. Zeal. 82(5):
1149-1156.

Severin, H. H. P.

1911. The life history of the walkingstick Diapheromera femorata. J. of Econ.
Entomol. 4: 307-320.

Sivinski, J. M.

1977. Factors affecting mating duration in the walkingstick Diapheromera
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Mexico.

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Stockard, C. R.

1908. II Habits, reactions and mating instincts of the “walking stick”: Aplopus
mayeri. Papers from the Tortugas Lab. of the Carn. Inst, of Wash.
2: 43-59.

Strohecker, H. F.

1966. New Timema from Nevada and Arizona. The Pan-Pac. Entomol. 42:
25-26.

Thornhill, R.

1976. Reproductive behavior of the lovebug, Plecia nearctica (Diptera: Bibi-
onidae). Ann. Ent. Soc. Amer. 69(5): 843-847.

Trivers, R. L.

1972. Parental investment and sexual selection. In P. Campbell (ed.). Sexual
selection and the descent of man, 1871-1971. Aldine-Atherton, Chicago,
pp. 136-179.

Uvarov, B.

1966. Grasshoppers and locusts: A handbook of general acrid ology, vol. 1,
481 pp. Cambridge University Press.

Wegner, A. M. R.

1955. Biological notes on Megacrania wegneri Willemse and M. alpheus
Westwood (Orthoptera, Phasmidae). Treubia 23(1): 47-52.

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1951. Notes on the giant walking stick Megaphasma dentricrus (Stal)
(Orthoptera: Phasmatidae). The Texas J. of Sci. 2: 305-310.

SEXUAL CALLING BEHAVIOR IN
HIGHLY PRIMITIVE ANTS*

By Caryl P. Haskins

Haskins Laboratories, New Haven, Connecticut 06510

In recent years an interesting mating pattern has been discovered
in a number of socially parasitic and dulotic Myrmicine ants,
including the related genera Leptothorax, Doronomyrmex, and
Harpagoxenus, and in the guest ant Formicoxenus nitidulus
(Buschinger, 1968, 1971a, 1971b, 1974; 1975). Typically the alate or
ergatoid young female leaves the parent colony, crawls to a
prominent position, and settles there motionless for a long period —
sometimes amounting to hours — with gaster raised, legs extended,
and sting extruded. Sooner or later males assemble about the
“calling” female, and mating takes place. Buschinger, who first
observed it in these genera, has described the pattern in detail for the
slave-making Harpagoxenus sublaevis (1968), and in the perma-
nently socially parasitic Doronomyrmex pads, Leptothorax kutteri,
and L. goesswaldi (1975). He also observed the pattern in fine detail
in populations of F. nitidulus (1975) maintained in the laboratory as
guests of Leptothorax acervorum, and found it generally similar to
the others, though differing in detail in ways to which we will later
allude. Buschinger was able to demonstrate that in all these forms a
sex-attractant pheromone was released from the poison gland, as
Holldobler (1971) had been the first to demonstrate in the Myrmi-
cine ant Xenomyrmex floridanus.

Recently Moglich, Maschwitz, and Holldobler (1974), in a par-
ticularly provocative study, have presented the results of detailed
analyses of “tandem-calling” behavior in the independently-living
Leptothorax acervorum, L. muscorum, and L. nylanderi, a pattern
by which workers of these independently-living species recruit
sisters to newly discovered food sources, one recruiter guiding only
one follower to the source at a time. A component of the initial
behavior of the recruiting worker is described as essentially identical
with the “calling” behavior in Harpagoxenus females, and the
authors call attention to the interesting hypothesis posited earlier by
Holldobler (1971) that, in at least some Myrmicine ants, sex

* Manuscript received by the editor May 15, 1979.

407

408

Psyche

[December

attractants and recruitment pheromones may have had a common
evolutionary origin.

In view of these findings and this challenging suggestion, it
becomes unusually interesting to query how early in Formicid
evolution the “calling” behavior may have originated. For this
reason we are led to put on record some observations of a similar
behavior pattern made several years ago in the highly archaic
Ponerine genus Amblyopone, and also to elaborate some further
observations of the same kind in the genus Rhytidoponera, a part
but not all of which has been noted priviously (Haskins and
Whelden, 1965), but without particular reference to this context.

The Genus Amblyopone

The Ponerine genus Amblyopone includes a rather large group of
species widely distributed over the world. In anatomy, colony
structure, and behavior patterns they must certainly rank among the
most archaic of surviving ants, rivalling or even exceeding in their
primitiveness, the very different but also immensely archaic genus
Myrmecia, as Wilson (1975) has recently emphasized. It was thus of
interest to observe what clearly seems to be “calling” behavior in
virgin females of two species — the North American A. pallipes and
the Australian A. australis, both widely distributed and fairly
common forms in their respective habitats.

