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PSYCHE

A Journal of Entomology

Volume 72
1965

Editorial Board

Frank M. Carpenter, Editor P. J. Darlington, Jr.

W. L. Brown, Jr. H. W. Levi

E. O. Wilson H. E. Evans

Published Quarterly by the Cambridge Entomological Club
Editorial Office: Biological Laboratories
1 6 Divinity Ave.

Cambridge, Mass., U. S. A.

The numbers of psyche issued during the past year were mailed on the
following dates:

Vol. 71, no. 4, Dec., 1964: March 6, 1965

Vol. 72, no. 1, March, 1965: June 25, 1965

Vol. 72, no. 2, June, 1965: September 25, 1965

Vol. 72, no. 3, Sept., 1965: January 18, 1966

PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol. 72 March, 1965 No, 1

W. M. Wheeler Memorial Issue

CONTENTS

CONTENTS

The William Morton Wheeler Memorial Issue 1

Trail Sharing in Ants. Edward O. Wilson 2

Simultaneous Care of More Than One Nest by A mmophila azteca

Cameron ( Hymenoptera, Sphecidae). Howard E. Evans 8

The Ant Larvae of the Subfamily Leptanillinae (Hymenoptera,

Formicidae). George C. Wheeler and Jeanette Wheeler 24

The Australian Ants of the Genus Pristomyrmex, with a Case of

Apparent Character Displacement. Robert W. Taylor 35

A New Species of Megalomyrmex from the Chilean Andes

(Formicidae, Hymenoptera). George Etter shank 55

The Habits and Distribution of Cryptocerus rohweri Wheeler (Hymen-
optera: Formicidae). William S. Creighton and William L. Nutting 59
Contributions to a Reclassification of the Formicidae. IV. Tribe

Typhlomyrmecini (Hymenoptera). William L. Brown, Jr 65

A Revision of the Ant Tribe Cardiocondylini (Hymenoptera,

Formicidae). Jonathan Reiskind 79

“Queenlessness,” Worker Sibship, and Colony Versus Population Struc-
ture in the Formicid Genus Rhytidoponera. Caryl P. Haskins and

Roy M. Whelden 87

Observations on the Nesting Site and Biology of the Arizona Damp-
wood Termite, Zootermopsis laticeps (Banks) (Hodotermitidae).

W . L. Nutting 113

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1964-65

President...................... . R. W. Taylor, Harvard University

Vice-President J. Reiskind, Harvard University

Secretary .........H. Reichardt, Harvard University

Treasurer ...F. M. Carpenter, Harvard University

Executive Committee.. N. S. Bailey , Bradford, Mass.

E. G. Macleod, Harvard University

EDITORIAL BOARD OF PSYCHE

F. M. Carpenter (Editor), Professor of Entomology , and Alexander
Agassiz Professor of Zoology, Harvard University

P. J. Darlington, Jr., Alexander Agassiz Professor of Zoology,
Harvard University

W. L. Brown, Jr., Associate Professor of Entomology, Cornell
University; Associate in Entomology, Museum of Comparative
Zoology

E. 0. Wilson, Professor of Zoology, Harvard University

H. W. Levi, Associate Curator of Arachnology, Museum of Com-
parative Zoology

H. E. Evans, Curator of Insects, Museum of Comparative Zoology

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: $4.50 to Club members, $5.00 to all other
subscribers. Single copies, $1.25,

Checks and remittances should be addressed to Treasurer, Cambridge Ento-
mological Club, 16 Divinity Avenue, Cambridge, Mass.

Orders for back volumes, missing numbers, notices of change of address, etc.,
should be sent to the Editorial Office of Psyche, Biological Laboratories, Har-
vard University, Cambridge, Mass.

IMPORTANT NOTICE TO CONTRIBUTORS

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

Authors contributing articles over 4 printed pages in length may be required
to bear a part of the extra expense, for additional pages. This expense will
be that of typesetting only, which is about $10.00 per page. The actual cost of
preparing cuts for all illustrations must be borne by contributors: the cost
for full page plates from line drawings is ordinarily $12.00 each, and the full
page half-tones, $18.00 each; smaller sizes in proportion.

AUTHORS SEPARATES

Reprints of articles may be secured by authors, if they are ordered at the
time proofs are received for corrections. A statement of their cost will be
furnished by the Editor on application.

The December, 1964 Psyche (Vol. 71, no. 4) was mailed March 6,

1965.

The Lexington Press. Inc., Lexington, Massachusetts

Psyche, 1965

Vol. 72, Plate 1

William Morton Wheeler

march 19, 1865 — APRIL 19, 1937
PROFESSOR OF ENTOMOLOGY, HARVARD UNIVERSITY

1908-1937

Vol. 72

No. 1

PSYCHE

March, 1965

WILLIAM MORTON WHEELER MEMORIAL ISSUE

March 19, 1965, is the centennial of the birth of William Morton
Wheeler, Professor of Entomology at Harvard University from
1908-1937 and an active member of the Cambridge Entomological
Club for that period. The Editorial Board of Psyche has designated
the present number of the journal as the William Morton Wheeler
Memorial Issue and has arranged to include in it articles on ants
and other social or subsocial insects. No attempt has been made to
obtain contributions by all of Professor Wheeler’s former students;
most of the papers published here were already in the editorial office
before plans for the memorial issue were definitely made. Included are
contributions by three generations of students whose interest in social
insects can clearly be traced to Professor Wheeler. We are indebted
to Professor W. L. Brown, Jr., of Cornell University for the use
of the accompanying photograph, which was taken about five years
after Professor Wheeler was appointed at Harvard University.

A biographical account of Professor Wheeler, with a complete
list of his publications, was published in Psyche, Volume 44, No. 3.,
1937.

F. M. Carpenter, Editor

I

JWTHSONIA* jlil |

WSTlTUt^

TRAIL SHARING IN ANTS

By Edward O. Wilson
Biological Laboratories, Harvard University

introduction: the kinds of trail sharing

A very few cases have been recorded of ant workers regularly
utilizing the trails of other ant species. Forel (1898) designated as
“parabiosis” the following complex behavior that includes trail
sharing. Colonies of the Neotropical rain forest species Crematog aster
limata parabiotica Forel and Monacis debilis (Emery) [ —Doli -
choderus debilis var. parabiotica Forel] commonly nest in close
association, with the nest chambers kept separate but interconnected
by passable openings; while the workers forage along common odor
trails. Wheeler (1921) confirmed the phenomenon and showed that,
in the one instance where he observed food gathering, the two species
were attending membracids together. Wheeler also discovered a
similar association between Crematogaster parabiotica and Campo-
notus femoratus (Fabricius). Both species were observed utilizing
common trails and gathering honeydew from jassids and membracids
on the same plants, as well as nectar from the same extrafloral
nectaries of Inga. Not only were the Crematogaster and Camponotus
workers tolerant of each other in this potentially competitive situation,
they were on quite intimate terms. They “greeted” each other with
calm antennation on the trails, and on three occasions Wheeler
observed Camponotus actually regurgitating to Crematogaster.

It has not been established whether parabiosis is mutualistic or
parasitic in nature. The distinction must be a subtle one in such a
complicated relationship. The form <( parabiotica” of Crematogaster
limata is evidently always associated with other ants. If future
taxonomic studies prove it to be a species distinct from limata , it is a
likely parasite. It would then be shown to be dependent on its associ-
ates, while the latter species often nest and forage by themselves. But
the prima facie case for mutualism seems even stronger. The broods
are never mixed, and as Weber (1943) points out on the basis of his
own studies, all of the parabiotic species participate vigorously to-
gether in nest defense. There is no evidence that the presence of the
Crematogaster harms the other species, except possibly by competition
for the same food resources. On the contrary, Camponotus femoratus

Manuscript received by the editor January 5, 1965.

2

1965]

Wilson — Trail Sharing

3

maintains flourishing populations in localities where virtually every
colony lives in parabiosis with Crematogaster.

While the Neotropical parabionts are doubtfully mutualistic, the
relationship of the European Camponotus lateralis (Olivier) and
Crematogaster scutellaris (Olivier) can be classified as weakly
parasitic. Goetsch (1953) and Kaudewitz (1955) have described
instances in which Camponotus workers followed the Crematogaster
trails in large numbers to the Crematogaster feeding grounds and
exploited the same food resources during the same time of day. The
Crematogaster were hostile to the Camponotus, which assumed a
crouching, conciliatory “Wartestellung” on meeting the host workers.
Unlike the Neotropical parabionts, the two species nest separately.
Moreover, the relationship is not obligatory on the Camponotus
lateralis , since the colonies of that species are often found far removed
from Crematogaster colonies.

I will now describe a third example of trail sharing which I
recently discovered between the dolichoderine Azteca chartifex Forel
and formicine Camponotus heehei Wheeler. This case is of additional
interest in that it seems to illustrate a close approach to the third or
neutral class of symbiosis, namely commensalism.

AZTECA CHARTIFEX AND CAMPONOTUS BEEBEI

During a trip to Trinidad, West Indies, in 1961, my attention was
drawn to Camponotus heehei, a formicine ant previously known from
only several specimens collected in Trinidad and British Guiana. On
each of three occasions on which the species was encountered, twice at
Spring Hill, Arima Valley, and once near Cumuto Village on the
Aripo Savanna, workers were found running over tree trunks along
the odor trails of the much more abundant and aggressive dolichode-
rine Azteca chartifex. The Camponotus were never found away
from the Azteca trails. Extended observations at Spring Hill revealed
that the Camponotus always followed the Azteca trails for long
distances with fidelity equal to that maintained by the Azteca them-
selves. That this was true trail symbiosis was further evidenced by
the fact that no other alien species remotely approximated such
behavior. Workers of several other arboreal species occasionally
blundered into the same Azteca files but ran abruptly away without
tracing the main route of the files.

One of the Spring Hill Camponotus nests was located. It was in a
dead, hard branch of a mango tree that had fallen and lodged in the
crown of a three-meter-tall grapefruit tree in a citrus plantation. The
Camponotus workers were seen to emerge from their nest holes, run

[March

Psyche

Minor worker of Camponotus beebei Wheeler from Spring Hill, Trinidad.

1965]

Wilson — Trail Sharing

5

down the mango branch to the branches of the grapefruit tree, which
held an Azteca colony, and follow the Azteca trails to the ground.
The Azteca workers seldom ventured up to the Camponotus nest.
The Camponotus occupied scattered flat galleries in the mango branch.
When cut apart the nest yielded 2 winged queens, 16 males, 6 major
workers, 36 minor workers, and several larvae and pupae in various
stages of development. The mango tree, from which the Camponotus
colony fragment had evidently recently fallen, was also occupied by
Azteca chartifex. In a second locality at Spring Hill, Camponotus
workers were tracked up into the foliage of a tonka bean tree
( Dipteryx sp.) beyond a large Azteca nest, but the Camponotus nest
was not found. Nevertheless, it was evidently separate from the
Azteca nest.

Both the Azteca and Camponotus followed the Azteca trails to the
bases of the nest trees. Presumably both foraged extensively on the
herbaceous ground vegetation, but their diets were not determined.
Regardless of the nature of the diets, competition between the two
species was reduced by the existence of opposite diel schedules. The
Camponotus foraged apparently exclusively during the day, at the
time the Azteca files were at their lowest ebb. In the early evening the
number of Azteca workers on the trails were seen to increase by as
much as a hundred-fold, but not a single Camponotus worker was
found through several (hours of searching during this time.

The Camponotus workers, then, “borrow” the Azteca trails when
the owners put them to minimal use. The Azteca workers on the
Spring Hill trails were hostile to the Camponotus workers and
attacked them on the rare occasions when the latter slowed in their
running, but the Camponotus were larger and faster and usually
easily avoided their hosts without causing any visible disturbance.
The Camponotus were never observed to interfere with the Azteca
in any other way.

On the basis of the first observations it could still be legitimately
asked whether the Camponotus were merely using the same visual
or tactile “landmarks” on the tree trunks as the Azteca, rather than
following their odor trails. This possibility was eliminated by the
following experimental result. A freshly killed insect was pinned to
the trunk of a tree one meter beneath the trail along which both
species were running but within the range of occasional Azteca
scouts. Within ten minutes, two Azteca workers had found the
insect and laid odor trails from it back to the main trail. In the next
five minutes over 100 Azteca workers moved back and forth along the
new trail to the insect. In the same interval three Camponotus

6

Psyche

[March

workers, a major and two minors, approached along the main trail
and, on reaching the junctures of the new trails, departed down them
for various distances. The major went all the way to the insect and
prowled around it for several minutes before returning to the main
trail. In two subsequent replications of the experiment, two of
thirteen and one of five Camponotus workers passing along the main
trail were deflected onto the Azteca side trails during the period of
peak Azteca response to the baits. Such deviations from the main
trail were never observed except at this time. It was concluded that
the Camponotus respond to the Azteca communication.

The following observation led to the further conclusion that the
Camponotus were tracking the Azteca olfactorially rather than
visually. Occasionally around midday the Azteca were unusually
scarce on the main trail, while the Camponotus remained moderately
common. Stretches of 30 to 50 cm. of the trail were often bare of
Azteca, but many individual Camponotus followed the established
track just as well. On close examination I found no alterations in
the surface structure of the main trail, other than the postulated
chemical one, that could have supplied the Camponotus with a clue.

Although the Camponotus beebei utilize Azteca trails extensively,
the following observation shows that they have maintained their own,
private trail system. On a single occasion in February a line of seven
Camponotus were seen moving along the main Azteca trail. Four of
the workers ran in a tight group directly behind the leader, frequently
advancing enough to touch the abdomen of the ant ahead. When the
leader was touched, it dashed forward at a faster pace over a short
distance. This part of the behavior was typical of communication by
“tandem running”, which I have described earlier in a paper on the
genera Cardicondyla and Camponotus (Wilson, 1959). The re-
maining two workers followed at a greater distance, tracing each
twist and turn taken by the leader. During the next 15 minutes
several other Camponotus workers passed the same way, again tracing
parts of the route of the leader with close fidelity. After that time,
new Camponotus workers continued to run on the Azteca trail but
ignored the Camponotus trail. There could be no doubt that the
lead ant had secreted an odor trail of the recruitment type (see
Wilson, 1963). It was laid on top of the Azteca trunk trail, which
for most of its length was about 10 centimeters wide. Equally
interesting was the fact that only the Camponotus responded to it.
The Azteca workers continued to pass along their own trail during
the episode but failed to orient to the inner track followed so closely

1965]

Wilson — Trail Sharing

7

by the Camponotns. Thus the Camponotus workers appear to respond
to two odor trails, while the host Azteca respond only to one.

Acknowledgements

This study was supported by a grant from the National Science
Foundation. The figure was prepared by Mrs. H. C. Lyman.

ABSTRACT

Trail sharing is a rare event in ants. Of two previously described
cases, one is interpreted as part of a relationship that is either mutu-
alistic or weakly parasitic, probably the former, and the other as part
of a weakly parasitic relationship.

A third, new case has been discovered which appears to be com-
mensalistic. On Trinidad, West Indies, workers of the rather scarce
formicine Camponotus beebei utilize the arboreal odor trails of the
abundant dolichoderine, Azteca chartifex. The Camponotus “borrow”
the latter’s trails during the day, when Azteca foraging is at a low
ebb. The Camponotus workers are treated hostilely by the Azteca
workers but are too swift and agile to be caught; their presence does
not disturb the Azteca seriously. On a single occasion Camponotus
workers were observed to lay their own private recruitment odor
trail on top of the Azteca trails. The Camponotus trail lasted for
about fifteen minutes and had no visible effect on the Azteca .

Literature Cited

Forel, A.

1898. La Parabiose chez les Fourmis. Bull. Soc. Vaud. Sci. Nat., 34:
380-384.

Goetsch, W.

1953. Vergleichende Biologie der Insekt-Staaten. Geest und Portig
K.-G., Leipzig. 482 pp.

Kaudewitz, F.

1955. Zum Gastverhaltnis zwischen Crematogaster scutellaris 01. mit
Crematogaster lateralis bicolor 01. Biol. Zentralbl., 74: 69-87.

Weber, N. A.

1943. Parabiosis in Neotropical “ant gardens.” Ecology, 24: 400-404.

Wheeler, W. M.

1921. A new case of parabiosis and the “ant gardens” of British
Guiana. Ecology, 2:89-103.

Wilson, E. O.

1959. Communication by tandem running in the ant genus Cardio-
condyla. Psyche, 66: 29-34.

1963. The social biology of ants. Ann. Rev. Entomol, 8:345-368.

SIMULTANEOUS CARE OF MORE THAN ONE NEST
BY AMMOPHILA AZTEC A CAMERON
(HYMENOPTERA, SPHECIDAE)1

By Howard E. Evans
Museum of Comparative Zoology

In attempting to trace the origin of social behavior among wasps,
in his Social Life Among the Insects (1923), William, Morton
Wheeler selected Arnmophila as “a paradigm of the whole group of
Sphecoids and solitary Vespoids”. Were he alive today, and able to
utilize all the considerable knowledge of this genus gained in the last
three decades, it seems likely that he would embrace Ammophila even
more enthusiastically as a paradigm not only of the solitary wasps
but of several preliminary stages in the origin of sociality.

It has been shown by Evans (1958, 1959) and by Powell (1964)
that the North American species of this genus which have been
studied can be arranged in series as follows: (1) strictly solitary

species which utilize a single large caterpillar per nest, (2) species
which mass-provision with two to several small caterpillars, (3)
species in which provisioning is commonly “delayed” such that the
last prey is brought in after the egg has hatched, and (4) species
employing progressive provisioning regularly. Several other aspects
of behavior are roughly correlated with this progression : for example,
species using smaller caterpillars usually carry the prey in flight, and
these same species generally carry the soil of excavation away from
the nest in flight. Also, most records of gregarious nesting pertain to
species employing progressive provisioning.

It is apparent that the European species can be arranged in a very
similar series (Adriaanse, 1947; Teschner, 1959). One European
species, A. puhescens Curtis, illustrates still a fifth stage in this
ethocline: the female maintains two or three nests at one time,

remembering the location of each of them accurately and behaving in
accordance with the status of the egg or larva in each nest as deter-
mined during frequent inspections (Baerends, 1941). Simultaneous
care of more than one nest is otherwise virtually unknown among
digger wasps, although a few species of Bembicini which make more
than one cell per nest are reputed to begin provisioning a second cell

Supported by a grant from the National Science Foundation, no. G17497.
Most of the studies reported here were conducted at the Jackson Hole
Biological Research Station, Moran, Wyoming.

Manuscript received by the editor January 5, 1965

8

1965]

Evans — Armnophila azteca

9

before the previous cell is fully stocked (Janvier, 1928; Tsuneki,
1956).

Powell (1964) suggested that studies with marked wasps might
well reveal that some of our North American A?n?nophila maintain
more than one nest simultaneously. By coincidence, during the same
week that Powell’s paper appeared, I was able to establish that this
is, in fact, the case. Working along the Snake River in Jackson Hole,
Wyoming, I found that Ammophila azteca Cameron behaves in a
manner strikingly like pubescens in almost every detail relating to
provisioning. My studies were relatively brief and leave a number
of questions unanswered, but they seem worth putting on record at
this time with the thought that others may be able to extend our
knowledge of this widely distributed species before I am able to do so.

Ammophila azteca has been the subject of three published notes, all
of them brief. Hicks (1935) observed several wasps digging and
closing nests near Boulder, Colorado (identified as aculeatus Fernald,
a synonym ) . Evans ( 1 963 ) reported on a single nest found in
Yellowstone Park, Wyoming, and Powell (1964) presented prey
records from California and Baja California. The last two authors
both identified the species as pilosa Fernald, a name now regarded as
a junior synonym of azteca Cameron.2 I also found one female of
this species nesting at Great Sand Dunes National Monument,
Colorado (elevation 7800 feet), in August 1964, and have included
this record below. This is chiefly a montane species ; Powell’s records
are from 6000 feet elevation in Baja California and over 10000 feet
in California; the Jackson Hole and Yellowstone localities where I
have worked are both at about 6800 feet. However, Menke (in
lift.) reports the species from near sea level in California and various
places in Canada.

General aspects of ecology and behavior. — Most of my studies
were conducted in a small area of flat, bare soil along the Snake
River at the Cattlemen’s Bridge, about one kilometer east of the
Jackson Hole Biological Research Station, Moran, Wyoming. The
first observations were made on July 18, 1964, the last observations
on August 14. My impression is that this species became active only
a few days before my initial observations and that it had nearly
completed its nesting season by August 14. In this area, where the
active season for most wasps is no more than 4-6 weeks, progressive

2This is a new synonymy, and should be credited to Arnold Menke, who is
currently revising the genus Ammophila and who has studied Cameron’s
type. I am indebted to Dr. Menke for identifying the specimens collected in
the course of these studies and for critical reading of the manuscript.

IO

Psyche

[March

provisioning would seem uneconomical unless the wasp is able to
maintain two or more nests simultaneously. However, two progressive
provisioned which maintain only one nest at a time nested in some
abundance in the same area; these were Bembix spinolae and Steniolia
obliqua. The only other species of Ammophila found here was a
single female A. macra Cresson which nested on the edge of the bare
soil on August 14; I have presented a few notes on this wasp below,
as it presented a striking contrast to azteca.

In Jackson Hole, A. azteca appeared to be strictly confined to
patches of bare, moderately firm, sandy loam along the river; none
were seen or taken in general collecting in other habitats. The major
nesting area was about 20 meters long and varied from 5 to 8 meters
in width, paralleling the river and separated from it only by a narrow,
oblique bank which was not used for nesting. Parts of this area were
covered with grass and herbs, chiefly around several trees, but these
places were not utilized by the Ammophila. Also, the Ammophila
did not nest in a small plot of very loose sand which was occupied by
a colony of Bembix spinolae. The area was surrounded on the three
sides away from the river by open woodland in which the dominant
tree was lodgepole pine (Pinus contorta latifolia) ; near the river
there were also narrow-leaved cottonwoods (Populus angustifolia)
and willows ( Salix spp.). I estimated that there were about 50
females scattered about the nesting area. My notes cover 20 females,
but only a few of these were marked, and only four of the marked
individuals were followed over a period of days. The most complete
observations pertain to no. 2030, which was marked with a red spot
on the second day of study ( July 19) and followed until August first,
including one complete day of observation (July 31) and observations
for several hours on five other days.

That it was impossible to make continuous records of more than a
very few females was a consequence of the fact that females spaced
themselves widely, each maintaining a small nesting arena, no more
than 30 cm. in diameter, where all her nests were prepared (Figs. 6,
7). These arenas were separated, for the most part, by half a meter
or more. On one occasion an unmarked female shared a circumscribed
area with a marked female (no. 2028C), the nests of the two being
more or less intermingled, but this was exceptional. I observed no
aggression among females provisioning their nests, but females digging
in proximity would sometimes attack one another. It seems possible
that there is a measure of territoriality among females of this species,
but since the wasps spend only a small portion of their time in the
nesting arenas, contacts between neighboring females are few.

1965] Evans — Ammophila azteca II

Both males and females were seen frequently on the flowers of
Solidago and Erigeron growing in and near the nesting area, and it is
probable that they obtain nourishment throughout the season from
this source. Early in the season males were observed in considerable
numbers flying close to the ground in the nesting area. On one
occasion three males were seen digging intermittently and trying to
enter a vertical hole, presumably in anticipation of the emergence of
a female. I did not observe mating, chiefly because most of my studies
were made a week or more after nesting had begun, when most
mating had probably ceased and the males were declining in numbers.
Mating in several species of Ammophila has been well described (e.g.,
Baerends, 1941; Olberg, 1959).

As compared to other wasps in this area, Ammophila azteca made
its appearance rather early each morning (0830-0900) and was
active until late in the afternoon (1630). Active wasps were often
noted in partial shade or during periods of partial cloudiness, but they
invariably disappeared when the sun was heavily obscured for more
than a few minutes. I did not discover where they went during these
inactive periods or at night. Hicks (1935) found a female of this
species “sleeping” on a stem in late afternoon, presumably solitarily.
The wasp held her body at nearly a right angle to the stem, grasping
the support with her mandibles and some of her legs. A number of
species of Ammophila are known to sleep in somewhat this manner,
often more or less gregariously (e.g., Evans and Linsley, i960).

Digging the nest. — Nests appear to be started at any time of day,
but more especially in the mid-morning hours or late in the afternoon.
Digging females periodically produce a “chirping” sound as they
break through the soil with their mandibles. One female starting a
new nest dug 1 cm in 13 minutes, the second cm in another 7 minutes;
she completed the burrow and cell in a total of 58 minutes. This
female had previously been digging at another point 10 cm away, but
she abandoned this nest after apparently striking a stone; the aban-
doned burrow was left open.

When the wasp has loosened a small lump of earth, she holds it
between her mouthparts and front legs, backs from the burrow, rises
into the air a short distance, and with a slight jerk flings it onto the
ground. She may fly in any direction from the entrance, but most
wasps make the majority of flights on the same side, such that the
earth comes to form a diffuse but discernible mound in the form of a
small arc. The mound is rarely more than 0.5 cm deep, and varies
from 2 to 4 cm wide by at least twice that long, although its borders
are ill-defined. In some nests no mound can be seen, as the female

Psyche

[March

Figs. 1-3. Typical nests of Ammophila azteca (all from Jackson Hole,
Wyo.) showing the three types of closures. Fig. 1. No. 2018, an empty nest
closed with a single pebble and a small amount of sand. Fig. 2. No. 1993, a
nest with a single caterpillar bearing the egg, showing a typical temporary
closure. Fig. 3. No. 1996, a fully provisioned nest which had received the
final closure.

scatters the soil rather widely. In any case it weathers away in a few
days, and some of the soil may be used in closures.

There appears to be considerable variation in the distance the
female flies with the soil, and consequently the distance from the nest
entrance the mound accumulates. In Jackson Hole, most mounds
were only a few cm from the entrance (2-6, rarely up to 20). Hicks
(1935) observed very similar digging behavior in Colorado, but he
found that the soil was carried a considerable distance from the nest,
in one case four feet (1.2 m). Hicks timed several females and found
that they carried from five to seven loads per minute. “One load”,
he reports, “was so large and heavy that the wasp was not able to fly
with it after a typical and normal fashion and only reached the refuse
spot by short flights and hops.” Great variation in the distance the
soil is carried was noted by Hicks (1932) in another species, A.
aherti.

Burrow diameter in A. azteca is about 5 mm; the terminal cell is
horizontal and has a diameter of about 10 mm, a length of 20-25 mm.
All burrows at Jackson Hole were vertical or very nearly so; they
varied in length from 3 to 6 cm, nearly all of them measuring between
4 and 5 cm (Figs. 1-4). The single nest excavated at the Great Sand
Dunes, Colorado, was very similar, having a vertical burrow 3.2 cm
long reaching a horizontal cell 2.2 cm long. The nest from Yellow-
stone was very similar in size but was dug into the sloping of a foot-
path and had an oblique burrow (Fig. 5 in Evans, 1963).

Closure of the nest. — As in many species of this genus, three types
of closure' can be distinguished : initial closure of the empty nest,
temporary closures during provisioning, and final closure of the fully
provisioned nest. The initial closure consists of a single pebble or
hard lump of earth which is placed in the burrow entrance with the

1965]

Evans — A mmophila azteca

13

mandibles, often after “trying for size” several such lumps and
rejecting each of them until one is found which fits the mouth of
the burrow (Fig. 1). A small amount of sand is often scraped over
this closure, but no pounding occurs. The wasp normally returns to
this nest within a few hours or early the next day with the first prey,
on which the egg is laid. The single nest studied at the Great Sand
Dunes, Colorado, was like the many observed at Jackson Hole: the
female removed a single small stone from the entrance before intro-
ducing the first caterpillar and laying an egg upon it.

Following oviposition, and after supplying each additional cater-
pillar, a very different type of closure is made (Fig. 2). Again,
various lumps and pebbles are “tried for size”, but this time the
initial lump is placed well down in the burrow, at least half way
down and often near the bottom; the object selected must lodge
perfectly in the burrow so that material may be packed above it (this
is the hauptverschlussklumpchen of Baerends and other European
workers) . When the initial lump is in place, the wasp scrapes sand
into the nest with her front legs, facing of course away from the hole.
She may utilize some of the soil of the mound if this is close by ; or if
loose soil is not available she may loosen some by biting the ground
around the entrance. Periodically small pebbles or bits or earth are
picked up and placed in the burrow with the mandibles. From time
to time the wasp remains with her head down in the burrow and
moves her body up and down, packing the soil in place with blows of
the antero-ventral surface of the head in the usual manner of wasps
of this genus. When the burrow is full, additional packing occurs,
the wasp sometimes retaining her grasp on a small pebble while so
doing (“using a tool”) and then leaving the pebble in place in the
fill (Fig. 8). Finally, sand is scraped in various directions over the
top, resulting in thorough concealment of the entrance. The closure
is prepared very rapidly and requires only a minute or two; one wasp
completed a typical temporary closure in only thirty seconds. It is
probable that the same pebbles and lumps of earth serve in successive
closures, so one would expect a reduction in the length of time re-
quired after the first temporary closure.

Final closure may be indistinguishable from temporary closure
unless one follows the provisioning of the nest to determine that this
is, in fact, the final closure. My limited data suggest that in the
final closure ( 1 ) the initial lump is always placed at or near the
bottom of the burrow (Fig. 3), and (2) packing with the head is
relatively prolonged and some packing while holding a pebble in the
mandibles always occurs toward the end of the closure (Fig. 8).

14

Psyche

[March

Further studies will be needed to quantify these factors and to deter-
mine if there is a real difference between temporary and final closure.
The closure observed by Hicks (1935) in Colorado was presumably
temporary, since no “tool using” was noted. The wasp studied by
Hicks used “an old hackberry seed to close the upper end of the shaft.
Sand was scraped in over this, and some score or more of objects were
brought to further cover and conceal the nest site”.

Provisioning the nest. — Females were observed on several oc-
casions flying closely about low branches of willows and cottonwoods,
apparently searching for prey. On one occasion a wasp seized a
caterpillar on a willow branch, but the latter thrashed violently and
the wasp left without stinging it; this was a considerably larger
caterpillar than was ever found in the nests of the wasp. The 44 prey
taken from various nests were all of roughly the same size (slender,
12-18 mm in length) ; all were “naked” larvae, and all were green in
color except for a few gray or reddish geometers. Specific identifi-
cation of the prey was not possible, but many (perhaps all) belonged
to groups which feed upon broadleafed trees rather than upon conifers.
The following were recovered from the various nests dug out:
Hymenoptera [det. B.D. Burks]

Tenthredinidae : Neimatinae: Nematus (Pteronidia)

sp. and Amauronematus sp. 28

Lepidoptera [det. D.M. Weisman]

Geometridae (four spp.) 7

Gelechiidae (apparently all one sp.) 8

Sphingidae: Smerinthus sp. (early instar) 1

The use of sawfly larvae in considerable numbers is of interest,
since Adriaanse (1947) found that A. pubescens restricts itself to
caterpillars, the closely related A. campestris to sawflies. In the
population of A. azteca studied, several nests contained nothing but
sawfly larvae, a few nothing but moth larvae, and a very few (e.g.,
no. 2012) both sawfly and moth larvae. Individual wasps tended to
stock successive nests wholly with sawfly larvae (rarely wholly with
caterpillars), but there were numerous exceptions. I noted no shift
in type of prey concordant with the progress of the season.

Prey records from other parts of the range of this species indicate
use of lepidopterous larvae of several different groups, including
relatively hairy forms, but there are no further records of use of
sawfly larvae. The single female studied at the Great Sand Dunes
brought in a gelechiid larva very similar to those used in Jackson
Hole. The nest studied at Yellowstone contained one noctuid, one
geometer, and five larvae of lycaenid butterflies (Evans, 1963).

1965]

Evans — A?nmophila azteca

15

Powell (1964) found a lycaenid larva to be used in Baja California,
while in Mono County, California, a larva of the moth family
Pterophoridae was employed. As Powell points out, lycaenid larvae
are covered with short, secondary setae, while pterophorid larvae
have tufts of elongate setae. The caterpillars taken by Powell
measured 14 and 15.5 mm in length, while those taken from the nest
in Yellowstone measured from 8 to 15 mm in length. Thus it appears
that size of the prey is of critical importance, but not its vestiture or
taxonomic affinities.

Powell reported that the lycaenid larva was carried in flight, about
a meter above the ground. I observed prey carriage in flight once in
Colorado and many times at Jackson Hole; in no case did I observe
prey carriage over the ground. The female straddles the caterpillar,
grasping it with her mandibles a short distance behind the head and
during flight embracing the body of the caterpillar with her legs, the
prey being venter-up. She lands a short distance from the nest and
walks to the entrance, usually without circling or hesitation, then
drops the caterpillar at the threshold while she removes the closure.
At Jackson Hole, I experienced much difficulty approaching females
closely for photography at this time; they would very readily take
flight with their prey, often disappearing for several minutes.

Removal of the closure occurs very rapidly, taking only a few
seconds if it is an initial closure, generally less than 30 seconds in any
case. Soil particles and objects removed from the burrow are deposited
within a few cm of the entrance and may be reused when the closure
is restored. When the burrow is clear, the wasp grasps the anterior
end of the prey with her mandibles and backs into the nest with it.
The entire process (arriving with prey — clearing entrance — draw-
ing in prey — reclosing entrance) occurs with remarkable speed, some
individuals requiring less than a minute. When it is considered that
only from one to four prey are introduced per day, it will be appreci-
ated that even prolonged periods of observation afford one only
fleeting opportunities to study this behavior.

In addition to entry with prey, each nest is entered periodically
when the wasp is without prey. There are presumably “inspections”
which function to impress upon the wasp the size and food require-
ments of the larva, as Baerends (1941) found to be true in A.
pubescens. Inspections may occur at any time of day, but most
characteristically occur early in the morning (0830-1030). In fact,
the first act of each Ammophila arriving in the morning consists in
entering the nest and closing it again. I did not observe any one
female inspect more than one nest at this time; apparently when there

Psyche

[March

16

4

NEST DEPTH

T3 A I A2 A3 A4 A5
SEGMENT OF PREY

Fig. 4. Number of nests of Ammophila azteca (all Jackson Hole, Wyo.)
exhibiting various depths (in centimeters). In each case the measurement
was rounded off to the nearest half cm in the field. Fig. 5. Number of
examples of oviposition on various segments of the prey (all Jackson Hole,
Wyo.). T3 — third thoracic segment; Al . . . = various abdominal seg-
ments.

is more than one active nest, she inspects the one containing a larger
larva (see detailed data on no. 2030, below). However, other in-
spections may occur later in the day (see also below). The entry and
reclosure resemble closely the behavior occurring when the wasp is
bringing prey.

The maximum number of prey found in any nest at Jackson Hole
was six (2030D), but the nest studied in Yellowstone contained
seven (Evans, 1963). It appears that about two days are required
for hatching of the egg, about five days for larval feeding. Apparently
females bring in the last prey and make the final closure when the
larva is in the last instar and at two-thirds its full size; thus final
closure usually occurs six or seven days after the nest is dug.

Oviposition. — The egg is laid on the first prey placed in the cell.
It is glued firmly by the anterior end to the side of the caterpillar, in
a more or less vertical position (Fig. 9, lower figure). As in many
species of A?nmophila, there is much variation in the placement of the
egg. At Jackson Hole, one egg was found on the third thoracic
segment, others on each of the first five abdominal segments, but more
on the second abdominal segment than elsewhere (Fig. 5). The single
nest studied in Colorado contained a caterpillar bearing an egg on the
fourth abdominal segment.

Successive nests of individual females. — As mentioned earlier, it
became apparent in the course of studies at Jackson Hole that each
female prepares all of her nests in a small nesting arena. It also
became apparent that each female prepares an unusual number of
nests. For example, when I dug out the nest of no. 2013, an un-

1965]

Evans — A mmophila azteca

7

marked female studied early in the season, I found not one nest but
six, all in an area measuring 8X14 cm. This was on July 27, and
I could not believe that females had been nesting for more than two
weeks (I began collecting in this area July 4, and took the first
A. azteca , a male, on July 7) . But if it takes six or seven days to rear
a larva, this female should have started nesting at least 30 days
earlier (one nest was still unprovisioned). This example and several
similar ones made it apparent that some overlapping of successive
nests must occur. I therefore marked several individuals and at-
tempted to follow them over several days. Since my observations were
not absolutely continuous, and since during the one complete day of
observation I could keep only two wasps under full surveillance,
because of the wide spacing mentioned earlier, my notes are not
entirely convincing except in two cases. The arrangement of suc-
cessive nests of these two individuals is shown in Figs. 6 and 7> and
I here present, in abbreviated form, my field notes concerning one of
them, no. 2030, which I painted with a red spot on July 19.

July 19. Brought prey at 1100 hrs to site A (Fig. 6).

July 20. Working on a nest at B.

July 23. Prey to B at 1530.

July 25. Prey to nest C at 1630.

July 29. Inspection followed by final closure at C.

July 31. Complete day of observation. Arrived in area at 0815.
0845 : Wasp appeared at site D and made inspection, followed

by temporary closure.

1022: Prey to D, followed by what appeared to be a final

enclosure, including “use of tool” and scraping of sand
over top for several minutes. Left at 1036.

1037: Reappeared at same site, walked about, flew off.

1047: Reappeared at same site, walked over old nest and around

area until 1050.

1053 : Back again, walking over nesting arena with her abdomen

held high; off at 1055.

1 1 13: Reappeared and began to remove closure at E. This

appeared to be a rather thorough temporary closure. After
a brief “inspection” she made a hasty temporary closure
and left at 1120.

1237: Back to E with a sawfly larva; opened nest and drew it

in within one minute of arrival. At closure three pebbles
were “tried for size” before finding one which fit the bore
of the burrow properly. Left at 1240.

1 355 : Back to E with another sawfly larva. After introducing

i8

Psyche

[March

the prey made a fairly prolonged closure which I took to
be a final closure but which proved not to be (see Aug. I ).
During this she drove away two chrysidids (see below,
under “parasites”).

1400, 1409, 1414, 1445: Each time she reappeared in the nesting
arena, walked about for a few seconds to a minute, then
flew off.

Started digging at G.

Stopped digging, leaving hole open, and is now walking
about the nesting arena.

Digging at G1.

Closed burrow with a single pebble, scraped a small
amount of sand over top, and flew off.

Landed at nest, then flew to a Solidago in blossom two
meters away, remained for a few minutes, then was not
seen again today.

August 1. 1020: Landed at G1, remained two minutes and flew off.

This is a cool, partly cloudy morning.

1040: Now cloudy and windy (began to rain at 1100). I dug

out the nesting arena, eventually finding 12 nests and one
incompleted burrow (G) (see Fig. 6). Eight of these
had cocoons, one was an empty nest (G1), one had a
single caterpillar with an egg (F), and two had wasp
larvae (D and E, closed yesterday, E apparently tempo-
rarily, since the wasp larva was small and there were only
3 sawfly larvae in the cell).

Apparently nest F had been initiated on July 30 and was not
visited on July 31 (the wasp presumably remembering that it con-
tained an egg and did not require additional prey). Thus we can say
that this wasp very definitely maintained two nests simultaneously (D
and E ) , very probably three ( if we include F ) , and that she was
associated with three nests in the course of one day (D, E, G1) while
presumably retaining a fourth in her memory. Clearly this calls for a
much more detailed study, but it would appear that the models
provided by Baerends (1941) for A. pubescens apply very well to
this species.

Some unexplained features of behavior. — On two different oc-
casions I dug out a series of nests in a single nesting arena and found
that all or most of the nests contained eggs. In one case there were six
nests, five of them containing a single prey with an egg, the remaining
one being empty. In another case there were four nests, all of them
containing only one prey; in three cases the prey bore an egg, in the

1450:

1457:

1502 :
1600:

1632:

1965]

Evans — A mmophila azteca

19

Fig. 6. Arrangement of successive nests of no. 2030. Nests A-G1 were
prepared in that order (see text) ; unlettered nests were not seen to be dug
or provisioned and were presumably completed before the beginning of
intensive observations (at least in part). Fig. 7. Arrangement of nests of
no. 2031, which nested about 2 meters from the preceding at Jackson Hole.

fourth case a small larva. Unfortunately in both instances the wasp
associated with the nesting arena was unmarked, and I did not observe
the site for a long enough time to be certain that only one female was
responsible for each set of nests. As mentioned earlier, I did find one
case in which an unmarked female shared a nesting arena with a
marked female. If the above two cases did involve a single female
each, then that female must have prepared several nests on the same
day, or over no more than two or three days, and laid several eggs in
rather rapid succession. When the larvae in these nests approached
maturity, the female would have to obtain many prey in order to
provision all of these nests. Whether truly synchronous nesting (as
opposed to the overlapping of successive nests) occurs in this species
remains to be determined.

On one occasion (2007D) I dug out a nesting arena and found
three nests, one of which contained one caterpillar, the other two two
sawflies each : but none of the nests contained an egg. This suggests
that these might have been storage burrows, the prey to be exhumed
later and used for oviposition or for feeding a larva. If in fact this
species does at times maintain several nests synchronously, then the
maintenance of storage nests would seem of great adaptive value,
since it would spread out the requisite hunting over a greater number
of days. It is possible that the maintenance of storage burrows is not
a rare phenomenon in A?nmophila. Hartman (1905) found two

20

Psyche

[March

Fig. 8. Female Ammophila azteca at final closure of her nest. A small
pebble is being held in the mandibles and used for packing soil in the filled
burrow.

nests of A. procera in Texas which did not contain eggs, Tilden
(1953) had the same experience with this species in California, and
Criddle (1924) observed a female of this same species in Manitoba
place a caterpillar in a nest one day and then dig it out the next day,
return it to the nest, and lay an egg upon it This is an aspect of
behavior requiring much further study.

Parasites. — Chrysidid wasps (Ceratochrysis perpulchra Cresson)
were observed commonly in the nesting area of A. azteca at Jackson
Hole, but I am unable to state definitely that they were parasitizing
the Ammophila. The chrysidids would often land on the soil near
Ammophila nests, and on several occasions they were seen to enter
nests either partially dug or partially closed, in which case they were
normally driven away by the Ammophila. On one occasion a chrysidid
returned after the Ammophila had completed her closure and spent
several minutes around the nest, but I did not see her dig into it.

1965]

Evans — Ammophila azteca

21

Notes on Ammophila macra. — As noted earlier, a single female
of the large species A. macra Cresson was found nesting on the
periphery of the azteca nesting area at Jackson Hole. She was seen
digging her nest at noo on August 14. The soil was taken from the
burrow in lumps much as in azteca , but it was carried over the ground
and deposited in a diffuse pile 6-14 cm from the hole. At 1115 the
wasp selected a large stone and placed it in the burrow (which was
fully 1 cm in diameter). She then dropped in several small pebbles
and proceeded to scrape in sand, from time to time turning around
and packing the sand in place with her head while making a loud
buzzing sound. When the burrow was nearly full, she added several
more stones, pressing each into place. Finally, she dug soil from a
small, semicircular hole 3 cm from the entrance, making a shallow
quarry or “false burrow”, and scraping this soil into the nest entrance.
This was packed into place by holding a stone in her mandibles and
pounding, finally leaving the stone in place. The wasp then picked
up a stick 4 cm long and dragged it over the top of the covered nest
entrance, then added a second stick 7 cm long. She completed the
closure at 1135 and was not seen again. The nest was dug out two
days later and found to contain a single large sphingid larva ( Smerin -
thus geminatus Say) bearing an egg on the side of the fourth ab-
dominal segment (Fig. 9, upper figure). The burrow was vertical,
6 cm long, the cell horizontal and 5 cm long.

Thus this species differs from azteca in several important features:
the soil is carried from the nest on the ground ( and the prey doubtless
carried on the ground) ; some of the soil for closure is obtained from
a quarry and various objects are placed on top of the closed nest; and
the nest is stocked with a single large caterpillar, thus eliminating the
possibility of progressive provisioning or of the overlappng of pro-
visioning of successive nests. A. macra bears a close resemblance to
the well-studied species procera Dahlbom, and the behavior is similar
to that species, although procera more commonly flies with the soil
when digging.

Summary. — Ammophila azteca differs from other known North
American species of its genus in that several nests are maintained
simultaneously; in this respect it bears a close resemblance to the
European species puhescens. Data indicate that at least two and at
times probably three or more nests in various stages are maintained
at one time, the female remembering the precise location of each and
inspecting each nest, or at least most nests, each day before provision-
ing. The total duration of each nest is six or seven days.

The prey consists of caterpillars and sawfly larva of many diverse

22

Psyche

[March

Fig. 9. Above, larva of sphingid bearing the egg of Ammophila macra.
Below, larva of a geometrid bearing the egg of A. azteca. These were
extracted from nests in close proximity at Jackson Hole.

groups, all of them diurnal leaf-feeders of about the same size (8 to
1 8 mm, usually 12 to 16 mm). The egg is laid on the first prey in
each nest and shows much variation with respect to the segment on
which it is laid. Up to seven prey are provided per nest.

Soil is carried from the burrow in flight, and the prey is carried to
the nest in flight.

Initial closure of the nest consists of a single object placed at the
top of the burrow. Temporary closures after oviposition consist of a
large object blocking the lumen of the burrow on top of which loose
soil and smaller objects are packed, with or without “use of a tool”.
At final closure, a “tool” is apparently always employed.

Fragmentary data suggest the possibility that some females maintain
storage burrows, this perhaps being correlated with maintenance of
synchronous nests, all at about the same stage, by some females. More
data on these and many other points are needed.

Brief observations on one female A. macra nesting in the same area
revealed several important differences between this species and azteca.

References

Adriaanse, A.

1947. Ammophila campestris Latr. und Ammophila adriaansei Wilcke:
ein Beitrag zur vergleichenden Verhaltensforschung. Behaviour,

1: 1-34.

1965]

Evans — Ammophila azteca

23

Baerends, G. P.

1941. Fortpflanzungsverhalten und Orientierung der Grabwespe
Ammophila campestris Jur. Tijdschr. Ent., 84: 68-275.

Criddle, N,

1924. Observations on the habits of Sphex procera in Manitoba. Canad.
Field Nat., 38: 121-123.

Evans, H. E.

1958. The evolution of social life in wasps. Prox. Xth Internat.
Congress Ent., 2: 449-457.

1959. Observations on the nesting behavior of digger wasps of the
genus Ammophila. Amer. Midi. Nat., 62: 449-473.

1963. Notes on the prey and nesting behavior of some solitary wasps of
Jackson Hole, Wyoming. Ent. News, 74: 233-239.

Evans, H. E. and E. G. Linsley

1960. Notes on a sleeping aggregation of solitary bees and wasps. Bull.
So. Calif. Acad. Sci., 59: 30-37.

Hartman, C.

1905. Observations on the habits of some solitary wasps of Texas. Bull.
Univ. Texas, 65: 1-73.

Hicks, C. H.

1932. Notes on Sphex aberti (Hald.). Canad. Ent., 64: 145-151.

1935. Notes on rare western sphecid wasps. Pan-Pac. Ent., 11 : 97-102.
Janvier, H. (F. Claude-Joseph)

1928. Recherches biologiques sur les predateurs du Chili. Ann. Sci.
Nat, Zool., (10) 11: 67-207.

Olberg, G.

1959. Das Verhalten der solitaren Wespen Mitteleuropas (Vespidae,
Pompilidae, Sphecidae). Berlin, Deutscher Verlag Wissenschaf-
ten, 402 pp.

Powell, J.

1964. Additions to the knowledge of the nesting behavior of North
American Ammophila (Hymenoptera : Sphecidae). Jour. Kansas
Ent. Soc., 37: 240-258.

Teschner, W.

1959. Starrheit und Variabilitat im Verhalten von Sandwespen. Zool.
Beitrage, (n.f.)4: 411-472.

Tilde n, J. W.

1953. The digging and provisioning behavior of Ammophila saeva
Smith (Hymenoptera: Sphecidae). Pan-Pac. Ent., 29: 211-218.

Tsuneki, K.

1956. Ethological studies on Bembix niponica Smith, with emphasis on
the psychobiological analysis of behaviour inside the nest
(Hymenoptera, Sphecidae). Mem. Fac. Lib. Arts, Fukui Univ.,
Ser. II, Nat. Sci., no. 6, pp. 77-172.

Wheeler, W. M.

1923. Social life among the insects. New York, Harcourt Brace & Co.,
375 pp.

THE ANT LARVAE OF THE SUBFAMILY
LEPTANILLINAE (HYMENOPTERA, FORMICIDAE)

By George C. Wheeler and Jeanette Wheeler
University of North Dakota, Grand Forks

The Leptanillinae are a small subfamily comprising 14 species in
3 genera: Leptanilla Emery, 11 species; Leptomesites Kutter, 1
species; Phaulomyrma G. C. & E. W. Wheeler, 2 species.

This subfamily has been recorded only from the warmer parts of
the Old World: Corsica, Sardinia, North Africa, India, Malaya,

Java, Queensland, Western Australia and Japan. Seemingly it is
rare, but the paucity of the records may be due to minute size and
hypogeic habits. As W. M. Wheeler optimistically remarked (1932,
p. 54) : “We should expect careful collecting with the Berlese funnel
to bring additional forms to light in South Africa, Madagascar, Asia
Minor and India, or even, perhaps, in the warmer parts of the New
World.’’ Brown (1954, p. 28) noted: “The habits of the species
are such as to render their discovery highly fortuitous under present
collecting methods.’’ In the 30 years since Wheeler wrote, only two
species have been discovered (one in India and one in Japan), which
would rather support another statement by Wheeler in the same
article (p. 57-58) : “The Leptanillinae . . . must be very ancient,
like many other components of the microgenton. . . . L. swani is
particularly interesting in this connection, because the extreme south-
western corner of Australia, in which it was taken, is known to possess
the oldest and least disturbed fauna of any portion of the continent.’’
Antiquity is compatible with a disjunct distribution and with rarity.

The genus Leptanilla was established by Emery in 1870 and then
for half a century was kicked about over the taxonomic table. Origi-
nally Emery placed it in the “Dorylidae” near Typhlopone. Mayr,
however, in a letter to Emery (date not given — see Emery 1904)
dissented, maintaining that it belonged with the Myrmicinae. Emery
was evidently convinced, for in 1875 he removed it to the “Mirmi-
cidei” in the neighborhood of Stenamma and Liomyrmex. In 1877 he
moved it to the vicinity of Monomoriuin and Leptothorax in the
“Myrmicidei genuini” but marked it with a query to signify un-
certain position. In 1881 Ern. Andre had it in the first tribe,
“Myrmicidae verae,” of the “Myrmicidae” but mentioned its affini-
ties with the “Dorylides.”

Forel in 1893 did not mention the genus but might have meant

24

1965]

Wheeler and Wheeler — Ant Larvae

25

to include it in the subfamily Myrmicinae, when he said that the
fourth tribe (“Myrmicii”) included “les autres genres de la sous-
famille des Myrmicinae.” And later (1901) he excluded it from the
Dorylinae when he says, “Done, je maintiens la sous-famille Dory-
linae limitee aux genres Dorylus, Aenictus, Eciton et Cheliomyrmex
In Dalla Torre’s “Catalogus” (1893) it was still in the Myrmicinae
but near Trigonogaster and Pheidologeton. In 1895 Emery was still
of the opinion that it belonged in the subfamily “Myrmicini” in the
second tribe (“Myrmicii”) near Huberia and Phacota. But nine
years later (1904), after describing the female of L. revelierei, he
returned it to its original subfamily (Dorylinae). In the following
year in Ashmead’s skeleton it stood between “ fLiomyrmex” and
Epipheidole in the tribe Stenammini, subfamily Myrmicinae, family
“Myrmicidae.” In 1907 Santschi described males of 3 species, which
he referred to the genus Leptanilla and claimed that their doryline
affinities justified Emery’s original allocation of the genus. It should
be noted, however, that males of Leptanilla have never been taken
with females or workers; hence it is not certain that those described
by Santschi belong to this genus. In the “Genera Insectorum” Emery
(1910) established for the genus a separate tribe (Leptanillini) in
the subfamily Dorylinae, where it seemed destined to abide in iso-
lation: Wheeler (1910), Forel (1917 and 1923), and Wheeler
(1922) did not disturb it. Wheeler (1923, p. 335) suggested that
even further isolation might be necessary: “I believe that the tribe
Leptanillini, which Emery includes among the Dorylinae, will have
to be separated out as a distinct subfamily (Leptanillinae) . Dr.
George C. Wheeler finds that the larva of Leptanilla is very aberrant,
and the characters of the adult are either quite unlike those of other
Dorylinae or only superficially similar and due to convergence, or
similarity of subterranean habits.” By 1932 (p. 57) the separation
had been effected : “Emery, as is well known, regarded the Leptanil-
linae as constituting a special tribe of the Dorylinae, but Dr. G. C.
Wheeler and I have raised the group to subfamily rank.” Bernard
(1951) raised the taxon to family rank.

Brown and Nutting (1950, p. 124) in their study of the wing
venation for the family Formicidae wrote: “The position of the
leptanillines is debatable. They are usually treated as a subfamily by
modern authors, and the venation is so highly reduced in the forms
we have seen that little may be deduced from them concerning
relationships to the other subfamilies.” Brown was apparently still
puzzled in 1954 (p. 28) for he wrote: “This little subfamily has
suffered such drastic anatomical reduction in most of the usually

26

Psyche

[March

valuable phylogenetic characters that it is doubtful whether we shall
ever be certain of its true affinities. ... It is possible that forms as yet
unknown will reveal their ancestry more clearly. Until that time,
however, subfamily rank for the Leptanillinae may as well be main-
tained. Present opinion seems to favor relating this group to the
Dorylinae ”

The larvae of the Leptanillinae do bear certain resemblances to the
known doryline larvae (Dorylus, Aenictus , Eciton, Cheliomyrmex) :
the long slender body and the small feebly sclerotized mandibles. On
the other hand, they differ in the constriction at the metathorax, the
long hairs on the abdomen, the absence of hairs on the head and the
shape of the head. But these differences become trivial and insignificant
beside the three which not only differentiate them from the Dory-
linae, but also from all other known formicid larvae (160 genera) :

( 1) the peculiar projection from the ventral surface of the prothorax;

(2) the reduction of the spiracles from the normal ten pairs to a
single pair, which is located on abdominal somite III; and (3) the
shape and stance of the mandibles.

Subfamily leptanillinae Emery

Elongate and very slender; slightly constricted at the metathorax;
anterior end curved ventrally ; remainder of body straight and clavate.
With a curious complex structure projecting anteroventrally from the
ventral surface of the prothorax. Only one pair of spiracles, located
on abdominal somite III; each spiracle opening eccentrically on a
naked circular area. Body hairs simple ; the minute hairs very
abundant and uniformly distributed ; a few conspicuous long hairs
sparsely scattered. Head naked and elongate. Antennae small,
slightly^ raised, each with two sensilla. Labrum slightly broader than
long, with the ventral border semicircular; posterior surface spinulose,
with the spinules in rows. Mandibles apparently turned laterally
(instead of medially, as is usual with ant larvae) ; feebly sclerotized;
each with a rather long slender sharp-pointed apical tooth, which
curves laterally; lateral (=outer) border furnished with several long
slender sharp-pointed teeth; anterior surface with rows of spinules.
Labium thin, flap-like and narrowed basally; lateral surfaces sclero-
tized; each palp a low ventrolateral projection bearing five sensilla.

Of all the ant larvae studied we have found the Leptanillinae the
most difficult to process partly because of their small size but chiefly
because of their extreme slenderness. A slender larva is more apt to
collapse than a stout one. Furthermore the constriction at the
metathorax aggravates the difficulty of cleaning and predisposes to
breakage in that region. As a result processing has often left us only

1965]

Wheeler and Wheeler — Ant Larvae

27

fragments to mount on slides. Fragments are all right — in fact,
necessary for some parts — if none is lost and if they can be correctly
oriented.

References. — W. M. Wheeler (1923, p. 335) used larval
characters in establishing the subfamily (quoted above and also by
G. C. Wheeler, 1928, p. 88-89 and referred to by G. C. and E.
Wheeler, 1930, p. 198).

G. C. Wheeler (1928, p. 89) justified W. M. Wheeler’s establish-
ment of the subfamily. (Repeated G. C. and E. Wheeler, 1930,
p. 199.)

Kutter 1948 p. 294: “Alle bis jetzt bekannt gewordenen Larven
der Leptanillinae haben den teilweise chitinisierten, ventralen
Thorakalanhang gemein, wie offenbar auch die Senkrechtstellung der
Mandibeln, wahrend der Besitz des als Tympanalorgan bezeichneten
Organs noch nicht als typisch fiir alle Larven der Unterfamilie
bezeichnet werden darf.”

Bernard, 1951, p. 1041 : “Larves eucephales, carnivores; nourries
par les ouvrieres.”

Genus Leptanilla Emery

We are unable to separate Leptanilla generically from Leptome-
sites: the difference between the larvae of the two known species of
the former are as great as the difference between either species and
the larva of the latter. Therefore the subfamilial description will
suffice for the genus.

Bernard (1951, p. 1017) described primitive larvae and mentioned
the larva of Leptanilla as an example.

Kutter ( 1948, p. 292) differentiated the two genera by the absence
of the naked area around the spiracle (“tympanum”) and the structure
of the ventral prothoracic projection. As we show below, this dis-
tinction is no longer tenable.

The two species of Leptanilla differ with respect to the following
characters: size and shape of terminal boss; complexity of ventral
prothoracic projection; size and arrangement of long body hairs;
shape of head ; number of teeth and spinules on the mandibles; and the
sclerotized band bordering the spiracular area.

Leptanilla revelierei sardoa Emery

(Fig. 1-8)

Length approximately 1.3 mm. Body elongate and slender; thorax
slightly curved ventrally, slightly constricted at the metathorax;
abdomen straight and clavate, the diameter increasing gradually to
abdominal somite V and decreasing to the posterior end which is

Psyche, 1965

Vol. 72, Plate 2

Wheeler — Ant Larvae

1965]

Wheeler and Wheeler — Ant Larvae

29

rounded and terminates in a small naked hemispherical boss. With
a complex structure projecting anteroventrally from the ventral
surface of the prothorax (see Fig. 4-5). With only one pair of
spiracles, located on the third abdominal somite near the posterior
border; each spiracle opening eccentrically on a naked circular area,
which is bordered by a narrow heavily sclerotized band. Body hairs
simple. Of three types : ( 1 ) numerous, increasing in length from

0.005 mm on the thorax to 0.02 mm at the posterior end, without
alveolus and articular membrane, uniformly distributed but lacking
from the anterior portion of the prothorax, the circumspiracular area
and the terminal boss; (2) 0.01-0.15 mm long (shortest on the
prothorax), few, conspicuous, the longest attenuated and flexible
distally, with alveolus and articular membrane, absent from the dorsal
surface of abdominal somites III-IX; (3) about 0.3 mm long, with
attenuated flexible tip, two (one dorsal and one ventral) on the
posterior end near the terminal boss. Head naked; elongate; widest
above the antennal level ; cranium subpyriform in anterior view.
Antennae small ; slightly raised ; each with two sensilla, each of which
bears a spinule. Labrum slightly broader than long; the ventral
border semicircular; posterior surface spinulose, the spinules in long
rows, the rows concentric with the ventral border. Mandibles with
the toothed border directed laterally; feebly sclerotized; each with a
rather long slender sharp-pointed apical tooth, which is curved
laterally; lateral border with four long slender sharp-pointed teeth;
a few rows of rather large spinules on the anterior surface. Maxillae
adnate to the labium; palp a stalked hemispherical knob directed
laterally and bearing five sensilla; no galea seen. Labium a thin flap,
narrowed basally; lateral surfaces sclerotized; each palp a low
ventrolateral projection bearing five sensilla; an isolated sensillum
between each palp and the opening of the sericteries; the latter a
transverse impression on the anterior surface.

Explanation of Plate 2

Figs. 1-8. Leptanilla revelierei sardoa. 1, head in anterior view, X417;
2, larva in side view, X76; 3, left mandible in anterior view, X847; 4,
ventral prothoracic structure in side view, X423 ; 5, ventral prothoracic
structure: left half in posterior view, right half in anterior view, X423 ;
6-8, three body hairs, X500.

Figs. 9-18. Leptanilla swani. 9, head in anterior view, X417 ; 10, larva
in side view, X76; 11, left mandible in anterior view, X 1333 ; 12, left
maxilla in anterior view, X847; 13 and 14, two body hairs, X500; 15 and
16, spiracle and circumspiracular area in side and surface phantom views
(hairs omitted) X333; 17 and 18, ventral prothoracic structure in anterior
view and in side view, X667.

30

Psyche

[March

Material Studied. — Three larvae labeled “Sardegna: Golfo

Aranci. I. 1909 A. Dodero.” These are the specimens studied by G.
C. Wheeler (1928). We studied them first with a phase microscope;
then they were dismounted, stained lightly, remounted and studied
under both phase and light microscopes.

Literature. The description and figures by G. C. Wheeler (1928,
p. 85-87) have been completely revised for this article. Bernard,
1951, Fig. 949 C, D after G. C. Wheeler, 1928.

Leptanilla swani Wheeler
(Fig. 9-18)

Length about 1.4 mm. Body elongate and very slender; anterior
end curved ventrally, slightly constricted at the metathorax ; remainder
of body straight and clavate; diameter increasing gradually to
abdominal somite VII and diminishing to the posterior end, which is
round-pointed and terminates in a small naked boss. With a complex
structure projecting anteroventrally from the ventral surface of the
prothorax (see Fig. 17-18). With only one pair of spiracles, located
near the posterior border of abdominal somite III; each spiracle
opening eccentrically on a naked circular area. Body hairs simple.
Of two types: (1) abundant and uniformly distributed (except on
the terminal boss and the circumspiracular area), minute (0.005-0.024
mm long), longest near the posterior end, without alveolus and
articular membrane; (2) long (0.084-0.23 mm), slender, with the
apical portion fine and flexible, with alveolus and articular membrane,
few, conspicuous, absent from the dorsal surface of the abdomen.
Head naked; elongate; widest at the level of the antennae; cranium
subpyriform in anterior view. Antennae small and slightly raised;
each with two sensilla, each of which bears a spinule. Labrum slightly
broader than long, with the ventral border semicircular; posterior
surface spinulose, the spinules rather numerous and long, arranged in
rows concentric with the ventral border. Mandibles with the toothed
border directed laterally; feebly sclerotized; each with a rather long
slender sharp-pointed apical tooth, which curves laterally; lateral
border with six long slender sharp-pointed teeth; anterior surface
spinulose, the spinules numerous, rather long and arranged in rows;
posterior surface with one row of long spinules. Maxillae adnate to
the labium; palp an irregular knob projecting laterally and bearing
five sensilla; no galea seen. Labium a thin flap, narrowed basally;
lateral surfaces sclerotized; each palp a low ventrolateral projection
bearing five sensilla; an isolated sensillum between each palp and the
opening of the sericteries; the latter a transverse slit on the ventral
border.

1965]

Wheeler and Wheeler — Ant Larvae

31

Material Studied. — Three larvae from Glittering, Western
Australia collected by D. C. Swan. These are the specimens referred
to by W. M. Wheeler in 1932 (p. 56-57). We studied them first
with a phase microscope; then they were dismounted, stained lightly,
remounted and studied under both phase and light microscopes.

In 1963 Rev. B. B. Lowery of St. Ignatius College (Sydney,
Australia) generously sent us 40 larvae of this species, which he had
collected at Cunningham’s Gap in southern Queensland. The two
best specimens have been kept in alcohol. Many others were processed
according to our standard technique (i960) and studied under both
phase and light microscopes.

Literature. W. M. Wheeler (1932, p. 56-57) compared the larva
of L. swani with that of L. sardoa by quoting from a letter from G.
C. Wheeler to whom he had sent the larvae for study.

Genus Leptomesites Kutter

As explained above under the genus Leptanilla we are unable to
separate Leptomesites generically from Leptanilla. Therefore our
subfamilial description will have to suffice for this genus.

Kutter (1948, p. 287) has characterized the genus thus: “Korper
sehr langgestreckt-zylindrisch, gegen das Ende keulenformig aufge-
trieben. Labrum ohne gezahnte, laterale Lappchen. Mandibeln
abwarts gerichtet, gezahnt. Auf der Ventralseite des Prothorax mit
breitem, in der Mittelpartie mit chitinisiertem, quergerilltem, lappigem
Anhang. Beiderseits dieses einzigartigen Anhanges je eine sehr lange,
sichelformig ventralwarts gebogene Borste. (Ein Tympanalorgan,
wie es von der Leptanilla-Larve gemeldet wird, konnte nicht mit
Sicherheit nachgewiesen werden.)”

Leptomesites escheri Kutter
(Fig. 19-28)

Length about 1.5 mm. Body elongate and slender, apparently
constricted at the metathorax; anterior end slightly curved ventrally;
abdomen clavate, with the posterior end narrowly rounded. Project-
ing ventrally from the anterior portion of the prothorax is a curious
flap-like structure furnished with 14 ridges on the posterior surface.
Only one pair of spiracles present, on abdominal somite III (or IV?).
Body hairs simple. Of two types: (1) minute (0.009-0.026 mm

long), longest near the spiracle, numerous, uniformly distributed,
without alveolus and articular membrane; (2) short to very long
(0.042-0.096 mm), a few on each somite, the apical portion fine and
flexible, with alveolus and articular membrane. Head naked; elongate;
widest above the antennal level ; cranium subovoidal. Antennae small

32

Psyche

[March

Figures 19-28. Leptomesites escheri. 19, head in side view, X278; 20,
head in anterior view, X278; 21, left mandible in anterior view, X423; 22,
left maxilla in anterior view, X415; 23, restoration of larva in side view
(minute hairs omitted), X33; 24-26, three body hairs, X139; 27 and 28,
ventral prothoracic structure in side view and in anterior view, X272.

and slightly raised ; each with two sensilla, each of which bears a
spinule. Labrum slightly broader than long; the ventral border
semicircular; posterior surface spinulose, the spinules in long rows,
the rows concentric with the ventral border. Mandibles with the
toothed border directed laterally; feebly sclerotized; each with a
rather long slender sharp-pointed apical tooth, which is curved
laterally; lateral border with nine long slender sharp-pointed teeth;
anterior surface furnished with numerous short rows of long spinules,
the rows so close together that the spinules overlap. Maxillae conoidal,

1965]

Wheeler and Wheeler — Ant Larvae

33

pointing laterally; palp apical and consisting of five sensilla; galea
represented by two sensilla, each of which bears a digitiform spinule.
Labium feebly bilobed, flap-like, narrowed basally; palp a low ventro-
lateral projection bearing five sensilla; an isolated sensillum between
each palp and the opening of the sericteries; the latter a transverse
slit on the ventral surface. Hypopharynx spinulose, the spinules in
short transverse rows, the rows so close together that the spinules
overlap.

Material Studied: — Two semipupae (?) from southern India.
These are the specimens studied by Dr. Heinrich Kutter (1948,
p. 290-292) of Mannedorf, Switzerland. In 1963 we asked Dr.
Kutter whether he would be willing to let us examine them. His
generous response was to send them to us as a gift. We studied them
first with the phase microscope; then they were dismounted, stained
lightly, remounted and studied under both phase and light microscopes.
We were not able to inflate the wrinkled specimens; therefore our
drawing of the larva in side view is a restoration.

Literature. Kutter (1948, p. 290-292) : a description of the

species; Fig. 6, larva in side view; Fig. 7, anterior end enlarged.

Literature Cited

Andre, Ern.

1881-1882. Species des hymenopteres composant le groupe des formicides
d’Europe, etc.

Ashmead, W. H.

1905. A skeleton of a new arrangement of the families, subfamilies,
tribes and genera of the ants, or the superfamily Formicoidea.
Canad. Ent. 37: 381-384.

Bernard, F.

1951. Super-famille des Formicoidea. In: P. P. Grasse (ed.), Traite
de Zoologie, 10 (2) : 997-1104. Masson et Cie, Paris.

Brown, W. L.

1954. Remarks on the internal phylogeny and subfamily classification of
the family Formicidae. Insectes Sociaux 1: 21-31.

Brown, W. L., and W. L. Nutting

1950. Wing venation and the phylogeny of the Formicidae. Trans.
Amer. Ent. Soc. 75: 113-132, 2 pi.

Dalla Torre, K. W. von

1893. Catalogus Hymenopterorum hucusque descriptorum systematicus
et synonymicus. Vol. VII: Formicidae (Heterogyna) . W.

Engelmann, Leipzig. 289 p.

Emery, C.

1870. Studi mirmecologici. Boll. Soc. Ent. Ital. 2: 9 p, 1 pi.

1875. Le formiche ipogee con descrizioni di sp. nuove o poco note. Ann.
Mus. Civ. Genoa 7: 465-474.

1877. Saggio di un ordinamento naturale dei myrmicidei e consider-
azioni sulla filogenesi delle formiche. Boll. Soc. Ent. Ital. 9:
1-17, 1 pi.

34

Psyche

[March

1895. Die Gattung Dorylus Fabr. und die systematische Eintheilung der
Formiciden. Zool. Jahrb. Abt. Syst. 8: 685-788.

1904. Le affinita del genere Leptanilla e i limiti delle Dorylinae. Arch.
Zool. Napoli 2: 107-116.

1910. Fam. Formicidae, Subfam. Dorylinae in Wytsman’s “Genera
Insectorum.” Fascicle 102: 34 p. 1 pi.

Forel, A.

1893. Sur la classification de la famille des formicides, avec remarques
synonymiques. Ann. Soc. Ent. Belg. 37: 161-167.

1901. A propos de la classification des fourmis. Ann. Soc. Ent. Belg.
45: 136-141.

1917. Cadre synoptique actuel de la faune universelle des fourmis.

Bull. Soc. Vaud. Sci. Nat. 51: 229-253.

1923. Le monde social des fourmis du globe. 5 : 174 p., 3 pis. Librairie
Kundig, Geneva.

Kutter, H.

1948. Beitrag zur Kenntnis der Leptanillinae. Eine neue Ameisengat-
tung aus sud-Indien. Mit. Schweiz. Ent. Ges. 21: 286-295.

Santschi, F.

1907. Fourmis de Tunisie capturees en 1906. Rev. Suisse Zool. 15:

305-334.

Wheeler, G. C.

1928. The larva of Leptanilla. Psyche 35 : 85-91.

Wheeler, G. C., and Esther W. Wheeler

1930. Two new ants from Java. Psyche 37: 193-201.

Wheeler, G. C., and Jeanette Wheeler

1960. Techniques for the study of ant larvae. Psyche 67 : 87-94.
Wheeler, W. M.

1910. Ants: Their structure, development and behavior. Columbia

University Press, New York. 663 p.

1922. Ants of the American Museum Congo Expedition. Bull. Amer.
Mus. Nat. Hist. 45: 1-1139.

1923. Social life among the insects. Harcourt, Brace & Co., New York.
375 p.

1932. An Australian Leptanilla. Psyche 39: 53-58.

THE AUSTRALIAN ANTS OF THE GENUS
PRISTOMYRMEX , WITH A CASE OF APPARENT
CHARACTER DISPLACEMENT1

By Robert W. Taylor
Biological Laboratories, Harvard University

Introduction

The Old World myrmicine ant genus, Pristomyrmex Mayr 1866,
contains about 40 named forms, mostly from rain forests of the
Ethiopian, Oriental and Australian regions. The Australian mainland
fauna is small compared to that of New Guinea; it includes four
species, three of which are newly described here. There are two
species-groups, those of P. foveolatus n. sp., and P. quadridentatus
(Andre) 1905, each containing two species.

The former group includes P. foveolatus and P. thoracicus n. sp.,
closely related species with strong foveolate-reticulate sculpturation,
having clear affinities with components of the New Guinean fauna,
and known only from rain forests in the Cairns district of North
Queensland. Foveolatus seems to be restricted to the Atherton Table-
land, at elevations greater than 1000 ft, while thoracicus is apparently
more widespread, but is not known from the Tableland proper.

The affinities of the quadridentatus group are less clear; its species
occur further south, in extreme S.E. Queensland, and N.E. New
South Wales, and they may be derived from a separate, older, Mela-
nesian stock. The group includes P. quadridentatus and P. wheeleri
n. sp. Quadridentatus was originally placed in a separate monotypic
genus, Odontomyrmex Andre 1905, which was synonymized with
Pristomyrmex by Emery ( 1922). The lack of foveolate sculpturation
on the head and body, and the presence of pronotal spines were thought
by Andre to taxonomically characterize Odontomyrmex . But these
characters occur together or separately in many Pristomyrmex species,
and cannot be used to logically define a separate taxon (Brown, 1953).
Nonetheless the “Odontomyrmex” habitus seems to characterize a
natural phyletic group within the Australian fauna, and quadriden -
tatus and wheeleri are undoubtedly cognate, or at least closely related.
The former species is typically self-colored golden-brown, but it has
an apparently conspecific bicolored variant, quadridentatus variety

Research supported by U.S. National Science Foundation Grant No. GB
1634 (1963-1965). Australian field studies in 1962, reported here, were
supported by the Committee on Evolutionary Biology, Harvard University.

35

36

Psyche

[March

queenslandensis Forel 1915, which is synonymized below. These two
forms differ only slightly, except in coloration, and distributional
evidence suggests that the bicolored form is a character-displaced
variant, produced wherever the range of quadridentatus overlaps that
of wheeleri (see fig. 15, plate 3, and discussion on p. 53). Like
their northern congeners the quadridentatus-g roup species are known
only from rain forest, usually at elevations of ■+ 1000 ft.

Available biological information is given in the taxonomic section
to follow. Nesting apparently occurs in rotting logs in all species
except wheeleri, colonies of which are usually located in the soil, under
stones. Feeding and foraging behavior have not been extensively
studied, but the bicolored form of quadridentatus forages nocturnally,
in the open, on logs, tree trunks and low foliage. It appears that
wheeleri may restrict its foraging activity to the soil and leaf litter,
since strays have not been taken elsewhere, in spite of extensive search
by the author.

All Australian Pristomyrmex types are workers, and the females
are known for all species except foveolatus ; I have seen no male
specimens. Larvae of a North Queensland species were described by
G. C. and J. Wheeler (1954). I have been unable to locate adults
from the series involved ; they would conform to the foveolatus-group
diagnosis according to their collector, Dr. W. L. Brown, Jr.

This study is based on material from the Museum of Comparative
Zoology collection at Harvard University, from Entomology Division,
CSIRO, Canberra, The National Museum of Victoria, Melbourne,
and the collection of Rev. B. B. Lowery, S.J., Sydney.

MEASUREMENTS AND INDICES

The following abbreviations are used in the descriptions below.
Measurements were prepared using a stereomicroscope fitted with an
eyepiece scale, reading directly in units of 0.1 and 0.01 mm at 100 X
magnification. The measurements cited are the maxima obtained after
a series of trials, a fact of special importance when dealing with HL
and WL.

HL Maximum head length, frontal view, from apex of median
clypeal denticle to median occipital border.

HW Plead width, frontal view, including the eyes.

Cl Cephalic index (HW X ioo)/HL

SL Scape length, excluding articular boss and condyle ( hut in-
cluding the expanded lamella at base of shaft).

SI Scape index (SL X ioo)/HW

PW Pronotum width, anterior view, immediately below the bases
of the pronotal spines.

1965]

T ay lor — Pristomyrmex

37

Scutum Width (females only), dorsal view, across the widest part of
the sclerite.

WL Weber s length of mesosoma, lateral view; in worker , maximum
diagonal distance from junction of anterior and dorsal faces of
pronotum to apex of inferior metapleural lobe; in female, the
anterior limit of measurement is the point where the pro-mesono-
tal suture meets the anterodorsal mesosomal profile.

Pronotal Spine L(ength), as seen in PW measuring position, true
vertical distance between base of spine on its mesial side, and the
level of its apex.

Propodeal Spine L(ength), lateral view, maximum distance from
apex of spine to opposite edge of propodeal spiracle.

Petiole L(ength), lateral view, from tip of metapleural spine to
posterodorsal edge of petiolar tergite.

Postpetiole Height, lateral view, maximum distance from apex of
subpostpetiolar process to postpetiolar dorsum.

Eye Diameter, maximum diameter of eye.

Ocular Index (Eye Diameter X ioo)/HW.

Key to the Australian Pristomyrmex Species
(Based on the workers)

1. Front of head, between the antennal scrobes, with numerous large
shallow umbilicate foveae, their average diameter about 0.05-0.06

mm (group of P. foveolatus) 2

Area of frons between antennal scrobes entirely smooth, lacking
sculpturation apart from a few minute, pilosity-bearing punctures
(group of P. quad rid en tat us ) 3

2. Scapes barely reaching level of occipital border when at rest
in the antennal scrobes, with the head held in full-face view

(fig. 1 ) ; Scape Index 78-83 P. foveolatus n. sp.

Scapes exceeding level of occipital border by 1-1.5 X their
maximum thickness when at rest in the antennal scrobes, with the

head viewed full-face (fig. 3), Scape Index 96-101

P. thoracicus n. sp.

3. Head relatively narrow (Cephalic Index 97-103) (fig. 7), either
concolorous with mesosoma or much lighter in color (plate 3) ;
pronotal spines, in side view, distinctly longer and stronger than

those of propodeum (fig. 8) P. quadridentatus (Andre)

Head relatively much broader (Cephalic Index 110-116) (fig.

1 1 ) and usually much darker in color than the mesosoma ( plate
3) ; pronotal spines subequal in size to propodeal ones, or very
slightly smaller (fig. 12) P. wheel eri n. so.

38

Psyche

[March

Species-level Taxonomy
i. Pristomyrmex foveolatus new species
(Figs, i, 2)

Holotype worker. HL 0.84 mm; HW 0.92 mm; Cl no; SL 0.75
mm; SI 81 ; PW 0.55 mm; WL 0.79 mm; Propodeal Spine L 0.35
mm; Petiole L 0.30 mm; Postpetiole Height 0.30 mm. General
features as in figures 1 and 2. Mandibles with two adjacent strong
acute apical teeth and a truncated basal one, separated by a broad,
shallowly concave diastema. Clypeus with a median longitudinal
Carina, its anterior border with 5 obtuse denticles, the median and
outer lateral ones largest. Antennal scrobes shallow, their ventral
borders obscure. Occipital area of cranium enclosed dorsally and
laterally by a fine arched carina. Scapes barely attaining level of
occipital border when lying in the antennal scrobes, with the head
viewed in HL measuring position (see p. 36). Funiculus with a
3-segmented club, proportions of its segments as in Figure 1. Eyes
moderately large, maximum diameter 0.12 mm, their anterior edges
almost twice this distance from the anterior genal borders. Mesosomal
profile as in Figure 2. Pronotum truncated anteriorly, its inferior
angles edentate. Humeri each with a small acute tooth about 0.05
mm long, inclined dorsolaterally to project beyond the main lateral
outline of the pronotum, when viewed from above. Posterolateral
corners of pronotal dorsum forming blunt angles of about 90 degrees
in dorsal view. Anterodorsal edge of mesepisternum extended anterior-
ly as a strong, moderately acute process which partially overlaps the
fore-coxa. Propodeal spines long, almost straight, each diverging from
the dorsal midline at an angle of about 20 degrees. Metapleural spines
small, moderately acute, each about 0.07 mm long, slightly wider at
base. Petiolar and postpetiolar profiles as in Figure 2. Petiole slightly
narrower than postpetiole when viewed from above, its crest transverse
in posterior view and feebly convex.

Mandibles generally smooth, with several longitudinal ribs.
Clypeus strongly shining. Head polished and reflective, with numer-
ous large shallow subcircular umbilicate foveae, averaging about
0.06 mm in diameter, largest on the posterior genal surfaces, lacking
on a small smooth area between eye and antennal socket on each side,
generally separated by distances averaging about one-third their mean
diameter. Postgenae obscurely foveolate. Antennal scrobes smooth,
with an almost effaced rugulosity and several arched transverse striae
anteriorly. Pronotal dorsum obscurely foveolate, with several median
longitudinal costulae. Posteriorly these meet a transverse carina which
apparently represents the mesonotum. Propodeal dorsum smooth,

1965]

Taylor — Pristo??iyrmex

39

framed by lateral and posterior carinae. Sides of mesosoma generally
smooth, ventral parts of mesepisternum, metepisternal area, and sides
of propodeum obscurely roughened. Petiole, postpetiole and gaster,
antennae and legs, smooth and shining.

Pilosity moderately long and abundant on head and mesosoma,
erect to sub-erect, longest on clypeus, crown of head, and pronotal
dorsum; shorter erert hairs on scape, sub-erect ones on mandibles,
funiculus and legs. Hairs lacking on sides of mesosoma. Petiolar and
postpetiolar tergites each with three bilaterally symmetrical pairs of
hairs (positions as in Figure 2), postpetiolar sternite with a single
pair. Gaster very finely pilose apically, its first tergite completely
bare. Pubescence lacking.

Color rich golden sienna-brown, mandibles, clypeus, antennae and
legs lighter, eyes black, pilosity whitish.

Type-locality. Clump Point, west of Tully, N. Queensland.

The holotype was taken in a rotting wood fragment on the floor
of very disturbed, depauperate rain forest, a few feet above sea level
(June 25, 1962; R. W. Taylor acc. 1746).

Paratype workers. Three specimens collected with the holotype
resemble it closely; they have the following dimensions: HL 0.82-
0.88 mm; HW 0.87-0.96 mm; Cl 106-109; SL 0.72-0.75 mm; SI
78-83; PW 0.52-0.57 mm; WL 0.77-0.79 mm; Propodeal Spine L
0.32-0.35 mm; Petiole L 0.29-0.33 mm; Postpetiole Height 0.28-0.30
mm.

An additional specimen from rain forest at Thornton Peak (Mt.
Alexander), north-west of Daintree, N. Queensland (elevation not
specified but probably + 1000 ft (Darlington, i960: 1 1 7 ) , December
20-23, I957i P- F. Darlington) has: HL 0.85 mm; HW 0.95 mm;
Cl 1 12; SL 0.75 mm; SI 79; PW 0.53 mm; WL 0.78 mm; Pro-
podeal Spine L 0.32 mm; Petiole L 0.28 mm; Postpetiole Height
0.29 mm.

This species may show slight unimodal polymorphism. In the short
series available SL, and the mesosomal dimensions PW and WL are
negatively allometric relative to HW, while HL is positively allo-
metric.

Distribution and Material Examined. Known only from the five
workers listed above. Clump Point and Thorton Peak are within the
area of the main “base-of-peninsula” rain forest system of North
Queensland.

Type deposition. Holotype and three paratypes in Museum of
Comparative Zoology (MCZ) at Harvard University (Type No.

40

Psyche

[March

Figures 1-6: North Queensland species of Pristomyrmex foveolatus species-
group — P. foveolatus n. sp., holotype worker: fig. 1, head, frontal view;
fig. 2, mesosoma, petiole and postpetiole, lateral view. — P. thoracicus n. sp.,
holotype worker: fig. 3, head, frontal view; fig. 4, mesosoma, petiole and
postpetiole, lateral view. Paratype female: fig. 5, head, frontal view; fig. 6,
mesosoma, petiole and postpetiole, lateral view.

1965]

Taylor — Pristomyrmex

41

31 152); one paratype (from Clump Point) in Australian National
Insect Collection (CSIRO), Canberra.

2. Pristomyrmex thoracicus new species
(Figs. 3-6)

Worker. The following description is based on the holotype and
14 paratypes collected with it. Dimensions (holotype cited first) :
HL 0.95, 0.89-0.96 mm; HW 0.95, 0.89-0.99 mm; Cl 100, 98-104;
SL 0.93, 0.90-0.96 mm; SI 98, 96-101; PW 0.49, 0.45-0.51 mm;
WL 0.91, 0.87-0.94 mm; Propodeal Spine L 0.30, 0.28-0.32 mm;
Petiole L 0.34, 0.33-0.36 mm ; Postpetiole Height 0.32, 0.28-0.32 mm.
General features as in Figures 3 and 4. Similar to P. foveolatus,
differentiated from it by the following characters:

1. Relatively narrow head (Cl 98-105 against 106-110 in foveo-
latus) .

2. Much longer scapes, which surpass the level of the occipital
border by 1-1.5 X their maximum thickness when lying in the
antennal scrobes, with the head viewed in the HL measuring position
(SI 96-101 against 78-83).

3. Proportionately much longer mesosoma (WL 0.94-1.00 X
HW, against 0.84-0.89 X in foveolatus) , which is relatively very
narrow (PW 0.5 1-0.53 X HW, and 0.50-0.55 X WL against
0.56-0.62 X and 0.67-0.73 X respectively in foveolatus) .

4. Eyes relatively slightly smaller; antennal scrobes more distinctly
outlined. Humeral teeth more erect, failing to project beyond lateral
pronotal outline when viewed from above. Posterolateral corners of
pronotal dorsum obtuse, broadly rounded in dorsal view. Propodeal
spines slightly sinuate, their apices turned upwards; diverging from
the dorsal midline at angles of about 20 degrees. Metapleural spines
more acute. Petiolar crest strongly convex in dorsal view, its outline
almost a half circle.

5. Sculpturation virtually identical. Cephalic foveae a little more
spaced ; sides of mesosoma less sculptured, generally smooth and
shining.

6. Pilosity similarly distributed, the hairs shorter, especially on the
crown of the head, and the pronotal dorsum; less abundant on pro-
podeal dorsum. Hairs of petiole, postpetiole and gaster placed exactly
as in foveolatus.

Type-locality. Vision Falls, Lake Eacham National Park, N.
Queensland. The type nest series was taken from a rotting log in
rain forest, elevation c. 2500 ft (June 6, 1962; R. W. Taylor acc.
1399)-

42

Psyche

[March

Additional paratype workers. Three workers from Beatrice River,
Crawford’s Lookout, N. Queensland (elevation probably c. 1000 ft.
(Darlington, i960: 118), rain forest, February 21-22, 1958;

Darlingtons) and a single worker from Kuranda, N. Queensland
(1100 ft, rain forest, July 4, 1962, R. W. Taylor acc. 1358) have
the following dimensions (Crawford’s Lookout material cited first) :
HL 0.86-0.89, 0.89 mm; HW 0.87-0.90, 0.93 mm; Cl 101-102, 105;
SL 0.88-0.89, 0.90 mm; SI 99-101, 97; PW 0.45-0.47, 0.47 mm;
WL 0.84-0.85, 0.88 mm; Propodeal Spine L 0.27-0.28, 0.31 mm;
Petiole L 0.31-0.33, 0.33 mm; Postpetiole Height 0.29-0.31, 0.31 mm.
Polymorphism weak, as in P. foveolatus — SL, PW and WL nega-
tively allometric relative to HW, HL positively allometric.

Paratype females. Two females are present in the type series, one
from the holotype nest, the other from the Crawford’s Lookout
collection. Dimensions (Lake Eacham specimen cited first) : HL
0.96, 0.95 mm; HW 1.04, 1.02 mm; Cl 109, 108; SL 0.95, 0.92
mm; SI 91, 90; Scutum W 0.67, 0.65 mm; WL 1.09, 1.04 mm;
Propodeal Spine L 0.32, 0.3 1 mm; Petiole L 0.38, 0.38 mm; Post-
petiole Height 0.35, 0.35 mm; Eye Diameter 0.17, 0.17 mm; Ocular
Index 16, 17. General features as in Figures 5 and 6. Pronotum
without humeral spines; mesoscutum lacking parapsidal lines, pro-
podeal spines diverging from dorsal midline at angles of 30 and 33
degrees. Mesonotal sclerites moderately coarsely longitudinally
costulate, with vestiges of interstitial foveae. Pilosity as in worker;
petiolar and postpetiolar hairs similarly arrayed, the petiolar peduncle
with an additional pair of lateral hairs.

Color as in worker; each ocellus with an adjacent black spot; the
mesosomal suture lines darkly infuscated.

Distribution and material examined. Known only from the 19
workers and two females discussed above. Lake Eacham, Crawford’s
Lookout and Kuranda are all at moderately high elevations on the
Atherton Tableland, within the compass of the main “base-of-
peninsula” rain forest system of N. Queensland.

Type deposition. Holotype, 11 paratype workers and paratype
female (from holotype nest) in MCZ (Type No. 31 153); four
paratype workers and female in CSIRO; single paratypes in Queens-
land Museum, Brisbane; National Museum of Victoria, Melbourne;
and British Museum (Nat. Hist.), London.

3. Pristomyrmex quadridentatus (Ern. Andre)

(Figs. 7-10, Plate 3)

Odontomyrmex quadridentatus Ern. Andre, 1905, Rev. Ent. Caen., 24: 207,
worker. Type-locality: Sydney.

1965]

Taylor — Pristomyrmex

43

Pristomyrmex quadridentatus : Emery, 1922, Gen. Ins., 174c: 234.
Pristomyrmex (Odontomyrmex) quadridentatus v. queenslandensis Forel,
1915, Arkiv F. Zool., 9 (16): 53, worker. Type-locality: Mt. Tambou-
rine (— Tamborine Mountain). NEW SYNONYMY.2
Pristomyrmex quadridentatus var. queenslandensis : Emery, 1922, Gen. In9.,
174c: 234.

Syntype workers. Two quadridentatus syntypes, each labeled
“Sydney/Duchaussoy/TYPE” have kindly been loaned for study by
Mile. S. Kelner-Pillault of the Museum National D’Histoire Natu-
relle, Paris. These specimens are here designated lectotype and
paralectotype and are so labeled. The paralectotype gaster is missing.

Judging from subsequent records the Sydney type-locality may be
outside the true range of this species and should be regarded with
reservation as a distributional record until confirmed.

Dimensions as given in Table i (lectotype cited first). General
features as in Figures 7 and 8. Mandibular dentition as in P. foveo-
latus. Clypeus with a median longitudinal carina, its anterior border
with three separated, moderately acute denticles (apparently repre-
senting the median and outer-lateral ones of foveolatus) . Antennal
scrobes open posteriorly but well defined dorsally and ventrally by
distinct sharp sub-parallel carinae. Occipital carina distinct. Scapes,
in frontal view, laid in the scrobes, exceeding the level of the occipital
border by about 1.5 X their maximum thickness. Funicular pro-
portions as in Fig. 7. Ocular Index 17-18.

Mesosomal profile as in Figure 8. Pronotum with a distinct
transverse anterior carina. Pronotal spines almost vertical, consider-
ably stronger than propodeal ones. Mesonotum represented by a
transverse carina on mesosomal dorsum. Propodeal spines each
diverging from the midline at an angle of about 12 degrees; about
equal in size to metapleural spines in lectotype, somewhat smaller in
paralectotype. Petiole and postpetiole as in Figure 9; petiolar crest
strongly arched in posterior view ; postpetiole very slightly wider than
petiole in dorsal view.

Head and body almost completely lacking sculpturation, smooth
and strongly shining, except for a little fine shagreening on the
pronotal collar and slight obscure roughening on the sides of the
propodeum. Pilosity generally as in P. foveolatus ; petiolar and post-
petiolar hairs similarly distributed, but with an additional pair on the
petiolar peduncle (as in the thoracicus female). First gastric tergite
completely naked. Pubescence virtually absent.

2I have not seen the queenslandensis type, but specimens from the Mt.
D’Aguilar series, listed below, have been directly compared with it for me
by Dr. W. L. Brown, Jr.

[March

44

Psyche

j I'Omm

Figures 7-10: Pristomyrmex quadridentatus Andre. — lectotype worker:
fig. 7, head and pronotal outline in frontal view ; fig. 8, mesosoma, petiole
and postpetiole in lateral view. Female (self-colored form) : fig. 9, head,
frontal view; fig. 10, mesosoma, petiole and postpetiole, lateral view.

Color almost even rich light-golden sienna-brown, mesosoma a
shade darker, masticatory border of mandible, and segments 2-8 of
antennal funiculus darkly infuscated, less dark infuscation on scapes
and the various carinae and spines of head and mesosoma. Eyes dark
brown, almost black.