Amblyopone pallipes: From 1924 to 1927 an extensive and
continuous series of observations was undertaken in the laboratory
and field in an attempt to elucidate the nuptial flight and colony-
founding behavior patterns in A. pallipes. This species appears to be
ordinarily completely hypogeaic in habit, foraging entirely under-
ground to satisfy its wholly entomophagous and highly specialized
dietary requirements, normally if not invariably confined to a very
limited range of Myriopoda. During late summer in the regions of
New York State and Massachusetts where the work was done the
galleries and chambers of the colonies are concentrated near the
surface of the soil, commonly under logs or stones, and are
characteristically crammed with cocoons of both sexes and castes.
The alate females and males eclose there, and shortly escape
aboveground, singly or in small groups. The males, fully pigmented,
fly off at once. The females, still of the red callow coloration, climb
prominent objects in the vicinity. They may or may not make short

1978]

Haskins — Highly Primitive Ants

409

flights from these vantage points before coming to rest. They then
adopt a posture closely similar to that described for the Leptothora-
cine ants, resting motionless with legs and antennae extended, gaster
arched, and sting protruded or slowly alternately exserted and
withdrawn. Characteristically, the arched gaster is rubbed by the
tibia of one or the other of the third pair of legs. At the height of the
flight season in the localities investigated (typically shady wood-
lands) numbers of low-flying males soon appeared, landed on
vegetation near the female in a high state of arousal, and after some
running, one quickly copulated. Often, as in cases described by
Buschinger, males attempted to mount one another. Following
copulation, females characteristically exhibited reversed photo-
tropism and geotropism and promptly retired underground. This
behavior was observed repeatedly both with alate females naturally
emerging from wild colonies, and with females reared in the
laboratory and subsequently released in appropriate localities.
Unfortunately, no determinations were made of the source of the
pheromone. The prominence of the sting movements suggest that
the source may well, as in Leptothorax, have been the sting gland,
but the Dufours gland has not been experimentally ruled out at this
time, as the source.

Amblyopone australis: A. australis, widely distributed in Aus-
tralia and in the Pacific Islands as far as New Zealand, is larger and
“bolder” in demeanor and habitus than A. pallipes, but in general
resembles it quite closely. It is, however, partly epigeaic and a much
more generalized Arthropod feeder. In this species colony founda-
tion by isolated dealated females has been demonstrated (Haskins
and Haskins, 1951). During this period the foundresses forage
episodically from their frequently-opened cells in the general pattern
of colony-founding females of Myrmeeia. In the course of work
designed to delineate the details of this pattern, alate young females
eclosing from pupae collected in the wild were released under
observation in appropriate locations. Some of these released fe-
males flew briefly, but all soon adopted the “calling” pattern,
although, on the occasions of the experiments, no males appeared.

The Genus Rhvtidoponera

Rhytidoponera is a generalized Ectatommine genus widely dis-
tributed in Australia (its apparent headquarters), but extending to

[December

410

Colony no.
2

Colony no.

3

(Male flights
in this colony
only)

Psyche

Table I

Date of

77mc of

Number of

Observation

Observation

Workers Involved

4/29/68
4/ 8/75

7:45 a.m.

1

15

8:45 a.m.

11

8:55 a.m.

2

4/12/75

7:00 a.m.

11

4/16/75

5:30 a.m.

10

4/21/75

7:25 a.m.

6

4/29/75

7:05 a.m.

2

4/30/75

7:05 a.m.

1

5/ 2/75

7:00 a.m.

2

5/ 6/75

7:10 a.m.

6

5/ 7/75

7:10 a.m.

2

5/ 8/75

6:30 a.m.

3

5/13/75

7:40 a.m.

2

5/14/75

5:50 a.m.

5

5/15/75

6:55 a.m.

9

5/16/75

7:00 a.m.

2

5/22/75

8:00 a.m.

2

6/ 3/75

6:45 a.m.

2

6/ 7/75

6:45 a.m.

2

7/22/75

6:30 a.m.

9

10/ 1/75

7:50 a.m.

1

1/17/76

7:00 a.m.

6

3/22/76

7:25 a.m.

3

3/23/76

5:55 a.m.

2

3/25/76

6:25 a.m.

3

Dare of

Time of

Number of

Observation

Observation

Workers Involved

5/22/75

8:10 a.m.

1

4/16/75

5:30 a.m.

5/14/75

5:50 a.m.

1978]

Haskins — Highly Primitive Ants

411

Table 1 ( continued )

Date of

Time of

Number of

Colony no.

Observation

Observation

Workers Involved

4

4/ 8/75

7:45 a.m.

3

8:45 a.m.

2

8:55 a.m.