Additional material examined , distribution. Additional distri-
butional records fall into two geographical clusters (Fig. 15) which
coincide with the distribution of two morphological forms, apparently
representing geographical variants of a single species. These variants
are discussed in detail below and illustrated in plate 3 ; essentially there
is a southern, self-colored golden-brown form (to which the lectotype
and paralectotype belong), and a more northern bicolored form with
dark blackish-brown mesosoma, petiole and postpetiole (equivalent
to Forel's suppressed variety queenslandensis) . They are distributed
as follows:

1965]

Taylor — Pristomyrmex

45

1. Self-colored form: N.E. NEW SOUTH WALES between
about S. lat. 29.40 and 30.20 (not including Sydney — see above)
(open circles in Fig. 15).

Specific records are: Point Lookout to Grafton , dealate female,
April 1958, (P. F. Darlington). New England National Park ,
Platypus Creek (elevation not given, probably 2-3000 ft — see LeGay
Brereton, 1963, fig. 1), in log, February 27, 1961 (Nicholls). The
Dorrigo 3000 ft (W. Heron).

2. Bicolored form: Extreme S.E. QUEENSLAND, between

about S. lat. 27.20 and 28.20 (closed circles in Fig. 15).

Specific records are: Mt. D’ Aguilar Range , 2000 ft, nest in

rotting log, May 4, 1962 (RWT acc. 799). Cunningham's Gap ,
2500 ft, nest in rotting log, May 1, 1962 (RWT). Stradhroke
Island, dealate queen, December 3, 1912 (H. Hacker). Tamborine
Mountain: no further data (A. M. Lea) ; Cedar Creek, ex rotting
logs — dealate female, May 15, 1951, workers November 13, 1950
and May 12, 1951 (W. L. Brown) ; worker under log, February 27,
1956 (B. B. Lowery). Binna Burra: c. 2600 ft: nests in rotting
logs, one with alate female, May 22-23, 1962 (RWT accs. 1145,
1182); several series of workers collected straying at night (see
below under “ecology”) , May 21-23, 1962 (RWT). Lamington
National Park, Coomera Gorge: leafmold berlesate, October 29,
1955 (T. E. Woodward) ; 2200 ft, nest in rotting log, May 23, 1962
(RWT acc. 1168).

All the above records are specifically cited as rain forest collections,
except the dealate female from Stradbroke Island, which has no
ecological data.

IV orker variation. Measurements and indices of workers from the
above series are summarized in Table 1. The similarity of ranges and
means in the two color variants is obvious and reflects their consider-
able resemblance in all characters except color.

The coloration of the types (Plate 3, lower figure) is typical of the
self-colored form. The bicolored form has the mesosoma, petiole and
postpetiole very dark blackish-brown, contrasting sharply with the
golden-brown head, gaster and appendage's (Plate 3, middle figure),
and the antennal infuscation is much less intense. The pronotum and
postpetiole may be medium to dark reddish-brown, usually in the
larger specimens of a given series. Such individuals are probably
incompletely colored ; since these areas are the last to become fully
pigmented in callow specimens.

The mesosomal spines vary in thickness, length and inclination. The
pronotal pair are typically longer and thicker than the propodeal ones,

46

Psyche

[March

SELF-COLORED FORM

BICOLORED FORM

Series No.

Types

1

2

Total

3

4

5

Total

Number

2

4

11

17

10

14

31

55

HL

0.95, 0.99

1.03-1.07

(1M)

1.00-1.06

(1403)

0.95-1.07

(1.00)

0.88-1.06

(0.97)

0.92-1.11

(1.03)

0.97-1.11

(i.oi)

0.88-1.11

(1.00)

HW

0.95, 1.00

1.01-1.06

(1.04)

0.97-1.0 6
(1.03)

0.95-1.06

am

0.88-1.03

(0.97)

0.92-1.12

(1.03)

0.89-1.14

(0.98)

0.88-1.14

(0.99)

Cl

100. 100

98-100

(99)

100-103

(101)

98-103

(100)

97-103

(100)

98-101

(100)

98-103

(101)

97-103

(100)

SL

0.90, 0.93

1.01-1.06

(1.04)

1.00-1.03

(1.01)

0.90-1.0 6
(0.97)

0.88-1.04

(0.97)

0.87-1.09

(1.02)

0.86-1.15

(1.01)

0.86-1.15

(1.00)

SI

95, 93

100

97-103

(99)

93-103

(97)

97-103

(99)

95-105

(99)

95-103

(98)

95-105

(99)

PW

0.46, 0.50

0.48-0.53

(0.51)

0.48-0.52

(0.50)

0.46-0.53

(0.49)

0.43-0.50

(0.48)

0.44-0.53

(0.50)

0.45-0.56

(0.50)

0.43-0.56

(0.49)

WL

0.95, 0.99

1.03-1.06

(1.04)

1.01-1.06

(1.03)

0.95-1.06

(1.00)

0.92-1.03

(0.99)

0.92-1.05

(1.00)

0.92-1.15

(1.03)

0.92-1.15

(1.01)

WL X 100
HW

100, 99

100

100-105

(101)

99-105

(100)

100-104

(101)

100-101

(100)

98-103

(101)

98-104

(101)

PW x 100
HW

48, 50

48-50

(49)

46-50

(48)

46-50

(49)

47-51

(48)

47-50

(48)

47-51

(49)

47-51

(48)

PW x ioo

WL

48, 50

47-50

(48)

46-48

(48)

46-50

(48)

47-50

(48)

47-50

(48)

47-51

(48)

47-51

(48)

Pronotal
Spine L.

0.15, 0.15

0.17-0.19

0.15-0.20

0.15-0.20

0.13-0.18

0.13-0.17

0.13-0.17

0.13-0.18

Propodeal
Spine L.

0.23, 0.22

0.18-0.20

0.20-0.24

0.18-0.24

0.15-0.20

0.16-0.19

0.16-0.20

0.15-0.20

Petiole L.

0.33, 0.33

0.34-0.37

0.33-0.37

0.33-0.37

0.32-0.37

0.30-0.37

0.30-0.38

0.30-0.38

Postpet. H.

0.32, 0.33

0.34-0.35

0.33-0.37

0.32-0.37

0.32-0.35

0.31-0.36

0.28-0.35

0.28-0.36

Table 1. Pristomyrmex quadridentatus Andre. Comparison of standard
measurements (in mm) and indices, yielded by specimens from all known
series. Mean values italicized. The types are ostensibly from Sydney ( hut
see text). Numbers in the “series” column refer to the following localities:
1. New England National Park; 2. The Dorrigo ; 3. Mt. D’Aguilar; 4.
Tamborine Mt. ; 5. Binna Burra.

which are usually about sub-equal to the metapleurals. The bicolored
form tends to have thicker pronotal and smaller propodeal spines than
the self-colored form; but the overall variation is such that the avail-
able quadridentatus material cannot be divided into two sets, co-
extensive with the color-variants, on the basis of spinational characters.

1965]

Taylor — Pristomyrmex

47

A weak negative allometric relationship between spine development
and body size is evidenced ; this is best marked in the bicolored form,
probably because of its better representation in the study series.

Female (previously undescribed). Six female specimens are avail-
able for study; one self-colored, five bicolored.

1. Self-colored form. A single dealate labeled Point Lookout —
Grafton, N.S.W., has the following dimensions: HL 1.05 mm; HW
1.07 mm; Cl 102; SL 0.98 mm; SI 88; Scutum W 0.68 mm; WL
1.20 mm; Propodeal Spine L 0.22 mm; Petiole L 0.35 mm; Post-
petiole Height 0.39 mm; Eye Diameter 0.22 mm; Ocular Index 21.
General features as in Figures 9 and 10. Pronotum with a partly
broken fine, arched transverse carina, its humeri each with a low
obtuse tubercle. Parapsidal lines of mesoscutum very obscure, es-
sentially vestigial. Bases of propodeal spines connected by a transverse
carina. Metapleural spines slightly longer and more acute than
propodeal ones.

Color and pilosity as in worker, the ocelli each with an adjacent
small dark brown spot, the mesosomal suture lines and axillary
sclerites infuscated dark brown.

2. Bicolored for?n. Two dealate specimens from Mt. D’Aguilar
Range, one each from Stradbroke Island and Tamborine Mt., and
an alate from Binna Burra, have the following dimensions: HL
0.95-1.22 mm; HW 0.99-1.30 mm; Cl 102-106; SL 0.94-1.18 mm;
SI 91-95; Scutum W 0.62-0.85 mm; WL 1.10-1.42 mm; Petiole L
0.32-0.46 mm; Postpetiole Height 0.32-0.43 mm; Eye Diameter
0.19-0.26 mm; Ocular Index 19-20. The smallest specimen is from
Mt. D’Aguilar Range, the largest is the Binna Burra alate (which
has a forewing length of 1.9 mm, hindwing length of 1.3 mm).
Distribution of indices in the series implies negative allometry of scape
length and head length, relative to head width. Pronotal spine
development variable, apparently negatively allometric, smallest
specimen with a low obtuse tubercle, largest one with an almost
imperceptible slightly raised convexity. Propodeal spines similarly
variable, best developed in smallest individuals (about comparable to
self-colored specimen), represented only by slightly raised propodeal
corners in largest specimen. Characters otherwise as in self-colored
form, with the scapes proportionately slightly longer.

Coloration as in worker, ocellar area darkly infuscated; pronotum
and scutum reddish brown in the Binna Burra alate, which may be
incompletely colored.

Ecology. All detailed records of quadridentatus show that nests
are usually located in rotting logs, a fact confirmed in the field by

48

Psyche

[March

the author, and by the experienced Australian collector B. B. Lowery,
S. J., (pers. comm.).

On May 21-23, 1962, I established that the bicolored form is
probably an exclusively nocturnal forager. During 4 days of intensive
collecting in rain forest near Binna Burra, workers were never
encountered abroad from nests during the day. At night, however,
22 workers in seven samples were taken as strays, during about four
hours total collecting. These specimens were all collected between
two and six hours after sunset (the limits of the observation period),
and all were in the open on the surfaces of widely separated rotting
logs. Several further individuals were taken near the ground on tree
trunks, and one was collected from foliage in a sweep-net. In several
cases these individuals appeared to be following trails, but they were
always well separated from each other, at distances of 14-20 cm. The
ants moved slowly, their rate of progress being estimated at 10-15 cm
per minute. The nature of the food was not determined.

Relationships. See discussion on p. 35.

4. Pristomyrmex wheeleri new species
(Figs. 11-15, Plate 3)

Type workers. The following description is based on the holotype
and numerous paratypes from all parts of the distributional range
given below.

Dimensions (holotype cited first) : HL 1.14, 0.97-1.23 mm; HW
1.26, 1. 04- 1. 40 mm; Cl 111, 108- 1 16; SL 1.11, 0.92-1. 14 mm; SI
88, 81-90; PW 0.60, 0.51-0.67 mm; WL 1.09, 0.89-1.20 mm;
Pronotal Spine L 0.17, 0.15-0.20 mm; Propodeal Spine L 0.35,

0.27-0.39 mm; Petiole L 0.41, 0.36-0.44 mm; Postpetiole Height
0.38, 0.31-0.43 mm. General features as in Figures 11 and 12, and
Plate 3 (upper figure). Similar to P. quadridentatus, as described
above, but differing in the following features:

1. Generally larger size (HW 1.04- 1.40 mm, opposed to 0.88-1.14
mm in quadridentatus) , with much broader head (Cl 108-116 against
97-103), and lower Scape Index (81-90 opposed to 93-105).

2. Inferior carina of antennal scrobe vestigial or absent.

3. Pronotal and propodeal spines longer and more slender, in side
view approximately sub-equal in length ; metapleural spines acute,
rarely more than half as long as propodeals.

4. Pilosity similar, the body hairs thicker and longer, petiolar
peduncle with a single hair on each side, first gastric tergite naked.

5. Color distinctive; head deep dark umber-brown. Mandibles,
clypeus and antennae rich golden-brown. Remainder of body golden-
brown, much like the self-colored form of quadridentatus , but more

1965]

Taylor — Pristomyrmex

49

Figures 11-14: Pristomyrmex wheeleri n. sp. — Holotype worker: fig. 11,
head and pronotal dorsum, frontal view; fig. 12, mesosoma, petiole and
postpetiole, lateral view. — Paratype female: fig. 13, head, frontal view;
fig. 14, mesosoma etc., lateral view.

yellowish. Posterior half of first gastric tergite shaded medium to dark
reddish brown.

Type locality. Vicinity of Binna Burra, S.E. Queensland. The
type colony was taken in soil under a stone in rain forest, elevation c.
2800 ft (May 24, 1962, R. W. Taylor acc. 1281).

Worker variation. Negative allometry of head length and scape
length, relative to head width, is suggested by the distribution of
cephalic and scape indices in the study series. Other characters,
including the clypeal structure and the development of the mesosomal
spines, show no apparently significant variation.

Paratype females. A female from the type-nest (cited first), and
six others, one from each locality given below except National Park
and Woodenbong, have the following dimensions: HL 1.11, 1 . 1 7-
1.25 mm; HW 1.23, 1.3 1- 1.42 mm; Cl 112, 112-114; SL 1.04,

50

Psyche

[March

Figure 15 (above) : Distribution of members of the Pristomyrmex quadri-
dentatus group. Land above 200 meters elevation shaded.

EXPLANATION OF PLATE 3 (opposite)

Workers of the Pristomyrmex quadridentatus group, to illustrate coloration.
Top: P. wheeleri n. sp., from Binna Burra, S. E. Queensland. Middle:
P. quadridentatus Andre, bicolored form, from Binna Burra (sympatric with
•wheeleri) . Bottom: P. quadridentatus Andre, self-colored form, from New
England National Park (Point Lookout) (allopatric with eivheeleri). Author
del

Psyche, 1965

Vol. 72, Plate 3

Taylor — Pristomyrmex

52

Psyche

[March

1.08-1.12 mm; SI 85, 79-82; Scutum W 0.67, 0.78-0.84 mm; WL
1. 1 9, 1.26-1.36 mm; Propodeal Spine L 0.33, 0.35-0.39 mm; Petiole
L 0.41, 0.44-0.48 mm; Postpetiole Height 0.39, 0.41-0.45 mm; Eye
Diameter 0.24, 0.26-0.27 mm; Ocular Index 20, 19-20. Negative
allometry of scape length and head length, relative to head width, is
indicated. General features as in Figures 13 and 14. Distinguished
from P. quadridentatus by very different proportions and coloration.
Structure of clypeal apron variable, usually as in Figure 13; the
median projection narrowed with reduced teeth in one specimen,
bidentate in another, and completely edentate in a third. Transverse
pronotal carina distinct. Pronotal spines variable, ranging from low
rounded tubercles to small acute teeth (maximally about 0.03 mm
long) , with no apparent allometric tendency. Parapsidal lines feeble
to moderately distinct. Propodeal spines almost invariable, their bases
bridged by a low carina. Metapleural spines acute, distinctly shorter
than propodeals.

Color and pilosity as in worker, mesosomal suture lines and alary
sclerites infuscated dark brown.

Distribution and Material Examined. All records of P. wheeleri
are from S. E. Queensland and N.E. New South Wales, between
about S. Lat. 27.20 and 29.00 (Fig. 15), as follows: QUEENS-
LAND: A It. D’ Aguilar Range, 2000 ft, colony, May 4, 1962
(RWT acc. 819). Cunninghams Gap: 2500 ft, six colonies,

January 22, 1961 (B. B. Lowery) ; 2000 ft, colony, May 1, 1962
(RWT acc. 721). T amborine Mountain : no further data (A. M.
Lea); colony, January 15, 1957 (B. B. Lowery); Curtis Falls,
workers, leafmold berlesate, May 8, 1953 (T. E. Woodward). Binna
Burra: 2800 ft, colonies, May 21-25, 1962 (RWT accs. 1192, 1281
( holotype nest series) , 1290, 1293) J stray, forest floor, 2 hours after
sunset, May 23, 1962 (RWT acc. 1210) ; 2600-3000 ft (P. F.
Darlington). National Park , December 1919 (H. Hacker). NEW
SOUTH WALES: W oodenbong ; JJnumgar Forest , c. 2000 ft;
Tooloom Range , c. 2000 ft, dealate female; all May 1958 (Darling-
tons). Mt. Warning, 2600-3000 ft, five colonies, September 13, 1962
(B. B. Lowery).

Type deposition. Holotype and numerous paratypes (worker and
female) in MCZ (Type No. 31 154), paratypes in CSIRO; Queens-
land Museum, Brisbane; Australian Museum, Sydney; National
Museum of Victoria, Melbourne, and other Australian collections,
also British Museum (NH), London, and B. P. Bishop Museum,
Honolulu.

Ecology. All the above records are from rain forest. Sixteen of the

1965]

Taylor — Pristomyrmex

53

1 8 colonies listed were found in soil under or between stones, one was
in a rotting log (RWT acc. 819), and one in a rotting wood frag-
ment (RWT acc. 721). Nesting behavior thus contrasts with that
of the sympatric bicolored form of quadridentatus , which has been
found nesting only in rotting logs. Lowery’s six Cunningham’s Gap
colonies each contained 18 to 36 workers and one queen. At Binna
Burra I searched extensively for foragers of wheeleri but encountered
only one (acc. 1210), which was taken two hours after sunset on top
of a small rock, later found to shelter a colony. Foraging behavior
presumably is unlike that of P. quadridentatus (see p. 48).

Relationships. See below.

DISTRIBUTION AND VARIATION IN THE QUADRIDENTATUS GROUP

All known bicolored series of Pristomyrmex quadridentatus were
collected in immediate geographical and temporal sympatric associ-
ation with P. wheeleri , while the self-colored form has never been
taken in such association. This correlation invites the hypothesis that
character displacement of quadridentatus has occurred wherever the
species is sympatric with wheeleri.

Character displacement is the situation, usually involving several
related species, where interspecific differences are more distinct in
areas of sympatry than in areas of allopatry. This phenomenon has
long been recognized, but was first crystalized by Brown and Wilson
( 1956) , who reported its occurrence in birds, frogs, fishes, crabs and
various insects. Subsequent reports involve many animal groups,
including the South African cyprinid fishes, Barbus marequensis and
B. natalensis ( Farquharson, 1962), the North American salamanders
Plethodon cinereus (Green) and P. richmondi Netting and Mittle-
man (Highton, 1962), the Central American iguanids. Basiliscus
plumifrons Cope and B. vittatus Weigmann (Maturana, 1962),
the East African gekkonids Holodactylus africanus Boettger and H.
cornii Scortecci (Laurent, 1964), several pairs of Indian nuthatches
of the genus Sitta (Ripley, 1959) and the New Zealand pelecypods
Bassinia yatei (Gray) and B. parva Marwick, which apparently
underwent displacement on becoming sympatric in the Lower Pleisto-
cene (Fleming, 1959). Few of these cases have been studied in detail
aimed at elucidating the selective significance of the displacement,
though it seems clear that secondary reinforcement of premating
isolating mechanisms, or special adaptation to greater niche specificity
resulting from competitive displacement, are involved in some cases.

This hypothesis is invoked to explain the origin of the bicolored
form of P. quadridentatus on limited distributional evidence, which
needs supplementation. A selective mechanism is difficult to envisage
since there is no evidence that color is involved in interspecific relations

54

Psyche

[March

between these ants. Ecological differences between the sympatric
forms are reported above; at Binna Burra quadridentatus nests in
logs and forages openly at night, while wheeleri nests under stones
and apparently forages cryptically. Nothing is known of the behavior
of self-colored quadridentatus and such information is required.
Bicolored quadridentatus and ivheeieri are perhaps the most spectacu-
larly colored members of their genus; their coincident distribution
suggests that the causal factors producing their color patterns may
be inter-related. The two “forms” of quadridentatus might prove
ultimately to be good biological species, but even if this is the case,
character displacement need not be rejected in explaining their color
differences. These ants are worthy of much further study.

Literature cited

Brereton, J. LeG.

1963. Point Lookout survey New England National Park. Aust. J. Sci.,
26(1): 10-14.

Brown, W. L., Jr.

1953. Characters and synonymies among the genera of ants, Part I.
Breviora, 11 :1-13.

Brown, W. L., Jr., and E. O. Wilson

1956. Character displacement. Syst. Zool., 5 : 49-64.

Darlington, P. J., Jr.

1960. Australian carabid beetles IV. List of localities, 1956-1958.
Psyche, 67(4): 111-126.

Emery, C.

1922. Formicidae, Subfamily Myrmicinae. Genera Insect., 174c: 207-
397.

Farquharson, F. L.

1962. The distribution of cyprinids in South Africa. Ann. Cape Prov.
Museums, 2: 233-251.

Fleming, C. A.

1962. Paleontological evidence for speciation preceded by geographic
isolation, in The Evolution of Living Organisms , G. W. Leeper
ed. pp. 1-459, Melbourne University Press, (pp. 225-241).
Highton, R.

1962. Revision of North American salamanders of the genus Plethodon.
Bull. Florida. State Mus., 6 (3) :235-367.

Laurent, R. F.

1964. Deplacement des characteres dans le genre Holodactylus (Repti-
lia, Gekkonidae). Senck. Biol., 45 (3/5): 417-423.

Maturana, H. R.

1962. A study of the species of the genus Basiliscus. Bull. Mus. Comp.
Zool., Harv., 128 (1): 1-33.

Ripley, S. D.

1959. Character displacement in Indian nuthatches (Sitta). Postilla,
Yale, 43: 1-11.

Wheeler, G. C., and Jeanette Wheeler

1954. The ant larvae of the myrmicine tribe Myrmecinini. Proc. Ent.
Soc. Washington, 56(3): 126-138.

A NEW SPECIES OF MEGALOMYRMEX
FROM THE CHILEAN ANDES
(FORMICIDAE, HYMENOPTERA)

By George Ettershank*

Department of Entomology, Cornell University

The genus Megalomyrmex has generally been thought of as
associated with the wet tropical rain-forests of the New World.
Members of the goeldii species-group in particular forage openly on
the rain-forest floor and tend Homoptera on low bushes. It was thus
a considerable surprise to receive a few specimens of a goeldii group
species collected only 25 miles from the crest of the Andes in Chile.
This is a new species, and its discovery extends the genus into a new
habitat.

The specimens under study were found by Mr. E. Ackerman at
Pachon Peak (Cerro Pachon), near La Serena, Coquimbo Province,
Chile, at an altitude of 8500 dz 100 feet. The collection site is a
rather flat hill crest with sparse low shrub cover; the substrate is
andesite boulders resting on andesite rubble and soil. The colonies,
Mr. Ackerman notes, were populous and by no means rare. A few
live workers were sent to Dr. Caryl P. Haskins of the Carnegie
Institution of Washington, who placed the workers in a laboratory
nest; a few eggs were laid, from which three larvae were reared. All
the material was later sent to me; this consisted of four live workers
and three preserved larvae. My thanks are offered to Dr. Haskins
for allowing me to study this interesting species.

Measuring techniques and abbreviations quoted below follow the
system of Brown ( 1 ) while certain other terminology follows Etter-
shank (2).

Megalomyrmex bicolor n. sp.

Holotype worker (measurements in millimeters; those for two
parat}q>e workers are enclosed in parentheses) : total length, TL 4.8
(4.6, 4.4), head length, HL 1.04 (0.98, 0.94), length closed man-
dibles, ML 0.24 (0.22, 0.20), alitrunk length, WL 1.34 (1.32,
1.26), head width, HW 1.00 (0.92, 0.88). Cephalic index, Cl =
100 HW/HL = 96 (94, 94), mandibular index = 100 ML/HL

^Present address: Department of Zoology and Comparative Physiology,

Monash University, Clayton, Victoria, Australia. This article is a partial
result of work done under U. S. National Science Foundation Grant GB-2175,
principal investigator, W. L. Brown, Jr.

Manuscript received by the editor January 4-, 1964

55

Psyche, 1965

Vol. 72, Plate 4

O- 5mm.

Ettershank — Megalomyrmex

1965]

Ettershank — Megalomyrmex

57

= 2 3 (22, 21). A fourth paratype worker was dissected and not
measured.

This species is smaller than the other goeldii group species. The
head (fig. 1) in dorsal view is rather more square in shape. The eyes
are large, about 15 ommatidia high and 8 wide, and their black color
contrasts strongly with the golden-brown ground color of the head.
Antennae 12-segmented with a weak 3-segmented club (fig. 2).
Clypeus strongly convex with a slight median sulcus, giving a weakly
bicarinate effect. Median seta present, but variable in position around
the midpoint of the oral margin of the clypeus ; one pair of intercarinal
setae present ; first paracarinal setae well developed, higher paracarinal
setae weaker. A few strong lateral setae present. Lateral areas of
clypeus and oral margin of median area concolorous with mandibles.
Mandibles with dental formula 1 + 4, masticatory margin more
darkly colored, brown black, rest of mandible red-brown. Palpal
formula 4, 3.

Alitrunk (fig. 3) of normal worker form. Promesonotal suture
very distinct on the pleurae to the height of the spiracle, curving
forward and then posteriorly; indistinct laterally above the spiracle
and on the dorsum. Metanotal groove distinct, impressed on the
dorsum and on the pleurae. Faint grooves separate the anepisternum
from the notum above and the katepisternum below. Mesothoracic
dorsum narrowing rapidly above, quite distinctly compressed at the
metathoracic groove. Propodeum with angle distinct but rounded;
declivity distinctly impressed. Propodeal spiracle small, round.
Inferior propodeal plate's large, semicircular. Metapleural glands
large, very distinct, surmounted postero-ventrally by a rounded
protuberance bearing the gland opening.

Petiole and postpetiole of normal Megalomyrmex form, with nodes
high and rounded (see fig. 3). Subpetiolar process dentiform, distinct;
anterior subpostpetiolar process distinct, posterior process slight ; nodes
not particularly broad above. Gaster biconvex in profile, attached at
its most anterior end, in life normally carried with its long axis
inclined downward so that the posterior end just clears the substrate.

Smooth shining ants, bearing a few faint and several distinct rugae
on the mesokatepisternum and metapleuron. Posterodorsal flange of
the postpetiole finely granulose. Head, alitrunk and gaster bearing
sparse, long, white setae. Head (except as noted earlier), alitrunk,

Explanation of Plate 4

Megalomyrmex bicolor, n. sp., Worker: Fig. 1, head, dorsal full-face

view. Fig. 2, antenna. Fig. 3, alitrunk and nodes, lateral view. Larva:
Fig. 4, lateral profile. Fig. 5, setae (see discussion in text) ; a, dorsal and
lateral abdominal; b, of head and prothorax; c, of ventral abdominal tufts.

58

Psyche

[March

petiole and postpetiole golden brown, with a narrow margining of
darker brown ventrally on the alitrunk ; gaster brown-black, the sheen
of the rest of the body being reduced noticeably by a fine shagreening
over all the gaster.

Malpighian tubules 5 (one worker dissected), the tips of the
tubules being attached to the rectum (cryptonephric) .

Larvae: Three larvae were reared from worker-laid eggs and were
presumably genetically male. A profile drawing of the larva is shown
in fig. 4, and is essentially similar to that of M. symmetochus shown
by G. C. and J. Wheeler (3). The head and prothoracic dorsum
bear simple, slightly curved setae of the type shown in fig. 5b. The
remaining body segments dorsally and laterally bear deeply cleft setae
of the type seen in fig. 5a, while ventrally on each abdominal segment
are paired clusters of setae of mixed types: short simple; long simple;
and long deeply bifid, as shown in fig. 5c. In their characterization of
Megalomyrmex larvae, the Wheelers (loc. cit.) state that only simple
setae are present — short ones on the head, long ones on the body —
and that the ventral and lateral surfaces of the abdomen are nearly
naked. This diagnosis is based on M. symmetochus , a member of the
modestus species group. Further comparative study is required, but
perhaps this setal character represents another fundamental difference
between the two species groups.

Disposition of Types: Holotype worker, three paratype workers (one
dissected) and three larvae have all been deposited in the collection of
the Museum of Comparative Zoology, Cambridge, Mass., under type
number 31 139.

References

1. Brown, W. L.

1953. Revisionary studies in the ant tribe Dacetini. Amer. Midi. Nat.
50: 1-137; cf. pp. 7-15.

2. Ettershank, G.

1965. A generic revision of the world Myrmicinae related to Solenopsis
and Pheidolog eton (Hymenoptera, Formicidae). Bull. Mus. comp.
Zool. Harv. (in press).

3. Wheeler, G. C. and J. Wheeler.

1955. The ant larvae of the myrmicine tribe Solenopsidini. Amer.
Midi. Nat. 54: 119-141.

THE HABITS AND DISTRIBUTION OF
CRYPTOCERUS ROHWERI WHEELER
(HYMENOPTERA: FOR MI Cl DAE)

By

William S. Creighton, City College, New York1

AND

William L. Nutting, University of Arizona2

When W. M. Wheeler described Cryptocerus rohweri in 1916 he
stated that the specimens which Rohwer sent him had been taken by
Chrisman in a canyon of the Santa Catalina Mountains of Arizona
and that they had been nesting in the dead limbs of a palo verde
tree ( 1 ) . There was no reason for Wheeler to question that the
host tree was Cercidium torreyanum , but this question has since
arisen and it is advisable to consider it here. The host plant identifi-
cation appears to have been made from the dead limb sent in with
the ants. Chrisman’s field notes stated only that his specimens were
nesting in “palo verde”, which would cover either of the two species
of Cercidium in the Santa Catalina area. But in this area C. torrey-
anum is scarce and it usually occurs at elevations below those where
rohweri has been taken. The abundant and widespread species is C.
microphyllum and all seven colonies of rohweri which we took in the
Santa Catalinas were nesting in this tree. The probability is that

Table 1. Distributional Data for Cryptocerus rohvoeri Wheeler

STATION

ELEVATION

NESTS

NEST SITE

COLLECTOR

STA. CATALINA MTS.
Buehman Canyon

_

1

palo verde

M. Chrisman

Brush Corrals

3700'

1

palo verde

M. Chrisman

Sabino Canyon

3000'

2

hackberry

E. D. Algert

Sabino Canyon

—

1

not given

W. D. Edmonton

Catalina Springs

—

1

not given

Hubbard & Schwarz

Catalina Foothills

2800'

1

palo verde

Wm. L. Nutting

Catalina Foothills

2800'

6

palo verde

Wm. S. Creighton

Saguaro Nat. Mon.

3100'

strays

unknown

F. G. Werner

BABOQUIVARI MTS.
Baboquivari Canyon

3500'

1

mesquite

Wm. S. Creighton

Baboquivari Canyon

3500'

1

mesquite

C. H. Musgrove

A JO MOUNTAINS
Alamo Canyon

stray

unknown

E. D. Ball

ATASCOSA MTS.

Pena Blanca Sprs.

3700'

1

live oak

Wm. S. Creighton

Tmeritus Professor, Department of Biology,
department of Entomology, College of Agriculture.
Manuscript received by the editor December 28, 1964.

59

6o

Psyche

[March

Chrisman’s specimens came from C. microphyllum and not from
C. torreyanum. The list in Table i is an expanded version of the list
of localities for rohweri which M. R. Smith presented in his 1947
study of Cryptocerus (2).

While this list is unbalanced by the preponderance of records from
the Santa Catalina Mountains it permits several conclusions. The
range of rohweri covers a number of mountainous areas in southern
Arizona. In each of these areas rohweri prefers to nest in canyon
bottoms or on foothills at comparatively low elevations (2800-3700').
Although as yet unconfirmed, it is certain that the range of rohweri
extends into northern Sonora. Pena Blanca Springs is (or perhaps
better “was”, for the area has been dammed and flooded) about five
miles north of the border of Sonora and the mountains there run south
into Mexico. It is clear that rohweri will accept at least four different
trees as nest sites ( Cercidium microphyllum , Prosopis juliflora,
Q'uercus emoryi and Celtis sp.). Despite the large number of records
from palo verde there are indications that rohweri has no special
preference for this tree. In 1954 Creighton and Gregg showed that
Crypt, texanus prefers to live in live oaks (3). This view was based
not only on a preponderance of records for nests in live oak limbs but
also on the fact that texanus nests in live oaks wherever these occur
within its range. If rohweri prefers to nest in palo verde limbs it
might be expected to do so over its entire range. As far as we have
been able to determine it does not do so. In January of 1963 the
senior author made an extensive survey of palo> verde trees in the
area between Benson and Ajo. This survey failed to produce a single
nest of rohweri. Negative evidence of this sort is not conclusive but
at least it may be said that the high incidence of rohweri nests in palo
verde limbs in the Santa Catalina Mountains is not maintained in
other parts of its range. It is possible that our present fragmentary
view of the range of rohweri is an outcome of the fact that the tree
which it prefers as a nest site has not yet been recognized.

Most of the observations in the remainder of this paper are based
on three captive colonies of rohweri. One of these was observed by
the junior author from October 1961 until August 1963. During
this period the colony was studied at Tucson, less than five miles from
its original nest site in the Santa Catalina Mountains. The senior
author was less fortunate for the two captive colonies which he
observed were carried far out of their range. From January to mid-
April of 1963 they were studied at Riverside, California. Thereafter
until June 1964 they were studied at Rockport, Ontario. It would
appear, however, that these expatriate colonies behaved in a normal

1965]

Creighton and Nutting — Cryptocerus

61

fashion, for there was no noticeable difference between their behavior
and that of the Tucson colony. This colony was housed in a plastic
petri dish 90 mm. in diameter. A short polyethylene tube connected
this petri dish to a second one which served as a feeding chamber.
This arrangement permitted a close watch on the development of
the brood. The other two colonies were kept in sealed aquaria which
contained oak block observation nests. This provided the ants with
passages similar to those which they normally use and at the same
time gave them the opportunity to forage outside the nest.

Most of the habits of rohweri are like those of texanus but there
are some significant differences in the behavior of the two species. At
maturity the rohweri colony is notably smaller than that of texanus.
There are seldom more than seventy-five workers present and in most
of the colonies that we have seen the total has been less than fifty
individuals. This is undoubtedly due to the fact that rohweri is
seldom, if ever, pleometrotic. Each of the nine colonies of rohweri
which we have examined had a single queen. The nests of rohweri
are established in abandoned burrows of wood-boring beetles (often
those of small buprestids) which are cleared of the detritus left in
them by the beetle larvae. While most of the branches selected by
rohiveri consist of hard, sound wood it will nest in rotten branches
as well. A limb housing one of the Santa Catalina colonies was so
badly decayed that the ants were extracted by crumbling the wood
between the fingers. As shown elsewhere (4) texanus ordinarily
rejects nest sites in rotten wood. The burrows chosen by rohweri are
of a size that permits the major to occlude the terminal opening. This
occlusion is like that of texanus; the opening is blocked by the head
and pronotum of the major, who crouches to admit the minor. An
interesting variation of this response was observed in the junior
author’s colony. This colony originally occupied burrows in a large,
dead palo verde branch. Just inside the entrance of one of these
burrows was a circular flange of detritus. The circular opening in
this flange was slightly more than 2 mm. in diameter. This opening
was occluded by the cephalic disc of the major, who stood in the
passage behind the flange. The workers of rohweri pack themselves
tightly into the outer portion of the nest passage, as do those of
texanus, but show one response under these conditions that texanus
does not display. The minor worker of rohweri can reverse its
position in the passage by a twisting somersault. This begins with a
lowering of the head, whose forward edge is thrust under the anterior
coxae. Thereafter the body is swung forward and downward and
during this arc it is twisted sidewise. The end result is that when the

62

Psyche

[March

minor regains its feet it is facing in the opposite direction. No major
of rohweri was ever seen to behave in this way.

The foraging activities of rohweri are of interest since there is
evidence that it deliberately forages on the ground. This is probably
true of texanus as well but it has not yet been conclusively proved in
that species. On one occasion Dr. F. G. Werner took six foraging
workers of rohweri from white cholla in the Saguaro National
Monument. Unless the ants were living in the cactus, which seems
completely unlikely, they must have reached it over the surface of the
soil. It is not clear why the foragers had visited the cholla. It was
not in bloom and efforts by both writers to interest the captive colonies
in cholla were unsuccessful. Workers in the aquarium colonies spent
much time crawling over leaves and twigs of various plants with
which they were kept supplied. When a worker fell from a leaf its
righting reaction was completely different from that of texanus.
When a worker of rohweri lands on its back it shows no fixed right-
ing reaction. The body is violently contorted and the legs are flailed
about until one of them anchors on something that enables the ant to
pull itself over. The stereotyped righting reaction of texanus has
been described elsewhere (4).

The junior author’s colony was fed on diluted honey, which was
supplied through a wick from a reservoir. They were also fed on the
juices of phalaenid caterpillars. It was necessary to tear the cater-
pillars open before the workers would feed on them. Entire insects,
either alive or dead, were avoided, as was pollen taken from honey
bees. The colonies of the senior author were fed on pollen from the
start. It was found that rohweri will accept a wide variety of pollen
if it is smeared on the surface of leaves, although they seldom take it
from the anthers of flowers. Of the various sorts of pollen fed to the
colonies that of Ouercus agrifolia was clearly the most relished. As
will be shown, the colonies were also fed with aphid honey dew. As a
rule they preferred this to pollen but on one occasion, while the ants
were feeding on honey dew, catkins of Quercus agrifolia were placed
in the aquaria. The response to these was immediate and spectacular.
The foragers feeding on honey dew deserted it for the oak catkins
and practically every worker turned out to take pollen from them.
They gathered such quantities of pollen that they returned to the nest
with masses of grains in their jaws. These masses were held against
the heads of the larvae who nibbled away the pollen grains. The
original experiments with honey dew were disappointing. The ants
paid little attention to coccids on the leaves of Quercus chrysolepis or
to the rims of liquid which surrounded them. They were definitely

1965]

Creighton and Nutting — Cryptocerus

63

interested in Aphis fabi> a species that is abundant on Mesembryan-
themum at Riverside. The ants tried to milk the aphids but the
latter, who were clearly afraid of the rohweri workers, failed to
cooperate. They would usually run away from the ants, which
resulted in some of them being1 killed when the ants tried to catch
them. Much better results were secured with Aphis sphaericola. This
aphis produces such large quantities of honey dew that it will drip
from the leaves on which the aphids are feeding. When leaves of
Viburnum suspensum bearing Aphis sphaericola and coated with its
honey dew were placed in the aquaria the foraging ants gorged
themselves until their intersegmental gastric membranes were visible.
This they did by lapping the honey dew from the surface of the
leaves. The ants paid little attention to the aphids and made no
attempt to milk them.

Since the colonies were well-supplied with food a number of eggs
were soon present. When the major or female handles an egg it is
positioned so that its long axis runs through the notch in the anterior
edge of the cephalic disc. The larger eggs fit the rim of this notch
closely when so positioned. Most of the eggs hatched in about 27
days. The voiding of the larval meconium was observed three times
and took between 60 and 75 minutes. The movement of the black
meconium, both within the larva and during its emergence is so slow
that it is difficult to follow. Infrequent contractions of the posterior
half of the larva probably reflect peristaltic movements of the gut
within. Once outside the larva the meconium invariably attracted
workers, both majors and minors, although they were never observed
to assist the larva in any way. It was only after the meconium was
presented that the larva was groomed. On one occasion the meconium
was eaten by the attendant worker. On the other two it was carried
to the dump in the feeding chamber. The period between the passing
of the meconium and pupation was from six to ten days. The pupal
moult was never observed. New pupae are ivory white and the first
suggestions of pigmentation appear in the compound eyes. The color
of the pupa darkens from yellow ochre to brown during the second
and third weeks. There is a four- or five- day callow period after the
adult emerges. In the senior author’s colonies worker brood developed
from egg to adult in about three months (egg to larva ±27 days;
larva to pupa ± 33 days; pupa to adult ± 23 days). The nests were
kept at room temperature and the range, for the most part, lay
between 6o°F and 70°F. Since the temperature range to which a
free colony is subjected is far wider, these figures are useful only as an
indication of the relative length of the several stages. Moreover, the

64

Psyche

[March

development of the larva may be drastically modified. In the junior
author’s colony eight males emerged during the period from July 3
to July 24, 1963. These males developed from eggs which had been
laid in June of 1962. These eggs developed into larvae in about a
month and the larvae grew for about two months. But from Septem-
ber 1962 to June 1963 the larvae showed no further development.
In short, these eight males over-wintered as larvae. They emerged in
July and it is interesting to note that the marriage flight of rohweri
occurs in that month. The senior author took a female of rohweri
which had just completed her marriage flight in Baboquivari Canyon
on July 28, 1951.

Literature Cited

1. Wheeler, W. M., Proc. New Eng. Zool Club, 6, pp. 29-35 (1916)

2. Smith, M. R., Proc. Ent. Soc. Wash. 49, No. 1, pp. 29-40 (1947)

3. Creighton, W. S. & Gregg, R. E., Psyche, 61, No. 2, pp. 41-57 (1954)

4. Creighton, W. S., Psyche, 70, No. 3, pp. 133-143 (1963)

CONTRIBUTIONS TO A RECLASSIFICATION OF THE
FORMICIDAE. IV. TRIBE TYPHLOMYRMECINI
(HYMENOPTERA)

By William L. Brown, Jr.

Department of Entomology, Cornell University

The Typhlomyrmecini (spelling here emended) are a tribe of
Ponerinae here considered to contain the single small genus Typhlo-
myrmex. In this sense the tribe dates only from Brown, 1953. The
name Typhlomyrmicini (sic) } however, goes back to Emery, 1 9 1 1 ,
who first proposed it as a subtribe of tribe Ectatommini to contain the
three genera Prionopelta , Typhlomyrmex , and Rhopalopone. Brown
(1950) showed that Prionopelta belongs to tribe Amblyoponini, while
Rhopalopone is a synonym of Gnamptogenys in tribe Ectatommini
(Brown, 1958). After these subtractions, the genus Typhlomyrmex
could not be placed comfortably in any existing tribe, and its present
taxonomic position is an expression of this fact.

At first sight, Typhlomyrmex workers look like rather ordinary
small cryptobiotic members of tribe Ponerini, although the frontal
lobes are not as prominently developed as in Ponerini, and the petiole
is never quite “right” in form. The males and larvae clearly conform
to Emery’s “Section Proponerinae,” including Amblyoponini, Ecta-
tommini, and Platythyreini in the modern sense; (the cerapachyines
all probably belong here as well), so that the resemblance of the
workers to those of certain Ponerini (in Emery’s “Section Eupone-
rinae”) is either convergent or else marks a side lineage from near
the base of the stock that led to the Ponerini.

Among “proponerines”, Typhlo?nyrmex shows some similarities to
Amblyoponini and to Ectatommini, but it can be distinguished from
both by the wing venation of the sexes and the larval mandibles. The
main similarity between Typhlomyrmecini and Amblyoponini, other
than in “basic ponerine” traits, lies with the shape of the petiolar node
of one Typhlomyrmex species, T. rogenhoferi. This node, because
of its elongate form without a distinct posterior face, resembles that
of an Amblyopone very closely in side view. In dorsal view, however,
T. rogenhoferi proves to have a much thinner (bilaterally compressed)
petiolar peduncle, and this makes it seem possible that its amblyoponine
features could have been convergently acquired. Whether or not this
is the correct interpretation, it is true that, aside from basic “pro-
ponerine” characters, the Typhlomyrmex adult has little in common

65

66

Psyche

[March

with the Amblyoponini. It shares more characters with the small
“degenerate” members of Gnamptogenys , but here again, though less
certainly, I feel that the similarities may be convergent ones. The
two main characters contributing to this opinion are the forewing
venation of the larger Typhlomyrmex species (Mfi arising basad of
cu-a) and the shape of the mandible in the larva (inflated basal part,
suddenly narrowed to an acute apical blade). Even these characters
do not weigh decisively against a possible origin of Typhlomyrmex
from ectatommine ancestors, and it must be admitted that the con-
vergence hypothesis is to some extent based on subjective impressions
that remain to be tested.

Tribe Typhlomyrmecini
Genus Typhlomyrmex

Typhlomyrmex Mayr, 1862, Verh. zool.-bot. Ges. Wien, 12: 736. Type
species: Typhlomyrmex rogenhoferi Mayr, 1862, monobasic.

Typhlomyrmex Emery, 1911, Gen. Insect., 1 18: 33-34, characterization and
catalog of species.

Nec Typhlomyrmex Gistel : J. Betrem and C. Jacot-Guillarmod have called
my attention to a generic name Typhlomyrmex obscurely published by
J. Gistel in his Mysterien der europaischen Insectennvelt (1856, cf. p.
447) for a “Myrmica typhlops L.” This species name is a nomen
nudum of Lund (not Linnaeus!), published in 1831 in Ann. Sci. Nat.,
23: 128. Because it was based on an unavailable species name, and is
itself without description, indication or figure, Typhlomyrmex Gistel is
considered to be a stillborn name (nomen nudum), and I am well
satisfied to let it subside into permanent nomenclatorial limbo.

Worker: Monophenic (“monomorphic”) or feebly polyphenic;

size small (full length under 2 mm to slightly over 5 mm) ; pigment-
poor, yellowish to ferruginous in color.

Head parallel-sided, or sides slightly converging anteriad, occipital
margin straight to slightly concave. Eyes reduced to minute vestiges
with or without pigment. Clypeus with a broad, convex median part
and narrow, concave side pieces, the anterior median border often
with a narrow translucent margin, in a minority of cases produced as
a variously-shaped small median process. Frontal carinae forming
small frontal lobes that lie close together and roof a small basal part
of the antennal scape insertion (but not the basal collar of the scape) ;
lobes not or only weakly pinched in behind, and not expanded as in
typical members of tribe Ponerini. Between the lobes lies a narrow,
often indistinct frontal fossa, and a shallow median furrow may run
back from the fossa to the vertex, or even to the occipital margin ; it
is not usually as distinct as in most Ponerini. Antennae short, with
thick scapes that either fail to reach the occipital margin, or else
surpass it just barely, When held straight back. Funiculus of 11

1965] Brown — Typhlomyrmecini 67

segments, with the last 3 or 4 enlarged and forming a more or less
distinct club.

Mandibles triangular in basic plan, their dorsal surfaces convex in
both directions; basal border distinct from masticatory border, al-
though they may meet either in an angle or a broad curve. Masticatory
border with minute crenulation, denticulation and/or small, uneven
teeth; apical tooth large to very large, and tending to cross with its
opposite number when the mandibles are fully closed. Shape and
dentition varying with the species.

Under-mouthparts relatively bulky; labrum bilobed, with a broad
median excision. Palpi segmented maxillary 1, labial 2.