3

4/12/75

7:25 a.m.

3

4/29/75

7:05 a.m.

1

4/30/75

7:05 a.m.

3

5/ 6/75

7:10 a.m.

4

5/ 8/75

6:35 a.m.

2

5/13/75

7:40 a.m.

2

5/14/75

5:50 a.m.

1

5/15/75

6:40 a.m.

5

6/ 3/75

6:45 a.m.

3

New Guinea, to neighboring regions of Melanesia, and to Timor,
the Moluccas, the Solomon Islands, and the Philippines (Brown,
1958; Wilson, 1958; 1959). The genus is a large one, and ranges from
tropical rain-forest environments at one latitudinal extreme of its
range to temperate and relatively wet habitats in southern Victoria
and Tasmania and the extreme Australian southwest, while one
group within the genus has invaded highly xerophytic environments
i n the Australian inland. A large number of species are particularly
interesting in that normal alate females are either absent or rare and
apparently do not take a normal part in colony functions. Instead a
proportion of workers, externally morphologically indistinguish-
able from their fellows, possess functional spermathecae, are
fertilized by the active, low-flying males, and serve as multiple
worker-producing reproductives in the colony (Whelden, 1957;
1960; Haskins and Whelden, 1965).

Rhytidoponera is a fairly dominant genus where it occurs, and, as
Brown (1958) has pointed out, its more abundant members appear
to occupy in 'Australia the ecological niche of such relatively
primitive general-feeding Myrmicines as the genus Myrmica in
palearctic and nearctic environments. Indeed, it has been suggested
that the primitive Ectatommines may be fairly close to the ancestral
stirp of the Myrmicinae.

The commonest and best-known member of the genus Rhytido-
ponera is R. metallica, the Australian “greenhead” ant. It is fairly

412

Psyche

[December

ubiquitous in eastern Australia, even commonly invading parks and
gardens. It is readily obtainable, easy of culture, and in general a
nearly ideal laboratory animal for the study of behavioral patterns
in that section of the genus in which normal alate females are
usually lacking.

In the course of a ten-year study of this species in the laboratory
and the field, particular attention was paid to the mode of mating of
the fertilized workers, and of the founding of new colonies. In the
course of this investigation, numerous observations have been made
of worker behavior closely resembling the “calling” pattern in
Leptothorax. The earliest of these observations were recorded
several years ago (Haskins and Whelden, 1965) but as they have
been considerably expanded since, and as they seem to “fit” so
accurately the pattern described for Leptothorax and its congeners,
it has seemed appropriate to expand the record.

In R. metallica males are produced irregularly throughout the
year, both in the laboratory and in the field. They are characteristi-
cally low-flying, and W. L. Brown, Jr. some years ago observed
them coursing closely over the ground, and entering the nests of
other colonies (unpublished observations).

In laboratory colonies, as earlier described (Haskins and
Whelden, 1965) groups of workers were observed in September,
1952, to emerge from the artificial nests and rest quietly grouped
near their entrances, with head and thorax closely appressed to the
substrate and gaster raised and arched. At the same time, the sting
was characteristically exserted. This immobile “pose” was usually
main-tained only briefly, but in the longest interval recorded, for
over twelve minutes. It was commonly interrupted eventually by
other foraging or recently “calling” workers nearby. During this
process, copulations might occur with recently emerged males,
although there was no clearly exhibited preference of males for
“calling” workers over those that were wandering or foraging
normally.1

Since these observations were published, this pattern has been
observed repeatedly in further colonies of R. metallica housed in
laboratory nests. The sets of observations shown below are typical.
They involved three colonies in all, originally collected at Nam-

'See Holldobler, B. and C. P. Haskins ( Science 195, 793-794) for illustrations of
this behavior.

1978]

Haskins — Highly Primitive Ants

413

hours, Northern Queensland, in December of 1963, and maintained
under constant conditions in the laboratory since that time.

Several details of interest emerged from this range of observa-
tions. The first was the virtually seasonally-independent nature of
the events. In 1952 ‘ calling” workers were seen in September and
October, in 1953 in July and November, in 1975 in April, May,
June, July, and October, in 1976 in January and March. Thus every
month was represented except February, August, and December.
This is particularly noteworthy since the observations of 1952 and
1953 were made on colonies from a population taken near Suther-
land, N.S.W. in essentially a temperate location, while those for
1975-76 were derived from a population in the vicinity of Nam-
bours, Queensland, many miles to the north and in essentially a
tropical rain forest area.

Flights of males, as noted both by Brown (1958) and ourselves
(1965), are likewise highly non-specific with respect to season.
Worker “calling” and male flights both appear to take place
predominantly during early to mid-morning hours. It is of interest,
also, that the presence of males in the nest, much less their
emergence, is not a prerequisite for worker “calling”. During the
observations of “calling” made in January, 1957 and May 1975
males were present in the nests, and in at least one case there was
simultaneous male emergence. However, worker “calling” took
place in the absence of male flights in October, 1952 and July 1953
(involving the same colonies in which males were present during
“calling” in 1957) and in all of the 34 instances recorded for Colonies
1 and 3. In both those colonies males had long been absent, and
examination of samples of pupae present in Colony #1 (3) made
during the height of the “calling” period in that small community
revealed only workers to be present in the brood.

The length of time of the “calling” behavior of workers of R.
metallica is typically brief, averaging 1-2 minutes (although one
record period of more than 12 minutes has been noted). Usually,
however, workers moved about after a short period, commonly
resuming the behavior at another location. In this respect, the
pattern resembled that observed by Buschinger for Formicoxenus
nitidulus.

It seems probable that this behavior in R. metallica (observed
also in three other species of the genus: R. tasmaniensis; R.
inornata, and R. violacea) does indeed correspond quite closely with

414

Psyche

[December

the typical “calling” pattern in Leptothorax and the related Myrmi-
cine genera where it has been observed. As in those cases, the sting is
clearly exserted during the “calling” process, but Holldobler (Holl-
dobler and Haskins, 1977) has clearly demonstrated that here the
newly-described tergal gland, strongly developed in Rhytidoponera
is an important, if not the sole, source of the pheromone.2

With Rhytidoponera, as with Amblyopone, the behavior is
evidently only of sexual significance. No evidence of any worker-
worker recruiting has been obtained for either genus.

It seems of real interest that the “calling” behavior seems to be
well established, entirely in a sexual context, at so primitive a grade
of Formicid evolution. It is particularly interesting that it should be
so clearly demonstrated within the Ectatommini, a division of the
Ponerinae which, on morphological grounds, is thought to be fairly
close to the main evolutionary stem leading from the Ponerinae to
the Myrmicinae. Finally, the nature of the colonies and the nuptial
flights of both Amblyopone and Rhytidoponera emphasizes a
reflection of Buschinger, who has suggested (1975) that there may
well be a correlation between the “use” of male mandibular-gland-
derived sex pheromones in the mediation of highly coordinated
massive mating swarms and of female sex pheromones, derived
from sting or Dufours glands, in smaller, poorly coordinated
“straggling” colony flights. The former are characteristic of ant
species forming large closely-knit communities, the latter of those
existing in smaller, more diffuse, less highly integrated communities
of species which are often more rare. Though none of the species
considered here is especially rare, all do form typically scattered,
rather feebly integrated communities having inconspicuous and
essentially uncoordinated nuptial flights.

Literature Cited

Buschinger, A.

1968. “Locksterzeln” begattungsbereiter ergatoider Weiber von Harpagoxenus
sublaevis NYL. (Hymenoptera Formicidae). Experientia 24: 297.

1971a. “Locksterzeln” und Kopula der sozialparasitischen Amiese Leptothorax
kutteri Buschiner (Hym., Form.). Zool. Anz. 186: 242-248.

1971b. Weitere Untersuchen zum Begattungsverhalten sozialparasitischer Amei-
sen ( Harpagoxenus sublaevis Nyl. und Doronomyrmex pads Kutter,
Hym., Formicidae). Zool. Anz. 187: 184-198.

2An extensive paper by B. Holldobler and H. Engel on tergal and sternal glands in
ants is included in this issue of Psyche.

1978]

Haskins — Highly Primitive Ants

415

1974. Zur Biologie der sozialparasitischen Ameise Leptothorax goesswaldi
Kutter (Hym., Formicidae). Insectes Sociaux 21: 133-143.

1975. Sexual pheromones in ants. International Union for Study of Social
Insects, Proc. of Symposium on Pheromones and Defensive Secretions
in Social Insects, September, 1975, Dijon, pp. 225-233.

Brown, W.L., Jr

1958. Contributions toward a reclassification of the Formicidae: II. Tribe
Ectatommini (Hymenoptera). Bull. Mus. Comp. Zool. Harvard 118(5):
175-362.

1954. Systematic and other notes on some of the smaller species of the ant
genus Rhytidoponera. Breviora: Mus. Comp. Zool. Harvard 33: 1-11.

Haskins, C.P. and E.F. Haskins

1951. Note on the method of colony foundation of the ponerine ant Amblyo-
pone australis. Amer. Midland Naturalist 45(2): 432-445.

Haskins, C.P. and R.M. Whelden

1965. Queenlessness, worker sibship, and colony versus population structure
in the formicid genus Rhytidoponera. Psyche 72: 87-111.

Holldobler, B.