Alitrunk robust, with rounded humeri and propodeum, slightly
constricted at posterior mesonotum ; promesonotal suture distinct and
apparently movable; metanotal groove distinct, only feebly impressed.
A distinct line curving posteroventrad from the propodeal spiracle
represents the upper edge of the metapleural gland atrium showing
through the integument. Legs short and thick; tarsal claws small,
those on the anterior legs toothed, the others simple; tibial calcariae
of middle and hind legs 1 or 2, indistinctly pectinate or simple.

Petiole briefly pedunculate; node distinct, variable in form (Figs.
1, 3) ; subpetiolar process well-developed, acute or rounded at apex.
Gaster porrect or slightly downcurved, of the usual ponerine type,
with slight but distinct constriction after postpetiole; sternum fused
to tergum in abdominal segment III (postpetiole) and IV, but not
fused in V ( T. rogenhoferi worker, female) . Sting well-developed
and acute, usually exserted in dried specimens.

Sculpture generally fine, varying in development with the species
and, within species, allometrically. Cranium longitudinally striate or
striolate above, especially mesad, mostly shading off to reticulate on
the sides. Mandibles and central part of clypeus usually smooth and
shining. Alitrunk, petiole and gaster smooth, with spaced punctation,
or partly reticulo-striate or otherwise roughened, the sculpture always
becoming weaker caudad. Pilosity fine, rather short, uneven, fairly
abundant and widely distributed. A pair of long fine sensory hairs
rises steeply from the clypeus.

Alate female, or gyne: Slightly (T. pusillus) to considerably

(T. rogenhoferi) larger than associated workers, and often darker in
color, at least around the ocelli ; darkest in T. rogenhoferi , which is
brown. Sculpture sometimes better developed than in workers, and
petiolar node distinctly shorter and more transverse. Compound eyes
large and hairy; ocelli developed.

Alitrunk somewhat box-like, with a rather flat dorsal surface;

68

Psyche

[March

notauli obsolete; parapsidal furrows present but inconspicuous. Vena-
tion nearly “complete/’ with the median abscissae of Rs (Rsf2#3)
missing, so that the cubital cell is undivided (single). The first
abscissa of M forks off from Cu basad of crossvein cu-a, as in the
army ants. (This pattern of venation does not hold for T. pusillus,
in which the veins are reduced and their relationships modified.) The
hind wing entirely lacks an anal lobe, but has the large discal cell,
usually with 2 or 3 stubs of apical abscissae corresponding to Rs, M
and Cu; another small cell may occur at the base of the discal cell
behind, or may be incorporated into the discal cell. The hamuli
number 3, and usually arise from a small darkened sclerotic patch a
little beyond the midlength of the costal margin.

In other characters, gynes resemble workers.

Male: (Based on T. rogenhoferi and T. clavicornis ) smaller and
more slender than the corresponding gyne, but the difference is slight
in the smaller species; dark brown to black in color; habitus typical
of proponerine males. Eyes large and hairy, occupying nearly half of
the sides of the head. Ocelli distinct. Scapes straight, of moderate
length, usually equal to about the basal 3 or 4 flagellar segments;
flagellum 12-segmented, the segments all longer than broad and
increasing very slightly in thickness toward the apex. Mandibles well-
developed, opposable or crossing at closure, dentition a variably
reduced copy of that of the corresponding worker. Palpi segmented
1, 2 (rogenhoferi) or 1, 1 (clavicornis). Clypeus with a large,
swollen mid section and small sunken side pieces.

Alitrunk with notauli developed only as the anterior arms of the
“Y” and obsolete medially; parapsidal furrows present but inconspicu-
ous. Wings as in gyne (see above). Legs slender, all three pairs with
tarsal claws toothed.

Petiole subclavate, i.e., with the peduncle rising gradually caudad
toward nodal summit, which is rounded; ventral tooth or process
present. Gaster with a slight constriction behind postpetiole; the
latter segment has tergum and sternum firmly fused, but the next
segment, abdominal IV, appears to have them connected only by thin
cuticle or membrane. Genitalia only partly retractile, with parameres
broadly rounded at apex; volsellae varying with the species (Figs. 7,
8) ; aedeagal valves ordinary, serrate. Hypopygium with a long, more
or less digitiform, upcurved, hairy, median process (ventral view,
Fig. 9)-

Head longitudinally striate, rest of body predominantly smooth
and shining, with fine scattered punctures. Pilosity fine, mostly erect
and rather short, abundant and generally distributed.

1965]

Broiun — Typhlomyr/necini

69

Larva: (After G. C. and J. Wheeler, 1952 (rogenhoferi —

robustus) and 1964 (pusillus). Thorax moderately stout and bent
ventrally; slightly constricted at first abdominal somite; remainder
of abdomen stout and ovoidal. Body densely covered with moderate-
sized branching (mostly trifid) hairs; head with a few bifid hairs.
Mandible distinctive, composed of a strongly inflated basal 2/5 and
a very narrow, acute apical 3/5, the latter with 2 small median teeth
in addition to the apical. This mandible is somewhat like those of
amblyoponine larvae, except that the basal portion is relatively much
wider than in Amblyopone.

Distribution and biology: So far as known, Typhlomyrmex is

restricted to the warmer parts of the Americas, from southern Mexico
to northern Argentina. Within this region, T. rogenhoferi is the most
widespread and by far the most often-collected species, being an in-
habitant of rotten logs in forest. This species is common in the
Amazon Basin, where I have seen nests of several hundred workers
moving in file through the rot zone just beneath the bark of a log. I
have examined several such aggregations in the field, but I was not
able to find definite indications of the prey of these undoubtedly
predaceous ants. In some sites in the Amazon Basin, where rogen-
hoferi is moderately common, I found the species in very close prox-
imity to termite colonies, but I never saw it actually taking or feeding
upon a termite.

T. pusillus appears to be a soil dweller in cultivated and pampas
areas as well as in forest (Kempf, 1961). It also seems to exist at
higher elevations (e. g., in a coffee plantation at Venecia, near Me-
dellin, Colombia). Probably it is strongly subterranean in foraging
and nesting habits. The small series taken by P. F. Darlington at
the mouth of the Amazon came from a rotten root in rain forest soil.

The remainder of the species are rare, and nothing is known of their
biology. Probably their habits are strongly cryptic; the large pro-
portion known from alate males and females indicates that most
samples are taken during or after nuptial flight. T. major may be
restricted to the south of Brazil and neighboring countries, while T.
clavicornis is widespread in South America. T. prolatus is known
only from the unique type, a female from Costa Rica.

Synonymic synopsis of Typhlomyrmex species
Typhlomyrmex clavicornis (Figures 4, 5, 7)
Typhlomyrmex clavicornis Emery, 1905, Bull. Soc. ent. Ital., 37: 112, nota,
alate female. Type locality: Mapiri, Bolivia. Holotype examined in
Coll. Emery, Museo Civico di Storia Naturale, Genoa.

Typhlomyrmex clavicornis var. divergens Forel, 1906, Ann. Soc. ent. Belg.,

0. 5 mm.

Psyche, 1965

Vol. 72, Plate 5

Brown — Typhlomyrmex

0.5 m nru

1965]

Brown — Typhlomyrmecini

71

50: 248, female, male (in cop.). Type locality: San Bernardino, Para-
guay. Type examined in Coll. Forel, Museum d'Histoire Naturelle,
Geneva. New synonymy.

Typhlomyrmex richardsi Donisthorpe, 1939, Ent. mon. Mag., 75: 161, male.
TL: Mazaruni Clearing, British Guiana. Type series examined in

British Museum (Natural History). New synonymy.

This species is distinguished in all castes by means of the wide
head; long, falcate apical mandibular tooth (Figs. 4, 5) ; and oblique
basal borders of the mandibles, which fail to meet the clypeus when
closed. The worker-female antennal club is prominent, as the name
suggests, and the petiolar node is short. A single worker (taken at
Bartica, British Guiana by H. O. Lang, together with winged
females) has a head length, without mandibles, of 0.67 and a head
width of 0.65 mm., which is within the size range of the smaller T.
rogenhoferi workers.

The type series of T. richardsi consists of numerous males, ac-
companied on one card by a female specimen ( the latter not mentioned
by Donisthorpe). The differences cited among the synonymous
species by Forel and Donisthorpe mainly concern mandibular form
and the proportions of the antennal segments. On examining all the
types and comparing them with digms from British Guiana, I was
impressed by the similarity of the mandibles between members of the
same caste from different series. The basal segments of the funiculus
show noticeable variation in length among males, even in those on
one card, and I do not think they make a good diagnostic character.

T. clavicornis is known from the above-mentioned widely separated
localities in South America, ranging from Bolivia and Paraguay north
to British Guiana. A female with forewings missing, probably fully
alate when captured, comes from the Floresta di Tijuca, near Rio de
Janeiro, February i960, C. A. Campos Seabra leg. The T. richardsi
types were a part of a large series (apparently nearly all males) taken
from a nest of the social vespid Polybia bistriata.

Typhlomyrmex major , new status
Typhlomyrmex pusillus st. major Santschi, 1923, Rev. Suisse Zool., 30: 246,
worker. Type locality: Blumenau, Santa Catarina, Brazil. Location of
type unknown (not in Santschi Collection).

Explanation of Plate 5

Figures 1-6, Typhlomyrmex spp. Fig. 1, T. pusillus worker from El Rey,
Salta, Argentina, lateral view of body. Fig. 2, Same, head in full-face view.
Fig. 3, T. rogenhoferi, large worker, lateral view of petiole. Fig. 4, T.
clavicornis, mandible of gyne from British Guiana, hairs omitted. Fig. 5,
T. clavicornis from British Guiana, full-face view of male head. Fig. 6,
T. prolatus sp. nov., fertiale holotype, anterodorsal view of right mandible.
Drawings by Nancy Buffler, D. Alsop and the author.

72

Psyche

[March

I refer to this species a worker from Agudos, Sao Paulo State,
Brazil (W. W. Kempf leg.) with head length (HL) 0.71 mm, head
width (HW) 0.63 mm. In habitus, this worker is like a small
specimen of T. rogenhoferi , but the short, pusillus- like petiolar node
separates it at once. It is distinguished from pusillus by its larger size
and longer antennae, the scapes of which reach or surpass the occipital
border when they are held straight back. The cephalic striation is
also coarser and more distinct than in pusillus.

A female from Petropolis, Guanabara, Brazil (T. Borgmeier
leg.), apparently belonging to this species, has HL 0.76 and HW
0.65. As has already been indicated, the female described by Santschi
as T. foreli may belong to T. major.

Typhlomyrmex prolatus species nov. (Figure 6)

Diagnosis (gyne) : A medium-sized Typhlojnyrmex with unusually
elongate, narrowly subtriangular mandibles; basal border short and
curving broadly into long, indistinctly denticulate masticatory border
(Fig. 6) ; apical tooth very long and acute. Petiolar node as seen
from above broader than long, with feebly concave (almost straight)
anterior border and strongly concave posterior border. Postpetiole
with a distinct, sharp median longitudinal carina on the anterior third
of its dorsal surface.

Holotype gyne, further description: Total outstretched length

(TL) 3.8 mm, head length (HL) 0.72, head width without eyes
(HW) 0.63, in full-face view, closed mandibles extend beyond
median clypeal margin (ML) 0.31, straight-line length of right
mandible from external point of insertion to apex 0.49, length of
alitrunk (WL) 1.13, scape length 0.52, greatest diameter of eye 0.18,
petiolar node length in dorsal view 0.20, width 0.33 mm. Cephalic
index (HW/HL X 100) 88.

Head with parallel, feebly convex sides; occipital border with a
shallow v-shaped concavity. Eyes feebly convex. Clypeus with the
raised median area very smooth, with a broad strip hairless. Mandibles
smooth, with small elongate punctures, becoming striatopunctate and
opaque laterally toward insertions. Antennal scapes just barely reach-
ing occipital margin in full-face view when held straight back from
insertions ; antennal club rather distinctly 3-segmented ; flagellar
segments 2-8 broader than long. Median furrow distinct, reaching
anterior ocellus. Ocelli small but distinct.

Remainder of body much as in other species. Middle and hind
tibiae each with a single simple calcar. Forewing as in T. rogenhoferi ,
but M forking from Cu even farther back toward wing base. In hind
wing, there is only a single large cell.

1965]

Brown — Typhlomyrmecini

73

The median carina on the postpetiole is a curious feature; it is
symmetrical and does not look like a pathological condition. It is
followed by a flat central area, which may even be slightly impressed.
It remains to be seen whether the corresponding worker also carries it.

On the head, the longitudinal striation is indistinct except on the
cheeks, and dense opaque reticulo-punctulation prevails. Dorsal
surface of alitrunk densely punctulate, opaque to subopaque; pro-
podeum mostly smooth and shining; sides of alitrunk weakly shining,
pronotal part densely punctulate, remainder finely longitudinally
striate, with scattered punctures. Petiole smooth and shining, with
sparse punctulation on sides. Gaster smooth and shining, but with
dense punctulation. Scapes densely punctulate, subopaque, as are also
most of legs; mesal surfaces of femora smooth and shining.

Pubescence short, reelinate or appressed, fairly abundant over most
dorsal body surfaces, gaster and appendages; longer fine hairs on
clypeus, mandibles, and sparse on gaster above and below (abundant
at gastric apex). Short oblique hairs extend beyond pubescence on
scapes, funiculi and legs.

Color light ferruginous, legs lighter and more yellowish; head
infuscated around ocelli.

Holotype (and only known specimen) from the vicinity of San
Jose, Costa Rica, in 1940 (H. Schmidt leg.). Deposited in the
collection of W. W. Kempf, Sao Paulo, Brazil.

Typhlomyrmex pusillus (Figures 1, 2)

Typhlomyrmex pusillus Emery, 1894, Bull. Soc. ent. Ital., 26: 141, pi. 1,
fig. 2, worker. Type locality: Bolivia.

Typhlomyrmex schmidti Menozzi, 1927, Ent. Mitt., 16: 268, female, male.
Type locality: vie. San Jose, Costa Rica. Syntypes in Istituto di Entomo-
logia della Universita, Bologna, Italy; 2 alate females examined. New
synonymy.

This is the smallest species of the genus. It will probably turn out
to be much more common and widespread than it seems at present ; its
habitat in the soil and its very small size have made it scarce in
collections.

Samples that I have referred to this species show so much variation
that they may actually represent more than one species. Specimens
from the south (northern Argentina, Santa Catarina) and the
Colombian Andes (Venecia, near Medellin) average larger and more
robust (HW 0.34-0.50 mm) than those from Amazon drainage and
the Guianas. Among the smaller forms, most samples (Surinam:
Dirkshoop and Maripaheuvel ; Brazil: near Belem do Para; Peru:
Finca Santa Beatriz, Chanchamayo) have the petiolar node and

74

Psyche

[March

postpetiolar dorsum very finely and densely sculptured, usually strio-
late with interspersed shallow punctulation, and opaque or nearly so,
but two samples from Tambahredjo in Surinam have the petiole and
postpetiole smooth and shining, with only the usual abundant fine
punctures. Of special interest is the occurrence, in two of the above
samples (Peru: Finca Santa Beatriz, Chanchamayo, io°57’S,

75°i2’N, iooo m, C. A. Portocarrero leg., No. 88-SB29, 11 July
1964. Brazil: Utinga tract, near Belem, Para, P. F. Darlington
leg., No. 335, Aug. 24, 1962) of a small, square to sharply trapezoi-
dal lamellar lobe or process on the median anterior clypeal margin.
This process is variable in shape and size, and is transparent and thus
very difficult to see unless the mandibles are open at least partly. It
appears to represent a modification of the lamellar free clypeal margin
seen in other samples. This margin is usually evenly convex, but some
Surinam specimens show a tendency for its most central part to form
a narrow, shallow truncate lobe. Such a character would normally
separate good species among ponerine ants, but the overall variation
in the small Typhlomyrmex calls for caution and more than the
present meager material before we draw new species boundaries.

The wings of the female of this species are atypical for Typhlomyr-
mex, in that M leaves Cu distad of cu-a, and the crossvein r-m is
missing, so that the cubital cell is open at its apex.

The Santa Beatriz collection was made from a single chamber about
4 mm in diameter, located in the ground; Portocarrero found there
21 workers and one queen, plus 8 pupae, 1 larva, and 10 eggs. The
Utinga collection came from a red-rotten root in rain forest.

Typhlomyrmex rogenhoferi (Figures 3, 8-1 1)

Typhlomyrmex rogenhoferi Mayr, 1862, Verh. Zool.-bot. Ges. Wien, 12: 737,
worker. Type locality: “Amazonas.”

Typhlomyrmex Rogenhoferi r. robustus Emery, 1890, Bull. Soc. ent. Ital., 22:
40, worker. Type locality: Alajuela, Costa Rica. Syn. Brown, 1957.
Typhlomyrmex robustus subsp. manco Wheeler, 1925, Ark. f. Zool., 17A
(8): 2, worker. Type locality: Pablobamba, Peru. Syn. Brown, 1957.
Centromyrmex sculpturatus Santschi, 1931, Revista Ent., Rio de Jan., 1: 266,
dealate female. Type locality: Panama Canal Zone. Holotype examined
in Naturhistorisches Museum, Basel. New synonymy.

Easily recognized by the relatively large size, triangular mandible
and shape of the petiolar node in worker and female. The male can
be recognized by size, the distinct angle between basal and masticatory
borders of the mandible, and by the distinctive volsella of the genitalia.

Interesting Brazilian records for the species have been furnished
by W. W. Kempf from his collection : Goias State, Goiaffiia,

Campinas ( Schwarzmaier leg.). Espirito Santo State, Santa Teresa

1965]

Brown — Typhlo?nyrinecini

75

Figures 7-11, T yphlomyrmex spp. Fig. 7, Right half of genital capsule of
T. clavicornis male from British Guiana, viewed from inside, ventral side
to right, semidiagrammatic. Fig. 8, same, T. rogenhoferi, aedeagal valve
omitted. Fig. 9, hypopygial process of T. rogenhoferi, ventral view. Fig. 10,
maxillary palpus (left) and labial palpus (right) of T. rogenhoferi. Fig. 11,
right wing of T. rogenhoferi Figs. 8-11 are drawn from a male from
Perene, Peru. Drawings by D. Alsop and the author.

76

Psyche

[March

(O. Conde leg.)- Minas Gerais State, Serra Caraga (K. Lenko leg.).
Sao Paulo State, Fazenda Itaquere, Nova Europa (K. Lenko leg.).
The species ranges from Bolivia to Veracruz State in Mexico.

Typhlomyrmex foreli

Typhlomyrmex foreli Santschi, 1924, Ann. Soc. ent. Belg., 64: 6, female.

Type locality: Rio Negro, Parana, Brazil. Location of type unknown

(not in Santschi Collection).

This species was described from a single gyne. I suspect it to be the
gyne of T. major , but some items in Santschi’s description will leave
doubts until the type can be re-examined. Probably the “court sillon
median” on the clypeus can be dismissed as an illusion caused by the
shiny surface here; I have noticed this in other species. Santschi says
that the external margins of the mandibles are slightly concave, a
description that will fit no specimen of any species of Typhlomyrmex
in full-face view, but will fit all of them if viewed obliquely from
above and slightly to the side.

The “bord terminal long, finement denticule avee une dent apicale
bien developee” would fit the new species prolatus (see above), but
would also fit T. major reasonably well.

Santschi’s statement, “The first article of the funiculus is not quite
as long as the three following ones taken together” fits prolatus fairly
well, while in a gyne I take to be major, the first funicular segment
is shorter than this.

In spite of these difficulties, the like of which often crop up in
connection with Santschi’s descriptions of ants, I think it probable
that T. foreli and T. major are conspecific. The distributional
evidence weighs for this decision, and Santschi mentions no posterior
concavity of petiole and no median postpetiolar carina such as prolatus
carries.

Summary of changes proposed
in species-level taxonomy of Typhlomyrmex

clavicornis Emery

— clavicornis var. divergens Forel, n. syn.

— richardsi Donisthorpe, n. syn.

f foreli Santschi (possibly a synonym of T. major )
major Santschi, raised to species level
prolatus sp. nov.
pusillus Emery

= schmidti Menozzi, n. syn.
rogenhoferi Mayr

— rohustus Emery

1965]

Brown — Typhlomyrmecini

77

= robustus manco Wheeler
= Centromyrmex sculpturatus Santschi, n. syn.

Note: Brown (1953) cited Prionopelta marthae Forel as a syno-
nym of Typhlomyrmex rogenhoferi. The synonymy was based on a
specimen labeled as a cotype of P. marthae , found in the Wheeler
Collection, and which is clearly a specimen of T. rogenhoferi. Studies
in European museums in 1963 and 1964 show that this specimen is
mislabeled, because P. marthae syntypes (“cotypes”) in the Forel
Collection and elsewhere are true Prionopelta close to, and possibly
conspecific with, the species currently called Prionopelta antillana.
P. marthae must, therefore, be deleted from the synonymy of T.
rogenhoferi and returned to genus Prionopelta.

Key to Typhlomyrmex species, workers

1. Petiolar node longer than high, without a differentiated posterior

face (Fig. 3) rogenhoferi

Petiolar node as high as or higher than long, with a differentiated
posterior face (Fig. 1) 2

2. Head width <0.55 mm pusillus

Head width >0.55 mm 3

3. Mandibles triangular, with basal border fitting tightly against
clypeus at full closure; apical tooth stout, not notably elongate

major

Mandibles more elongate, basal borders oblique and not closing
up against clypeus; apical tooth notably elongate and very acute

( Fig. 4) clavicornis

Note: The worker of T. prolatus, at present unknown, probably
would key to couplet 3, where it would undoubtedly be dis-
tinguished from both alternatives by having mandibles like those
of its gyne (Fig. 6).

Key to Typhlomyrmex species, gynes

1. Petiolar node without a differentiated posterior face; size large,

head width (without eyes) normally >0.85 mm rogenhoferi

Petiolar node with a differentiated posterior face; size smaller .. 2

2. Head width (without eyes) <0.58 mm; r-m crossvein missing

from forewing pusillus

Head width (without eyes) >0.58 mm; r-m present in forewing

; 3

3. Petiolar node as seen from above concave behind; mandibles

elongate but only weakly denticulate, of a particular form (Fig.
6) ; postpetiolar disc with a distinct anteromedian longitudinal
carina prolatus

7»

Psyche

[March

Petiolar node as seen from above with a straight or convex
posterior margin; mandibles not as in Fig. 6; no carina on post-

petiolar dorsum 4

4. Same as couplet 3 of worker key, above: major vs. clavicornis.
Note: T. foreli is not included in the key. It may be the female
of T. major.

Acknowledgements

I am indebted to my colleagues G. Grandi, F. Keiser, W. W.
Kempf and C. Portocarrero for the opportunity to review important
material in their care. This project was supported by National
Science Foundation Grants G-23680 and GB-2175, and publication
was aided by a grant from the Grace Griswold Fund, Department of
Entomology and Limnology, Cornell University.

References Cited

Brown, W. L., Jr.

1950. Morphological, taxonomic and other notes on ants. Wasmann
Jour. Biol., San Francisco, 8: 241-250.

1953 (1952). Composition of the ant tribe Typhlomyrmicini. Psyche,

59: 104.

1958. Contributions toward a reclassification of the Formicidae. II.
Tribe Ectatommini (Hymenoptera). Bull. Mus. comp. Zool.
Harv., 118: 173-362.

Emery, C.

1911. Ponerinae, in Wytsman, Gen. Insect., 118: 125 pp. and 3 pi.
Kempf, W.W.

1961. A survey of the ants of the soil fauna in Surinam (Hymenoptera:
Formicidae). Studia Ent., Petropolis, Brazil, 4: 481-524.
Wheeler, G. C., and J. Wheeler

1952. The ant larvae of the subfamily Ponerinae — Part I. Amer.

midi. Nat, 48: 111-144, 5 pi. Part II. Ibid., pp. 604-672, 2 pi.
1964. The ant larvae of the subfamily Ponerinae: Supplement. Ann.
ent. Soc. Amer., 57: 443-462, 19 figs.

A REVISION OF THE ANT TRIBE CARDIOCONDYLINI
(HYMENOPTERA, FORMICIDAE)

I. The Genus Prosopidris Wheeler1

By Jonathan Reiskind
Biological Laboratories, Harvard University

This is the first of a series of papers revising the tribe Cardio-
condylini, a group of small myrm'icine ants (up to 3.0 mm in length),
characterized by the possession of a pedunculate petiole, a wide
postpetiolar dorsum, propodeal spines, and a general lack of pilosity.
The tribe currently embraces the genera Cardiocondyla Emery
(virtually cosmopolitan, mainly Old World), Xenometra Emery
(West Indian) and Prosopidris (Papuasian). Xenometra includes a
single poorly known species which is apparently parasitic on Cardio-
condyla emeryi (Emery, 1917). Cardiocondyla and Prosopidris are
free living, and form small inconspicuous colonies usually in the soil
or in rotting logs.

In this contribution Wheeler’s subgenus Cardiocondyla (Prosopi-
dris) is elevated to full generic status. The worker and female types
of its single previously known species, P. sima Wheeler (Philippines),
are redescribed and a second species, P. papuana (New Guinea), is
newly described. The papuana material includes the first known male
of the genus, along with females and workers. The male is highly
ergatoid, like those of some Cardiocondyla species.

Measurements and Indices: In order to characterize properly cardio-
condyline ants the following measurements and indices (with their
abbreviations) will be used in this and succeeding papers:

Head Length (HL) — Maximum length of head in frontal view
from clypeal apex to posterior border of
occiput.

Head Width (HW) — Width of head measured in frontal view
immediately behind the eyes.

Scape Length (SL) — Maximum measurable length of scape, not
including its articular condyle.

Weber’s Length (WL) — Weber’s length of mesosoma ( = ali-
trunk), measured in lateral view along a
line connecting the place where the prono-
tum joins the cervix and the apices of the

Research supported by U.S. National Science Foundation Grant No.
GB1634.

79

8o

Psyche

[March

the flanges of the sides of the propodeal
declivity (Brown, 1953). The lateral view-
ing position of specimens must be standard-
ized by lining up the tips of the propodeal
spines.

Pronotum Width (PW) — Width of pronotum, measured in dorsal
view, disregarding humeral angles.
Propodeal Spine L(ength) — Maximum measurable length of pro-
podeal spine, in lateral view, from its apex
to the far edge of the propodeal spiracle.
Petiole L(ength) — Length of petiole, measured in lateral view,
from the apices of the flanges of the sides
of the propodeal declivity to the dorsal
posterior border of the petiolar tergite.
Petiole Height — Height of petiole, measured in lateral view, on a
line perpendicular to the petiole length ;
from the dorsalmost point on the petiolar
node to the ventral side.

Postpetiole W(idth) — Maximum width of postpetiolar dorsum.
Postpetiole L(ength) — Midline length of postpetiolar dorsum.
Cephalic Index (Cl) — Head width expressed as a percentage of

head length. HW X 100/HL
Scape Index (SI) — SL X 100/HW
Mesosomal Index — PW X 100/WL

Propodeal Spine Index (Spl) — Propodeal Spine L. X 100/WL
Petiolar Index — Petiole L. X ioo/Petiole Height
Postpetiolar Index — Postpetiole L. X ioo/Postpetiole W.

Genus Prosopidris Wheeler new status

Cardiocondyla subgenus (Prosopidris) Wheeler, 1935, Psyche 42: 40-41.

Type-species: Cardiocondyla (Prosopidris) sima Wheeler, Ibid: 42-43.

Wheeler’s subgenus Prosopidris is here raised to full generic status.
This change of rank is based on the distinctive clypeus, the 11-
segmented antennae, and the high mesosoma, which readily distinguish
Prosopidris from Cardiocondyla , though the two genera are obviously
closely related.

Wheeler’s (1935) subgeneric diagnosis requires some modification
in light of the new species, Prosopidris papuana, described below. In
the workers the mesometanotal impression is lacking or very feeble.
In the female wing venation the stub of vein Rs, extending beyond the
cubital cell, may or may not be present. The only known male of the
genus is described below.

1965]

Reiskind — Cardiocondylini

81

Prosopidris papuana new species
(Figures 1-8)

WORKER

Measurements: Based on the holotype and 4 paratype workers
(holotype cited first) : HL 0.59 mm, 0.59-0.60 mm; HW 0.44 mm,
0.44 mm; Cl 75, 73-75 ; SL 0.49 mm, 0.49-0.52 mm; SI 111, m-
1 1 8 ; PW 0.33 mm, 0.33 mm; WL 0.70 mm, 0.68-0.70 mm; Meso-
somal Index 47, 47-48; Propodeal Spine L. 0.18 mm, 0.18-0.20 mm;
Spl 26, 26-29; Petiole L. 0.28 mm, 0.28-0.29 mm; Petiole Height
0.17 mm, 0.17-0.18 mm; Petiolar Index 159, 155-164; Postpetiole
W. 0.24 mm, 0.24 mm; Postpetiole L. 0.15 mm, 0.15-0.16 mm;
Postpetiolar Index 60, 60-65. Palpal formula Maxillary 5 : Labial 3
(paratype dissected).

Description: General form is shown in Figures 1 and 2. Head
subrectangular, sides sub-parallel, slightly convex; posterior corners
rounded ; occipital border slightly concave. Scape slightly curved
distally, exceeding occipital border by a distance about equal to its
maximum diameter. Antennae 1 1 -segmented, with a 3-segmented
club, the apical segment about 3 times as long as the preapical. Eyes
strongly convex, maximum diameter (in all specimens) 0.13 mm;
their anterior edges at a distance of 0.7 X their diameter from the
lateral genal border. Outer borders of mandibles convex; masticatory
border with 5 teeth, the 2 apical enlarged. Clypeus produced ante-
riorly, forming a shelf over the mandibular bases and bulging out-
wards; anterior border convex, with thick lateral sections anterior to
each antennal sulcus; posterior portion extending broadly between
the frontal carinae.

Mesosomal profile as in Figure 2 ; lacking a promesonotal con-
striction, humeral angles indistinctly rounded. Mesometanotal im-
pression weakly developed on mesosomal dorsum as a very shallow
concavity, lacking an incised sutural trace. Propodeal spines moderate-
ly long, pointed, uncurved. Petiolar node from above, oval, distinctly
wider than long. Postpetiole with convex anterior and posterior
margins and convex sides. Dorsal width of first gastric segment 2.1 X
postpetiole width.

Head and mesosoma strongly reticulate-punctate and mat; petiole,
postpetiole and gaster moderately reticulate and shining.

Yellow pilosity on clypeus, mandibles and posterior end of gaster.
Yellowish adpressed pubescence dense on antennae, moderate on head,
petiolar node, postpetiolar dorsum, and gaster.

Entire body yellow to dark yellow.

82

Psyche

[March

Figs. 1-3. Prosopidris papuana new species. Figs. 1 and 2, worker
(Holotype). Fig. 1. Head, frontal view. Fig. 2. Mesosoma and petiole,
lateral view. Fig. 3, female (Paratype) ; mesosoma and petiole, lateral view.
Scale line: 0.40 mm.

Type locality: Bisianumu, near Sogeri, Papua, 15-20 March, 1955
(E. O. Wilson acc. 655).

FEMALE

Measurements : Based on 5 specimens collected with the workers:
HL 0.60-0.62 mm; HW 0.46-0.47 mm; Cl 76-78; SL 0.49-0.52
mm; SI 103-114; PW 0.46-0.47 mm; WL 0.83-0.85 mm; Meso-
somal Index 54-57 ; Propodeal Spine L. 0.20-0.21 mm; Spl 23-25;
Petiole L. 0.31-0.33 mm; Petiole Height 0.20-0.21 mm; Petiolar
Index 146-167; Postpetiole W. 0.28-0.29 mm; Postpetiole L. 0.16
mm; Postpetiolar Index 56-59.

Description : General form of mesosoma as in Figure 3. Head,
antennae, clypeus and mandibles similar to worker. Ocelli slightly
elevated. Eyes strongly convex, maximum diameter (in all specimens)
0.16 mm; their anterior edges at a distance of 0.5 X their diameter
from the lateral genal border. Propodeal spines longer and thinner
than worker. Petiolar node and postpetiolar dorsum as in worker.
Dorsal width of first gastric segment 2.2-24 X postpetiole width.

Head and mesosoma strongly and coarsely reticulate-punctate and
moderately shining; petiole, postpetiole and gaster moderately reti-
culate and shining.

Yellow pilosity as in worker. Yellowish adpressed pubescence dense

1965]

Reiskind — Cardiocondylini

8 3

on antennae, postpetiolar dorsum and gaster, and moderate on head
and mesosoma.

Color as in worker, pterostigma pale brown, wing veins pale
yellow; apical stub of wing vein Rs absent beyond cubital cell.

MALE

Measurements: Based on a single specimen collected with the

workers: HL 0.59 mm; HW 0.47 mm; Cl 81; SL 0.44 mm; SI
93; PW 0.36 mm; WL 0.75 mm; Mesosomal Index 48; Propodeal
Spine L. 0.15 mm; Spl 20; Petiole L. 0.34 mm; Petiole Height 0.21
mm; Petiolar Index 162; Postpetiole W. 0.29 mm; Postpetiole L.
O.15 mm; Postpetiolar Index 52.

Description : Ergatoid. General form as shown in Figures 4 and 5.

Head with sides slightly divergent behind eyes, then strongly
converging; posterior corners rounded; occipital border slightly
concave. Scapes straight, their apices falling slightly short of the
occipital border. Antennae 12-segmented. Funiculus and clypeus as in
worker. Eyes strongly convex, maximum diameter 0.13 mm; their
anterior edges at a distance of 0.7 X their diameter from the lateral
genal border. Mandibles narrowly falcate, ending in a point. Frontal
carinae more extensive than in the worker.

Mesosoma high, in lateral view, with no promesonotal constriction,
but with distinctly rounded humeral angles; mesometanotal im-
pression and constriction clearly developed, but lacking an incised
suture. Propodeal spines blunt; shorter and wider at base than in
worker. Petiolar node and postpetiole dorsally, as in worker. Dorsal
width of first gastric segment 2.0 X postpetiole width.

Pro- and mesonotum strongly reticulate-punctate; rest of mesosoma,
head, petiole, postpetiole and gaster moderately reticulate. Whole ant
very shining.

Yellowish pilosity on clypeus and posterior end of gaster. Yellowish
adpressed pubescence very dense on antennae, pronotum, mesonotum,
gaster, and dorsa of petiole and postpetiole.

Entire body yellow.

Genitalia: General form shown in Figures 6, 7, and 8. Basal ring
entire ; the genital foramen inclined ventrally. Gonoforceps each with
a lobate, hirsute ventrolateral subapical extension; a peculiar large
ventrally directed curved tooth arising on the ventral part of the
inner face (Figure 8). Volsella hook shaped, its apex directed
ventrally, the base swollen with a small acute ventrally directed tooth
(Figure 8). Penis valves rounded apically, each with a row of fine
teeth on its ventral edge.

84

Psyche

[March

Figs. 4-8. Prosopidris papuana new species. Figs. 4-8, male (Paratype).
Fig. 4. Head, frontal view. Fig. 5. Mesosoma and petiole, lateral view.
Fig. 6. Genitalia, dorsal and ventral views. Fig. 7. Genitalia, left half,
external lateral view, showing gonoforceps and basal ring. Fig. 8. Geni-
talia, right half, internal lateral view, showing penis valve, volsella,
gonoforceps and basal ring. Scale line: 0.40 mm., for Figs. 4 and 5; 0.15
mm., for Figs. 6, 7 and 8.

Type deposition: The holotype and most paratypes are deposited
in the Museum of Comparative Zoology at Harvard College, Cam-
bridge, Massachusetts (Type No. 31156), single worker and female
paratypes in the Australian National Insect Collection, CSIRO,
Canberra.

Prosopidris sima Wheeler

Cardiocondyla (Prosopidris) sima Wheeler, 1935, Psyche 42: 40-43, figs.

The worker and female are adequately described (with figures of
the worker) in Wheeler’s paper. The measurements and indices
applied above to Prosopidris papuana have also been applied to P.
sima and are listed here:

1965]

Reiskind — Cardiocondylini

85

WORKER

Based on 5 workers from Wheeler’s syntype series (MCZ Type
No. 20798), the lectotype (by present designation, so labelled) cited
first, the others paralectotypes: HL 0.57 mm, 0.55-0.57 mm; HW
0.46 mm, 0.44-0.46 mm ; Cl 80, 79-8o ; SL 0.5 1 mm, o.47"°*52 rnm ;
SI hi, 107-114; PW 0.34 mm, 0.33-0.34 mm; WL 0.68 mm,
0.67-0.68 mm; Mesosomal Index 50, 48-50; Propodeal Spine L. 0.20
mm, 0.18-0.20 mm; Spl 29, 26-29; Petiole L. 0.28 mm, 0.26-0.29
mm; Petiole Height 0.20 mm, 0.19-0.20 mm; Petiolar Index 141,
1 3 3- 1 54J Postpetiole W. 0.23 mm, 0.23-0.24 mm; Postpetiole L.
0.16 mm, 0.15-0.16 mm; Postpetiolar Index 69, 64-71. Dorsal width
of first gastric segment 2. 1-2.3 X as wide as postpetiole.

FEMALE

Based on three paralectotypes: HL 0.59 mm; HW 0.46-0.47 mm;
Cl 78-81; SL 0.49-0.51 mm; SI 104-m; PW 0.46-0.47 mm; WL
0.80 mm; Mesosomal Index 57-59; Propodeal Spine L. 0.21-0.22
mm; Spl 27; Petiole L. 0.31-0.33 mm; Petiole Height 0.20-0.21 mm;
Petiolar Index 148-167; Postpetiole W. 0.25-0.26 mm; Postpetiole
L. 0.16 mm; Postpetiolar Index 63-65. Dorsal width of first gastric
segment 2. 3-2. 4 X as wide as postpetiole.

Type Locality: Dansalan, on Lanao Island, Philippine Islands.

The following chart presents the major distinguishing character-
istics of the two Trosopidris species considered above:

Prosopidris sima Wheeler

Prosopidris papuana n.sp.

Worker

Anterior clypeal border straight.
Mesosoma almost smooth, shining.
Yellow-brown.

Anterior clypeal border evenly con-
vex.

Mesosoma mat, highly reticulate-
punctate.

Yellow to dark yellow.

Female

Anterior clypeal border flattened.

Propodeal spines thicker than in
worker.

Pronotum shining, almost smooth.
Pronounced lateral propodeal striae
converging on propodeal spine.
Stub of vein Rs extending beyond the
cubital cell.

Anterior clypeal border evenly con-
vex.

Propodeal spines thinner than in
worker.

Pronotum highly reticulate-punctate.
No clear propodeal striae.

Stub of vein Rs beyond cubital cell
lacking.

86

Psyche

[March

The author wishes to thank Dr. Robert W. Taylor of Harvard
University for his invaluable assistance.

References

Brown, W. L. Jr.

1953. A Revision of The Dacetine Ant Genus Orectognathus. Mem.
Queensland Mus. 13 (1): 84-104.

Emery, C.

1917. Questions de nomenclature et synonymies relatives a quelques
genres et especes de Formicides. Bull. Soc. ent. Fr. 1917: 94-97.
Wheeler, W. M.

1935. New Ants from the Philippines, Psyche 42: 38-52.

“QUEENLESSNESS,” WORKER SIBSHIP, AND COLONY
VERSUS POPULATION STRUCTURE
IN THE FORMICID GENUS RHY T'lD OPONERA

By Caryl P. Haskins1 and Roy M. Whelden2

INTRODUCTION

William Morton Wheeler, in his Colony Founding Among Ants
(!933)> called special attention to the fact that in a number of
formicid genera, and particularly in the socially primitive subfamilies
Ponerinae and Cerapachyinae, typical alate female forms have never
been described. In such genera as Onychomyrmex, Eusphinctus,
Acanthostichus, Megaponera , and Plectroctena, this normal female
may be replaced by a wingless ergatogyne, intermediate in structure
between queen and worker. The same condition obtains among
certain species of the archaic subfamily Myrmeciinae, as Wheeler also
pointed out. In some cerapachyine species, normal females typically
coexist with ergatogynes, and such caste duality occurs elsewhere also.

In certain respects, the shift from a “queen-like” toward a “worker-
like” form of reproductive female is curious and striking. At first
sight it may even suggest a reversal of the trend tending to emphasize
the dichotomy between more “vegetative” and more actively “for-
aging” forms which, as Wigglesworth (1954) and Kennedy (1961)
have pointed out, is so characteristic of evolution in insects generally,
whether at the level of “successive polymorphism” in the juvenile and
adult phases of the individual, or of “alternative polymorphism”
among adult populations of such forms as aphids, migratory locusts,
and the social insects. Among the ant genera cited, however, evolution
from alate to ergatoid reproductive may only superficially appear to
lie in that direction, for the ergatoid is clearly at least as effective a
reproductive as the winged female. In most species with such females,
moreover, it seems likely that the ergatoid has been directly derived
by a stabilization of a queen-worker intercaste, as Brown (i960) has
suggested, and merely replaces the normal queen, with no drastic
change in the general economy or structure of the colony. Even in the
ponerine genus Leptogenys sens, str., where the laying female is no
longer morphologically distinguishable from the worker, the course
of evolution still seems relatively clear. As Wheeler (1933) pointed
out, a normal female is present in the related Lobopelta langii , and

’Carnegie Institution, Washington, D.C.

2Haskins Laboratories, New Durham, New Hampshire

87

88

Psyche

[March

in L. ergatogyna the wingless laying female still possesses well de-
veloped ocelli and a normal female thorax. In the genus Myrmecia
similar series, ranging through increasing female microptery toward
total aptery, can be assembled, culminating in the ergatoid-like gynes
of, for instance, M. tarsata. In all these species, however, there is
little evidence of any significant modification in the social structure of
the colony, which remains at base a typically matrifilial community.

Certain other ponerine genera referred to by Wheeler, however,
may present a rather different picture. In the genera Diacamma ,
Streblognathus , and Dinoponera, and in some species of Rkytido-
ponera, for example, distinguishable ergatogynes have not been
reported. A male of a Philippine species of Diacamma has been
described in copula with a form indistinguishable from a worker by
Wheeler and Chapman (1922), suggesting that in this organism the
normal reproductive may have been replaced by what is in effect a
worker-producing worker. Our overall knowledge of the first three
genera, however, is hardly sufficient to support even speculation about
their social situations at present. For the genus Rhytidoponera,
however, something more is known or knowable, and it may quite
possibly represent a situation of considerable interest to the student of
social evolution in the Formicidae.

The Ponerine genus Rhytidoponera comprises an extensive but
relatively compact series of species inhabiting the Australian and
parts of the Melanesian and Malaysian areas, ranging from New
Caledonia in the east through New Guinea and neighboring parts of
Melanesia to Timor, the Moluccas, and the southern Philippines in
the west and occcupying a large portion of the Australian continent
and of Tasmania (Brown, 1954, 1958; Wilson, 1958, 1959a). They
are members of the widely distributed ponerine tribe Ectatommini,
bearing considerable resemblance in many respects to the generalized
New World tropical genera Acanthoponera and Ectatomma , as well
as to the pantropical Heteroponera, with the Old World components
of which they may well have shared common ancestry. The females
of Ectatomma and Acanthoponera, so far known, are of the normal
winged form. Those of Heteroponera may be winged or ergatoid.

In at least three species of Rhytidoponera, R. impressa, R. purpurea ,
and R. chalybaeae , typical winged queens are the rule. Normally a
single queen is found in each colony examined in the field, and com-
munities appear to be initiated by isolated dealated females following
a normal Ponerine dispersion and mating flight. These species are
confined in distribution to well-watered and warmer areas, ranging
from New Guinea and eastern Queensland rain forest southward

1965]

Haskins and Whelden — Rhytidoponera

89

along a belt down the eastern edge of Australia at least as far as
south central Victoria (Brown, 1958).

In a second and much larger group of Rhytidoponera species, an
exactly opposite situation obtains with respect to the queen. Many of
these species include among the largest and most conspicuous members
of the genus, are widely distributed, abundant, and well known,
especially in the drier areas of Australia, and have been extensively
collected over long periods. Yet in none of them has a reproductive
morphologically or functionally distinguishable from a normal worker
ever been described.

Finally there is a third group, designated by Brown (1958) the
Rhytidoponera metallica complex, which may be the most interesting
from the standpoint of social evolution. The type species is one of the
most widely distributed and ubiquitous of Australian ants; an inhabit-
ant of thickly populated as well as remote situations over a very large
area both temperate and subtropical; and so familiar as to have been
known to a wide public for many years by the popular name of
“greenhead” ant. Alate typical queens of this species have been
described, and are represented in limited numbers in some collections,
notably that of the Harvard Museum of Comparative Zoology.
Wheeler described a single dealate and possibly colony-founding
female of R. inornata, a member of the complex from southwestern
Australia in 1931 (Brown, 1958). Brown (1958) has described a
dealate female of another related species, R. aspera , collected by
H. Hacker in southeastern Queensland and also in the Harvard
Museum of Comparative Zoology. A single perfect female of R.
victoriae, taken by Brown at Seaford, Victoria, is in the same col-
lection. But it is striking that so few typical females have been
identified in a complex of species as extraordinarily abundant and
well-collected. It is clear that the vast majority of colonies in nature
must exist without such females.

Even more interesting is the fact that in no species of Rhytidoponera ,
including those of the metallica complex, has a queen-worker inter-
mediate ever been recorded. This could suggest that evolution to the
loss of the typical female took a somewhat different course from that
in the Lobopelta-Leptogenys complex or even in Heteroponera.
Instead of the alate female reproductive being morphologically modi-
fied toward a stabilized intermediate between queen and worker while
continuing the same functional role in the colony, the original queen
caste may have disappeared entirely and one or more laying workers
substituted as the usual reproductives. If, as Carroll Williams (in
Brown, i960) has suggested, worker development in ants is due to a

90

Psyche

[March

precocious decline of juvenile hormone titer in the maturing larva, or
if on the other hand as Brian (1959, 1961) proposes, to a sharp rise
in the concentration of ecdyson near the critical period or periods of
caste determination in larval ontogeny, it seems conceivable that
mutations have accumulated in evolution affecting neurosecretory
products or processes or timing, such that the threshold for worker-
queen determination is passed only rarely in species of the R. metallic a
complex, and never in many of the larger species. Alternatively, it is
conceivable that dimorphic female reproductive forms originally
existed, as they presently do, for example, in species of N eophyrcaces,
one form being ergatoid and the other unmodified, and that further
evolution resulted in the loss of the latter and so close an approxi-
mation of the worker form, by the former that it is no longer dis-
tinguishable except through its reproductive capacity. If the latter
course has been the actual one, however, we should perhaps suspect
that the fertile “workers” would be comparatively rare in the Rhyti-
doponera colony, at least as rare as are the laying true queens in
colonies of relatively primitive pleometrotic species. It appears, as
will later be indicated, that they are in fact much more abundant.