1971. Sex pheromone in the ant Xenomyrmex floridanus. J. Insect Physiology
17: 1497-1499.

Holldobler, B. and C.P. Haskins

1977. Sexual calling in a primitive ant. Science 195: 793-794.

Moglich, M., U. Maschwitz and B. Holldobler

1974. Tandem calling: a new kind of signal in ant communication. Science 186:
1046-1047.

Whelden, R.M.

1957. Anatomy of Rhytidoponera convexa. Ann. Ent. Soc. Am. 50: 271-282.

1960. Anatomy of Rhytidoponera metallica. Ann. Ent. Soc. Am. 53: 793-808.

Wilson, E.O.

1958. Studies on the ant fauna of Melanesia III. Rhytidoponera in western
Melanesia and the Moluccas. Bull. Mus. Comp. Zool. Harvard 119(4):
303-320.

1959. Adaptive shift and dispersal in a tropical ant fauna. Evolution 13(1):
122-144.

1975. Sociobiology. Belknap Press of the Harvard University Press. Cam-
bridge, Massachusetts 02138.

A RESTUDY OF TWO ANTS
FROM THE SICILIAN AMBER

By William L. Brown, Jr.1 and Frank M. Carpenter2
Introduction

The ants of the (presumably Miocene) Sicilian Amber were mono-
graphed by Emery (1891), and, except for corrections published by
Emery himself (1913), this faunule has not again been subjected to
critical study. Since 1891, of course, formicid taxonomy has under-
gone radical changes, some of them affecting genera found in this
amber. Ectatomma gracile, for example, was described from a male
specimen that would not today be placed in Ectatomma , but in-
stead, as based on Emery’s description ( 1 89 1 :57 1 ) and figures (PI. 1 ,
fig. 1, 2) is assignable to Gnamptogenys (Kugler and Brown, in
prep.).

It is not our purpose here, however, to review all of Emery’s
Sicilian Amber ants. Rather, we want to present the results of our
study of just two of his type specimens that are particularly signifi-
cant for ant taxonomy. The specimens, in two separate pieces of
amber belonging to the Museo Mineralogico dell’Universita degli
Studi, Bologna, Italy, were lent through the kindness of Prof.
Gianfranco Simboli, Director of the Museo Mineralogico, who has
our thanks. The new preparation of the specimens and their photo-
graphs were done by FMC, while WLB is responsible for the
taxonomic interpretation of the material.

Hypopomyrmex bombiccii
Emery, 1891:574-575, pi. 1, fig. 10, 11, alate queen.

This specimen (figs. 1, 2) is a badly collapsed winged queen
closely involved in the same piece of amber with a worker specimen
of Cataulacus planiceps. Emery, in his fig. 10, and especially fig. 11,
portrays the H. bombiccii specimen as a Strumigenys-like individ-

1 Department of Entomology, Cornell University, Ithaca, New York, 14853. Study
aided by National Science Foundation Grant GB-31662.

2Museum of Comparative Zoology, Harvard University, Cambridge, Mass. Study
aided by National Science Foundation Grant DEB78-09947, F.M. Carpenter, Prin-
cipal Investigator.

Manuscript received by the editor May 30, 1979.

417

418

Psyche

[December

Figure 1 . Hypopomyrmex bombiccii, type queen viewed from left side. Dark
mass below queen is the curled type of Cataulacus planiceps. The view is essentially
the same as portrayed by Emery (1891: pi. 1, fig. 10). Length of fore wing, 3.7 mm.

1978]

Brown & Carpenter — Ants from Sicilian Amber 419

Figure 2. Hypopomyrmex bombiccii, type queen viewed from right side. Length
of fore wing, 3.7 mm.

420

Psyche

[December

mm.

1978] Brown & Carpenter — Ants from Sicilian Amber 421

Figure 4. Sicilomyrmex corniger, type worker, dorsal (facial) view of head,
tilted slightly forward.

422

Psyche

[December

ual; a small eye is shown arising from beneath a scrobe-like head
groove, and elongate mandibles are suggested in vague outline. The
10-segmented antenna is depicted by Emery with a clearly 2-merous
club and an apically thickened and sharply bent scape.

The amber piece has now been cleaned, partly re-ground, and
somewhat cleared by injection of a small amount of Canada balsam.
Figures 1 and 2 are photographs of the Hypopomyrmex bombiccii
type, a winged queen, in the new preparation. The specimen is badly
shrivelled and compressed, especially from side to side, and the
petiolar and postpetiolar nodes are strongly compressed anteropos-
teriorly. It can now be seen that the Strumigenys-\ike cranial shape
portrayed by Emery is really only his free interpretation of the
crumpled head; the deformed left eye protrudes from the dorso-
lateral margin of the head, not from any scrobe, and the mandibles
do not extend as Emery’s figure 11 vaguely suggests they do. The
right side view (fig. 2) of the head now available shows the right
compound eye also distorted, but larger, more elliptical and less
protruding than the left eye. The right antennal scape has its apex
flattened, but not sharply bent like that of the left scape, indicating
that the latter was distorted after death.