Whatever the channel of evolution at the level of individual
morphology, its end result poses some intriguing questions about the
direction of evolution in Rhytidoponera at the level of the society.
Some years ago Sturtevant (1938), and later Williams and Williams
( 1 95 7 ) > emphasized the evolutionary significance of the close family
relationships which typically obtain among the members of the
matrifilial colony so characteristic of the higher social Hymenoptera.
Very recently W. D. Hamilton (1964), in two important papers,
beginning with Haldane’s (1955) concept of the evolutionary sig-
nificance of genetically based altruistic behavior, has derived a quantity
in social evolution that he defines as “inclusive fitness.” It may be
regarded in certain respects as an analogue at the social level of the
concept of Darwinian reproductive fitness at the level of the individu-
al. Like the latter it should be found to maximize in evolution. This
maximization, in the social insects, has obviously involved an extra-
ordinary evolution of worker altruism, at both structural and
behavioral levels. Now as Hamilton demonstrates, it can be expected
that such an evolution will be positively correlated with the
maintenance of close genetic relatedness among the members of a
colony. In the absence of parthenogenesis, the closest genetic relation-
ship between queen and worker possible in a colony is that of mother
to daughter, and among workers that of siblings. The pronounced
evolutionary trends toward the inclusion of but two generations in the

1965]

Haskins and Whelden — Rhytidoponera

91

colony structure and toward the restriction of fertilized reproductives
in the colony to a single or a few individuals, so evident in a great
number of ant species, both evidently contribute to maintaining this
maximal degree of relatedness within the community.

Exceptions to both trends, to be sure, are known. On the one hand,
they are well illustrated by the puzzling situations explored in species
of the Formica obscuriventris group (King, 1949, 1955 j King and
Sallee, 1951) and the Formica rufa group (Sturtevant, 1938;
Chauvin, Courtois, and Lecompt, 1961) where it appears that young
reproductives, even of different species, may be adopted by large
colonies, thus prolonging the existence of the colony well beyond the
second generation and introducing both multiple queens and what
must be a remarkable degree of unrelatedness among the worker
personnel of individual colonies. Exceptions to the second trend may
be presented by the numerous pleometrotic species of ants, though it
is still unclear, as in the Polybiine and Polistine wasps, how usually
such multiple reproductives are in fact sisters, and how often or how
elaborately special behavior patterns may be adapted to restricting
reproduction in practice to a single dominant female, or to preventing
non-sisters from coexisting in the colony.

In this context, the course of social evolution in the genus Rhyti-
doponera is of special interest, as is the probable degree of average
relatedness among the workers of a single community of both normal
and “queenless” forms. In the so-called “queenless” species, do
workers in fact regularly give rise to workers? Is the same true of
those species possessing functional typical queens? Is worker pro-
duction accomplished in one or both groups through parthenogenetic
thelytoky, so common among lower nonsocial Hymenoptera, and
frequently reported in the Cape honey bee? Such thelytoky has been
described in ants by several investigators over a long period of years,
including Reichenbach (1902), Crawley (1912), and Comstock
(1903) for Lasius niger, Haskins and Enzmann ( 1945) for Aphaeno-
gaster picea and A. lamellidens, Soulie (i960) for Cremastog aster ,
and Otto (i960) for Formica polyctena, while Ledoux (1949, 1954)
has reported extensively on a specialized social adaptation of thely-
toky in the workers of Oecophylla. If such thelytoky obtains in
Rhytidoponera the pattern of relatedness among colony members
might be quite different than if laying workers possess developed
spermatheca and are fertilized in the manner normal to ordinary
ergatoid queens, by active low-flying males from other colonies. Such
males are indeed characteristic of all species of Rhytidoponera , both
those possessing and those lacking typical queens. Do laying workers

92

Psyche

[March

of the “queenless” species occur only one to a colony, or are they
present in some numbers? If the latter, are they commonly actual
siblings, or how closely are they, on average, related? Is there a
tendency, in the “queenless” species, to confine worker production to
a single individual even if a number of potential worker-producers
are present? If such laying workers are in fact fertilized, are their
mates normally derived from the same or from other colonies? Are
such workers singly or multiply inseminated? Is a single individual
inseminated more than once during its lifetime? What is the average
contribution of male progeny by the non-fertilized members of the
colony, and how is the production of males regulated ? How resistant
is the genus to extreme inbreeding? Is it the rule that a proportion of
each successive brood of workers brought to maturity in a colony is
fertilized and that these individuals remain with the parent colony, so
prolonging the life of the community well beyond the normal two
generations, or do newly fecundated workers typically leave the
parental nest? How indeed are new colonies normally formed?

Such questions as these must be answered before any critical
assessment of the direction of social evolution in Rhytidoponera can
be undertaken. The results reported in the present paper, derived in
the course of some ten years of investigation of the genus both in the
field and in the artificial nest, represent the early stages of an obser-
vational attempt to provide answers to a very few of them.

SOURCE OF WORKER AND MALE BROOD IN A SPECIES OF
RHYTIDOPONERA POSSESSING NORMAL QUEENS
Rhytodoponera purpurea

Rhytidoponera purpurea is a typical member of the R. impressa
group, in which normal queens are characteristic. A single such queen
is typically found in each colony taken in the field. According to
Brown (1954) the species occurs in New Guinea and ranges in
Australia through the rain forests of the Cairns-Atherton Tableland
region of northern Queensland.

On December 27, 1963, a typical, populous colony of R. purpurea ,
comprising the parent female, some 250 workers, and numerous larval
and pupal brood including sexual males and females, was collected
near Kuranda in northern Queensland. The following day a similar
colony was taken at Millaa Millaa on the Atherton Tableland. In
early January these colonies were housed in a type of modified earth-
containing glass Lubbock nest used throughout these investigations.
The colony from Kuranda was divided at the time of nesting into
several isolated groups of workers with broods of cocoons and larvae.
Only one such group had access to the brood female. The colony

1965]

Haskins and Whelden — Rhytidoponera

93

from Millaa Millaa was divided into two portions, one with and one
without the brood queen. An interval was then allowed to permit
the maturation of larvae and pupae in the nests at the time of capture.
In the fragments of both colonies lacking parent queens, as well as ia
those where the normal females were present, oviposition soon oc-
curred, and fairly copious broods were shortly reared. By late March
abundant pupae were present in several groups. In that month, and
over a following period until mid- January, 1965, samples of cocoons
were regularly withdrawn from these colony-fragments and opened,
and their contained pupae scored for sex and caste. The results, in
which the numbers of pupae in all the queenless fragments of Colony
No. 1 are summed for each date, appear below:

No. of Colony

Dates of Assay of Pupal

1 and Young Adult Samples

Fragment

Examination Workers

Males

1 A (Kuranda —

4/29/64

23 pupae ; 3 callows 0

with parent

6/ 4/64

14 pupae ;

0

female)

7/12/64

15 pupae; 1 callow 0

9/20/64

15 pupae

0

12/ 5/64

34 pupae

0

1/19/65

30 pupae

0

Total: 131

worker pupae;

4 callow workers; 0 males.

1 B-E (Kuranda —

4/29/64

0

47 pupae; 26 adults

fragments of

5/10/64

0

3 pupae

colony without

6/ 4/64

0

12 pupae

brood queen)

7/12/64

0

10 pupae

9/20/64

0

27 pupae

12/ 5/64

0

34 pupae

1/19/65

0

47 pupae

Total: 180 male pupae;

26 male adults; 0 workers.

2 A (Millaa-Millaa

— 4/29/64

23 pupae; 2 callows 0

with parent

6/ 4/64

39 pupae

0

female)

6/22/64

1 pupa

0

12/ 5/64

32 pupae

0

1/19/65

29 pupae

0

Total: 124

worker pupae ;

2 callow workers; 0 males.

2 B (Millaa-Millaa -

- 4/29/64

3 callows*

15 pupae

fragment without

6/ 4/64

0

5 pupae

parent female)

6/22/64

0

4 pupae

9/20/64

0

8 pupae

12/ 5/64

0

14 pupae

1/19/65

0

14 pupae

Total: 60 male

pupae; 3 callow workers.*

*These callow workers, found fresh-hatched on April 29, 1964, almost
certainly represent the final fragment of maturing brood collected with the
original colony, and seem with little doubt to have been progeny of the
fertilized brood female.

94

Psyche

[March

Thus, summing the output of the two colonies together over the
period of approximately a year after observation was begun, those
fragments containing a brood queen produced a total of 255 pupae
which were identified as workers, 6 identifiably callow workers, and
no males. Those fragments of both colonies containing workers only
brought to maturity a total of 240 pupae identified as males and 26
identified young adult males, a total of 266 males. In Colony No. 2 B,
three callow workers were also brought to maturity at the time of an
early count. It seems a safe assumption, however, that these indi-
viduals represented the final, delayed residual of queen-laid brood
“inherited” from the partitioning of the colony some three months
earlier.

Thus it seems very clear that in R. purpurea worker brood is
entirely derived from the fertilized brood female in typical Formicid
fashion. Workers, however, can produce and rear a prolific male
brood, at least in the absence of the parent female. Whether the male
brood which typically appears seasonally in large numbers in normal
wild colonies is queen- or worker-derived, or both, is an interesting
and important question for future investigation. It applies with
equal cogency, of course, to the vast range of “normal” Formicid
species.

SOURCE OF WORKER AND MALE BROOD IN SPECIES WHERE
NORMAL QUEENS ARE RARE OR ABSENT

Rhytidoponera metallica

Between December 23 and 25, 1963, a number of vigorous colonies
of Rhytidoponera metallica were collected at various points in the
Blackall Range in Queensland, Australia, some sixty miles north and
thirty miles east of Brisbane. No perfect females were found. These
colonies were housed and maintained in modified glass Lubbock nests
of the same design as those used for R. purpurea. After a preliminary
incubation period of approximately six months, to allow brood resident
in the colonies at the time of capture to mature, samples of cocoons
were withdrawn at intervals, opened, and the contained pupae scored
for sex and caste. Callow workers that were obviously fresh-hatched
were scored at the same time. The result are given in the table below
(p. 95)-

Thus a total of 644 worker pupae or young adults were produced
in the five “queenless” colonies of R. metallica over a period of little
more than six months, and only 1 1 males. It seems clear that worker
production by morphological workers is a normal feature of this
species.

1965]

Haskins and Whelden — Rhytidoponera

95

Dates of

Assay of Pupal and Young Adult S;

arnples

Colony No.

Examination

Workers

Males

!

7/12/64

20

pupae ; 1 fresh-hatched

callow

0

12/15/64

24

pupae; 13 fresh-hatched

callows

0

1/23/65

33

pupae ; 1 fresh-hatched

callow

2 pupae

2.

7/12/64

43

pupae

2 pupae

12/15/64

44

pupae; 36 fresh-hatched

callows

0

1/23/65

46

pupae

4 pupae

3.

7/12/64

20

pupae

0

12/15/64

38

pupae; 6 fresh-hatched

callows

0

1/23/65

69

pupae ; 4 fresh-hatched

callows

0

4.

7/12/64

27

pupae

0

12/15/64

67

pupae ; 28 fresh-hatched

callows

0

1/23/65

66

pupae ; 4 fresh-hatched

callows

2

5.

12/15/64

31

pupae ; 23 fresh-hatched

callows

1

Queensland appears

to lie near the northern limit of the

natural

range of R. me tallica. Ample confirmation that the same situation
obtains elsewhere in its range, however, was provided by counts made
from a single colony of the species taken at Sutherland, N. S. W., on
June 2, 1952, and observed continuously in the artificial nest over a
ten-year period. This colony contained no typical females when
collected, though much later in its history some were produced, as
will be described later. It was kept as a single unit in the standard
glass modified Lubbock type of nest until July 8, 1956, when it was
split into three portions, one of which perished rather shortly. The
second and third were maintained in Lubbock nests until January 1,
1962, when the second also died out. The third portion survived
somewhat longer, but eventually perished on July 30, 1962.

Throughout the ten years of observation, these two colony frag-
ments were kept in closed foraging arenas, to which no males could
enter from outside and from which no individuals matured within
could escape. The ants established and maintained regular kitchen-
middens within these arenas, outside the nests proper. The ambient
humidity of the arenas was maintained low, and the contents of the
middens therefore remained well preserved and readily recognizable
for considerable periods. Thus periodic removal of the middens and
examination of their contents could provide a rather accurate picture
of the quality and type of brood produced.

Until June 13, 1954 (two years after observation was begun),
only workers were brought to maturity. A count of cocoon fragments
accumulated in the middens at intervals during this period thus

96

Psyche

[March

offered a measure of the number of workers produced from worker
parentage. Only cocoon fragments were tallied which either were
nearly intact or which included the larval meconium, thus ensuring
that each fragment represented no more than one individual.

Between January 30, 1954, and March 13, 1954, 1472 such
cocoon-fragments were counted, representing the workers arising
from worker parentage in a typical one and one-half months’ pro-
duction for that period.

Rhytidoponera tasmaniemis

R. tasmaniensis is a species closely allied to R. metallica and
resembling it closely in both habitus and ethology. It is characterized
by a relatively southern distribution in Australia, including south
Victoria and Tasmania.

A colony of R. tasmaniensis was collected at Lower Fern Tree
Gully Station, Victoria, on January 2, 1962, and a number of further
colonies were taken in the Domain at Hobart, Tasmania, on January
10 and 11.* None contained perfect females. All these colonies were
housed in the usual glass modified Lubbock nests. On May 19 and
on subsequent dates samples of cocoons were removed, and opened
and assayed for sex and c^ste as usual. The results follow:

Colony

Dates of

Assay of Pupal and Young Adult S

amples

No.

Examination

Workers

Males

1.

5/19/62

17 pupae

0

9/17/62

37 pupae; 4 fresh-hatched callows

0

1/ 5/63

24 pupae

0

2/ 9/63

27 pupae

0

3/10/63

18 pupae

0

3/ 1/64

24 pupae; 7 fresh-hatched callows

0

9/20/64

13 pupae

0

Other

9/ 9/62

2 pupae ; 1 fresh-hatched callow

0

colonies

10/22/62

8 pupae

0

grouped

1/ 5/63

0 (2 male-producing nests)

10 pupae

Thus over a period of two years and four months, beginning four
and one-half months after capture, a single colony of R. tasmaniensis
produced 1 7 1 worker pupae or fresh-hatched adults, another 8, and a
third contributed two worker pupae and a single adult. Two further
(and small) groups contributed 10 male pupae. Once again the
evidence for worker production is very clear.

*1 am especially appreciative of the help given in obtaining the Hobart
material by Mr. John Hickman of the University of Tasmania.

1965]

Haskins and kf7 helden — Rhytidoponera

97

Fig. 1. Alate Female, Male, and Worker of Rhytidoponera metallica.

Rhytidoponera inornata

This species, also allied to R. metallica but normally, according
to Brown (1958) having a range allopatric to it, is a typical resident
of extreme southwestern Australia. Two colonies without typical
queens were collected at Manjimup, West Australia, between Decem-
ber 25 and 27, 1959, and housed in the usual glass modified Lubbock
nests. As in the preceding instances, cocoons were removed at inter-
vals after a suitable beginning period, opened and scored for sex and
caste. The totals from the two colonies on each examination have
been pooled. The results follow:

98

Psyche

[March

Assay of Pupal and Young Adult Samples
Dates of Examination Workers Males

5/ 5/60
5/14/60

1 pupa

1 fresh-hatched callow

5/18/60

1 pupa

6/15/60

1 pupa

6/20/60

1 pupa

7/23/60

51 pupae

3/ 1/64

33 pupae;

13 fresh-hatched callows

5 pupae

6/22/64

7 pupae

4 pupae

Thus over a period of approximately four and one-half years after
being taken from the field, and beginning five and one-half months
after capture, these colonies were observed to produce a total of 109
worker pupae or fresh-hatched callows, and 9 male pupae. The
evidence of consistent production of workers by workers in this
species seems clear.

Rhytidoponera victoriae

This small form is a common eastern Australian species, especially
abundant in Victoria. On January 2, 1962, a colony was collected at
Lower Fern Tree Gully Station, near Victoria National Park, and
on January 3 a number of colonies was collected within the Park
itself. All were without typical females. They were again housed
in glass modified Lubbock nests, and maintained until May, 1962, in
open arenas. Samples of cocoons were then removed and opened and
the pupae scored for sex and caste in the usual manner, together with
freshly hatched callow workers. The results, summed over all the
colonies examined, are shown below:

Assay of Pupal and Young Adult Samples

Dates of Examination

Workers

Males

5/19/62

42

pupae

11 pupae

8/ 3/62

1 pupa

8/24/62

1

pupa

9/16/62

1

pupa

10/14/62

2

pupae

10/22/62

55

pupae

13 pupae

7/ 2/64

31

pupae ;

2 fresh-hatched callows

46 pupae

7/15/63

22

pupae ;

3 fresh-hatched callows

1 pupa

9/21/64

17

pupae

7 pupae

Thus over a period of twenty-eight months a total of 176 workers is
known to have been produced and a total of 79 males — good evidence
once again that in this species worker production by workers is the
rule.

1965]

99

Haskins and H/helden — Rhytidoponera

Rhytidoponera violacea

This is an example of one of the larger species of the genus, in
which alate females have never been observed. It is a typical inhabitant
of rather dry country in southwest Australia, and closely resembles in
appearance and habit the better known R. convexa of the eastern half
of the continent. A single colony of R. violacea taken in Kings Park,
Perth, W. A., on December 30, 1959, was maintained in an earth-
filled modified glass Lubbock nest through January, 1965. Through-
out this five-year period abundant broods of young workers were
produced, together with a few males. On October 26, 1964, 3
cocoons were extracted from this colony and opened and their con-
tained pupae scored for caste and sex, and a second sample of 8 was
similarly examined on October 27. All eleven pupae were of workers.
Thus worker production from workers was continuing four years and
ten months after the colony was taken.

In three of the five species of Rhytidoponera investigated (R.
metallica , R. inornata, and R. victoriae) perfect queens are repre-
sented in collections, though with great rarity in all except R. metal-
lica. A fourth, R. tasmaniensis, is so closely similar to R. metallica
that it is hard to imagine that similar alate females do not occur with
it too, and that their absence from collections does not simply reflect
the fact that this species has not been particularly extensively taken.
Yet in all these species (together with the fifth, R. violacea, where
alate females have never been found and, from all the circumstantial
evidence, probably do not occur) worker production by individuals
themselves morphologically indistinguishable from normal workers is
clearly the rule. Males of the species, of course, must also be derived
from workers. The question thus is posed whether worker production
by workers results from thelytokous parthenogeny, or whether
fertilization of workers, in the pattern of Diacamma, is the rule.

THE MODE OF PRODUCTION OF WORKERS BY WORKERS IN THE
NORMALLY “QUEENLESS” SPECIES OF RHYTIDOPONERA

In a series of careful and extensive histological studies of workers
of Rhytidoponera metallica , R. inornata , and R. violacea collected in
the field, R. M. Whelden ( 1 95 7> i960, and unpublished data) has
conclusively demonstrated that a normal fertilization mechanism is
involved in the production of workers by “workers.” Examination of
the spermathecae of a total of 836 workers of these three species
yielded the results shown below:

100

Psyche

[March

Number

Per Cent

Number

Individuals

Individuals

in

with

with

Species

Sample

Sperm

Sperm

R. metallica

386

21

54

R. inornata

176

4

2.3

R. violacea

274

22

8.0

It thus seems clear that thelytokous parthenogenesis at least is not
usual in these species of Rhytidoponera , and that a normal mechanism
of fertilization is involved in the production of worker progeny. Other
questions concerned with an estimate of the average relatedness among
workers in a single Rhytidoponera colony, earlier posed, are thus
raised. It becomes important, for example, to ascertain whether
single or multiple insemination of workers is the rule, whether
individual workers may be inseminated more than once during their
lifetimes, and to what degree outcrossing among different colonies
obtains. These issues, of course, can only be determined by careful
and complete observations of the mating flight. Our observations are
still very incomplete. Since, however, they have provided answers
to one or two such questions, even at this preliminary stage, they are
presented here.

CHARACTER OF THE MATING FLIGHT IN
RHYTIDOPONERA METALLICA

Some field evidence of the fertilization of workers by males in
Rhytidoponera metallica has been obtained by W. L. Brown, Jr.
(unpublished observations). He noticed males in low flight which
alighted and entered the main gallery of an established nest of the
species. There seems to be no observational record, however, of actual
fertilization of workers. We have been so fortunate as to witness this.

A colony of R. metallica cited earlier in another connection, was
collected at Sutherland, N. S. W., Australia, on June i, 1952. As
already described, it was housed in a modified earth-containing glass
Lubbock nest, and was then maintained for three months at I5-I7°C,
within the normal winter temperature range of its environment.
During this period workers were allowed to forage from the nest in
a closed arena, and were supplied dilute honey continuously and
provided frequently with a considerable variety of living insect prey.
On September 3, 1952, the nocturnal temperature was raised to 2Q°C,
and diurnal temperatures were allowed to reach a maximum of 25 °C.
Two days after this temperature rise, a number of males which had
matured in the nest during the cool period left it and wandered over

1965]

Haskins and Whelden — Rhytidoponera

IOI

the crater. Simultaneously, several workers were observed resting
quietly outside the nest and in its near vicinity, with head and thorax
rather closely appressed to the substrate on which they rested, but
with the gaster held high and arched. The impression was strong that
an attractant was being disseminated. Males which had emerged
from the nest but had not yet flown paid no special attention to these
individuals. Freshly emerged males, indeed, neither flew nor ran to
any great extent, but went through extensive cleaning motions near the
nest, and were frequently deported back into it by foraging workers
which met them by chance. Nothing further occurred at this time.

On the morning of September 6, with an ambient temperature of
22 °C, ten workers were observed resting quietly near the nest crater
with head and thorax appressed and gaster raised. They remained
immobile in this position for a measured minimum of i minute 5
seconds to a maximum of 12 minutes io seconds. Six were finally
deported back to the nest by foraging workers encountering them
apparently by chance. At the same time a few males emerged, and,
after preliminary cleaning and running about, flew briefly. Nothing
further occurred.

On September 7, with ambient temperature of 24°C, males emerged
in some numbers and flew actively. Foraging workers encountering
such males became greatly excited and commonly attacked them at
once. After approximately one half hour of more or less continuous
flight, a few males alighted near the nest crater and, running rapidly,
sought out nearby workers. No obvious selection was made of workers
resting immobile with raised gasters, although a few of these were
still present. As soon as a male encountered a worker, it grasped the
worker in the cervical region with the mandibles and, thus firmly
attached, at once inserted the stipes. The mandibular grip was then
relaxed at once, and the male, with wings folded, was maintained in
a nearly vertical position by the stipes alone. The first and third pairs
of legs of the male were characteristically folded, while the second
pair projected upward so that they were pressed against the costal
margins of the wings. Both members of the pair usually rested quietly
in this position for a few seconds. Thereafter the worker character-
istically executed grooming movements and began to move about, and
almost at once seized the male by the thorax with the mandibles and
forcibly disengaged it. On a few occasions the worker began to move
almost as soon as copulation began, dragging the male and dislodging
it after progressing for an inch or two.

The first copulation was observed at 8 150 A.M. and lasted for 40
seconds. Between that time and 10:50 A.M., ten matings were

Psyche, 1965

Vol. 72, Plate 6

Haskins — Rhytidoponera

1965]

Haskins and Whelden — Rhytidoponera

103

closely watched and timed. Pairs remained in copula from a minimum
of 30 seconds to a maximum of 52 seconds. It was noted with special
interest that in one case the same pair copulated twice; in another an
individual male paired successively with two workers. In two instances
workers reentered the parent nest while still in copula.

All males observed sustained active flight for periods up to one
half hour before seeking to copulate. The final stages of search,
however, were accomplished in every case on the ground. The whole
process was suggestive of the mating pattern observed in certain
Ponerine species (such as Amblyopone australis and A. ( Stigmatom -
ma) pallipes — Haskins and Haskins, 1951; Haskins, 1928) where
the alate female flies only occasionally and with reluctance but
emerges from the parent colony to mate on the ground or on low
vegetation. The vigorous pattern of dispersive flight and the general
hyperactivity of the males during the flights seemed conspicuously
adapted to securing outcrossing between colonies of the species. Such
outcrossing, indeed, may be biologically essential, as will later be
indicated.

No detectable morphological features distinguished workers which
mated from those which did not. The group which was fertilized
included both among the smallest and among the largest individuals
in the population, as well as many in an intermediate range. This
observation confirms the conclusion of Whelden (1957; i960) derived
from his histological investigations, that workers showing sperm
content in the spermatheca included individuals of a wide range of size.

A second nuptial flight of identical pattern was observed in this
nest on January 27, 1957. Shortly after 9:00 A.M. of that day male
exodus and flight began in typical fashion, reaching a peak by 10:30
A.M. On October 17, 1952, July 17, 1953, and November 22, 1953
workers of this colony were again observed resting outside the nest
with gasters elevated, but on these occasions there was no correspond-
ing flight of males. Precisely this same behavior of workers has been
observed with colonies of Rhytidoponera tasmaniensis , R. inornata,
and R. violacea , all housed in the usual type of modified, earth-
containing Lubbock nests. The intervals for which the position was
maintained were similar to those in R. metallica. Males have been
seen to fly actively from a nest of R. violacea , closely simulating the
pattern of flight in metallica. Actual mating has not been observed
in these species, but it seems a probable inference that the pattern
is similar.

Explanation of Plate 6

Late male Pupae of Rhytidoponera purpurea, from Brood Matured in
Artificial Nest in Fragment Without Brood Queen.

104

Psyche

[March

Thus it is clear that both female and male behavior patterns in R.
jnetallica are well adapted to secure outcrossing among colonies as a
usual condition of the mating flight. Furthermore, multiple insemi-
nation of one worker, and insemination of more than one worker by a
single male, can occur. The related question of whether a single
worker can be inseminated several separate times during its life span
has not yet been answered, nor has the question of whether new-
hatched workers, after insemination, may return permanently to the
parent colony to add to its reproductive potential and prolong its life
beyond the limit of two generations so usual in Formicid communities.
It may be suggestive in this connection, however, that not all of the
worker spermathecae that contained sperm were found by Whelden to
be filled. Indeed, of 22 individuals of R. violacea found to contain
sperm, among a sample of 274 examined, in only two were the sperma-
thecae completely occupied by a dense sperm ball of cells. In three
individuals the sperm mass was very small and loose, comprising only
an estimated 50-100 cells. In the remaining cases, the spermathecae
were partially filled, though fully inflated.

All the evidence seems very suggestive that a rather high degree of
heterozygosity must indeed be regularly maintained in colonies of R .
met allica, involving the germplasm of populations greatly exceeding
the single colony in numbers. It became of considerable interest,
therefore, to enquire what might be the limits of viability in a single
colony where strict inbreeding was enforced over several worker
generations.

TOLERANCE OF RHYTIDPONERA METALLICA TO
EXTENSIVE INBREEDING

In this context, certain further observations made on the colony of
R. metallica already cited may be of interest. As already described,
this colony was maintained throughout the ten years of its existence
as a set of “closed” populations, from which no individuals could
escape and which none could enter from the outside. By the time that
its two longer-lived sections died out, at ages respectively of 9 years
6 months and 10 years 2 months, many new generations of workers
had been matured, to be fertilized in turn by successive generations of
males of the same colony. It is not possible to assess quantitatively the
degree of homozygosity finally attained in these two populations. It
must have increased very considerably, however, over the years. It is
therefore of special interest to note that in the last months of the lives
of both sections ova, although continuing to be produced in numbers,
characteristically failed to hatch larvae. A cytological examination of
these eggs by Whelden (unpublished) showed a high degree of

1965] Haskins and Whelden — Rhytidoponera 105

abnormality in embryonic development. Examination of 119 eggs
from the two colony fragments showed the greater number to be
uninucleate. A few contained 2-4 small nuclei, tending to be clumped
abnormally at one pole of the egg. In a very few cases embryonic
development was sufficiently advanced to show some segmentation and
to indicate the polarity of the embryo, but this was all. By contrast,
a control sample of eggs taken at the same time from a colony housed
and treated in the same manner but much more recently collected
showed clear evidence of development in over 80 per cent of ova
examined, ranging from early stages showing masses of well-defined
cell nuclei to late and well-formed embryos.

Such evidence suggests that the maintenance of a rather high degree
of female heterozygosity is no less a biological imperative for Rhyti-
doponera than for Formicid species possessing alate queens and ex-
hibiting highly organized mating and dispersion flights involving both
sexes. Indeed, the evidence for multiple insemination of workers
suggests that it may be considerably more dependent than some species.
It seems probable that a system of sex-locus lethal or semilethal alleles
exists similar to that described by Mackensen (1951, 1955) and
Rothenbuhler (1957) for Apis , and originally in Habrobracon by
Whiting ( 1943). It would seem that such a requirement of heterozy-
gosity, in the context of the kind of colony organization displayed by
the “queenless” species of Rhytidoponera , must complicate the main-
tenance of close genetic relationships among workers within a colony.

THE FORMATION OF NEW COLONIES IN R. METALLICA

Central to the problem of whether successive generations of fertile
workers remain in older colonies is the question of how new colonies
are formed. At present it is uncertain what may be the normal life
course of young workers subsequent to fertilization in a parent colony.
Some observations have accumulated, however, which may bear on
the mode of formation of new colonies in R. metallica.

Workers taken at random from established colonies in the artificial
nest and isolated in new nests regularly attempted to initiate fresh
communities. They secreted themselves in partly closed cells, from
which they emerged occasionally to forage in the fashion typical of
normal colony-founding Ponerine queens, laid eggs and tended them
carefully, reared larvae, and in a number of cases matured adult
progeny. Unfortunately, it has not so far proved possible to distinguish
and select fertilized workers for these experiments, and all progeny
reared by such isolated workers have been males. Had worker progeny
been reared, however, a new colony might easily have arisen.

Thus it seems at least plausible that isolated workers of R. metal-

io6

Psyche

[March

lica, if fertilized, may be capable of founding new colonies. It seems
somewhat more likely, however, that under natural conditions new
colonies are normally formed by the detachment from the parent nest
of worker groups which include one or more fertilized individuals.
Even though initially very small, such parties may expand rapidly in
numbers and reach the status of full-fledged colonies in a remarkably
short time. Thus on December 21, 1963, two isolated groups of
workers were taken at Sutherland, N. S. W. One consisted of 7
workers, about 6 cocoons, and a few young larvae; the second, also of
7 workers, with 2 cocoons and about 6 young larvae. These two
groups were colonized in the usual fashion in modified earth-contain-
ing nests, and housed throughout their history in adjacent arenas on
the same laboratory table. On January 17, 1965, the first group
included the following:

74 living adult workers.

127 dead workers found on the kitchen middens.

* 4 dead males found on the kitchen middens.

55 cocoons, which were opened and examined, and found to contain:

26 worker pupae.

9 semipupae, believed to be of workers from the size and shape
of the cocoon.

* 11 male pupae.

7 semipupae, believed to be of males from the size and shape of
the cocoon.

2 fresh-spun cocoons with larva present.

54 larvae of various sizes.

Several groups of eggs, totaling about 100.

Thus at least 227 workers and 15 males had matured to the pupal
stage and beyond. In sharp contrast, the second group at the same
date was found to contain 5 adult workers (almost certainly of the
original 7 ) , 2 males, and 3 cocoons, 2 of which contained male pupae
and the third a semipupa. Thus this group as collected apparently
had not included a fertilized worker.

It is particularly interesting, in view of the normally nonseasonal
production of males in R. metallica , that at least 15 males were ma-
tured within the first year of life of this colony. This situation may
be contrasted with the case of a colony of Ectatomma ruidum* ** kept
under observation, from its establishment by the original female, over
a ten-year period in the artificial nest. Here no males whatever
appeared until near the close of the fourth year of community life.

*Note presence of males at this early stage in colony development.

**An abundant New World Ectatomiine species with a well-defined alate
female caste, forming communities which are commonly if not always
monogynic.

1965]

Haskins and Whelden — Rhytidoponera

107

THE PRODUCTION AND FUNCTION OF ALATE QUEENS IN
R. METALLICA

As earlier mentioned, fully developed alate queens of at least four
normally “queenless” species of Rhytidoponera, with bulky thoraces
and typically large compound eyes have been collected — though
rarely — in the field. Under what conditions are they produced? And
what is their function?

On September 25, 1955, and again on October 23, 1955, a single
typical female was found in a colony of R. metallica described earlier,
which had been collected near Sutherland, N. S. W., in June, 1952.
When discovered, each individual was already dealate, was actively
foraging and feeding outside the nest with workers, and generally was
behaving precisely like a worker. When isolated in an earth-contain-
ing modified Lubbock nest each female at first made an imperfect
beginning of an isolated cell, behaving in the general fashion of an
isolated worker. Neither female formed a true cell, however, and
neither showed any sign of oviposition, so typical of isolated metallica
workers. One female remained passive for a full month, and when
at the end of that period it was lost, no trace of an established nest
could be detected. The second female behaved in precisely the same
manner and was ultimately killed and fixed for histological and
cytological examination.

On October 7, 1954, during one of the flights of males from the
same colony, two alate females, matured within the nest shortly before,
emerged with the males and workers and ran actively about outside
the nest. They showed no disposition to mate, however, and were
ultimately captured and preserved.

In January, 1957, the same colony produced about twenty perfect
females. On January 19 one of these emerged from the nest and
wandered about for a short time. It shortly reentered without having
flown. Eight days later, between 9:00 A.M. and 10:30 A.M., while
a flight of males was in progress and several workers were resting
motionless with elevated gasters, a dealate female made a number of
trips together with workers to and from the nest carrying excavated
soil and quite unaffected by the flight in progress. At 1 1 :oo A.M. an
alate female emerged from the nest and wandered briefly before
returning, but gave no evidence either of dispersal flight or of any
attempt to mate.

On February 2 a further dealate queen emerged and wandered.
On March 15 yet another young alate female emerged simultaneously
with three males, wandered, vibrated the wings briefly, then, without
mating or making further attempts at flight, dealated in the open and

ic8

Psyche

[March

returned to the parent nest. During the same day four additional
females behaved in a similar pattern.

Another colony of R. metallica , taken at Ashton Park, Sydney,
N. S. W., on January 8, 1964, had by May 20 produced two perfect
females. Both emerged from the parent nest while still alate and
perished shortly thereafter, having shown no disposition to found new
colonies.

This sketchy evidence of the production and behavior of perfect
females in R. metallica surely needs to be augmented. It does suggest,
however, that in this species the true female is indeed an evolutionary
remnant, in which innate behavior patterns governing emergence from
the nest, the undertaking of at least rudimentary flight, and dealation
are retained. Behavior leading to colony foundation, however, seems
feeble, or actually absent, and replaced by a marked tendency to rejoin
the parent colony and assume a worker-like function within it. It is
of course impossible yet to be certain that abnormal environmental
conditions may not have been responsible for this aberrant behavior,
and the isolated dealate female of R. inornata found by Wheeler may
be suggestive. But the virtual absence of true brood queens in normal
wild colonies surely suggests both that colony foundation by isolated
fertile females, if it occurs at all, must be seldom successful in com-
petition with the mode observed, and that the longevity of such
queens, even in the parent nest, can hardly be considerable. They
indeed appear to be of relict-like character.

What brings about the occasional production of perfect females?
In the two colonies where such production has been observed, it
occurred at the height of the brood-rearing season, and at a period
when the colony was most active in foraging and was most abundantly
supplied with food. In each colony, the total brood of females was
produced over a rather short interval, suggesting that trophic in-
fluences may have been important in effecting the crossing of a
“difficult” developmental barrier. Quantitative studies of the phe-
nomenon, analogous to those of Brian in Myrmica, are needed.

DISCUSSION AND SUMMARY

The work reported has shown that workers are regularly produced
from worker eggs in five species of Rhytidoponera in which alate
females are rare or unknown. On the other hand, worker-laid ova
give rise only to males (though prolifically) in R. purpurea, a species
in which normal queens are the rule. Males are also regularly and
copiously produced from worker eggs in artificially maintained
colonies of R. metallica which appear to lack worker-producing
individuals. These observations suggest that sex determination in

1965] Haskins and Whelden — Rhytidoponera 109

Rhytidoponera is of the normal Formicid type, and thelytokous
parthenogenesis, if it occurs at all, does not make a significant contri-
bution to the production of females. Such a conclusion has been
strengthened by the observation of nuptial flights and actual mating
of males with workers in R. metallica. It has been physically proven
in the finding by R. M. Whelden of sperm in the spermathecas of
workers in R. metallica , R. inornata, and R. violacea.

It is clear from the evidence of collections that, at least in R.
metallica , R. inornata, R. victoriae, R. aspera, R. croesus , and R.
strigosa, in addition to the members of the R. impressa group, normal
alate queens are produced under natural conditions. Numerous such
queens of R. metallica have been matured in the artificial nest. These
undertook some flight and dealated but neither mated nor made any
attempt to found new colonies, although conditions were made as
nearly optimal as possible. They proved uniformly very short-lived.
This evidence, taken with the fact that perfect females have never
been reported in a considerable number of species of Rhytidoponera
including some of the largest and most conspicuous, and the fact that
no queenworker intermediate has been reported for any species,
suggest that the queen-form has been entirely dropped out in the
“queenless” species, and that members of the R. metallica group (and
possibly some other forms as well) may be in an intermediate stage
of evolution, the queen persisting as a rare morphological form but
being virtually without biological significance. This impression is
strengthened by evidence both that individual workers, in contrast to
queens, do show strong instincts of colony formation, and evidence
that groups of as few as seven workers collected in the field can
quickly expand to full colony size.

While such observations answer some questions about the general
biology and social structure of the “queenless” species of Rhytidopo-
nera, they raise a great many more. Whelden’s finding of sperm in
worker spermathecae in R. metallica, R. inornata, and R. violacea
brought out two additional points of interest. First, sperm-containing
individuals were relatively abundant in typical colonies of all three
species collected in the field, ranging from 2.3 per cent of total
workers in R. inornata to 8.0 per cent in R. violacea. Second, in only
a few cases were worker spermathecae replete with sperm. Usually
they were but partly filled, and in several instances only a very few
sperms were present. This raises the question of whether an individual
worker is fertilized more than once in its lifetime, or whether it is
normally exclusively male-producing in the latter part of life.

It is clear, at any rate, that numerous worker-producing workers

I IO

Psyche

[March

coexist in normal colonies of at least three species of Rhytidoponera.
Further, in R. metcillica at least, multiple insemination of workers,
and insemination by individual males of more than one worker are
not uncommon. It is also evident that workers can and do make rich
contributions of male progeny, which may begin to appear (in small
numbers) very early in the life of an isolated, expanding, worker
group. All these circumstances make it seem improbable, at first
glance, that an average degree of relatedness among members ap-
proaching sibship can be maintained in a typical Rhytidoponera
colony. The situation might be quite different, however, if a single
fertile individual normally dominated oviposition in a colony, rival
fertile individuals being suppressed by hormonal or behavioral means.
Whether any such situation obtains remains to be determined.

If the relatedness of workers within a single colony is indeed
relatively low (at least not consistently maintained at the sibship
level), it may well be that among the members of a continuous
population of the species it is more than usually close. For if new
communities normally arise from small parties containing at least one
fertilized worker which leave parent colonies, it seems probable that
the average distance traveled before settling down would be consider-
ably less than that covered by actively flying queens before founding
new communities. One might indeed expect the Rhytidoponera
population to be considerably more ‘Viscous,” in Hamilton’s term,
than those of such an ant genus as Lasius, and might perhaps reason-
ably expect a lesser degree of “colony integrity” than in a monogynous
species with actively flying alates of both sexes. Some measure of the
degree of colonial integrity developed among those Rhytidoponera
species normally without alate females might be gained by a careful
study of “colony recognition” between communities — of the degree
of worker-to-worker hostility among individual colonies of the same
population, as compared to that between colonies from populations
considerably separated geographically. Work of this character is
planned.

References

Brian, M. V.

1959. The ontogeny of insects. Acta symposii de evolutione insectorum,
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1961. Organ transplants between ant castes. Symposia Genetica et.
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Brown,,, W. L., Jr.

1954.- Systematic and other notes on some of the smaller species of the
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1965]

Haskins and Whelden — Rhytidoponera

1 1 1

1958. Contributions toward a reclassification of the Formicidae. II.
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Chauvin, R., G. Courtois, and J. Lecompt

1961. Sur la transmission d’isotopes radio-actifs entre deux foumi-
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Comstock, A. B.

1903. Quoted in Wheeler, W. M. (1903).

Crawley, W. C.

1912. Parthenogenesis in worker ants, with special reference to two
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Haldane, J. B. S.

1955. Population genetics. New Biology, 18: 34-51 (edited by M. L.
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Hamilton, W. D.

1964. The genetical evolution of social behavior. I. Jour. Theoretical
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Haskins, C. P.

1928. Notes on the behavior and habits of Stigmatomma pallipes
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1945. On the occurrence of impaternate females in the Formicidae.
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Haskins, C. P., and E. F. Haskins,

1951. Note on the method of colony foundation of the Ponerine ant
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1961. Continuous polymorphism in locusts — Insect Polymorphism —
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1951. More mixed colonies in ants. Proc. Iowa Acad. Sci. 58: 487-489.
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Mackensen, O.

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1960. Zur Erscheinung der Arbeiterinnenfertilitat und Parthenogenese
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1957. Diploid male tissue as new evidence on sex determination in
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1960. Des considerations ecologiques peuvent-elles apporter une contri-
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1938. Essays on evolution. II. On the effects of selection on social
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1933. Colony Founding Among Ants. Harvard University Press,
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1957. Anatomy of Rhytidoponera convexa. Ann. Ent. Soc. Am. 50:
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1943. Multiple alleles in complementary sex determination of Habro-
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1958. Studies on the ant fauna of Melanesia. III. Rhytidoponera in
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OBSERVATIONS ON THE NESTING SITE AND
BIOLOGY OF THE ARIZONA DAMP-WOOD TERMITE
ZOOTERMOPSIS LATICEPS (BANKS)

( HODOTERMITIDAE ) 1

By W. L. Nutting

Department of Entomology, University of Arizona

The genus Zootermopsis includes the largest termites to be found
in the United States. It contains but three species: angusticollis, the
common damp-wood (or light-colored, rotten-wood) termite, ranging
from southern British Columbia to northern Baja California and
most abundant in the more humid coastal areas; nevadensis , the small
(or dark-colored) damp-wood termite, roughly coextensive with the
former but generally preferring cooler, drier areas and extending
farther inland to western Montana; and laticeps, the Arizona damp-
wood termite, recorded from southeastern Arizona and southwestern
New Mexico (Castle, 1934). Because of its large size and relatively
easy availability in logs and stumps, angusticollis has become moderate-
ly well known and has proved a useful laboratory animal in a wide
variety of studies (Snyder, 1956, 1961). To a lesser extent this is
true for nevadensis. In contrast, no new biological information has
appeared on laticeps since the early contributions by Townsend
(1893), Banks (1906) and Banks and Snyder (1920). Although it
is by far the largest, with soldiers 16 to 22 mm. long and alates 25
to 30 mm. over all, it has remained the least known of the three.

The following account summarizes the data on 27 collections of
the alates of this spectacular termite made by various members,
students and friends of the Department of Entomology at the Uni-
versity of Arizona, mainly during the last ten years. It also includes
information gathered on four colonies which were located by the
author after several years of sporadic but diligent searching. When
it is realized that this termite stages its flights during the night and
that it colonizes living, often very large, trees, its rarity becomes
more understandable. It is thus not likely to be taken by the casual
collector or without considerable effort.

Distribution. The collections of alates, made under lights and
in light traps (both white and ultra violet) have considerably
broadened the range of the species within the state of Arizona (Fig.

Arizona Agricultural Experiment Station Journal, Article No. 967.
Manuscript received by the editor February 19, 1965.

r\

114

Psyche

[March

Figure 1. Map of the natural vegetation of Arizona, showing distribution
of Zootermopsis laticeps: open circles, previous records; solid circles, new
records.

1). Nineteen new localities are represented between Safford and the
Chiricahua Mts. in the east and Sedona and the Baboquivari Mts.
toward the center of the state. Except for Las Cruces, N. Mex., all
previously published records fall within this area (Table 1). Each
collection has been made in the vicinity of a sizeable watercourse —
in canyons such as Ramsey in the Huachuca Mts., along more or less
permanent streams as Sonoita Creek near Patagonia, in river bottoms

1965]

Nutting — Zootermopsis

1 15

as at Hereford or in heavily irrigated areas as in the Gila Valley at
Salford. Although the altitudinal distribution ranges from 1500 feet
in the creosote bush — salt brush desert around Florence to 55°° feet
in oak — pine woodland along Workman Creek, all of these situations
support at least thin stands of one or more large trees such as cotton-
wood, sycamore, ash, walnut and alder. Considering the intensity of
light-trapping which has been done in Arizona, it does not seem
likely that the range of laticeps will be extended much farther into
the higher plateaus and mountains to the north or into the lesser
ranges and desert to the west. However, it is highly probable that it
might be found in the northwestern reaches of the Rio Grande, at
least above and below Las Cruces, and south on both slopes of the
Sierra Madre Occidental well into Sonora and Chihuahua.