Hypopomyrmex is clearly not a member of tribe Dacetini . Habi-
tus, wing venation and the form of the waist do place it in the
subfamily Myrmicinae. The 10-merous antennae with 2-merous
club, the forewing venation and the propodeal teeth make it most
likely a member of the group of genera near Pheidologeton, and it
may be regarded as a doubtful synonym of Oligomyrmex. The
taxonomy of the living forms of this group is still so poorly known,
and the fossil is in such poor condition that formal synonymy here
would be premature.

It may be noted, however, that Oligomyrmex sophiae ( —Aero -
myrma sophiae ), based on male specimens, was described by Emery
from the Sicilian Amber in the same (1891) paper.

With the removal of Hypopomyrmex from the Dacetini, that
tribe loses its entire known fossil history.

Sicilomyrmex corniger

Gesomyrmex corniger Emery 1891:581, pi. 3, fig. 33-35, worker.

This extraordinary formicine is portrayed in the photographs
(figs. 3 and 4). Emery originally assigned it to Gesomyrmex, but the
bicornuate head and two-spined propodeum clearly put it into a

1978] Brown & Carpenter — Ants from Sicilian Amber 423

separate genus, as W.M. Wheeler realized in 1915, when he applied
a new generic name in the combination Sicelomyrmex corniger.
Unfortunately, the new genus name was one of several misspelled in
this German wartime publication, which Wheeler apparently did
not see in proof. He published the name in the emended form
Sicilomyrmex in 1926, and again in 1928, but by 1929 he had
reverted, perhaps absent-mindedly, to the spelling Sicelomyrmex,
and even suggested for it a new tribe, Sicelomyrmicini. The tribal
name was in any case improperly coined, since the stem involved is
myrmec-, not myrmic-.

It would seem proper to recognize the emended spelling Sicilo-
myrmex of 1926 and 1928, since it is clear that Wheeler in 1915 was
alluding to the provenience of the specimen from the Sicilian Am-
ber, and that the original spelling Sicelomyrmex was therefore
either a lapsus calami or a printer’s error, according to Article
33(a)(ii) of the International Code of Zoological Nomenclature. The
necessary emendation of the tribal name thus results in Sicilo-
myrmecini. Whether this tribe is worth retaining can only be de-
cided after full revisionary study of the tribal classification of sub-
family Formicinae.

Emery’s original drawings of S. corniger were good ones, but we
think that the first photographs of the type specimen (figs. 3 and 4)
give an excellent idea of its habitus.

References Cited

Emery, C.

1891. Le formiche dell’Ambra Siciliana. Mem. R. Accad. Sci. 1st. Bologna
(5)1:568-591, pi. 1-3.

1913. Le origini e le migrazioni della fauna mirmecologica d’Europa. Rend.
Accad. Sci. Bologna, 1912-1913:29-46.

Wheeler, W. M.

1915. The ants of the Baltic Amber. Schrift. Phys.-okon. Ges. Konigsberg
55:1-142.

1926. Les societes d’insects. Paris, G. Doin et Cie; xii + 468 pp.; cf. p. 136.

1928. The Social Insects. New York; Harcourt, Brace and Co., xviii + 378 pp.;
cf. pi. 18, fig. 27, facing p. 107.

1929. The identity of the ant genera Gesomyrmex Mayr and Dimorphomyrmex
Ernest Andre. Psyche 36:1-12.

PSYCHE

INDEX TO VOLUME 85, 1978

INDEX TO AUTHORS

Aiello, Annette and Robert E. Silberglied. Life History of Dynastor darius
(Lepidoptera: Nymphalidae) in Panama. 331

Ashley, D. L. See Lilly, C. K.

Brandao, Carlos Roberto F. Division of Labor within the Worker Caste of
Formica peripilosa Wheeler (Hymenoptera: Formicidae). 229

Brown, William L., Jr. and Ronald G. Boisvert. The Dacetine Ant Genus
Pentastruma (Hymenoptera: Formicidae). 201

Brown, William L., Jr. and Frank M. Carpenter. A Restudy of Two Ants from
the Sicilian Amber. 417

Burnham, Laurie. Survey of Social Insects in the Fossil Record. 85

Carpenter, Frank M. See Brown, William L., Jr.

Carpenter, Frank M. and Eugene S. Richardson, Jr. Structure and Relationships
of the Upper Carboniferous Insect, Prochoroptera calopteryx (Diaphonoptero-
dea, Prochoropteridae). 219

Eberhard, W. G. See Lubin, Y. D.

Engel, Hiltrud. See Holldobler, Bert.

Evans, Howard E. A Solitary Wasp that Preys upon Lacewings (Hymenoptera:
Sphecidae; Neuroptera: Chrysopidae). 81

Glorioso, Michael J. See Valentine, Barry D.

Gwynne, Darryl T. See Morris, Glenn K.

Haskins, Caryl P. Sexual Calling Behavior in Highly Primitive Ants. 407

Henry, Charles S. An Unusual Ascalaphid Larva (Neuroptera: Ascalaphidae)
from Southern Africa, with Comments on Larval Evolution within the
Myrmeleontoidea. 265

Holldobler, Bert and Hiltrud Engel. Tergal and Sternal Glands in Ants. 285

Hunter, Kenneth W., Jr. Searching Behavior of Hippodamia convergens Larvae
(Coccinellidae: Coleoptera). 249

Jackson, Robert R. and Sandra Smith. Aggregations of Mallos and Dictyna
(Araneae, Dictynidae): Population Characteristics. 65

Kugler, Charles. Description of the Ergatoid Queen of Pogonomyrmex mayri with
Notes on the Worker and Male (Hym. Formicidae). 169

427

Kugler, Charles. Further Studies of the Myrmicine Sting Apparatus: Eutetra-
morium, Oxyopomvrmex, and Terataner (Hymenoptera, Formicidae). 255

Lilly, C. K., D. L. Ashley, and D. C. Tarter. Observations on a Population of
Sialis itasca Ross in West Virginia (Megaloptera: Sialidae). 209

Lubin, Y. D., W. G. Eberhard, and G. G. Montgomery. Webs of Miagrammopes
(Araneae:Uloboridae) in the Neotropics. 1

Montgomery, G. G. See Lubin, Y. D.

Moore, Ian and R. E. Orth. Notes on Brvothinusa with Description of the Larva
of B. catalinae Casey (Coleoptera: Staphylinidae). 183

Morris, Glenn K. and Darryl T. Gwvnne. Geographical Distribution and Bio-
logical Observations of Cyphoderris (Orthoptera: Haglidae) with a Description
of a New Species. 147

New' ton, Alfred F. See Thayer, Margaret K.

Orth, R. E. See Moore, Ian.

Peek, Stewart B. Systematics and Evolution of Forest Litter Adelopsis in the
Southern Appalachians (Coleoptera: Leiodidae). 355

Richardson, Eugene S., Jr. See Carpenter, Frank M.

Robinson, Barbara. See Robinson, Michael H.

Robinson, Michael H. and Barbara Robinson. Culture Techniques for Acanthops
falcata, a Neotropical Mantid Suitable for Biological Studies (with Notes on
Raising Web Building Spiders). 239

Rutow'ski, Ronald I. See Vetter, Richard S.

Shapiro, Arthur M. The Evolutionary Significance of Redundancy and Variability
in Phenotypic-Induction Mechanisms of Pierid Butterflies (Lepidoptera). 275

Silberglied, Robert E. See Aiello, Annette.

Sivinski, John. Intrasexual Aggression in the Stick Insects, Diapheromera veliei
and D. covilleae, and Sexual Dimorphism in the Phasmatodea. 395

Smith, Sandra. See Jackson, Robert R.

Tarter, D. C. See Lilly, C. K.

Thayer, Margaret K. and Alfred F. New'ton, Jr. Revision of the South Temperate
Genus Glvpholoma Jeannel, with Four New Species (Coloptera: Staphylini-
dae). 35

Valentine, Barry D. and Michael J. Glorioso. Grooming Behavior in Diplura
(Insecta: Apterygota). 191

Vetter, Richard S. and Ronald /. Rutowski. External Sex Brand Morphology of
Three Sulphur Butterflies (Lepidoptera: Pieridae). 383

Vo/lrath, Fritz. A Close Relationship Between Two Spiders (Arachnida: Aranei-
dae): Curimagua bavano Synecious on a Diplura Species. 347

Wood, T. K. Parental Care in Guavaquila compressa Walker (Homoptera: Mem-
bracidae). 135

428

INDEX TO SUBJECTS

All new genera, new species and new names are printed in capital type.