Colonizing Flights. Lack of complete seasonal series of collections
from any of the localities in Table 1 prevents a detailed character-
ization of the flight schedule of Z. laticeps. Alates appeared in a
laboratory colony, maintained between 60 and 70°F, during the first
three weeks of June. Obviously it may be expected to fly from late
June into early August over most of its range in Arizona, apparently
with little regard for small differences in altitude. The records from
Salford and Madera Canyon indicate that the seasonal flight pattern
involves small groups of alates taking part in frequent, perhaps almost
daily, flights over a period of six weeks or more. Since the highest
monthly maximum temperature over most of this region is usually
reached in late June or early July, and prior to any significant amount
of rain (Sellers, i960), it is suggested that high temperature may
provide the major stimulus for initiating the flight period rather than
the advent of the summer rains. Both Werner and Burger reported
that no rain of any significance had fallen prior to the collections they
had made in Madera Canyon and at Workman Creek (Table 1).

Light and Weesner (1948, p. 55) took small numbers of alates
between 1 and 4 A.M. and between 11 P.M. and 12:45 A.M. on two
successive evenings in the Huachuca Mts. The complete absence of
observations on any of the 27 flights recorded here, plus the fact that
most collections were taken in light traps, further testifies to the fact
that this termite does indeed fly during the night. The night at Pena
Blanca (Table 1) might be cited as typical of most of the light-
trapping ventures. The trap was run from shortly after sunset until
sunrise and rather closely tended by four persons till midnight. No
alates of any species of termite were seen; however, one alate of
laticeps was taken from the trap the next morning. The only other
data closely associated with any of the flights are the temperature and

Table 1. Distributional and flight data for Zootermopsis laticeps. An * indicates a previously published record

(See Banks and Snyder, 1920).

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♦Sabino Basin, Sta. Catalina Mts. ca. 2700' VII-8-20 Lutz and Rehn

♦Edgar Can., Sta. Catalina Mts. <5000'? VII-3-14 Colony M. Chrisman

1965]

Nutting — Zootermopsis 117

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ELEVATION

DATE

Oak Creek Can., vie.

I

G. Butler, C. O’Brien

C. W. O'Brien

C. H. T. Tn

1 1 8

Psyche

Figure 2

1965]

Nutting — Zootermopsis

119

relative humidity taken that evening at Pena Blanca — 920 and 33%
at 6:30 P.M., and 78°F and 44% at midnight, following a typical
warm day with showers in the vicinity during late afternoon. This
pattern of flight behavior is considerably more restricted than the
evening flights of Z. angusticollis which have been reported during
every month of the year (Castle, 1934). The more even temperatures
in the coastal regions may be more conducive to a longer flight period
than the seasonal extremes common to the desert Southwest.

Nesting site. Colonies have previously been reported in Fremont
cottonwood (Populus Fremontii Wats.) (Townsend, 1893 ; Emerson,
i960, in litt.) , alder (probably Alnus oblongifolia Torr.) and willow
(Salix sp.) (Banks and Snyder, 1920) . Cazier ( 1962) has mentioned
exposing portions of colonies in burls on living sycamores ( Platanus
Wrightii Wats.) in Cave Creek Canyon between Portal and the
Southwestern Research Station. Three colonies described in this
report were found in willow (probably Salix Gooddingii Ball).
Velvet ash (Fraxinus velutina Torr.), Arizona walnut [. Juglans
major (Torr.) Heller] and perhaps netleaf hackberry (Celtis
reticulata Torr.) would also appear to provide suitable nesting sites
for this termite. In each case colonies have been found in unhealthy
trees or in those which have contained at least some dead portions.

The impregnability of the nesting site is dramatically illustrated
by the first colony which was taken in a large Fremont cottonwood,
on the flood plain 100 yards from Sonoita Creek and about two miles
southwest of Patagonia. Fortunately the tree had been blown down,
probably within six months or so, since it still bore many dried leaves.
It measured at least 90 feet high and 6 feet in diameter at chest
height. A few large nymphs of Z. laticeps were first discovered in a
burl formed around the base of a broken branch on one of the main
limbs. This limb, at the trunk, was 18 inches in diameter 40 feet
above the ground and had been broken off at 23.5 feet, apparently
some years before. The wet, fibrous bark was easily stripped off and
the limb was then sawed into two-foot lengths starting from the
broken tip. It was later sawed into shorter lengths, tracings were
made of the cross-sectional cuts (Fig. 2), and finally each section was
carefully split and the entire colony removed. The colony was thus

Figure 2. Longitudinal section shows a reconstruction of the galleries of
Zootermopsis laticeps in a cottonwood limb from Patagonia, Ariz. The
actual numbers of nymphs (N) and soldiers (S) are indicated in each
segment. Cross sections represent tracings of the galleries in the seven
cuts AA to GG.

120

Psyche

[March

found to occupy the terminal six feet of the limb, ending approximate-
ly 54 feet above the ground.

Figure 2 shows the extent and character of what must have been
several years’ work by this well-established and vigorous colony. The
configuration of the galleries indicates that the colony had originally
extended farther up the limb and probably weakened it sufficiently to
result in the break during a windstorm. The very large and extensive
central galleries in the upper half of the colony were almost completely
filled with hardened masses of fecal pellets, while the same situations
were filled approximately 50 percent to within 18 inches of the lower
end. It is apparent that this species often takes advantage of the
softer spring wood in driving new galleries (note cross sections in
Fig. 2 ) , but through continued working these concentrically arranged
chambers may be developed into cavities three or four inches in
diameter and much more in length. The surface of the galleries has
a coarse, velvety finish, characteristic of the other species of Zooter-
mopsis. Cottonwood is fairly soft and light but very fibrous and
tough. At the time the limb was cut, it was so wet from recent
winter rains that water was squeezed out by the wedges used in
splitting the sections. Although no other insects had penetrated the
wood, the termites had followed shrinkage cracks in several places,
particularly in the burls. The extremely tough and contorted grain
of the burls seemed to be especially attractive. Four burls in the limb
were extensively worked and each contained tunnels to the outside
which were plugged with fecal material. One other tunnel, just
below section C-C (Fig. 2) was driven directly to the outside. These
openings measured about 3 X 5 to 6 mm. and were presumably used
as exits for the alates and perhaps also for dumping fecal pellets. It
is interesting to note that the galleries of Z. angusticollis are pre-
dominantly peripheral in logs and stumps and that long, slit-like
openings are made for the escape of the alates (Castle, 1934).

Three other colonies were found in willows along the bank of the
San Pedro River at Hereford. The second complete colony was
taken from a living tree approximately 30 feet high and 12 inches in
diameter at the base. Individuals of this colony were discovered in
damp, fermenting wood at the base of a broken branch six inches in
diameter and four feet above the ground. The tree was felled and
handled in the same way as the limb from Patagonia. As shown in
figure 3, the galleries extended about two feet below the crotch of the
snag in a pattern similar to that in the cottonwood limb. Some large
cerambycid larvae had attacked this region prior to the termites, for
the latter had driven galleries through the damp and firmly packed

1965] Nutting — Zootermopsis 121

Table 2. Composition of two colonies of Zootermopsis laticeps.

NYMPHS SOLDIERS

Patagonia, Arizona; March, 1963
no wing pads pre-alates small pads pigmented unpigmented

10-13 mm. 10-12 mm. 7-10 mm. 4-7 mm. ca. 3 mm.

552 200 162 103 103 22

Hereford, Arizona ; November, 1963
no wing pads

10-13 mm. 7-10 mm. 5-7 mm. 4-5 mm. ca. 3 mm.

232 200 431 155 40 26

detritus in the larval burrows. The willow had also been invaded,
apparently at the base of the broken branch, by a bracket-fungus
[. Fomes igniarius (L. ex. Fr.) Kickx., kindly identified by Paul D.
Keener of the Dept, of Plant Pathology] . The termites had restricted
their galleries entirely to the column of softer, fungus-infected wood
extending below into the base of the tree. It is probable that the
cottonwood limb from Patagonia had been completely invaded by
such a fungus but that the lack of contrast with any uninfected wood
caused this point to go unnoticed. Unidentified Basidiomycetes and
several other fungi have been isolated from colonies of Z. angusticollis
and later experiments indicated that fungi play an essential role in its
natural diet (Hendee, 1933, 1935). Only one obvious tunnel to the
outside was found one foot below the crotch, its opening (sealed) in
a deep furrow of the bark.

A third colony was also located, but not taken, at Hereford in a
medium-sized, three-trunked willow. Superficial galleries, containing
a few large nymphs, were exposed by chopping into the partly healed
stub of a branch three inches in diameter and three feet above the
ground. The utility of a trail-laying mechanism (reported in Z.
nevadensis: Stuart, 1961) may be envisioned in the existence of a
colony inhabiting galleries as extensive and complex as those described
above.

Founding of the colony. A single incipient colony was found in the
stub of a broken branch three inches in diameter and six feet from
the ground on another small willow at Hereford. The de-alated king
and queen had carved out a cell approximately 5 X 10 X 25 mm. in
the sap-soaked, splintered wood at the base of the branch. The pair
had probably flown between late June and early August, 1963, and
by this date (Oct. 26) had produced a single nymph, three mm. long.
This colony was maintained in a room varying between 550 and 70°F
and six eggs were produced during early March, 1964, when obser-
vations had to be discontinued.

122

Psyche

[March

Figure 3

1965]

Netting — Zootermopsis

123

This example, together with data accompanying the collections of
other colonies, demonstrates what is probably the usual way in which
this termite initiates its attacks on living trees. Several of the larger
trees commonly suffer mutilations by windstorms, particularly cotton-
wood and willow. These open wounds may then provide focal points
for the colonizing alates. This approach is reminiscent of the method
by which N eotermes tectonae Damm. attacks living teak trees via
dead branches, snags and wounds (Kalshoven, 1959). It remains to
be determined, however, whether odors of freshly exposed wood or
fermenting sap are actually attractive, and whether a fungous invasion
is a prerequisite for the establishment of a colony.

Colony size and soldier-nymph ratio. Table 2 summarizes the
composition of the two colonies which were investigated in detail.
Since they were both established in laboratory nests, the individuals
were quickly sorted into rough size-classes and counted ; no attempt
was made to determine the number of instars represented. The king
and queen were not found in the Patagonia colony and no eggs were
found in either colony. Although supplementary reproductives have
been reported to be common in colonies of the other two species
(Castle, 1934), none were found here. In the Patagonia colony, three
months before the flight period, nymphs with wing pads of three
different lengths were conspicuous: 200 pre- alates or nymphs in the
penultimate instar and 162 with wing pads of short and intermediate
lengths. The total number of alates eventually produced by this
colony in the laboratory was not determined. The colony contained
a total of 22 soldiers (2 small) and 1120 nymphs, with a soldier-nymph
ratio of 1 to 51. In the colony from Hereford, three months after
the flight period, none of the nymphs bore conspicuous wing pads.
This colony contained 26 soldiers and 1058 nymphs, making a soldier-
nymph ratio of 1 to 41. These counts are only one third to one-half
the maximum colony size reported for Z. angusticollis , allhough the
soldier-nymph ratio is about the same (Castle, 1934).

Insects associated with the colony. Among many miscellaneous
insects and mites found within the galleries of the two large colonies,
two deserve mention because the association may be more than casual.
At least 19 small (2-5 mm.) cockroaches were found fairly evenly
distributed through the galleries in the limb from Patagonia. Ashley

Figure 3. Longitudinal section shows a reconstruction of the galleries of
Zootermopsis laticeps in the trunk of a living willow, Hereford, Ariz. The
actual numbers of nymphs (N) and soldiers (S) are indicated in each
segment. Cross sections represent tracings of the galleries in the five cuts
AA to EE.

124

Psyche

[March

B. Gurney (Insect Identification and Parasite Introduction Research
Branch, U.S.D.A.) has informed me that the specimens appear to be
nymphs of Compsodes schwarzi (Caudell). Practically nothing is
known of this insect although Ball et al. (1942) reported that a
winged male was taken in an ant nest in the Santa Rita Mountains.

Fifty- two syrphid larvae (4-15 mm. long) were taken near the
center of the colony from Hereford. Of several larger larvae which
were set aside, one eventually pupated and developed into an adult.
Willis W. Wirth (Insect Identification and Parasite Introduction
Research Branch, U.S.D.A.) kindly identified the specimen as
Tenthredomyia tridens (Loew). At least one member of this genus
has been recorded “breeding” in the wounds of large trees (Wirth,
1964, in litt.). The presence of so many larvae, closely applied to the
walls of the galleries containing the highest concentrations of the
termites, is indeed puzzling. A few of the cockroaches and larvae
were put into the large laboratory termite colonies, but it was im-
possible to observe anything beyond the fact that all three insects
appeared to be indifferent to each other.

Additional collections of Z. laticeps are planned so that these
associations can be studied further. In co-operation with Gordon B.
Castle of Arizona State University, biological studies on laboratory
colonies of the termite are also in progress.

References Cited

Ball, E. D., E. R. Tinkham, Robert Flock, and C. T. Vorhies.

1942. The grasshoppers and other Orthoptera of Arizona. Univ. Ariz.
Agric. Expt. Sta. Tech. Bull. No. 93, p. 266.

Banks, N.

1906. Two new termites. Ent. News, 17:336-337.

Banks, N., and T. E. Snyder.

1920. A revision of the nearctic termites (Banks) with notes on biology
and geographic distribution (Snyder). U. S. Nat. Mus. Bull.
No. 108, 228 pp.

Castle, G. B.

1934. The damp-wood termites of western United States, genus
Zootermopsis. I. General biology and ecology, pp. 273-291. In
Kofoid, C. A., et al. [eds.] Termites and Termite Control, 2nd
Ed., Univ. Calif. Press, Berkeley.

Cazier, M. A.

1962. Personal communication.

Emerson, A. E.

1933. A revision of the genera of fossil and recent Termopsinae
(Isoptera), Univ. Calif. Publ. Ent., 6: p. 187.

Emerson, A. E.

1960. Personal communication.

1965]

Nutting — Zootermopsis

25

Granger, B. H.

1960. Will C. Barnes’ Arizona Place Names, rev., Univ. Ariz. Press,
Tucson, p. 38.

Hendee, E. C.

1933. The association of the termites, Kalotermes minor, Reticulitermes
hesperus, and Zootermopsis angusticollis with fungi. Univ. Calif.
Publ. Zool., 39:111-134.

Hendee, E. C.

1935. The role of fungi in the diet of the common dampwood termite,
Zootermopsis angusticollis. Hilgardia, 9:499-525.

Kalshoven, L. G. E.

1959. Observations on the nests of initial colonies of Neotermes
tectonae Damm. in teak trees. Bull. Union Internat. Etude
Insectes Sociaux, 6:231-242.

Light, S. F., and F. M. Weesner.

1948. Biology of Arizona termites with emphasis on swarming. Pan-
Pacific Ent., 24: p. 55.

Sellers, W. D., Ed.

1960. Arizona Climate. Univ. Ariz. Press, Tucson, v + 60 pp., +
climatological summaries.

Snyder, T. E.

1956. Annotated, subject-heading bibliography of termites, 1350 B.C.
to A.D. 1954. Smithsonian Misc. Coll., 130: iii + 305 pp.
Snyder, T. E.

1960. Supplement to the annotated, subject-heading bibliography of
termites, 1955 to 1960. Smithsonian Misc. Coll., 143: iii + 137

pp.

Stuart, A. M.

1961. Mechanism of trail-laying in two species of termites. Nature
(London), 189:419.

Townsend, C. H. T.

1893. Note on Termopsis angusticollis Hagen. Zoe, 4:139-141.
WlRTH, W. W.

1964. Personal communication.

CAMBRIDGE ENTOMOLOGICAL CLUB

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PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol. 72 June, 1965 No. 2

CONTENTS

CONTENTS

Observations on Microtus Nesting in Ant Mounds. Gerald Scherba 127

Observations on Three Species of Phidippus Jumping Spiders (Araneae:
Salticidae). Beatrice T. Gardner 133

New Species of Chilocorini (Coleoptera: Coccinellidae) . E. A. Chapin 148

Techniques for the Study of Spider Genitalia. Herbert IV. Levi 152

Scolytoidea (Coleoptera) 5: Notes on Neotropical Platypodidae, Mainly
from Central America. Hans Reichardt 159

Picrocryptoides : A,New Genus of the Tribe Mesostenini from Southern
South America (Hymenoptera, Ichneumonidae) . Charles C. Porter .... 167

Studies on North American Carboniferous Insects. 4. The Genera
Metropator, Eubleptus, Hapaloptera and Hadentomum.

F. M. Carpenter 175

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1965-66

President H. Reich ardt, Harvard University

Vice-President S. Vogel, Harvard University

Secretary C. C. Porter, Harvard University

Treasurer F. M. Carpenter, Harvard University

Executive Committee W. L. Brown, Jr., Cornell University

J. Reiskind, Harvard University

EDITORIAL BOARD OF PSYCHE

F. M. Carpenter (Editor), Professor of Entomology , and Alexander
Agassiz Professor of Zoology , Harvard University

P. J. Darlington, Jr., Alexander Agassiz Professor of Zoology ,
Harvard University

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University; Associate in Entomology, Museum of Comparative
Zoology

E. 0. Wilson, Professor of Zoology, Harvard University

H. W. Levi, Associate Curator of Arachnology, Museum of Com-
parative Zoology

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Vol. 72

PSYCHE

No. 2

June, 1965

OBSERVATIONS ON MICROTUS NESTING IN
ANT MOUNDS* 1

By Gerald Scherba
California State College at San Bernardino

A large number of different kinds of organisms inhabit social insect
nests and their relationship to their host varies from simple sheltering
during winter, through predation, to complex interactions involving
specialized secretory glands and other structures. There is a rich
literature describing these myrmecophiles and termitophiles and an
elaborate terminolovv defines each of the kinds of relationships be-
tween symbiont and host.

The overwhelming majority of these nest inhabitants are arthro-
pods, especially beetles of the family Staphylinidae and allied families.
However, vertebrates also inhabit social insect nests. Nichols (i94°)
reported the synbranch eel Synbranchus marmoratus Block from
moist cavities within nests of the leaf-cutting ant Atta sexdens in
South America. In Illinois the author has observed wood frogs,
Rana sylvatica overwintering in nests of Formica ulkei. Donisthorpe
(1927) recorded the slow-worm Anguis fragilis L. within Formica
fusca mounds in England ; the blind lizard A mphisbaena within A tta
nests on the Amazon; the legless lizard Caecilia annulosa within
! fungus growing ant nests in Guiana; and several species of snakes
living with ants in Australia. The Nile Monitor Varanus niloticus
j (Linn.) incubates its eggs within the termitaria of N asutitermes in
1 South Africa (Snyder, 1929; Cowles, 1930). Snakes, lizards, and
I geckos use termite nests for egg-laying in Jamaica. These animals
occupy cavities within the termite nest that were originally excavated
by Parakets (Hindwood, 1959).

Among the birds, at least 8 species are known to breed in arboreal
or terrestrial nests of termites, or in some cases, ants. The list in-
j eludes 23 species of kingfishers, 13 species of parrots, 4 species of

1Project 80 of the Jackson Hole Biological Research Station, supported by

I funds from the National Science Foundation, Grant No. G-23423.

Manuscript received by the editor March 14, 1965.

127

SKSTHSORi*.'

mSTITUTIG*

128

Psyche

LJune

woodpeckers, 4 species of trogons, 2 species of puffbirds, 1 species of
jacamar and 1 species of cotinga (Von Hagen, 1938; Hindwood,
I959i Hardy, 1963). Most of these species hollow out a portion of
the nest, and the termites seal the exposed portions so that there is
no contact between bird and termite. However, the Rufous Wood-
pecker, Micropternus brachyurus , which nests in active Crematogaster
carton nests, feeds upon the ants which run all about the bird’s nest
and over the young, without apparent harm.

The Gartered Trogon of central South America nests within wasp
nests, but eats the adult wasps before hollowing a nest cavity (Hind-
wood, 1959).

Among the mammals, the aardvark Orycteropus is predatory upon
termites and lives within the termitaria (Bequaert, 1921, and Alfred
E. Emerson, personal communication). The dormouse Claviglis
has been found nesting with young in a cavity in the side of an
arboreal termite nest in the Belgian Congo (Chapin, 1939). The
following description adds to the scant record of mammals which
inhabit social insect nests.

During observations of a western mound building ant, Form'icci
opaciventris Emery in 1962, the author found the meadow vole,
Microtus montanus nanus (Merriam) living within these mounds
and both adults and young animals entering and leaving their burrow
frequently. This report will describe the burrow system and detail
an attempt to define the relationship between vole and ant.

The study area is located in western Wyoming, one mile east of
Jackson Lake, near the townsite of Moran, and at an altitude of
2065 meters. The ant nests are located on a mesic silver sagebrush,
Artemisia cana, meadow. The soil is a clay loam with an abundant
ground cover of herbaceous vegetation.

The nests of Formica opaciventris are earthen mounds with a light
covering of thatch. The mounds vary in height to 0.7 meters with
basal diameters as large as two meters. The ants are “aggressive,”
and when disturbed bite readily, exuding droplets of formic acid.
There are approximately 400 mounds at the study site and these
nests have been under observation since 1957 (Scherba, 1961, 1963,
1964).

In the Jackson Hole area Microtus montanus has been captured
from these habitats: alpine meadow, subalpine meadow, lush grass-
forb meadow, wet sedge meadow, willow savannah, timothy meadow,
aspen forest, sage-grass community, and willow-alder streamside com-
munity. Findley (1951) and Negus and Findley (1959) state that
“this is one of the most versatile mammals . . . from the standpoint
of habitat selection.”

1965]

Scherba — Microtus

129

While there are no data available on the Microtus population
density changes at the study site, casual observation suggests that
population levels fluctuate greatly from year to year. The voles are
seldom seen during some summers, while they are abundantly evident
moving on the ground during other years. 1962 Was such a year of
apparently high vole density in Jackson Hole.

THE BURROW SYSTEM

Multiple openings of tunnels are apparent at the base of a large
number of the mound nests. Nest 434, a moderately active mound,
had eight such openings and this nest was selected for examination

(Fig. 1).

The tunnel entrances measured 3.0 to 3.8 cm. in diameter. Exca-
vation indicated that two of the tunnels on opposite slopes led down-
ward, then inward, below the soil surface level, then upward, termi-
nating in an oval chamber 12.2 cm. in length and 4.6 cm. high.
Other tunnels ended blindly within the mound, joined this chamber
after a circuitous path or entered a second, larger, chamber described
below. The floor of this first chamber, which was 16.8 cm. below
the mound apex, consisted of fine soil and numerous particles of short
green and dried plant stems, especially the rush J uncus which grows
as a fringe on the mound exterior.

The second chamber, larger and deeper than the one described
above, was abundantly lined with soft dried grasses and was accessible
by three tunnels which entered it internally from the North, South,
and West. These tunnels terminated, eventually, in openings at the
ant nest surface. The chamber measured 13.7 cm. in length, 10.7 cm.
in heighth, and its floor was located 13.7 cm. below the level of the
soil surface and beneath the center of the mound apex. (See Fig. 2).
Presumably this chamber serves as the bedding site for Microtus.

Excavation of a second, more active ant mound, nest 460, revealed
seven tunnel entrances with tunnels extending circuitously into the
mound for distances of several centimeters. All of these tunnels
terminated blindly.

In summary, Microtus activity results in a series of tunnels opening
at or just above the level of the soil surface. These tunnels, turning
frequently, either end blindly or lead into lined chambers or cavities
located within active ant mounds.

POSSIBLE RELATIONSHIPS

The unexpected presence of a rodent living within active ant nests
prompts the inquiry: Are the voles adapted to living within the
mounds, either by virtue of being protected from ant bites, or because
of avoidance of the voles by the ants? Further, are the ants serving

Psyche, 1965

Vol. 72, Plate 7

SCHERBA — MlCROTUS

1965]

Scherba — Microtus

I3i

as a source of food as well as providing shelter for Microtus?

A Microtus was live trapped and placed in an observation box 60
cm. long by 30 cm. wide by 30 cm. high. A full shovel of nest con-
tents and ants from the same nest as the Microtus was added tO' the
box.

Observations under these circumstances indicate that the worker
ants did not avoid Microtus J but instead would attack it whenever
the Microtus came into contact with the ants. When contacted by
an ant, the Microtus responded by hopping quickly a few centimeters
away. Ants which began biting a limb or the tail were usually quickly
dislodged by flicking the leg or tail, or by cleaning the leg with the
teeth. Some workers were not easily dislodged from the tip of the tail
or nares. When bitten in these places, Microtus appeared to be “in
j distress.” On another occasion, a newborn Microtus was seen
writhing when attacked by opaciventris.

It, therefore, seems evident that the ants do not avoid Microtus
\ and that Microtus is not insensitive to, or completely protected against,
ant bites.

Content analysis of three stomachs examined under a dissecting
microscope revealed only freshly chewed bright green plant parts.
No recognizable insect fragments were found.

DISCUSSION

Clearly, Microtus exhibits few of the adaptations one might expect
of a rodent which inhabits ant mounds. On this basis, we might
speculate, as Professor Negus (personal communication) suggests,
that for Microtus montanus ant mounds form a marginal micro-
11 habitat utilized only after other, more suitable, sites are already
\ occupied. In a year of apparent high Microtus density, this explana-
j tion appears reasonable.

We may categorize the relationship of this vole to the ant as that
of a myrmecocole. (Park, 1929).

ACKNOWLEDGEMENTS

Mr. Norman Heryford, Smithsonian Institution, ably assisted this
j work and his contribution is acknowledged with pleasure. Professor
Norman Negus, Tulane University, kindly provided the determina-
ij tion of the species of Microtus under discussion.

Explanation of Plate 7

Figure 1 — External openings of Microtus montanus tunnels at the base
of Formica opaciventris mounds. A covering of vegetation has been removed.

! Note forceps for scale.

Figure 2 — Cross section of mound showing large central Microtus nest
chamber. Dense, soft nesting material of plants has been removed. Note
tunnel into chamber at rear.

132

Psyche

[June

The author wishes to express his appreciation to the National Park
Service for permission to carry out this investigation within the
boundaries of Grand Teton National Park.

References Cited

Bequaert, J.

1921. The predacious enemies of ants. Bull. Amer. Mus. Nat. Hist.,
45: 271-331.

Chapin, James P.

1939. The birds of the Belgian Congo, pt. II. Bull. Amer. Mus. Nat.
Hist., 75: 1-632.

Colli as, Nicholas E.

1964. The evolution of nests and nest-building in birds. Amer. Zool.,
4: 175-190.

Cowles, R. B.

1930. The life history of Varanus niloticus (Linn.) as observed in
Natal, South Africa. J. Ent. and Zool., 22: 1-31.

1959. Zulu Journal. Field Notes of a Naturalist in South Africa. Uni-
versity of California Press, pp. i-xiv T 267.

Donisthorpe, H. St. J. K.

1927. The Guests of British Ants. George Routledge and Sons, Ltd.,
London, pp. xxiii + 244.

Findley, James S.

1951. Habitat preferences of four species of Microtus in Jackson Hole,
Wyoming. J. Mamm., 32: 118-120.

Hardy, John William

1963. Epigamic and reproductive behavior of the orange-fronted para-
keet. Condor, 65 : 169-199.

Haverschmidt, F.

1960. Some further notes on the nesting of birds in termites’ nests.
Emu, 60 : 53-54.

Hindwood, K. A.

1959. The nesting of birds in the nests of social insects. Emu, 59: 1-36.

Negus, Norman and James S. Findley

1959. Mammals of Jackson Hole, Wyoming. J. Mamm., 40: 371-381.

Nichols, J. T.

1940. Synbranch eel in ant nest. Copeia, 1940 (3) : 202.

Park, O.

1929. Ecological observations upon the myrmecocoles of Formica ulkei
Emery, especially Leptinus testaceus Mueller. Psyche, 36: 195-215.

Scherba, Gerald

1961. Nest structure and reproduction in the mound-building ant Formica
opaciventris Emery in Wyoming. J. N. Y. Ent. Soc., 69: 71-87.

1963. Population characteristics among colonies of the ant Formica

opaciventris Emery (Hymenoptera : Formicidae). Jour. N. Y.

Ent. Soc., 71 : 219-232.

1964. Species replacement as a factor affecting distribution of Formica
opaciventris Emery. Jour. N. Y. Ent. Soc., 72: 231-237.

Snyder, T. E.

1929. Friends and foes of termites or white ants. Zool. Anz., 82: 40-46.

Von Hagen, W.

1938. A contribution to the biology of N asutitermes (sensu stricto).
Proc. Zool. Soc. London, 100: 39-49.

OBSERVATIONS ON THREE SPECIES OF
PHIDIPPUS JUMPING SPIDERS
(ARANEAE: SALTICIDAE)*

By Beatrice T. Gardner
Department of Psychology, University of Nevada

The large, brightly colored jumping spiders of the genus Phidippus
are among the most conspicuous American spiders. In many ways,
they are ideal subjects for behavioral studies : the spiders are diurnal
in habit and are highly responsive to visual stimuli, they pursue prey
actively and the males perform elaborate courtship displays.

The present paper compares the behavior and the taxonomy of three
Phidippus species that are found in the same fields in the vicinity of
Reno, Nevada. The behavioral description emphasizes Phidippus
coccineus , as this species is being used in the laboratory for experiments
which continue a previous line of investigation on hunting responses
(Gardner, 1964). The other two species of Phidippus J P. apacheanus
and P. opifex , were collected and maintained in the laboratory for
comparative observations.

The diagnostic features for the three species have been illustrated
(Figs. 1 -10) since there are few records other than the original
description of each species. A modern taxonomic revision of the
genus would be of considerable value in helping research workers
determine specimens and in stimulating comparative studies.

Habitat

Phidippus coccineus Peckham and Peckham, 1909, occurs in the
sagebrush community of the Great Basin desert. These large jumping
spiders are found on bushes such as the sagebrush ( Artemesia triden-
tata) , the rabbitbrush (Chrysothamnus nauseous ), and the four-
winged saltbrush (A triplex canescens) . Phidippus coccineus prefers
bushes that grow on slopes with thin, stony soils, and appears to
avoid conifers and moist habitats (e.g., the proximity of irrigation
ditches). In the same habitat, often on the same bush, two other
Phidippus species are found : P. apacheanus Chamberlin and Gertsch,
1929, and P. opifex (McCook, 1883).

Description

The total size of adult P. coccineus ranges from 7-1 1 mm (median
— 9 mm) for males and from 10-12 mm (median — 10.5 mm) for
females. Both males and females have blue-green irridescent cheli-

*Manuscript received by the editor February 3, 1965

133

134

Psyche

[June

Figs. 1-3. Phidippus opifcx (McCook). 1. left palpus. 2. Female geni-
talia, dorsal view. 3. Epigynum.

Figs. 4-6. P. apacheanus Chamberlin and Gertsch. 4. left palpus. 5.
Female genitalia, dorsal view. 6. Epigynum.

Figs. 7 - 10. P. coccineus Peckham and Peckham. 7, 8. Left palpus. 7.
Ventral view. 8. Ectal view. 9. Female genitalia, dorsal view. 10. Epigynum.

cerae, a black cephalothorax and black limbs, and a bright red abdo-
men with a median black stripe. There are two pairs of minute white
spots between the black and red areas on the posterior part of the
dorsum of the abdomen. At the sides of the abdomen there are light
diagonal bands, and there is also a light transverse anterior band on
the dorsum. These markings vary in conspicuousness : the bands and
spots may be only a somewhat lighter shade of red than the remainder
of the abdomen, while the median black stripe may be so reduced in
width and length that the abdomen appears solid red. Sometimes, the
basic color is orange rather than red, and very old spiders may even
be yellow. In this species, the adult males and the adult females are

1965]

Gardner — Phidippus

135

similar in coloration, and this is also true of P. apacheanus. It is
more usual for the males of Phidippus to have irridescent chelicerae
and a distinctive adult coloration while the females of this genus
remain similar to the immature spiders (e.g., P. clarus Keyserling,
P. opifex, P. whitmanii Peckham and Peckham).

From the second instar on, Phidippus coccineus spiderlings have a
brownish-gray cephalothorax and limbs, and a red abdomen with
markings limited to the posterior portion. These consist of a pair of
black stripes, each bearing two conspicuous white dots, separated by a
light region. The light region may be gray, white, or even golden;
immediately anterior to the black stripes it is enlarged into a con-
spicuous light dot. While the abdomen of very young spiderlings
appear bronze, the red color of the abdomen is striking in later instars.
In one of the later instars, the 5th or perhaps the 6th, a red cap
appears in the eye region, but this marking disappears in the following
molt. The light basal band and side bands of the adult are also
present in immature spiders, and, in the two instars preceeding ma-
turity, the chelicerae are also irridescent.

Methods of collection

Phidippus coccineus can be collected by sweeping bushes with an
insect net. These hunting spiders are active from mid-morning until
dusk, and can be seen in bushes running along branches or poised near
their tips. Running is interrupted frequently: the spider stops, turns
to one side and then the other, apparently scanning its surroundings.
In the laboratory, the spiders are active between io A. M. and 4
P. M., and are still in a closely-woven retreat at other times.

Adults and large immatures are found on heavier branches near
the center of the brush, and are not accessible to the sweeping net.
Certain characteristic aspects of their escape behavior can be used in
capturing them. When approached slowly by a forefinger or a pencil,
the spider backs away while facing the oncoming object, and will
back right into a collecting tube held behind it on the branch. How-
ever, once the spider backs sufficiently far from the object (about
8 cm), it turns away. Then it will run a short distance along the
branch, attach a thread of silk, and plunge down toward the leafy
debris at the base of the bush, where it becomes extremely difficult
to find.

Phidippus coccineus can also be collected in its retreat, or nest.
The spiders will be found within a slightly flattened tube of silk, which
is surrounded by guy-lines that attach it to the twigs or leaves of the
bush. The molting nest and the nest that houses the female and her
cocoon of eggs are of the same basic shape as the sleeping nest, but
much more silk is used in their construction. Such prominent white

136

Psyche

[June

webs are found under stones at the base of the bush. Usually, the
spider places its nest under one of the few stones that remains shaded
from the strong desert sun even at mid-day. Occasionally, P. coccineus
builds nests inside or under cardboard containers dumped in their
habitat. Phidippus opifex makes frequent use of the insulation pro-
vided by these cardboard boxes; alternatively, and unlike coccineus ,
opifex builds a large and prominent nest among the branches of a
bush to souse its egg cocoon.

Life history

The breeding season of P. coccineus appears to be a very early one.
In 1964, six adult males, nine adult females, and about 90 immature
spiders of this species were collected near Reno, Nevada. The males
were found first, even before the final snowstorms of early June; adult
males were found from early April to early July, adult females,
from early May to mid-July. All but one of the females collected
in May were already gravid and laid eggs within the following
fortnight; all females obtained later in the summer (mainly, in July),
were found in a nest with an egg cocoon. Two spiders in the penulti-
mate instar were found in late May, and these became mature in the
laboratory toward the end of June. But from mid-July on, only
immature spiders could be found and none of these attained maturity
during the summer or fall of 1964, though they have molted as many
as four times since coming to the laboratory. It is of some interest
to note that the females survived the males by almost two months
when kept under identical conditions in the laboratory, and that,
for each sex, all the adults died within a short span of time (e.g., all
the females between 18 Sept. 1964 and 6 Oct. 1964).

The early breeding season of P. coccineus contrasts with that of
P. opifex and P. apache anus found in the same habitat. Adult males,
adult unmated females, and females in the penultimate instar of
P. opifex were collected in the first half of July; the nests containing
the egg cocoons of this species were abundant in the sagebrush late
in August. Phidippus apacheanus had the latest breeding season
among the three species: while immature spiders of this species were
abundant throughout the spring and summer, adults were only found
after the third week of September. Six immature P. apacheanus kept
in the laboratory since early July molted three times and became
mature around the end of October (see Table I).

The seven P. coccineus females kept in the laboratory during the
summer of 1964 laid either two or three successive batches of eggs,
each batch containing fewer eggs. A median of 44 spiderlings hatched
from the first batch of eggs, 30 from the second, and three from the
single third batch that did hatch. By contrast with females that

1965]

Gardner — Phidippus

137

Table I

The 1964 breeding season for the three species of Phidippus at Reno, Nevada

Collection dates for:*

P. coccineus

P. opifex

P. apacheanus

juveniles in pen-
ultimate instar

Oct. - June

? - July

(Aug.) - Oct.

adult males

April - July

July - Aug.

Sept. - Oct. |

adult females:
unmated
gravid

w. egg coccoon

May

May - July
July

July

?

Aug. - Sept.

Oct.

Oct., Nov. - ?
(Dec. - Jan.)**

^Parentheses indicate that field collection dates are unavailable and dates

for spiders maintained in the laboratory have been used.

**Possibly, abnormally early. In the laboratory, warmth and abundance of
prey during the winter produces anomalies, e.g., penultimate P. coccineus
juveniles become mature in Dec. -Jan.

bear only one batch of eggs, such as P. clarus Keyserling of the
northeastern United States, coccineus females came out of their nests
to capture the prey that were offered daily. Phidippus clarus remains
with the egg cocoon continually, and generally dies within a few days
of the spiderlings’ emergence from the nest. For P. coccineus, the
median time from oviposition to hatching was 21 days. The young
remained in the web through a first molt some 16 days later, and
emerged from the nest 21 days after hatching. These times show
little variation : for 1 1 batches of spiderlings the range of time from
egg-laying to emergence from the nest was 37-46 days.

The spiderlings were self-sufficient when they left the nest. If
moved to an individual glass vial, almost all spiderlings built a minia-
ture of the adult sleeping nest within two days. Flightless Drosophila
placed in the vial are soon captured : for one batch of 24 spiderlings,
the median catching time was 3.7 min. and all but two of these
spiderlings had caught their first prey within 60 min. However, it is
not easy to rear the spiderlings : only 30 of almost 180 young, kept
in individual vials over wet cotton wicks and fed every two days,
have survived to date (i.e., approximately five months and four post-
emergence molts). Most of the losses occurred in the first fortnight
after leaving the nest ; later deaths generally occurred while the
spider was molting.

Courtship

Method of observation

Complete mating behavior can be observed by placing the male
together with a female reared in the laboratory from the penultimate
instar to maturity, that is, a female known not to have mated pre-

138

Psyche

[June

viously. In such cases, courtship is likely to be very brief : the female
will accept the male after 30-60 sec. of courtship. The initial phase
of mating behavior, the courtship dance, can be observed with a wider
variety of partners: females laden with eggs, females that have laid
erggs, females of a different Phidippus species, females in glass vials,
and even simple models of females such as hemispheres of modelling
clay with a pair of wires as appendages (Drees, 1952). When a pre-
viously mated female is used, courtship is not likely to last long and
may end violently: the female will run away from the male or jump
at him as she does at prey. Prolonged courtship can be seen with
females confined in transparent vials, and with models of the female.

A glass or plastic hemisphere 15-20 cm in diameter provides a
suitably-sized observation chamber in which the spiders are always
in full view. The chamber should have a rough textured floor as
adult P. coccineus refuse to move or move unnaturally on a slippery
surface. It is possible to use the spider’s preference for moving upward
to introduce animals into the chamber with a minimum of disturbance:
the spider climbs into a vial placed over it in the home jar, and
climbs into the chamber when the open end of the vial is fitted into
a hole in its floor. As the spider is likely to orient toward the observer
when he is moving, it is advisable to shield writing movements and
to observe from a constant position, with a light intensity differential
to further obscure the observer. The Dazor Floating Magnifier unit,
which combines a wide-angle lens and fluorescent light, is very suitable
for this purpose.

Results

The courtship display of various Phidippus males ( apacheanus ,
clarus, coccineus , opifex) has a common basic pattern. The male
begins his display by holding the carapace very high, shifting the
abdomen to one side, and raising the first pair of legs. In this position,
he moves before the female, stopping after each few steps. The male
advances in a zig-zag pathway, shifting his abdomen to the other side
at the end of each oblique approach. Throughout, the dancing male
flicks his forelegs up and down, holding them wide apart at first and
bringing them closer and closer together as he nears the female.
Then, with forelegs held almost parallel before him, he touches the
female cautiously once or twice. If the female remains still at this

Explanation of Plate 8

Figs. 11-12. Phidippus opifex (McCook), female, total length 14 mm.
The female is tan, the male is black with light gray dorsum of carapace
and abdomen. (The white rings in the eyes are reflections of the light used.)

Psyche, 1965

Vol. 72, Plate 8

Gardner — Phidippus

140

Psyche

[June

stage, the male climbs over her, and uses the forelegs to help turn
her abdomen to the side. When the genital pore, which lies on the
ventral abdomen, is thus exposed, the male inserts his palpus. After
2-3 min., the male withdraws this pedipalp, turns the female’s abdo-
men in the other direction, and inserts the other pedipalp (fig. 2010,
Kaston, 1948).

Raising the forelegs and holding the abdomen to one side are by
no means specific to courtship. This display occurs to a wide variety
of objects that are approaching too close to the spider: other spiders
■ — regardless of sex (Crane, 1949), houseflies or other large prey,
models of prey that are about the size of the spider, such as 9-12 mm
spheres, or even the end of a finger or a pencil. In these situations,
however, the spider with its forelegs raised and waving backs off while
facing the moving object and, when 5-8 cm away, turns away and
flees. In intraspecific encounters, the effect of raising the forelegs is
to bring the other spider to an abrupt halt, whether it be a wandering
cagemate, a courting male preparing to touch the female, or a female
stalking or about to jump at the male. Indeed, raising of the forelegs
appears to have this ‘stop-sign’ effect even in encounters between
congenerics of the three Phidippus species. Drees, working with
female Salticus scenicus, was able to initiate hunting behavior by
moving a black dot along a white wall, and to stop the pursuing
spider by moving wires projecting from the side of the model through
an angle that imitated the waving of the forelegs (Drees, 1952).

Female P. coccineus and P. apacheanus are unusual in that they
perform an acceptance dance just before the male touches them. With
forelegs high and wide apart and abdomen bent to the side, the
female sways before the male, sometimes with a few steps to one
side and then the other. In other Phidippus species, the female rejects
the male by extending the first pair of legs whenever he approaches
too closely, and merely fails to ward him off when she is ready to
accept him.

Differences in the display of the three Phidippus species remain to
be investigated. At this stage, it is apparent that subtle quantitative
differences are involved. For example, while P. apacheanus moves
his forelegs both closer and higher as he nears the female until the

Explanation of Plate 9

Figs. 13 - 16. Phidippus apacheanus Chamberlin and Gertsch. 13. Juve-
nile, total length 9 mm. 14. Adult male below, female above, total length of
each 11.5 mm. Both male and females are black with bright orange dorsum
of carapace and abdomen, and with iridescent green chelicerae. 15. Male
signalling female. 16. Male signalling and facing female.

1965]

Psyche, 1965

Vol. 72, Plate 9

Gardner — Phidippus

Gardner — ■ Phidippus

142

Psyche

[June

tips touch in a circle above his head, P. coccineus keeps his forelegs
in the same vertical plane while bringing them together; again, P.
apacheanus is only 5 to 8 cm away from the female when he begins
to dance, and hence performs less extensive side-to-side movements
than does P. coccineus (or P. clarus) . There are also variations in
the tempo of flicking the forelegs, and in the synchrony of the two
forelegs, but further study is needed to sort out individual, drive-
level, and species differences.

As was previously noted, the adult apacheanus , coccineus , and
opifex appear in the same habitat at different times, so that the time
at which each species was courting did not overlap in the summer
of 1964. However, adults of one species were kept in the laboratory
until adults of the species succeeding them in the fields were collected.
In the few relevant observations, the male courted the female of a
different species as eagerly as his own (indeed, more eagerly if she
were larger, as in the case of P. opifex females), while the female
ran away or initiated hunting behavior as soon as she was released
from the transparent vial.

Hunting

Method of observation

Investigation of hunting behavior depends on techniques for con-
trolling the hunger-level of the spider and the stimuli received from
the prey.

As would be expected of a predator notorious for withstanding
long fasts, the three Phidippus spp. will molt, reproduce, and survive
many months in the laboratory, whether offered prey daily, bi-weekly,
or once a week. To control hunger-level, a behavioral criterion for
satiation was used : the spider was presented with successive Droso-
phila until it failed to accept a fly within 10 min. of its presentation.
Hunting was then tested at a specifiable level of food deprivation;
tests at 1, 7, and 12 days deprivation yielded considerable variation
in the intensity of hunting behavior (Gardner, 1964).

Explanation of Plate 10

Figs. 17 - 18. Phidippus coccineus Peckham and Peckham. 17. Juvenile,
total length 5 mm. The color is gray except for the abdomen. The abdomen
dorsum has a white basal band and the reddish area is separated posterior
by black bands broken by white spots. 18. Female, total length 10.5 mm.
Both males and females are black with bright red dorsum of the abdomen
and iridescent blue-green chelicerae. The longitudinal black stripe on the
abdomen is more distinct in females. The males (not illustrated) have a
mean total length of 9 mm.

Psyche, 1965

Vol. 72, Plate 10

Phidippus

144

Psyche

[June

In hunting, as in courtship, visual stimuli are the most important,
and complete hunting behavior can be observed with live prey in
transparent containers, dead prey dangled by means of an attached
fine gauge wire (or otherwise made to move), and simple models of
prey, such as moving dots or spheres. In hunting, the spider is
essentially tracking a moving target, and full control over the stimuli
received from the prey is possible with a moving model whose speed,
distance, and angular displacement can be regulated (Drees, 1952;
Precht, 1952). The main difficulty in working with actual prey lies
in the highly variable movements of living animals. To some extent,
this can be overcome: flightless Drosophila mutants with a uniform
manner of locomotion have obvious advantages over a prey that can
fly, and hence sporadically disrupts an ongoing bout of hunting
behavior by an abrupt change in angle and distance from the spider.
It is important to arrange a situation in which the prey is not some-
times moving and sometimes still, as the spider seldom initiates hunt-
ing behavior to still prey, and interrupts ongoing hunting behavior
when the prey ceases to move. Curly-type Drosophila melanogaster
are suitable for this purpose as they crawl upward on a vertical sur-
face and move continually until they have reached the top. The
pathway of the prey in relation to the predator can also be made
uniform. Jumping spiders will readily climb a slender pole, and
position themselves, facing down, near the top ; the curly-type Droso-
phila also climb such a pole, moving continually and at a constant
speed straight toward the spider. With such a technique, the distance
at which the spider initiates hunting can be measured, and can be
compared for different species or for different periods of deprivation.