A cant hops falcata, 239
Adelopsis alleghenyensis, 363
A de lop sis alta, 373
Adelopsis appalachiana, 366
Adelopsis bedfordensis, 369
Adelopsis cumberlanda, 369
Adelopsis fumosa, 371
Adelopsis joanna, 374
Adelopsis jonesi, 367
Adelopsis mitchellensis, 361
Adelopsis nashvillensis, 371
Adelopsis orichchalcum, 376
Adelopsis pisgahensis, 376
Adelopsis richlandensis, 365
Adelopsis scottboroensis, 370
Adelopsis steevesi, 362
Adelopsis suteri, 363

Adelopsis, systematics and evolution,
355

Amblyopone, 408

Ants, tergal and sternal glands, 285
Apoidea, fossil, 107
Ascalaphid larva, 265
Behavior in primitive ants, 407
Bembix stenebdoma, 81
Biology of Cyphoderris, 147
Bryothinusa catalinae, larva, 183
Calling behavior in primitive ants, 407
Chrvsoperla comanche, 83
Colias cesonia, 383
Curimagua bayano, 347
Cyphoderris, 147
Cyphoderris buckelli, 154

Cyphoderris monstrosa, 154
Cyphoderris strepitans, 149
Dacetine ant genus Pentastruma, 201
Diapheromera covilleae, 395
Diapheromera veliei, 395
Dictyna albopilosa, 65
Dictyna calcarata, 65
Diplura (Arachnida), 347
Diplura, grooming behavior, 191
Dynastor darius, life history
Eremochrysa punctinervis, 83
Eremoehrysa tibialis, 83
Eurema nicippe, 383
Eutetramorium, 255

External sex brand morphology of
butterflies, 383

Forest litter Adelopsis, 385
Formica perpilosa, 229
Formicidae, fossil, 98
Glands in ants, 285
Glypholoma pecki, 50
Glypholoma pustuliferum, 46
Glypholoma, revision, 25
Glypholoma rotundulum, 54
Glypholoma temporale, 49
Glypholoma tenuicorne, 53
Grooming behavior in Diplura, 191

Guayaquila compressa, parental care,
135

Hippodamia convergens, larvae, 249
Hypopomyrmex bombiccii, 417

Intrasexual aggression in stick insects,
395

429

Isoptera, in fossil record, 85
Lacewings, 81

Life History of Dynast or darius, 331

Mallos and Dictvna, aggregations, 65

M alios gregalis, 67

Mallos niveus, 68

Mallos trivittatus, 68

Mantid, 239

Myrmeleontoidea, 265

Myrmicine sting, 255

Nathalis iole, 383

Observations of Cvphoderris, 147

Oxypomyrmex, 255

Parental care in Guavaquila, 135

Pentastruma, 203

Pentastruma can in a, 203

Pentastruma sauteri, 203

Phasmatodea, 395

Phenotypic-induction mechanisms,
Pieridae, 275

Pierid butterflies, 275

Pogonomvrmex mayri, ergatoid queen,
169

Population of Sialis itasca, 209
Prochoroptera calopteryx, 219
Relationship between two spiders, 347
Restudy of two Sicilian amber ants, 417
Rhytidoponera, 409

Searching behavior of Hippodamia, 249
Sex brand morphology of Pieridae, 383

Sexual dimorphism in the Phasmatodea,
395

Sialis itasca, 209
Sicilomyrmecini, 423
Sicilomyrmex corniger, 422
Social insects in fossil record, 85
Stick insects, 395

Studies of the myrmicine sting appa-
ratus, 255

Sulphur butterflies, 383
Terataner, 255

Tergal and sternal glands in ants, 285
Vespoidea, fossil, 94
Web building spiders, 239
Webs of Miagrammopes, 1

430

The illustration on the front cover of this issue of Psyche
is a reproduction of the published figure of the minute diapriid,
Solenopsia americana (1.3 mm. long), described by C. T. Brues
in Psyche (1936, vol. 43, p. 17). The insect was taken in the nest
of an ant, Paratrechina parvula, in eastern Tennessee.

CAMBRIDGE ENTOMOLOGICAL CLUB

A regular meeting of the Club is held on the second Tuesday
of each month October through May at 7:30 p.m. in Room 154,
Biological Laboratories, Divinity Avenue, Cambridge. Entomolo-
gists visiting the vicinity are cordially invited to attend.

BACK VOLUMES OF PSYCHE

Requests for information about back volumes of Psyche should
be sent directly to the editor.

F. M. Carpenter
Editorial Office, Psyche
16 Divinity Avenue
Cambridge, Mass. 02138

FOR SALE

Reprints of articles by W. M. Wheeler

The Cambridge Entomological Club has for sale numerous reprints
of Dr. Wheeler’s articles that were filed in his office at Harvard
University at the time of his death in 1937. Included are about
12,700 individual reprints of 250 publications. The cost of the re-
prints has been set at 5c a page, including postage; for orders under
$5 there will be an additional handling charge of 50c. A list of the
reprints is available for $1.00 from the W. M Wheeler Reprint
Committee, Cambridge Entomological Club, 16 Divinity Avenue,
Cambridge, Mass. 02138. Checks should be made payable to the
Cambridge Entomological Club.

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