Results

Hunting behavior in salticid spiders can be described as a sequence
of stimuli and responses: the spider performs a given response until
there is some crucial change in the stimuli received from the prey
which then evokes the succeeding response. Thus, when introduced
into a hemispherical testing chamber, the spider generally climbs to
the top. A few seconds after the prey enters, the spider orients by
turning so that the large frontal eyes come in line with the prey, and
this relative position is maintained throughout the remaining hunting
responses. Once lined up, the spider pursues , moving rapidly at first
but more slowly as it comes near the prey. When P. coccineus is
within 5 cm of the prey, the spider presses its body close to the
ground, and draws the legs in toward the body. At about 1.5 cm,
the spider becomes still in this crouched position, attaches a thread

1965]

Gardner — Phidippus

145

of silk to the substratum, and jumps at the prey. Somewhat more
elaborate tracking occurs when a small P . coccineus pursues a much
larger prey, such as a blue-bottle fly: the spider may take a curved
course so that the final jump is from behind the prey.

It is possible to vary the frequency and duration of each hunting
response by withholding the stimuli for the succeeding response
(Drees, 1952; Precht, 1952; Gardner, 1964). For example, if the
model is kept about 10 cm from the spider by moving it at the same
speed as the pursuing predator, pursuit is prolonged far beyond its
normal duration as the next response, crouching, requires a very
small distance between prey and predator. Again, a model moving
slowly within a transparent tube 2.5 cm in diameter seldom evokes
jumping for the crouched spider cannot get close enough, while with
a narrower tube jumping occurs readily.

Having captured the prey, the spider settles in one spot and does
not move again until it has discarded the undigestible hard remains
of the prey — a small mass in the case of the Drosophila, or the
perfect empty shell of a large prey such as Calliphora . The time
required to consume the prey reflects the relative size of the spider
and its prey. For example, a group of small (4-6 mm) immature
P. coccineus, tested at 7 days food deprivation, required a median
time of 29.O min. to consume a Drosophila, while larger individuals
(8-10 mm) required only 8.6 min. Even the large adult females
required almost one hour to consume large prey such as Musca
domestica. For P. coccineus (and P. clarus as well), consuming time
increases reliably with days of food deprivation. Thus, the same
group of small spiders required only 21.8 min. to consume a Droso-
phila when tested at 1 day food deprivation.

Hungry P. coccineus will readily capture more than one prey at a
time, provided the second prey comes very near (the spider will not
move away from the place at which it has settled with the first prey).
This situation was explored with the pole technique and perhaps it is
common in the natural counterpart of the pole — the branch tips
where hunting Phidippus can be found. The spiders accepted several
pairs of prey on the pole, but as they became satiated, it was much
less likely that they would accept the second prey. Instead, the spiders
responded by extending the forelegs toward the Drosophila — once
again using the ubiquitous display that occurs in courtship, intra-
specific encounters, etc.

Given the choice between two prey that differ in size, such as
Musca and Drosophila or Calliphora and Drosophila, P. coccineus
chooses the large prey less consistently than does P. opifex and P.

146

Psyche

[June

apacheanus. In the field, both opifex and apacheanus have been seen
pursuing huge prey, such as grasshoppers and bees. A difference in
preferred size of prey is also found between smaller and larger P.
coccineus — although a 4 mm spiderling that pursued and caught a
9 mm housefly provides a notable exception. Both these factors would
reduce competition for food among the three Phidippus species. At
the present time, preferred size of prey is being investigated with
models (spheres 3, 6, 9 and 12 mm in diameter, moving at 8mm/sec.).

Nest building

As Phidippus spends the daylight hours wandering from branch to
branch while hunting prey, it seems likely that a new nest is built
every evening. In their laboratory containers, P. coccineus only occa-
sionally built more than one nest; ordinarily, the spiders returned to
the same nest, night after night. If however, the nests are removed
in the morning, the spiders will build a new nest late that afternoon.
On a typical test day, 19 spiders had their nests removed and were
exposed to natural light conditions. Of these, 16 built a nest within
the next 24 hrs. All nest building began in the afternoon, at a
median time of 55 min. before sunset. (Such a generalization is
limited to the sleeping nests of immature P. coccineus , for the molting
nest is begun much earlier in the day, and the adult often lets many
days pass without building.)

The spider begins to build by moving within a limited area in the
angle formed between the sides and top of the container. As it moves,
it attaches a framework of threads of silk, each about two or three
times its own length. Suspended within these lines, the spider starts
to weave by moving the abdomen from side to side so that the
spinnerets describe an arc of a circle. Moving forward and back a
step at a time, sometimes suspended on its back and sometimes on its
venter, the spider spins about itself a closely woven tunnel, roughly one
and one-half times its own length. Once started, weaving continues
uninterrupted, and by 30 min. after sunset most of the builders are
quiet in their nests. The roles of light and temperature in the choice
of nest-site and in the rather precise timing of this distinctive diurnal
behavior are presently being investigated.

Distribution of the species

The male and female syntypes of P. coccineus came from Clare-
mont, California and are kept in the Museum of Comparative
Zoology. As far as known the species occurs in southern and eastern
California and western Arizona. Phidippus opifex came from San

1965]

Gardner — Phidipp.us

147

Bernardino, California, the type should be in the Academy of Natural
Sciences, Philadelphia but could not be found. Its distribution is in
the southwestern United States. The male holotype of P. apacheanus
came from Black Rock, Utah; it is common in the southwestern
United States across the southern states to Kansas, Louisiana and
possibly to the east coast.

Acknowledgements

Dr. W. J. Gertsch provided the distribution data and identified
P. opifex , Dr. Herbert W. Levi identified the other two species by
comparing to the types and prepared the diagnostic illustrations.
Jane E. Thompson, an NSF undergraduate research participant, made
available her data on nest building. The photographs of P. apacheanus
displaying are by Dr. G. P. Ginsburg; all other photographs are by
Dr. Herbert W. Levi. The research was in part supported by Public
Health Service Research Grants MH-07200 and MH-08938 from
the National Institute of Mental Health.

References

Chamberlin, R. V. and W. J. Gertsch

1929. New spiders from Utah and California, Pomona College, J. Entom.
Zool., 21: 101-112.

Crane, J.

1949. Analysis of display of Salticid spiders. Zoologica, New York,
34: 159-212.

Drees, O.

1952. Untersuchungen uber die angeborenen Verhaltensweisen bei
Springspinnen ( Salticidae) . Z. Tierpsychol., 9: 167-207.

Gardner, B. T.

1964. Hunger and sequential responses in the hunting behavior of
Salticid spiders. J. Comp. Physiol. Psychol., 58 : 164-173.

Kaston, B. J.

1948. Spiders of Connecticut. Hartford: State Geological and Natural
History Survey, Bull. 70.

McCook, H. C.

1883. Proc. Acad. Nat. Sci., Philadelphia, p. 276.

Peckham, G. W. and E. G. Peckham

1909. Revision of the Attidae of North America. Trans. Wisconsin
Acad. Sci. Arts Lett. 16: 355-646.

Precht, H.

1952. Uber das angeborene Verhalten von Tieren. Versuche an Spring-
spinnen (Salticidae). Z. Tierpsychol., 9 : 207-230.

NEW SPECIES OF CHILOCORINI
(COLEOPTERA: COCCINELLIDAE)*

By Edward A. Chapin

Museum of Comparative Zoology, Harvard University

In the course of a study of the genera of this tribe a few un-
described species were found and are herein named. It was found
desirable to elevate Zagreus Mulsant from subgeneric to generic rank
because of certain structural differences that exist between that group
and Exochomus Redtenbacher. It seems likely that Zagreus , with
eight-segmented antennae, has replaced Exochomus , whose antennae
are ten-segmented, in the New World south of Panama.

Zagreus costalimai n. sp.

(Fig. 2)

Length: 3. 2-3. 5 mm. Similar to Z. jordani (Mulsant) (Fig. 1)
but slightly larger and with two additional subapical sutural spots
common to the two elytra. Head pale yellow (male) or piceous
black (female), finely and sparsely punctulate, sparsely set with fine
short pubescence, the surface between the punctures alutaceous.
Pronotum yellowish with a large blackish spot which extends over
five-sixths of the basal margin and which fails narrowly to reach the
anterior margin. The surface sculpture similar to that of the head
but without pubescence. Scutellum small, triangular, black. Elytra
yellowish with three common sutural spots and each with two others;
the first and largest of the sutural spots lies in the basal half of the
conjoined elytra, is almost quadrate and in length along suture is
two-sevenths of the total length of the beetle. It is flanked on either
side by large subtriangular spots, each slightly smaller than the
sutural spot. In the apical half of the elytra there are the second
and third sutural spots, each smaller than the preceding. On either
side of the second sutural spot is an almost oval spot which is about
two-thirds as large as the sutural spot. Both lateral spots are sepa-
rated from the lateral margin by a distance equal to the separation
between the lateral and sutural spots. The apical sutural spot is
about half as long and half as broad as the second sutural. Under
parts pale yellow, except for the blackish metasternum and first and
second abdominal sternites. Metacoxal arcs complete. Elytral
epipleura not foveolate. Legs pale except that the femora (posterior
in male, all in female) are partially dark.

* Manuscript received by the editor January 26, 1965

1965]

Chapin — Chilocorini

149

Type: a male, U. S. N. M. No. 67801, from Brazil, Rio de
Janeiro Province, Campo Grande, July 31, 1957, P. A. Berry. Two
female paratypes with same data as type.

I name this species in honor of the late Dr. A. da Costa Lima, in
recognition of his many contributions to our knowledge of South
American entomology.

Zagreus tetraspilus n. sp.

(Fig. 3)

Length: 3.0-3. 2 mm., superficially resembling Exochomus bisbino-
tatus Gorham but larger, black with bluish reflections and with all
spots smaller and the basal spots smaller than the apical spots. Head
pale yellow (male) or black except for the clypeal margin and genal
extensions which are brown (female), surface finely alutaceous and
very finely and rather sparsely punctulate, pronotum twice as wide

Figures 1-4. 1. Zagreus jordani (Mulsant). 2. Zagreus costalimai n.sp.

3. Zagreus tetraspilus n.sp. 4. Phaenochilus renipunctus n.sp.

as length on median line, black except for the very narrow brown
anterior and lateral margins, surface as described for head but with
stronger and slightly denser punctulation. Scutellum black, polished.
Elytra black with two yellow spots on each. The one on humeral
callus is oval and reaches from the basal margin backward and out-
ward a distance equal to the length of the pronotum at the point
adjacent to the spot. The apical spot is more nearly circular, is
slightly larger than the basal spot and fails to reach either lateral or
sutural margins. Surface as described for head but with much
stronger punctation. Inner half of epipleura yellow. Thoracic ster-
nites and most of the first two abdominal sternites of female piceous
black, rest of abdomen and legs brownish-yellow, the legs slightly
darker; sternites of pro- and mesothorax and side pieces of meta-
thorax, sides of first two abdominal sternites and rest of abdomen
and legs brownish yellow in male.

150

Psyche

[June

Type: a male, U. S. N. M. No. 67802, from Colombia, Int.
Choco, Istmina, Aug. 21, 1940. L. M. Murillo No. 5335. Two
female paratypes from type locality, same data as type.

Exochomus hisbinotatus Gorh. from Mexico to Guatemala is a
smaller species (2 mm.) with proportionately larger spots and with
the lateral margins of the elytra yellow.

Exchomus hisbinotatus Gorham is not known to me in nature but
only from the original description and figure in the Biologia Centrali-
Americana. Therefore I am not sure that the species does belong to
Exochomus. However I have not yet seen a species of Zagreus from
a mainland locality north of Panama. Since E. hisbinotatus is re-
corded from Mexico, British Honduras, and Guatemala it is probably
not a Zagreus. The most northerly species of Zagreus is Z. ritchiei
(Sicard) from the island of Jamaica. Exochomus jamaicensis Sicard,
also from Jamaica, is correctly placed as to genus.

Gurinus colombianus n. sp.

Length : 4.3-4. 8 mm. Similar in dorsal coloration to C. coeruleus
Mulsant but with the under parts and legs entirely pale reddish
yellow. Head steel-blue above, piceous beneath and around mouth-
parts, surface finely alutaceous, finely, moderately, and evenly punctu-
late. Pronotum slightly more than twice as wide as length on median
line (male) or twice as wide as long (female), median third steel-
blue with lateral thirds reddish-yellow. Surface sculpture same as
that of head except that the sides become more densely punctulate
toward lateral margins. Scutellum small, black and polished. Elytra
uniformly steel-blue above, the epipleura piceous. Surface sculpture
same as that of median area of pronotum except toward apices where
the punctulation becomes so minute as to be almost invisible under
high magnification. Under parts pale reddish-yellow, shining and
very sparsely punctulate, the punctures slightly coarser toward sides
and apex of abdomen. Metacoxal arcs as in C. coeruleus Muls. Fifth
sternite of male broadly and shallowly emarginate, the sixth with a
V-shaped notch at apex; that of female evenly rounded at apex.

Type: a male, U. S. N. M. No. 67803, from Colombia, Dept.
Valle del Cauca, Palmira, taken May 18, 1939, by L. M. Murillo
(5020). Paratypes (54) from type locality : same data as type (22);
Oct. 29, 1943, F. R. Fosberg (25); April 1942, B. Losada (1);
April 1943, B. losada (3) ; August 1943, B. Losada (1) ; Feb. 1944.
B. Losada (2).

Other specimens seen: Colombia. Valle del Cauca. Cali, 1939,
B. Losada; Cauca Valley, 1937, F. L. Gallego; Buga, Feb. 3-7,
1941, Murillo (14, 32, 5409). Antioquia. Valle Medellin, May-

1965]

Chapin — Chilocorini

151

Nov., 1943, 1944, i945j Gallego; Venecia, July 1942, 1943, Gallego;
Segovia, Sept. 1944. Gallego; El Picacho, May 1942, Gallego;
Sopetran, July 1943, Gallego. Cundinamarca. Gacheta, Aug. 27,
1941, Murillo (5220). Boyaca. Guayata, Oct. 18, 1940, Murillo
(5297). Santander. San Vicenti de Chucun, June 8, 1935, Murillo

(83).

While Curinus coeruleus Mulsant does occur in Colombia, it seems
to be quite rare and it is generally replaced by C. colombianus , which
has a wide distribution in the country. Large specimens of C. colom-
bianus can be immediately distinguished from small specimens of
C. coeruleus by the uniform reddish-yellow color of the underside.
In addition to the Colombian material listed above, one specimen
believed to have come from Costa Rica, and one specimen from
British Honduras, Belize, Sept. 1959, N. L. H. Krauss have been
seen.

Phaenochilus renipunctus n. sp.

(Fig. 4)

Length: 3.7 mm. Upper surface pale yellowish-brown, each

elytron with a piceous black, oblique, reniform spot just behind the
humeral callus. Under surface and legs pale yellowish brown except
for the metasternum which is somewhat darker. Front of head
slightly concave between the large eyes, finely and sparsely punctulate
near eyes, impunctate medially. Pronotum polished, very finely and
shallowly punctulate, basal angles obsolete, lateral margins near
obtuse frontal angles finely reflexed. Scutellum very small, triangular.
Elytra with lateral margins broadly reflexed, surface polished, finely,
shallowly and sparsely punctulate, the punctures near sutural and
lateral margins deeper and more evident than those on disc. The
reniform spot is two and a half times longer than its width at mid-
point; it commences two-thirds its least width behind the basal
margin and extends backward and outward toward but not reaching
the reflexed portion of the elytron. Elytral epipleura shallowly but
distinctly foveolate for reception of femoral apices of second and
third legs. Abdomen with five visible sternites (female), metacoxal
arcs strong, running backward and outward nearly to margin of
sternite, then following sternite margin toward lateral margin of
abdomen. Fifth sternite (female) evenly rounded. Tibiae each with
a sinuate denticle much as in Chilocorus. Tarsal claw with elongate,
triangular basal tooth which is two-thirds as long as the apical half
of claw.

Type: a female. M. C. Z. No. 31 143, from Morotai I., Indonesia,
Sept., 1944, P. J. Darlington.

TECHNIQUES FOR THE STUDY OF
SPIDER GENITALIA

By Herbert W. Levi *

Museum of Comparative Zoology

In response to many inquiries, I want to describe the techniques
used for studying genitalia in the spider family Theridiidae. These
procedures are not original, but have been adapted from methods used
by several colleagues and students. One technique used in the family
Araneidae is new and, I hope, helpful to students of that family.

I would like to urge strongly against making permanent microscopic
slides of genitalia. Most slides are not permanent; even balsam may
crystallize after fifty years. In dehydration, sclerotized parts may warp
and, as the slide dries, the cover glass may crush anatomical struc-
tures. The slides often become separated from the parts of the
specimen kept in alcohol and are lost. However, mounting the geni-
talia of common species on slides often saves time in their study. The
medium found most useful is Hoyer’s fluid (Baker and Wharton).

Luckily, most male spiders have two palpi. One, usually the left,
is removed so that it can be turned to the desired angle. Palpi should
always be examined completely submerged in 75 to 90% alcohol,
never dry. Drying shrinks and distorts softer parts; partial submer-
sion produces undesirable reflections. To prevent the parts from
floating away in convection currents caused by the heat of the micro-
scope light, the light is equipped with heat absorbing glass and
the pedipalpus anchored in a piece of fibrous paper tissue at the
bottom of the dish. Vaseline, sometimes recommended to keep the
palpus in position, may cling to it, smear over it, and is difficult to
remove. Examination is by binocular dissecting microscope, at magni-
fications of 150 to 240 times.

Small, translucent, weakly sclerotized palpi are common among
theridiid spiders. A transfer to glycerine, after blotting off alcohol,
may help to make visible the borders of transparent structures. A
temporary slide mount in glycerine may be made; for examination
under a compound or phase microscope. The palpus should always
be returned to alcohol in a microvial (4X10 mm) stoppered with
cotton, to be kept with the spider in a larger vial ( Fig. 1 ) . A per-
manent mounting generally ruins the palpus for study, as sclerites
become too transparent, may become distorted, and can not be turned

* Manuscript received by the editor November 4, 1964

152

1965]

Levi — Spider Genitalia

153

Figures 1-5. Techniques for the Study of Spider Genitalia. Fig. 1. Vial
containing two smaller vials, all stoppered with cotton. Fig. 2. Abdomen,
showing place of incision. Fig. 3. Abdomen with genital area bent over.
Fig. 4. Abdomen positioned for drawing internal genitalia. Fig. 5. Part of
cephalothorax with pedicel and abdomen integument including genitalia.

and placed conveniently. I have examined a few such slides of geni-
talia of holotypes only twenty years old ; they were almost useless.

Expanding the palpus to examine the position of sclerites is of
value in phylogenetic studies, but is rarely useful for descriptions in-
tended to facilitate species recognition. After the alcohol is blotted
off, the palpus is brought to boiling in 10% NaOH solution. Then
the palpus is transferred directly to distilled water where the hema-
todocha rapidly expands. The palpus can then be transferred to
alcohol for storage. Unfortunately, boiling in NaOH warps, distorts
and damages sclerites. Full strength lactic acid can be substituted
for sodium hydroxide. Sclerotized palpi must be boiled longer than
soft ones.

154

Psyche

Dune

The epigynum of holotypes or rare species should not be removed
from the body of the specimen. In many small spiders, especially if
the epigynum is weakly sclerotized, the whole spider can be dropped
into clove oil, after first blotting to remove excess alcohol. This
procedure is satisfactory for routine examination of Clubiona , Dipoena
and other small spiders having little pigment. After a few minutes’
clearing, the ducts can be seen ; longer clearing makes them too trans-
parent. In these two genera, specific determination is based on charac-
teristics of the duct that loops between the seminal receptacle and the
external plate. In other genera the ducts are behind the seminal re-
ceptacles. For description of new species, a more careful examination
is necessary. After removal from clove oil, the spiders are again
blotted and returned to alcohol for storage.**

The procedure used for more careful examination is to make two
cuts around the genital area (Fig. 2). The tools used are mounted,
sharpened insect pins (minuten nadeln). The epigynal plate with
seminal receptacles is folded back (Fig. 3) before or during submer-
sion in clove oil. The specimen is then anchored on fibrous paper
tissue for examination (Fig. 4). After examination the genitalia are
bent back, like closing a door. In poorly preserved specimens! the
structure sometimes breaks off. In that case it should be stored in a
microvial. On very small spiders it is advantageous at times to sepa-
rate cephalothorax and abdomen, but the epigynum and neighboring
integument should remain attached to the pedicel (Fig. 5). Some-
times a temporary slide can be made of the whole cephalothorax with
genitalia.

If there are numerous specimens and difficulties in their study, it
is best to make a temporary microscope slide of the epigynum with
its accompanying structures. The epigynum is first cleared in clove
oil and then, with a medicine dropper, it is transferred in a drop of
clove oil to a microscope slide and covered with a coverglass. This
preparation can be examined under a compound or phase microscope
and the parts later returned to alcohol to be stored in a microvial.
But there may be difficulties in this procedure. In Helvibis (Levi,
1964) it was possible to follow the course of the tortuous (but diag-
nostic) epigynum ducts in only two species: one because the ducts
were short and simple, the other because I could destroy one of the
numerous specimens available and tease the ducts apart with needles.

**Prof. M. E. Galiano informed me that examination under oil may damage
iridescence or structural colors.

1965]

Levi — Spider Genitalia

155

Boiling the female genitalia in 10% NaOH may be necessary to
clear heavily sclerotized parts, but this method invariably distorts
and warps structures, swelling softer parts. It should only be used
as an additional method if there are many specimens.

Among the numerous difficulties of taxonomic research in the
family Araneidae (= Argiopidae) is the problem of matching males
and females. They are often collected separately and may be different
in appearance. Abalos recently (1963) observed, in his study of
reproductive behavior of spiders, that during the mating of many
species the tip of the male palpal embolus, presumably carrying sperm,
breaks off and remains in the female genital ducts. Though this had
been noticed before in black widows, it appears to be widespread in
theridiid spiders and orb weavers. The reason it has been overlooked
is that the majority of males collected are virgin, in search of a
female ; the tip of the embolus is therefore still attached in most males
preserved in collections. Because they die soon after mating, mated
males are rarely collected. On the other hand, probably the majority
of females collected have mated. In the genus ArgyopeJ now being
studied, and presumably in other argiopids, the tip of the male em-
bolus can readily be “fished out” of the epigynum by jiggling the
projecting parts with a needle. Among related species of the genus
Argyope , these tips differ strikingly in structure, athough the female
genitalia are quite similar. An illustration of a broken off tip was
supplied by Petrunkevitch (1930) with the hope of finding a male
to match it. These tips not only permit matching males with females,
but facilitate identification of females that have similar epigyna. Of
course, the assumption is made that the male chose a mate of his
own species. But the same method, applied to species in which the
female is as easily determined as the male, may provide some data on
the frequency of mating between species in nature.

I would like to thank Dr. A. M. Chickering, Mr. Jon Reiskind
and my wife for editing, and Miss Vida Kenk and Mr. F. Vuilleumier
for translating the summary. The investigations were supported by
Public Health Service Research Grant AI-01944.

Resumen

Preparaciones microscopicas permanentes de los palpos y epiginios
de aranas son desvantajosas ya que, al deshidratar las partes escle-
rozadas se tuercen, se pueden destruir al secarse, y a menudo la
preparacion es separada del resto del especimen guardado en alcohol
y se pierde. Es mejor guardar los genitales en tubos pequenos (micro-

156

Psyche

[June

vials) (Fig. i), con el especimen. En especies comunes con numerosos
ejemplares, las preparaciones pueden hacerse usando Hoyer o un medio
de montaje similar para Acari (Baker and Wharton).

Para ilustrar un palpo, a veces es necesario separarlo de la arana
a objeto de orientarlo mejor. En tal caso deberia examinarse comple-
tamente sumergido en etanol de 75 a 95%. A1 secarse las partes
blandas se danan. El palpo se fija en el fondo del recipiente con un
pedazo de papel fibroso. El examen se hace con un microscopio
binocular de diseccion entre 150 y 240 aumentos. Los palpos pequenos
y transparentes pueden estudiarse transladandolos a glicerina 0, si se
desea examinarlos con el microscopio compuesto o de fase, a un
montaje temporal en glicerina; pero el palpo debe siempre vol verse a
guardar en alcohol.

Para dilatar el palpo, se hierve por unos minutos en NaOH al
10%, entonces se transfiere a agua destilada, y luego se guarda en
alcohol. Sin embargo, los escleritos se tuercen y se danan cuando se
hierven en NaOH. Por el momenta yo uso acido lactico en vez de
NaOH.

No se debe quitar el epiginio de los ejemplares raros. En especi-
menes pequenos puede estudiarse sumergiendo todo el animal en
aceite de clavo. Dos incisiones (Fig. 2) permiten doblar la region
de los genitales hacia atras (Fig. 3) y, entonces, puede estudiarse
despues de orientara cuidadosamente (Fig. 4). Mas tarde la
“puertecita” puede volver a cerrarse. En las aranas pequenas, el cefa-
lotorax puede separarse a veces con el epiginio todavia fijo al pedicelo
(Fig. 5). Algunas veces, en ejemplares que no estan bien preservados,
los genitales se desprenden 0 es necesario separarlos para examinarlos
en detalle. En tal caso, el epiginio se guarda en un tubo pequeno

(Fig. 1).

Cuando se trata de preparaciones transitorias, el cefalotorax entero
y el epiginio (Fig. 5) pueden montarse en aceite de clavo; o, una
preparacion temporal del epiginio, para su examen bajo un micro-
scopio compuesto o de fase, puede hacerse con una gota de aceite de
clavo cubierta con un cubre-objeto. Para el estudio detallado de
especies comunes es necesario, algunas veces, saerificar un ejemplar
y separar con agujas los conductos genitales. Luego, los genitales
vuelven a guardarse en alcohol. Solo epiginios con estructuras muy
esclerozadas pueden hervirse en NaOH al 10%. Este metodo deforma
las estructuras e hincha las partes blandas.

Los machos y las hembras de Araneidae son a menudo muy distintos
en tamano y apariencia. Abalos (1963), obsei"\r6 que en muchas
especies el extremo del estilo del palpo del macho se desprende durante

1965]

Levi — Spider Genitalia

157

la copula. El extrema del estilo palpal puede separarse del epiginio
de la hembra para aparejar machos y hembras de la misma especie.
Este metodo se aplica con buen exito en el genero Argyope, en el cual
la mayor parte de los machos colectados estan todavia virgenes, pues
los machos mueren poco despues de la copula, y la mayor parte de las
hembras estan fecundadas. Por lo tanto, es muy importante ilustrar
cuidadosamente estos extremos de los estilos masculinos.

Resume frangais

Les preparations microscopiques permanentes sur lame de palpes
et epigynes d’araignees ne sont pas desirables. Les parties sclerifiees
se deforment par deshydratation et peuvent etre ecrasees par assecbe-
ment et il arrive souvent que la lame soit separee du reste du specimen
conserve en alcool et se perde. II est preferable de garder palpes et
epigynes dans de petits tubes (4 X 10 mm) avec le specimen (Fig.
1 ) . Pour les especes communes dont on a de nombreux exemplaires
on peut monter des lames en utilisant la solution de Hoyer ou bien
des solutions semblables pour Acari (Baker et Wharton).

II se pent qu’on doive separer un palpe de Taraignee si Ton veut
1’orienter correctement pour une illustration. Dans ce cas il faut
1’examiner completement immerge dans de l’ethanol 75 a 90%. Le
dessechement ablme les parties molles. O11 attache le palpe au fond
de la cuvette au moyen d’un morceau de papier fibreux. On procede
a Texamen avec une loupe binoculaire grossissant de 150 a 240 fois.
Les palpes de petite taille et transparents peuvent etre etudies apres
etre transvases dans de la glycerine ou bien apres montage te?nporaire
sur lame, en glycerine, pour examen au microscope optique ou au
microscope a contraste de phase. Le palpe doit cependant etre remis
en alcool pour conservation.

Pour etendre le palpe on le fait bouillir quelques minutes dans
NaOH 10%, puis on le transpose dans de l’eau distillee, et enfin on
le met dans Palcool pour conservation. Cependant 1’ebullition dans
NaOH deforme et endommage les sclerites. J’emploie maintenant
Facide lactique a la place de NaOH.

L’epigyne ne doit pas etre detache des specimens rares. Il peut
etre etudie sur les petits exemplaires en submergeant Faraignee entiere
dans de l’huile de girofle. Deux incisions (Fig. 2) permettent de
courber en arriere la region genitale (Fig. 3), qu’on peut etudier
apres l’avoir soigneusement mise en place (Fig. 4). Par la suite on
peut fermer la “porte” ainsi pratiquee. Chez les petites araignees on
peut parfois detacher le cephalothorax avec l’epigyne encore fixe au
pedicule (Fig. 5). Quelquefois Fepigyne de specimens mal conserves

158

Psyche

[June

se casse, ou doit etre enleve pour une etude ulterieure plus detaillee.
On conserve ensuite Pepigyne dans un petit tube (Fig. i).

Pour des preparations microscopiques temporaires sur lame on peut
monter le cephalothorax et Pepigyne entiers dans Phuile de girofle.
On peut aussi preparer une lame temporaire en recouvrant Pepigyne
avec une goutte d’huile de girofle; une telle preparation protegee par
une lamelle peut etre alors examinee au microscope optique ou a
contraste de phase. Pour les especes communes dont on fait une etude
detaillee il se peut que Ton doive sacrifier un specimen afin d’ecarter
les conduits des genitalia au moyen d’aiguilles. On remet ensuite les
genitalia en alcool pour conservation. On ne peut faire bouillir
Pepigyne dans NaOH 10% que si les structures sont fortement
sclerifiees. Cette methode abfme les structures et fait gonfler les
parties molles.

Abalos ( 1963) a observe que dans de nombreuses especes l’extremite
de Pembolus du male se casse pendant Paccouplement. L’extremite
de Pembolus palpal peut etre enleve de Pepigyne femelle pour assortir
males et femelles (Chez les Araneidae les males et les femelles sont
souvent tres differents de taille et d’aspect). Ceci se fait maintenant
avec succes pour le genre Argyope, chez lequel la plupart des males
recoltes son vierges, car les males meurent peu apres Paccouplement,
et la plupart des femelles sont accouplees. Par consequent il est
important d’illustrer ces extremites de Pembolus male.

References Cited

Abalos, J. W. and E. C. Baez

1963. On Spermatic Transmission in Spiders. Psyche 70: 197-207.
Baker, E. W., and G. W. Wharton

1952. An Introduction to Acarology. Macmillan, New York.

Levi, H. W.

1964. The Spider Genus Helvibis (Aradeae, Theridiidae) . Trans.

Amer. Microscop. Soc. 83: 133-143.

Petrunkevitch, A.

1930. The Spiders of Por.to Rico. Trans. Connecticut Acad. Arts Sci.

30: 159-355.

SCOLYTOIDEA (COLEOPTERA) 5: NOTES ON
NEOTROPICAL PLATYPODIDAE, MAINLY FROM
CENTRAL AMERICA

By Hans Reichardt*

The present paper, fifth in a series of contributions towards the
knowledge of Neotropical Platypodidae (see Reichardt, 1964b, for
previous papers), deals mainly with the Platypodidae I was able to
study in the Museum of Comparative Zoology, Cambridge, Mass.
In this collection I recognized a new species from British Honduras,
along with interesting new distributional records of some previously
known species. The new data are summarized in a map (fig. 9),
together with the records available in the literature. The result is
a fairly good summary of the distribution of the genus T esserocerus
in Central America. It is discussed below, under that genus. Platypus
pini Hopkins, previously known only from the type, is redescribed,
figured, and assigned to another species group within the genus.

One specimen of Tesserocranulus nevermanni Schedl, interesting
not only for the rareness of material of this genus but also for the
locality, was kindly loaned for study by its collector, Dr. A. Prosen,
Argentina, to whom I am greatly indebted. To Dr. O. L. Cart-
wright, United States National Museum, I am grateful for having
made the type of Platypus pini available for study. Finally I am
also indebted to Prof. P. J. Darlington, Jr., after whom I name the
new species described below, for his assistance during my studies at
Harvard University.

Genus Platypus Herbst, 1793-
“caudati” group.

Platypus pini Hopkins, 1905.

(figs. 1-2).

Platypus pini Hopkins, 1905, Proc. Ent. Soc. Wash., 7:71. Holotype, $,
Chaleo, Mexico, United States National Museum; examined.

Male: head with flat front, with big, areolate punctures; vertex
with scattered normal punctures; covered with some long hairs.
Antennae normal for the group. Prothorax with big, sparse punctures
over its surface; median sulcus well marked in the posterior third;
femoral grooves normally developed. Elytra punctate-striate, with
all striae formed by a single row of punctures; base forming a sharp

*Departamento de Zoologia, Secretaria da Agricultural Sao Paulo, Brazil;
presently at Harvard University. Manuscript received by the editor January

22, 1965

159

i6o

Psyche

[June

carina, running from suture to fifth interstice; interstices normally
convex; base of third slightly wider, with a few granules (3-4).
Interstices alternate on the last third of elytra; first one ending at
declivity, without formation of tooth or spine, but with some few
granules; third with long spine at declivity; fifth and seventh with
very much shorter spines, the fifth being the longest of the two; even
interstices and ninth ending like the first, with a few granules at
beginning of declivity. Lateral margin prolonged posteriorly to form
the outer projection of the declivity; between this and the suture,
another projection, triangular in shape. Abdomen normal.

Measurements: total length, 4.9 mm.; maximum (elytral) width,
1.54 mm.; elytral length, 3 mm.; pronotal length, 1.3 mm.; maxi-
mum pronotal width, 1.45 mm.

Examined material : 1 cT , collected 7 miles north of San Cristobal
Las Casas, Chiapas, Mexico (altitude: 2,100 m. ; Km. 1,152 of
Pan American Highway), 2. VIII. 1964, by J. Shetterly.

Hopkins did not assign his new species to any of the groups in
which the genus was divided by Chapuis; Strohmeyer (1912, Coleop-
terorum Catalogus, Pars 44), lists P. pini as a member of the
“ trispinati” group, and to this group it has been referred since. The
study of the type and the present specimen, however, showed that
P. pini is a member of the “caudati” group. By the presence of two
posterior projections on each elytron, this species is easily distinguished
from all the other members of the group. Both localities where P.
pini has been collected (Chaleo, Mexico — misspelled Chaleo by
Hopkins — and San Cristobal Las Casas, Chiapas) are situated at
or above 2,100 m., suggesting that the species may be restricted to
higher altitudes.

“ costellati” group
Platypus pulchellus Chapuis, 1865.

Platypus pulchellus Chapuis, 1865, Monogr. Platyp. :230, fig. 134. Holotype
$, San Juan Baptista (Villa Hermosa), Tabasco, Mexico; Blandford,
1896, Biol. Centr. Amer., Col., 4(6) :1 12. Allotype 9, El Zapote, Guate-
mala.

This species is widespread over Central America, and one record
is known from Venezuela (Schedl, 1960b). It has already been
recorded from several localities (summarized on fig. 9) in Mexico

Explanation of Plate 11

Fig. 1, Platypus pini Hopk., lateral view ; fig. 2, P. pini Hopk., dorsal
view; fig. 3, P. darlingtoni, n. sp., lateral view; fig. 4, P. darlingtoni, n. sp.,
dorsal view; fig. 5, T. nevermanni Schedl, lateral view; fig. 6, T. never-
manni Schedl, dorsal view.

Psyche, 1965

Vol. 72, Plate 11

Reichardt — Platypodidae

Psyche

[June

162

(Schedl, 1940; 1956; 1963), Guatemala (Allotype), Costa Rica
(Schedl, 1933).

New records from the Museum of Comparative Zoology collection
are: 1 Toledo District, British Honduras, 20. IX. 1906, Peck
col.; 2<J', 1 9, Lancetilla, Tela, Honduras, Stadelmann col.

Platypus darlingtoni, n. sp.

(figs. 3-4).

Male: head with front slightly concave in the middle, densely
rugose on all its surface; vertex as rugose as front; with few scattered
hairs. Antennae of the typical form of the “ costellati” group. Pro-
thorax densely and finely punctured all over its surface; femoral
grooves well developed ; median sulcus well marked in the posterior
third. Elytra: base raised to form sharp ridge, running from suture
to fifth interstice; interstices carinate in the posterior third of elytra;
sulci regularly and finely punctured from base to beginning of de-
clivity ; uneven interstices ending on the upper margin of the declivity
in blunt spines, the first and third being the longest and best developed,
subequal in length; the fifth, seventh and ninth less well developed,
also subequal in length ; the even interstices end on upper limit of
the declivity, without formation of blunt spines; posterior margin
of declivity with a well developed tooth on the latero-posterior angle;
declivity rugose and setose, with an additional tooth between the
third interstice and the posterior margin. Last abdominal sternite
with two well developed teeth. Anterior tibiae with 5-7 transverse
carinae.

Measurements: total length, 5.5 mm.; maximum width, 1.75

mm.; elytral length, 3.2 mm.; pronotal width, 1.4 mm.; pronotal
length, 1.5 mm.

Holotype, icT, from Toledo District, British Honduras, 20. IX.
1906, Peck col., in the Museum of Comparative Zoology (M.C.Z.
n. 3II47)-

Platypus darlingtoni, n. sp., is the second species of the “costellati”
group to be recorded from Central America. It is readily distin-
guished from the other species, mentioned before, which occurs at
the same locality, by its larger size (P. pulchellus Chapuis is barely
3 mm. long), and by its differently shaped and armed posterior
declivity.

Genus Tesserocerus Saunders, 1836.

This genus has a typical Neotropical pattern of distribution, with
its main dispersal center in South America. Of the 22 known species,

1965]

Reichardt - — Neotropical Platypodidae

63

only five occur in Central America (only two of them endemic).
It appears that the limits of the genus in the Mexican Nearctic-
Neotropical transition zone lie in the State of Vera Cruz on the
Atlantic side, and in the State of Nayarit on the Pacific side. How-
ever, the records are not continuous on the Pacific side: none is yet
known between the Tehuantepec Isthmus and Nayarit. Of the five
Central American species, three are represented in the Museum of
Comparative Zoology, as follows.

Tesserocerus belti Sharp, 1880.

Tesserocerus belti Sharp, 1880, Ent. Monthl. Mag., 17:112. Types, $ and
9, Chontales, Nicaragua.

Besides the type-locality, this species is known from several locali-
ties in Costa Rica (Schedl, 1933; 1941) and Quito Ecuador (Schedl,
1941 ) , as shown in fig. 9.

New records from the Museum of Comparative Zoology collec-
tion are: icf, i?, Esquinas, near Golfito, Dept. Puntarenas, Costa
Rica, 1948, P. & D. Allen col.

Tesserocerus ericeus Blandford, 1896.

Tesserocerus ericeus Blandford, 1896, Biol. Centr. Amer., Col., 4(6) : 1 1 5 - 1 1 6.
Types, $ and 9 , from several localities in Mexico, Guatemala and
Panama

In describing the female of this species, Blandford points to the
fact that it is almost indistinguishable from T. linearis Chapuis, a
species described from a single female collected in Spain, and which
Chapuis guessed had been imported from Brazil. In his revision of
Chapuis’ types of Platypodidae, Schedl (1960a) was unable to locate
the type of linearis. Its real identity remains to be cleared. It may
actually correspond to Blandford’s species, since the locality given
by Chapuis for linearis , Brazil, as said above, is based on pure guess.

T. ericeus is known only from Central America, from several
localities in Mexico (Schedl, 1940), Guatemala (Blandford, 1896),
Costa Rica (Schedl, 1933), and Panama (Blandford, 1896), as
summarized in fig. 9.

New records from the Museum of Comparative Zoology collection
are: icf, Lancetilla, Tela, Honduras, Stadelmann col.; i$, Barro
Colorado Island, Canal Zone, Panama, 1924.

Tesserocerus dewalquei Chapuis, 1865.

Tesserocerus dewalquei Chapuis, 1865, Monogr. Platyp. :3 00, fig. 184. Types,
$ and 9, Ega, Amazonas, Brazil.

Tesserocerus aubei Chapuis, 1865, l.c. :301, fig. 185. Types, $ and 9, Santa
Catarina, Brazil.

Tesserocerus dejeani Chapuis, 1865, l.c. :303, fig. 187. Types, $ and 9,
Colombia, Mexico.

Psyche, 1965

Vol. 72, Plate 12

0.5 mm

0.2 mm

Reichardt — Platypodidae

1965] Reichardt — Neotropical Platypodidae 165

T. dewalquei is widely spread over South and Central America.
Chapuis’ T. dejeant has been recognized as a junior synonym of
dewalquei by Schedl (1960a); the same author (1963), has subse-
quently used dejeani for Mexican specimens. Having seen specimens
from South and Central America, I believe these species must be
considered synonyms, as originally proposed by Schedl.

In Central America, T. dewalquei (recorded always as dejeani )
is known from several localities in Mexico (Blandford, 1896;
Schedl, 1933; 1940; 1952; 1960a; 1963), Guatemala and Nicaragua
(Blandford, 1896), Costa Rica (Blandford, 1896; Schedl, 1933),
and Panama (Blandford, 1896) ; as shown in fig. 9.

New records from the Museum of Comparative Zoolorgy collec-
tion are: 2cf , Barro Colorado Island, Canal Zone, Panama, 24. VII.
1924, W. M. Wheeler & Nathan Banks col.; 1 cT , Esquinas, near
Golfito, Dept. Puntarenas, Costa Rica, 1948, P. & D. Allen col.

Genus Tesserocranulus Schedl, 1933.

Very little is known about this genus. Only very few specimens
from wide apart localities have been studied; both sexes are known
only in T. nevermanni, type of the genus. It seems, however, basically
South American in distribution, with at least one of the three
species ( T. nevermanni Schedl) reaching into Central America.
T. ftexibilis Schedl, known only from females, was recently re-
described and illustrated (Reichardt, 1964a). Study of additional
material, especially of males, may necessitate removal of this species
from Tesserocranulus. The female, at least, has very peculiar anten-
nal structure and head shape. T. ornatus Schedl, based on one female,
from an unspecified locality in Brazil, is known only from the type.

Tesserocranulus nevermanni Schedl, 1933.

(figs. 5-8).

T esserocranulus nevermanni Schedl, 1933, Rev. Ent., 3:164-165. Holotype $,
Hamburgfarm, Costa Rica; 1935, Ent. Nachrbl., 9:151. Allotype $,
Cayenne.

Examined material: icf, Espejo, Santa Cruz, Bolivia, II. 1962,
A. F. Prosen col. (Prosen collection, n. 23440).

T. nevermanni , previously known only from the two original
localities, is now recorded from Bolivia. This record extends the
range of the species far to the south, and may indicate that never-

Explanation of Plate 12

Fig. 7, T. nevermanni Schedl, antenna ; fig. 8, T. nevermanni Schedl,
anterior leg; fig. 9, distribution of some species of Tesserocerus and Platypus
in Central America.

Psyche

[June

1 66

manni is distributed throughout the Amazonian Basin and adjacent
areas. The original description seems detailed enough to permit
identification of the species. I have not therefore prepared a new
description of the available specimen, which is illustrated. The
antenna (fig. 7), which is strikingly different from that of T. flexibilis
(see Reichardt, 1964a, fig. 5), shows an interesting small hairless
patch on the club, around the funicle insertion point. This character
has not yet been observed in any other genus of Platypodidae. It is
also absent in T. flexibilis.

References

Blandford, W.

1896. Platypodidae, in Biologia Centr. Amer., Col., 4(6): 97-144.
Reichardt, H.

1964a. Scolytoidea (Coleoptera) 2: Sobre Platypodidae neotropicais, com
descricab de um alotipo. Pap. Avuls. Dep. Zool., 16: 145-151, 10
figs.

1964b. Scolytoidea (Coleoptera) 4: Notas sobre a sistematica e biologia
de Platypodidae neotropicais. Rev. Bras. Ent., 11: 85-90, 12 figs.
SCHEDL, K. E.

1933. New Platypodidae from Central and South America, Rev. Ent.,
3 : 163-177.

1940. Fauna Mexicana, I. An. Esc. Nac. Cienc. Biol., 1: 317-376, 3 pis.

1941. Platypodidae und Scolytidae. Rev. franc. d’Ent., 7: 152-157.

1952. Neotropical Scolytoidea. V. Pan-Pac. Ent., 28: 122-124.

1956. Some bark- and ambrosia-beetles from the Tres Marias Island,
Mexico. 1. c., 32: 30-32, 1 fig.

1960a. Chapuis Platypodidae: eine Revision mit Ergaenzungen. Mem.
Inst. R. Sci. Nat. Belg., (Ser. 2) 62: 3-68, 1 fig.

1960b. Bark- and timber-beetles from the Neotropical region. Col. Bull.,
14: 74-80.

1963. Fauna Mexicana, II. Ent. Arb. Mus. Frey, 14: 156-167.

PICROCRYPTOIDES: A NEW GENUS OF THE
TRIBE MESOSTENINI FROM SOUTHERN SOUTH
AMERICA (HYMENOPTERA, ICHNEUMONIDAE)*

By Charles C. Porter
Biological Laboratories, Harvard University

The ichneumonid genus Trachysphyrus Haliday and its close rela-
tives within the huge tribe Mesostenini are found in almost every part
of the world. With one exception, however, the greatest number of
these species seems limited to the Holarctic region. This exception
is temperate and subtropical South America.

The ancestors of the present-day South American Trachysphyrus
fauna appear to have entered the southern continent from the north.
In tropical countries, such as Ecuador and Peru, they radiated only
at high altitudes in the Andes (approximately 8,000 to 14,000 feet)
and, to a lesser extent, in the coastal desert and foothills on the west.
None, so far as known, penetrated the tropical cloud-forest and rain-
forest at lower elevations to the east. In the temperate regions of
central and southern Argentina and Chile, on the contrary, this
generic group of north-temperate extraction found a more suitable
area in which to expand and produced a very complex radiation of
close to 100 species, probably more than occur in any area of similar
size and, certainly, in greater variety than in any other region which
the Trachysphyrus Group inhabits.

Among this Argentine and Chilean radiation are many species-
groups which depart more or less radically from the customary
definition of the genus Trachysphyrus. Some, although superficially
bizarre in aspect, are not clearly to be distinguished from the main
genus, to which they are often connected by species of intermediate
character. Much of the group is still evolving rapidly and generic
lines are not always clearly defined. Other series, however, present
a combination of unusual features which so isolates them from their
relatives that it would seem natural and convenient to place them in
separate genera. One such group, the new genus Picrocryptoides, is
diagnosed below together with a description of its two known species.

Picrocryptoides, new genus

Antenna: unusually short and stout: in 9 subtly enlarged and slightly
flattened below toward apex, first flagellomere about 3.2 to 3.8 times

* Manuscript received by the editor April 19, 1965

167

Psyche

[June

1 68

as long as wide apically, segments in apical half of flagellum only
slightly or not distinctly longer than wide; in cf also unusually stout
but considerably tapered toward apex, first flagellomere about 2.4
to 2.6 times as long as wide apically. Ocellar-area: very strongly
swollen and prominently raised above level of rest of vertex. Clypeus:
strongly assymmetric blunt-pyramidal in profile, the long basal face
gently sloping, the short apical face abruptly declivous; apical margin
almost truncate to weakly convex on median 2/3, more strongly
convex toward sides; corners not reflexed. Malar-space : unusually
long, in 9 about 1.5 to 1.8 times as long as basal width of mandible,
in cf about 1.2 to 1.5 times as long. Fore tibia: not at all inflated.
Pronotum : epomia produced above into a broad, conspicuous lappet-
like plate. Mesoscutum: notauli reaching about apical 2/3 to 3/4,
strong and rather deep throughout or shallower and reduced to a line
of large punctures for much of their length. Wing-venation: areolet
very large and broad, intercubiti parallel, slightly convergent or a
little differentially inclivous above; radial cell unusually short and
broad, ending distally at a point distinctly less than half the distance
from second intercubitus to tip of wing; nervulus interstitial to some-
what antef ureal; disco-cu'bitus only slightly angled near middle;
ramellus rather long to completely absent ; second recurrent a little
reclivous on lower 24 > straightened out or a bit curved or angled
above ; mediella straight except near base ; nervellus broken far below
middle, upper part more than twice as long as lower; axillus, over
most of its length, a little closer to sub-mediella than to hind-margin
of wing. Propodeum : in 9 rather elongate and gently curved in
profile without strong discontinuity between dorsal and apical faces,
a little bulging and sphereoid in shape; often with a weak rectangular
area-basalis and fine but distinct basal and apical trans-carinae, the
latter slightly broadened into low, elongate sublateral cristae, areo-
lation frequently much weaker or entirely absent; in cf longer and
lower in profile with the areolation only weakly suggested ; spiracle
long oval. First gastric segment: 9: a stout, triangular expansion
near base; ventro-lateral carina strong and sharp throughout; dorso-
lateral carina distinct, sometimes weaker toward base; dorsal carinae

Explanation of Plate 13

Figures 1, 4, and 6. Picrocryptoides hesperios n. sp. male. 1. Dorsal view
of head. 4. Propodeal profile. 6. Dorsal outline of postpetiole and apex of
petiole. Figures 2, 3, 5, 7, and 8. Picrocryptoides nvillinki n. sp. 2. Dorsal
view of head in male. 3. Propodeal profile of male. 5. Dorsal outline of
postpetiole and apex of petiole in male. 7. Anterior view of head of female.
8. Lateral view of apex of ovipositor. Scale: Figures 1 to 7, X 28 ; figure 8,
X 52.

Psyche, 1965

Vol. 72, Plate 13

L_l_

_L_

_L_J

©

Porter — Picrocryptoides

170

Psyche

[June

variable, usually traceable but not sharp toward apex of petiole and
on base of postpetiole; postpetiole broadly expanded; <$ : basal

expansion as in female or a little blunter; ventro-lateral carina be-
coming very weak on petiole; dorso-lateral carina not clearly defined;
dorsal carinae absent ; postpetiole moderately expanded. Second gastric
tergite: $: mostly covered with dense, strong adjacent to subadjacent
punctures emitting abundant short setae; lateral margin carinate and
somewhat reflexed, especially toward base; <$ : smooth and shining
with many well-separated medium-sized to large more or less super-
ficial punctures emitting long and conspicuous but not extensively
overlapping setae. Ovipositor : straight, stout, scarcely compressed;
dorsal valve without nodus or notch, straight in profile to near apex,
then gently decurved to tip; ventral valve considerably depressed,
especially toward apex, its ridges strong and approximately vertical;
sheathed portion about 0.28 to 0.30 times as long as fore wing.

TYPE species: Picrocryptoides willinki n. sp.

affinities: Picrocryptoides, because of its very large areolet, elon-
gate propodeal spiracle, straight mediella, and the broad postpetiole
of the female is a typical member of the Trachysphyrus Group of
the tribe Mesostenini. Within this category it is distinguishable from

Map 1. Distribution of Picrocryptoides.

1965]

Porter — Picrocryptoides

171

all other genera most conspicuously by the following combination of
characters: the strongly-raised, pyramidal, nose-like clypeus; the

inflated ocellar area; the broad, lappet-like dorsal prolongation of
the epomia; the unusually short radial cell and the strong basal expan-
sion of the petiole. Other characteristic features, restricted however to
the female, are the very long malar space, grossly punctate second
gastric tergite, and the short, stout ovipositor without nodus or notch.
Various other groups may have one or two of these characters but
none approaches having the whole series. Indeed, Picrocryptoides ,
although obviously a derivative of Trachysphyrus combines so many
distinctive features that it occupies an isolated position within its
generic group.

generic name: Picrocryptoides is derived from the Greek adjective
pikros or sharp and from cryptoides or Cryptus- like, in reference to
the pointed clypeus and to Cryptus , a generic name which has com-
monly been used for many species of the Trachysphyrus Group.
species: Two species, both new, are available in the material before
me. These are described below.

1. Picrocryptoides willinki n. sp.

Figures 2, 3, 5, 7, and 8 ; map 1.

types: Holotype: Cordoba, Argentina, Davis. In Museum of

Comparative Zoology, Cambridge, Massachusetts. MCZ. No. 31165.
Paratypes: (4 cf cf and 9$$) from Republica Argentina (Cordoba:
Cosquin, Sierra de Cordoba, March 1-9, 1920, Cornell University
Expedition; Dto. San Martin, January 22, I95°> D. Lopez; Yac.
Calamuchita, January 17, 195b, A. Willink; Argiiello, November 14,
1955, A. Giorgetta; Capital, December 1955, A. Giorgetta; San
Luis: San Martin, January 31 to February 3, 1958, Willink and
Tomsic; Santa Fe: Rosario, Davis; Buenos Aires: Felipe Sala,
January 1954, F. H. Walz; La Pampa: Macachia, January 1951,
F. H. Walz) ; in Museum of Comparative Zoology, Cambridge,
Massachusetts; Cornell University Collection, Ithaca, New York;
Townes Collection, Ann Arbor, Michigan; Instituto Lillo, Tucuman,
Republica Argentina.

female: Color: head, thorax, coxae, propodeum, and gaster dull
metallic blue-green with scattered obscure purple reflections; antenna
dark blackish-brown, becoming somewhat paler toward apex, especially
below; apex of mandible brownish-piceous ; trochanters shining-black,
narrowly brownish apically; fore- and mid-legs bright, pale orange,
tarsi brownish or blackish-stained; hind-femur bright pale orange,
hind tibia and tarsus moderately dark brown to blackish ; wings rather

172

Psyche

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dark, with metallic reflections. Length of fore-wing: about 7-3 to
9.0 mm. Temples: smooth and shining with moderately large, shallow
subadjacent punctures above, which become much sparser below; setae
prominent, considerably longer than the interspaces of the punctures
above; strongly, a little roundedly, receding behind eyes, about 0.40
to 0.50 times as long as eye in dorsal view. Occipital carina: rather
broad and flange-like above, narrower on temples, especially toward
junction with the narrowly flange-like hypostomal carina. Meso-
pleuron : speculum smooth and shining with a variable number of
large, deep punctures; surface otherwise almost uniformly and rather
finely and granularly reticulo-punctate, ridges separating punctures a
little more broad-crested and shining above, sharper below. Meta-
pleuron: uniformly a little more finely reticulo-punctate than

mesopleuron. Propodeum : surface rather finely and granularly

reticulo-punctate, often with a patch of much finer granulation
medially just back of area-basalis, usually more shining basad of
basal trans-carina, where punctures are more widely separated by
polished intervals. First gastric segment: postpetiole with strong

adjacent to subadjacent punctures laterally and subapically, a narrow
apical band smooth and polished and the central area between the
dorsal carinae smooth with some scattered large punctures and often
with fine longitudinal wrinkling, especially behind.
male: Color: much as in female; metallic sheen less strongly
greenish, more nearly shining black; antenna blackish, scarcely paler
toward apex. Length of fore-wing: about 7.0 to 8.6 mm. Antenna:
first flagellomere about 2.2 to 2.6 times as long as wide apically;
tyloides linear, on flagellomeres 9 to 17, sometimes last three absent,
none of tyloidiferous segments excavated at base. Malar Space:
about 1.3 to 1.5 times as long as basal width of mandible. Temples:
about 0.50 to 0.60 times as long as eye in dorsal view; strongly
receding, a little rounded-off. Meso- and metapleuron: punctures
a little more widely spaced than in female, polished interspaces
distinct throughout. Propodeum: elongate; dorsal face rather high,
gently sloping basally and then more strongly curved to meet the
more or less clearly discrete but not sharply differentiated apical
face; with strong, dense punctures like those of metapleuron, but
about basal /2 smooth and shining with only irregularly

scattered punctures; laterally, toward apex of dorsal face, and on
apical face with long, bushy, erect, dark setae. First gastric segment:
postpetiole moderately and gradually expanded, sides, in dorsal view,
gently and evenly curved or almost straight in outline from spiracle

1965]

Porter — Picrocryptoides

73

to apex, strongly convex above, smooth and shining with a few widely
scattered small, shallow punctures emitting long, conspicuous setae,
a few larger punctures grouped above spiracle.

remarks : There is considerable variation in the female with regard
to development of the propodeal carinae and in the sculpture of the
postpetiole. These characters are not themselves correlated nor are
they related to other possible distinguishing features. There is thus
no basis for recognizing more than one species in the material at
hand.

specific name : This species is named in honor of Dr. Abraham
Willink of the Institute Lillo of the Universidad Nacional de Tucu-
man, Tucuman, Argentina in thanks for his kind and expert help
during my visit to Argentina in November and December of 1964
and in recognition of his valuable series of monographs on the neo-
tropical Hymenoptera.

2. Picrocryptoides hesperios n. sp.

Figures 1, 4, and 6; map 1

types: Holotype: cf, Concepcion, Chile, 3/12/1908, P. Herbst.

In Museum of Comparative Zoology, Cambridge, Massachusetts.
MCZ No. 31166. Paratypes: (2$$): Concepcion, Chile, 1903
and 10/11/1908, P. Herbst. In Museum of Comparative Zoology.
male: Color: Head, thorax, coxae, propodeum, and gaster dull

metallic blue-green, becoming almost black on head and dorsum of
thorax; antenna dull blackish-brown; mandible tipped with brownish-
piceous; trochanters shining blackish-piceous, narrowly brownish on
apices; fore-femur brown, broadly marked with pale testaceous toward
apex; hind-femur bright pale orange, a little stained with brown at
base and apically above; tibiae and tarsi brownish-black, fore-tibia
broadly testaceous below; wings rather dark, with metallic reflections.
Length of fore-wing: about 7.4 to 7.6 mm. Antenna: first flagello-
mere about 2.4 times as long as wide apically; tyloides linear, on
flagellomeres 9 to 17. Malar Space: about 1.2 to 1.3 times as long
as basal width of mandible. Temples: rounded-off, scarcely receding,
about 0.76 to 0.78 times as long as eye in dorsal view. Propodeum :
lower and flatter in profile than in P. willink i, dorsal face more
steeply sloping and more smoothly merging with apical face; mostly
smooth and shining with some large adjacent or subadjacent punc-
tures laterally and on apical Y ; with prominent long, erect setae,
which are conspicuously sparser than in P. willinki, laterally and to-
ward apex. First gastric segment: postpetiole moderately expanded,

174

Psyche

[June

sides, in dorsal view, almost straight for more than half the distance
from spiracle to apex, then rather abruptly rounded-off; a little less
strongly convex above than in P. willinki.

remarks : This species is intimately related to P. willinki but differs
especially in the scarcely receding temples and in the much more
broadly polished and shining, more sparsely setose, lower and more
strongly sloping propodeum. It is possible that future collecting will
show that P. hesperios is a geographic race of the Argentine species,
particularly since there is a wide overlap of the Argentine and Chilean
insect faunas from about the latitude of Concepcion southward. On
the other hand, the distinguishing characters are of a type that
generally has specific value in the Trachysphyrus Group so that the
most reasonable course at present seems to be the recognition of two
distinct species.

specific name: Hesperios is a Greek adjective meaning western.

Resumen

El autor describe Picrocryptoid.es, un genero nuevo argentino y chileno
de la tribu Mesostenini de la familia Ichneumonidae ( Hymenoptera) .
Picrocryptoides es muy parecido a Trachysphyrus Haliday ( Cryptus
auctorum) pero se diferencia de Trachysphyrus por tener el clipeo
elevado en forma de una piramide asimetrica, la area ocellar elevada
y ensanchada, la epomia prolongada hacia arriba en forma de una
placa ancha, por tener la celda radial de la ala anterior inusitadamente
corta, y por la expansion triangular de la base del peciolo. Se describen
dos especies, ambas nuevas: P. willinki de Argentina y P. hesperios
de Chile.

STUDIES ON NORTH AMERICAN CARBONIFEROUS

INSECTS.

4. THE GENERA METROPATOR, EUBLEPTUS ,
HAPALOPTERA AND HADENTOMUM*

By F. M. Carpenter

Harvard University

The four genera treated in this paper, belonging to three different
orders, have only one feature in common : all have been very

difficult to interpret and to classify. Metropator , originally placed in
the Palaeodictyoptera by Handlirsch (1906a), has subsequently been
regarded as protorthopterous by some investigators and as mecopterous
by others; Eubleptus, also placed by its author in the Palaeodic-
tyoptera, has been made the type of a new order, Eubleptidodea, by
Laurentiaux (1953); Plapaloptera and Hadentomum, originally
designated by Handlirsch 1906a) as types of two new orders (Hapa-
lopteroidea and Hadentomoidea) , have subsequently been either
assigned to these orders or placed with uncertainty in the Protorthop-
tera. Unfortunately, all of these genera are known only by their
type-species, which are still represented solely by the unique type-
specimens. From my study of these fossils, I am convinced that the
species are not nearly so peculiar as has formerly been thought and
that to a large extent their puzzling nature is the result of Hand-
lirsch’s unsatisfactory figures and descriptions. I believe that Metrop-
ator was based on the hind wing of a species of the order Miomoptera,
that Eubleptus is very close to the family Spilapteridae of the order
Palaeodictyoptera, and that Hapaloptera and Hadentomum are near
relatives of other genera in the order Prothorthoptera. In the follow-
ing account I have first redescribed the fossils in the taxa to which
I consider them to belong and then have given the reasons for my
conclusions on their affinities.

I am deeply indebted to Dr. G. A. Cooper of the U. S. National
Museum for placing these type-specimens at my disposal on the several
occasions during the past ten years when I have found it necessary
to examine them. They have been studied under optimum conditions,
with various types of illumination and with the use of alcohol-glycerine
and with ammonium chloride, which has proven to be of the greatest

*This research is aided by Grant No. GB 2038 from the National Science
Foundation. The previous part in this series was published in Psyche,
71 : 117-124.

175

176

Psyche

[June

aid in working out venational details. I am also grateful to Dr.
Jarmila Kukalova, of Charles University in Prague, who studied
these fossils with me during her visit to Harvard University in 1964
and who prepared several of the drawings which are included in the
present paper.

Order Miomoptera martynov

This is an order of small insects, apparently related to the Pro-
torthoptera. The fore wings were membranous and the hind wings,
which lacked an expanded anal area, had the media arising from the
cubitus and had CuA and CuP anastomosed for their entire lengths,
forming a strong concave vein. The order is known from Upper
Carboniferous and Permian strata.

Family Metropatoridae Handlirsch

Metropatoridae Handlirsch, 1906, Proc. U.S.N.M., 29: 681
Metropatridae Martynova, 1962, Osnovy Paleont. :2861

Related to the Archaemiopteridae and Palaeomanteidae. Hind
wing nearly oval ; Sc short, weakly developed and close to R, as in
Palaeomanteidae; Rs forking before mid-wing, forming 6 terminal
branches; MA arising independently of CuA at the base of the wing
and forked almost to the level of origin of Rs; CuA + CuP with
short terminal fork. Fore wing and body unknown.

Genus Metropator Handlirsch

Handlirsch, 1906, Proc. U.S.N.M., 29: 682

Hind wing: R4+5 more deeply forked than R2 + 3 ; R3 with a
deep fork, R2 with a very shallow one; Mi +2 forked distally,
M3 + 4 forked twice. Type-species: Metropator pusillus Hand-

lirsch

The generic name Metropator is obviously derived from the identical
Greek word for “maternal grandfather”. The genitive of this is Metropa-
toros , providing the root Metropator- and, therefore, the family name
Metropatoridae.

In changing the name to Metropatridae, Dr. Martynova was apparently
misled by the normal Greek word for father (pater), which ordinarily has
the stem patr- ; pater, however, as used in the compound metropator, does
not follow pater in declension, although it means the same thing and is
merely a collateral form of that word. Since Handlirsch used the generic
name Metropator, there is no question about the root or the spelling of the
family name. I am indebted to Mr. Charles C. Porter for providing me
with this etymological information.

1965]

Carpenter — Carboniferous Insects

77

Metropator pusillus Handlirsch
Figure i

Handlirsch, 1906, Proc. U.S.N.M., 29: 682, fig. 8; 1906, Foss. Ins. :1 12, pi.

12, fig. 12.

Tillyard, 1926, Amer. Journ. Sci., 11: 161, fig. 19.

Martynova, 1962, Osnovy Paleont., Arthropoda: 286, fig. 892.

This species is based on a unique specimen (type no. 38731,
U.S.N.M.), consisting of an isolated wing, 7 mm. long and 3 mm.
wide. It was collected near the Altamont Colliery, anthracite region,
Pennsylvania (Namurian age). The preservation is fair; most of
the main veins are clear, but the basal part of the wing is missing.
Since this is one of about a dozen insects known from the lower part
of the Upper Carboniferous, the oldest strata in which unquestionable
insects have been found, its structure and affinities are of unusual
interest. Some diversity of opinion exists about both aspects of the
fossil. Handlirsch, who originally placed Metropator in the Palaeo-
dictyoptera, believed that the anterior margin of the wing was
broken away, the front edge of the wing as preserved being the sub-
costa; he apparently reached that conclusion because he was unable
to discern the subcosta as a submarginal vein. Tillyard in 1926,
following his examination of the type specimen, concluded that the
anterior margin of the wing was actually preserved and that Sc was
discernible as a distinct vein between Ri and the wing margin. In
his description he points out that the subcosta is very faintly indicated,
and that he could follow it out only with care by examining the
fossil in a good oblique light. He also described and figured the
cubito-median “Y-vein”, this being much more strongly developed
than most of the other veins of the wing. His conclusions were that
Metropator was a mecopteron, closely related to the Permopanorpidae.

I He did not discuss the detailed evidence for this, but simply asserted
that the mecopterous affinities could readily be seen at once from the
figures. His view of the position of Metropator has been generally
accepted subsequently, and it is the one presented in the Osnovy
Paleontologii (Martynova, 1962).

The drawing included in figure 1 represents my own interpretation
of this fossil and shows only those structures which I confidently
believe are present. From my studies I am convinced that Tillyard
i was correct in his conclusion that the costal margin of the wing is
actually present in the fossil, but I am also convinced that he was
I incorrect in his interpretation of the subcostal and cubital areas. The
subcosta is discernible near the base of the wing, as noted by Tillyard,

| but that is the entire length of the vein; it extends only a short

1 78

Psyche

[June

distance beyond the origin of Rs. The ammonium chloride prepara-
tion brings this vein out clearly enough so that it is visible in photo-
graphs. On the other hand, the supposed branch of the media, which
Tillyard showed as one arm of the cubital-median “Y-vein” and
which was an important factor in his conclusions on the affinities of
the fossil, cannot be seen — at least not by any techniques used by
me. The cubital vein itself (labelled Cui in Tillyard’s drawing) is,
as mentioned by Tillyard, a distinct one, which stands out more
strongly than any of the others excepting Ri. It is, however, clearly
concave in the fossil. This is important, since the supposedly homol-
ogous vein (CuA) in the mecopterous wings is strongly convex. I
am convinced, therefore, that the venation of Metropator only super-
ficially resembles that of the Mecoptera and that it does not have
the essential features of the mecopterous venation.

I believe the type-specimen of M. pusillus can much more readily
be interpreted as a hind wing of a miomopteron. In these wings the
subcosta is very short (see figure 2), Rs arises close to the base of
the wing, and CuA and CuP are completely coalesced, forming a
strong concave vein. These are the outstanding features of pusillus.
Unfortunately, since the base of the wing is missing in the type of
pusillus, the precise relationship between Cu and M cannot be deter-
mined ; however, there is no reason to assume that M does not join
Cu near the basal part of the wing. The venation of pusillus shows
more extensive branching than in the miomopteron illustrated in figure
2 \P alaeomantis minuta (Sellards)] but in other genera of Mio-
moptera (e.g., Stefanomioptera Guthorl and Permonika Kukalova)
the radial sector has more branches than in Palaeomantis. It seems
to me, therefore, that the available evidence, such as it is, indicates
that Metropator is more likely a miomopteron than a mecopteron.
The occurrence of several genera of Miomoptera in the Carboniferous
deposits of Europe supports this probability. The Mecoptera, on the
other hand, are otherwise unknown from beds earlier than the
Permian and since these are endopterygote (holometabolous) insects,
evidence for their presence in the lowest strata of the Upper Carbonif-
erous should be really convincing before such a conclusion is reached.
At present I believe the evidence points to a very different conclusion.

Order Palaeodictyoptera goldenberg
Family Eubleptidae Handlirsch

Handlirsch, 1906, Proc. U.S.N.M., 29: 679 (Order Palaeodictyoptera).
Laurentiaux, 1953, In Piveteau, Traite de Paleontologie. 3: 423 (Order
Eubleptidodea) .

1965]

Carpenter — Carboniferous Insects

179

Fore wing: subcosta extending at least nearly to the wing apex;
Rs with 4 terminal branches; MA forked, MP with at least 3
terminal branches; CuA with a short terminal fork; CuP more
extensively developed, with a deep fork shortly after its origin ;
several anal veins; cross veins distributed generally over the wing,
not arranged in rows ; anterior margin of the fore wing nearly straight,
at most very slightly concave. Hind wing: little-known; slightly
broader than fore wing. Body structure: prothoracic lobes present;
abdomen slender.

This family seems closely related to the Spilapteridae, from which
it differs in having a less developed Rs and CuA. Lack of knowledge

Figures 1 and 2. Miomoptera. Figure 1. Metropator pusillus Handlirsch.
Drawing of holotype, no. 38731, U.S.N.M. ; hind wing (original). Figure 2.
Palaeomantis minuta (Sellards), hind wing (original). Lower Permian,
Kansas. Lettering: Sc, subcosta; Rl, radius; Rs, radial sector; M, media;
CuA, anterior cubitus; CuP, posterior cubitus, 1A, first anal vein; +, convex
veins ; — , concave veins.

i8o

Psyche

[June

of the hind wings prevents more definite determination of the affinities,
but all available evidence indicates that this is a group which fits
readily within the Palaeodictyoptera ; eventually the family may turn
out to be inseparable from the Spilaptaridae.

Genus Eubleptus Handlirsch

Handlirsch, 1906, Proc. U.S.N.M., 29: 680

Fore wing: Rs arising slightly beyond mid-wing; M forked before
the origin of Rs, and Cu forked even nearer the wing base ; Rs forked
and each of its branches forked; iA simple, 2A forked. Type-species:
Eubleptus danielsi Handlirsch.

Eubleptus danielsi Handlirsch
Figure 3

Handlirsch, 1906, Proc. U.S.N.M., 29: 680

Length of fore wing, as preserved 13 mm.; estimated total length
17 or 18 mm.; width of fore wing, 4 mm.; maximum width of hind
wing (as preserved), 4.8 mm. Type no. 35576, U.S.N.M., collected
near Morris, vicinity of Mazon Creek, Illinois (Westphalian age).

This species was originally based by Handlirsch on a single speci-
men consisting of obverse and reverse ; the obverse specimen, according
to Handlirsch’s description, was contained in the Daniel’s collection
and the reverse in the U. S. National Museum. The counterpart in
the National Museum has been studied in connection with the present
account and is depicted in figure 3 ; the specimen in the Daniel’s
collection has not been found.

Handlirsch’s figure, which has been reproduced many times • in
subsequent publications and which has been the basis for all discus-
sions of the relationships of this fossil, was probably based to some
extent on the counterpart in the Daniel’s collection; at any rate the
position of the body in Handlirsch’s figure is the reverse of that in
the counterpart in the National Museum. The Daniel’s specimen
presumably showed parts of the cerci, which are entirely missing in
the National Museum fossil; also the Daniel’s specimen probably
showed a little more of the apical regions of the fore wings than
the reverse half. The venation in the National Museum fossil is
distinctly preserved and can be brought out even more clearly by the
use of ammonium chloride. As shown in figure 3, it is only slightly
different from that given in Handlirsch’s figure; there are some
differences in the positions of branches of the veins, but in general
the patterns are very similar. Handlirsch apparently did not observe
the basal connection between CuA and C11P, although this is clearly
distinguishable in the National Museum specimen. His figure of the

1965]

Carpenter — Carboniferous Insects

1 8 1

abdomen is about as I have observed it, although this seems somewhat
broader in the fossil than his drawing shows. As noted above, the
National Museum specimen does not include the end of the abdomen
and therefore lacks the cerci. Handlirsch’s representation of the meso-
and metothoracic segments is in agreement with mine; of course,
considerable distortion undoubtedly occurred in the fossil and only
the general form is indicated. The major difference between Hand-
lirsch’s figure and mine is in the structures which are anterior to the
mesothoracic segment. Handlirsch was of the opinion that two large
globular eyes could be distinguished, these being separated from the
mesothorax by a structure which he interpreted as the prothorax. His
figure in this area is slightly out of proportion ; the structures which
he shows as eyes are actually much closer to the mesothorax than
indicated in his drawing. Furthermore the structures themselves do
not have the regular, globular appearance which he depicts and they
do not give any indication of being compound eyes. On the other
hand, there are clearly visible radiating lines similar to those which
occur on the paranotal lobes of many Palaeodictyoptera. The location
of these structures and their details have convinced me that they are
in fact small paranotal lobes. Between them and the mesothorax is
a short segmented appendage, almost certainly a part of one of the
legs; this is shown also in Handlirsch’s figure.

|

Figure 3. Eubleptus danielsi Handlirsch. Drawing of holotype, no. 35576,
U.S.N.M. (original), p, paranotal lobes; other lettering as in figure 1.

182

Psyche

[June

The venation of the wings is actually typical of that of many
Palaeodictyoptera, especially that of some of the Spilapteridae. The
convexities and concavities of the veins, which are well preserved,
have been marked in figure 3 in the usual manner. Perhaps the most
distinct feature of the venation is the reduction of CuA to a single
vein having a marginal fork; in the Spilapteridae this vein tends to
be somewhat more extensively developed. There is a slight difference
between the right and left wings so far as CuP is concerned; in one,
C11P2 is forked but in the other it is unbranched. The anal veins are
slightly recurved, having the arched form occurring in many Palaeo-
dictyoptera. The venation of the hind wing is very little-known but
it appears to show no marked differences from the pattern in the fore
wing; however, the wing itself is obviously somewhat broader than
the fore wing.

Although the specimen of Eubleptus in the National Museum does
not, presumably, show as much of the apical region of the wings as
the counterpart in the Daniel’s collection, I think there is no question
that Handlirsch’s figure is incorrect in showing the wings as very
broadly and bluntly rounded. In that figure the left fore wing is
completely restored, the apex being represented by dotted lines; but
the drawing of the right wing shows an irregularity of the apex,
which suggests that this is not the actual margin of the wing itself.
In all probability, the apical region of the wing was shaped like that
of spilapterids.

Handlirsch originally described Eubleptus in the Eubleptidae, as
a palaeodictyopteron. However, his figure and description emphasized
several peculiar features which actually do not exist in the fossil
(such as the supposedly large eyes and the bluntly rounded wings).
As a result of this, various workers on fossil insects who have not
examined the type specimen have come to regard Eubleptus as a more
peculiar and aberrant insect than it actually is. Martynov, in 1938,
although placing the family Eubleptidae in the Palaeodictyoptera,
stated that it could well belong to a distinct order; and in 1953
Laurentiaux established the order Eubleptidodea for it. He failed to
indicate any characteristics by which he separated the order from the
Palaeodictyoptera, although he referred to the eyes and the absence
of lobes on the prothorax. In the Osnovy Paleontologii, Rohdendorf
placed the Eubleptidae in a separate order, which he termed the
Eubleptodea, presumably accepting Laurentiaux’s ordinal status for
the group although no reference is made to Laurentiaux’s publication
or to the change of spelling of the name.

However, in view of the structure of Eubleptus danlelsi, as it now
seems to be, there is no justification for the isolation or separation of

1965]

Carpenter — Carboniferous Insects

183

Eubleptus into a distinct order or even into a distinct suborder. It
is, in fact, difficult to find significant differences in the venational
patterns of the Spilapteridae and the Eubleptidae; ultimately these
two families may turn out to be synonomous. However, I have not
indicated such synonomy at this time since the name Eubleptidae
would have priority, and to synonomize Spilapteridae with Eubleptidae
seems inadvisable until the evidence for this is conclusive.

Order Protorthoptera handlirsch
Family Hapalopteridae Handlirsch

Handlirsch, 1906, Die fossilen Insekten: 304 (Order Hapalopteroidea ) .

Fore wing: similar to that of the Cacurgidae but having fewer
branches on the main veins and having CuP forking much further
from the wing base; CuPi not branched except for forking at wing
margin ; cuticular swellings apparently absent. Hind wing unknown.

Genus Hapaloptera Handlirsch

Handlirsch, 1906, Proc. U.S.N.M., 29: 694

Fore wing: Sc extending nearly to wing apex; costal veinlets un-
branched; Rs with four branches, MP forked to about mid-wing;
CuA with a terminal fork only; cross veins numerous, weakly formed.
Type-species: Hapaloptera gracilis Handlirsch.

Hapaloptera gracilis Handlirsch
Figure 4

Handlirsch, 1906, Proc. U.S.N.M., 29: 694

Fore wing: length 14 mm., width 4.5 mm.; membranous and
delicate; costal margin slightly concave, apex broadly rounded; R2
forked, R3, R4+5, MPi and MP2 unbranched; cross veins tending
to be irregular, but not branched or forming a network. The holo-
type specimen, no. 38731, U.S.N.M., was collected at Sharp Moun-
tain Gap, near Tremont, Pennsylvania (Stephanian age). The details
of the venation are shown in figure 4.

This fossil consists of a fore wing, very nearly complete, with
portions of a second wing. The venation is not distinct but use of
ammonium chloride brings out most details clearly. Handlirsch had
difficulty interpreting the venation, mainly because he failed to note
that actually two wings are superimposed ; his figure shows some
veins which are in reality on the second wing. The distal part of
the costal margin of the second wing can be clearly seen near the end
of Sc of the complete wing, and part of its hind margin appears in
the region of the end of MP. Handlirsch correctly recognized that

184

Psyche

[June

Figures 4 and 5. Protorthoptera. Figure 4. Hapaloptera gracilis Hand-
lirsch. Drawing of holotype, no. 38731, U.S.N.M. (original) Figure 5.
Heterologies langfordorum Carpenter. Drawing of holotype, Illinois State
Museum (original). Upper Carboniferous, Illinois.

something was amiss with the venation for he represents one vein by
a dotted line, which crosses over the basal part of another vein. With
the use of ammonium chloride the actual venation of the upper wing
becomes distinct and the pattern turns out to be very close to that of
Heterologus , from the Upper Carboniferous of the Francis Creek
Shales (Mazon Creek), Illinois (See figure 5). In Hapaloptera gra-
cilis, as in Heterologus , the stem of CuA (which is strongly convex)
is anastomosed with MP, but diverges away at about the level of
the origin of Rs and then anastomoses with the concave CuP, only
to separate again a short distance further. The main feature which
distinguishes Plapaloptera from Heterologus and other Cacurgidae is
the late forking of CuP and the absence of a long basal branch on
CuP 1. My first thought on examining the fossil was that the wing
membrane was extensively wrinkled but further study indicated that
the wrinkles are in most cases actual cross veins between the veins.
Although only a few cross veins are shown in Handlirsch’s figure, they
are almost uniformly distributed over the wing.

1965]

Carpenter — Carboniferous Insects

185

Handlirsch placed the family Hapalopteridae in a separate order,
Hapalopteroidea, although only one species, H. gracilis , was known
at the time. His decision to establish this “provisional” order was
undoubtedly the result of his misinterpretation of the venation of the
unique specimen on which gracilis was based. In 1922 he placed
another family, Emphylopteridae Handlirsch, in the order; this group
was based on another monospecific genus, Emphyloptera. Pruvost,
from the Upper Carboniferous of Europe. The assignment of this
genus to the Hapalopteridae obviously resulted once again from Hand-
lirsch’s misinterpretation of the venation of the type of Hapaloptera.
Quite clearly, Emphyloptera shows no affinities with Hapaloptera ,
as now understood, and it is here assigned to family Incertae Sedis,
order Protorthoptera, until the fossil on which it is based can be
studied further. The genus Ampeliptera Pruvost (1927) from the
Upper Carboniferous of Holland was placed in the Hapalopteroidea
by Pruvost but removed to another extinct order, Protocicadida, by
Haupt in 1941. The fossil on which Ampeliptera was based was
studied by Kukalova (1958), who found that it was an unquestion-
able protorthopteron of the family Paoliidae.

As to the genus Hapaloptera itself, there is nothing known about
it which eliminates it from the Protorthoptera. In fact, as noted
above, it is very close to the Cacurgidae. Bolton (i934) described
two species in the genus Hapaloptera from the Upper Carboniferous
of South Wales. Neither of these fossils, however, has affinities with
Hapaloptera , as can readily be seen from an examination of his
figures; both of the species are known only from fragments of wings
which, far from belonging to the same genus, represent at least
separate families and may represent even separate orders. The order
Hapalopteroidea is accordingly now placed in synonomy with the
order Protorthoptera.

Family Protoperlidae Brongniart

Brongniart, 1893, Recherches l’histoire insectes fossiles: 407 [ nom . correct.

Lameere, 1917, p. 197 (pro Protoperlida Brongniart, 1892)]

— Palaeocixiidae Handlirsch, 1919, Denkschr. Acad. Wiss. Wein, 92: 29

— Fayoliellidae Handlirsch, 1919, ibid: 48

— Hadentomidae Handlirsch, 1906, Die fossilen Insekten: 303 (Order

Hadentomoidea)

Fore wing: costal area with numerous, simple veinlets; Sc extend-
ing well beyond mid-wing ; R 1 unbranched ; Rs arising at least
slightly before mid-wing, unbranched; M forked; MP usually weaker
than MA ; CuA extensively branched ; CuP straight or nearly so, un-
branched ; cross veins well developed ; no reticulation but rarely two

i86

Psyche

[June

rows of cells in a few areas. Hind wings (known only in Protoperla) :
Rs arising nearer the wing base than in the front wing; CuA fused
with the very base of M ; anal area expanded to form a distinct lobe.

From a study of the type material of Protoperla Palaeocixius ,
Fayoliella and Hadentomum, I am convinced that these genera belong
to one family, for which the oldest name is Protoperlidae. This
family is now known from Upper Carboniferous deposits in Europe
and North America.

Genus Hadentomum Handlirsch

Handlirsch, 1906, Proc. U.S.N.M., 29: 693

Fore wing: similar to that of Palaeocixius but having a coarse
reticulation between Ri and Rs. Hind wing: incompletely known,
but probably with a small, distinct anal lobe, as in Protoperla. Type-
species: Hadentomum americanwn Handlirsch.

Hadento?num americanum Handlirsch
Figure 6

Handlirsch, 1906, Proc. U.S.N.M., 29: 693, fig. 19-21.

Length of fore wing, 23 mm. ; width of fore wing, 7.3 mm. Length
of hind wing, 23 mm. Type, no. 35579, U.S.N.M., collected near
Morris, Illinois (Westphalian age).

This species was based on a unique specimen consisting of the
obverse, in the Daniel’s collection, and the reverse in the U.S.
National Museum. Since the location of the Daniel’s collection is
unknown, I have been able to study only the specimen in the National
Museum. The preservation of this fossil is not very satisfactory; the
two wings on one side, as shown in Handlirsch’s figure, overlap in
such a way as to interfere with the determination of the venational
pattern. However, by tracing on photographs the veins of one wing
with ink of a certain color and the veins which are apparently not
related to that wing with ink of another color, I have found it possible
to work out the venational patterns of the two wings saisfactorily.
My interpretation of the wings is shown in figure 6. In most respects,
the figure of the fore wing agrees with that of Handlirsch. However,
the base of M, which Handlirsch shows fused with R, is distinctly
free and independent; also, the fork of Cu is clearly preserved in the
fossil, although it is not represented in Handlirsch’s figure. The origin
of R4 + 5 is not visible in the National Museum specimen; possibly
it was preserved in the Daniel’s specimen. The convexities and con-
cavities of the veins are clearly preserved and are marked in figure 6.
It will be noted that Rs is concave, the media is neutral ( ± ) , CuA

1965]

Carpenter — Carboniferous Insects

187

Figure 6. Hadentomum amcricanum Handlirsch. Drawing of fore wing
(A) and hind wing (B). Holotype, no. 35579, U.S.N.M. (original).

strongly convex and CuP concave. Since there is no indication of a
distinctly convex MA or concave MP, I have designated the media
here simply as “M”. It is possible that the vein that has been
designated R4 + 5 is actually MA, which may be fused basally with
the radial sector; however, there is no indication of the free basal
part of such a vein.

The anterior margin of the hind wing, which is not shown in
Handlirsch’s figure, can be made out without difficulty in the fossil
by the use of ammonium chloride. The most significant difference
between Handlirsch’s interpretation of the hind wing and mine is
in the nature of the hind margin of the wing. Handlirsch shows the
hind margin continuing to the base with the uniform curvature of
the apical region of the wing — that is without an anal lobe. This
is particularly important, since the absence of an anal lobe would
virtually eliminate the species from the Protorthoptera. However,
the National Museum specimen does not show the hind margin basally
of the termination of Cui ; it is clearly broken away at this point.
There is no reason, therefore, to assume that the anal lobe was absent,
and in view of the affinities of the fossil as indicated by the fore

1 88

Psyche

[June

wing, there is every reason to assume that the anal lobe was present.

The body is only faintly indicated in the specimen of americanum.
Handlirsch’s figure depicts the abdomen and the thorax as they seem
to me to be in the fossil, except that the prothorax is slightly shorter
and somewhat broader than he has drawn it. I see no indications of
the head as it was drawn by him; there are some irregularities in the
rock which may possibly represent part of the head but no definite
form can be made out and there are no suggestions of the eyes, so
far as I can observe, in the National Museum specimen.

Handlirsch established the order Hadentomoidea (1906, p. 692)
for this genus. He gave no definite diagnosis of the order, his account
of the group being essentially a description of the individual specimen
of americanum. However, it is clear from his discussion that he
placed much emphasis on the apparent similarity of the fore and
hind wings and on his conviction that the hind wing lacked an anal
lobe. H is conclusion was that the Hadentomoidea were probably
closely related to the Embioptera, although showing some affinities
with the Perlaria. I believe that his conclusions based on the apparent
absence of the anal area are not valid. The reconstruction of Haden-
tomum americanum, which Handlirsch included in his account of
fossil insects in Schroder’s Handbuch der Entomologie (fig. 73,
p. 143), is highly imaginary, since it shows the legs, antennae
and mouth parts, none of which are even suggested in the fossil.
The general effect of this figure, of course, is to increase the
bizarre appearance of the insect, as conceived by Handlirsch. As a
matter of fact, the fore wings of Palaeocixius and of Hadento?nu?n
show striking similarities, which I believe can only be explained by
close relationship of these genera, at least to the family level. (See
figures 6 and 7) The media seems somewhat more reduced in Haden-
tomum than in Palaeocixius, but the vein which is labelled R4 + 5 in
the accompanying figure of Hadentomum may actually be the anterior
branch of the media (i.e., corresponding to the vein labelled MA in
Palaeocixius) . In the orthopteroids and the Perlaria there is much
individual variation in the amount of fusion between branches of M
and parts of the radial sector. Unfortunately, we do not know the
hind wing of Palaeocixius or that of any of the other genera which
I am now placing in the family Protoperlidae, with the exception of
the type-species of Protoperla itself. The latter genus is based upon
a single species known from the hind wing, which shows a definite
anal lobe, although the lobe is not as large as in most of the Protor-
thoptera. The venation of the Protoperla hind wing is difficult to
interpret on the basis of the single specimen known. The media is

1965]

Carpenter — Carboniferous Insects

Sc

Rs

Figure 7. Palaeocixius antiquues Handlirsch. Drawing of holotype, in
Laboratoire de Paleontologie, Paris, (original). Upper Carboniferous of
Commentry, France.

apparently coalesced at least in part with the radius or the radial
sector, giving the impression that the radial sector occupies a very
large area of the wing surface.

At the present time I believe that all the evidence at hand indi-
cates that Hadentomum is a member of the family Protoperlidae, as
here conceived. In any event, there is no evidence at hand to justify
the retention of the order Hadentomoidea, which is here placed in
synonomy with the Protorthoptera.

References

Bolton, H.

1934. New forms from insect fauna of the British Coal Measures.
Quart. Journ. Geol. Soc. 90: 278-301.

Brongniart, C.

1893. Recherches pour servi a l’histoire des insectes fossiles des temps
primaires. These Fac. Sci. Paris., No. 821: 1-494.

Handlirsch, A.

1906a Revision of American Palaeozoic insects. Proc. U.S.N.M., 29: 661-
820.

1906b Die fossilen Insekten. Leipzig.

1919. Revision der Palaozoischen Insekten. Denkschr. Acad. Wiss. Wien,
Math.-Naturwiss. Class, 96: 1-82.

1920. Palaontologie, in Schroder’s Handb. Ent. 3: 117-306.

1922. Fossilium catalogus (1): Animalia: Insecta palaeozoica. 16: 1-
230.

Haupt, H.

1941. Die alstesten geflugeten Insekten und ibre Beziehungen zur Fauna
der Jetztzert. Zeit. fur Natur. Halle, 94: 60-121.

Kukalova, J.

1958. On the systematic position of Ameliptera limburgica Pruvost,
1927. Vestnik U.U.G. 33: 377-379.

190

Psyche

[June

Lameere, A.

1917. Revision sommaire des insectes fossiles du Stephanien de Com-
mentry. Mus. Nat. Hist. Natur., Bull., 23: 141-201.

Laurentiaux, D.

1953. Insects, in Piveteau, Traite de Paleontologie, 3: 397-527.
Martynov, A. V.

1938. Etudes sur l’histoire geologique et de phylogenie des ordres des
insectes. Trav. Inst. Paleont. 7(4) : 1-148.

Martynova, O.

1962. Osnovy Paleontologii, Mecoptera, pp. 283-294.

Pruvost, P.

1927. Sur une aile d’insecte fossile trouvee au sondage de Gulpen.
Geol. Bur. Neded. Mijn. Heerlen: 76-77.

Rohdendorf, B. B.

1962. Osnovy Paleontologii, Eubleptodea, p. 154.

Tillyard, R. J.

1926. Kansas Permian Insects. Part 7. The order Mecoptera. Araer.
Jour. Sci. 11: 133-164.

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PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol. 72

September, 1965

No. 3

CONTENTS

A Redefinition of Ischyropalpus , and Six New Species (Coleoptera:
Anthicidae). Floyd G. Werner 191

Notes on Neotropical Tabanidae VI. A New Species of Lepiselaga Macq.
with Remarks on Related Genera. G. B. Fairchild 210

Self-Burying Behavior in the Genus Sicarius (Araneae, Sicariidae).
Jonathan Reis kind 218

A Second African Species of the Dacetine Ant Genus Codiomyrmex.

Robert W . Taylor 225

j Five New Species of the Genus T marus (Araneae, Thomisidae) from

the West Indies. Arthur M. Chickering 229

| A Key to the North American Statira (Coleoptera: Lagriidae).

Carl T. Parsons 241

i

l The Genus N eohermes (Megaloptera : Corydalidae) .

Oliver S. Flint, Jr 225

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1965-66

President H. Reichardt, Harvard University

Vice-President S. Vogel, Harvard University

Secretary C. C. Porter, Plarvard University

Treasurer F. M. Carpenter, Harvard University

Executive Committee W. L. Brown, Jr., Cornell University

J. Reiskind, Harvard University

EDITORIAL BOARD OF PSYCHE

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Agassiz Professor of Zoology , Harvard University

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University; Associate in Entomology , Museum of Comparative
Zoology

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parative Zoology

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PSYCHE

Vol. 72 September, 1965 No. 3

A REDEFINITION OF ISCHYROPALPUS , AND
SIX NEW SPECIES
(COLEOPTERA: ANTHICIDAE)1

By Floyd G. Werner

Department of Entomology, University of Arizona, Tucson

The principal features that have been used to distinguish Lappus
Casey from Ischyropalpus LaFerte are the generally smaller size, less
densely sculptured forebody and lack of sharp margins on head and
pronotum of the former. It has been evident for some time that the
division is somewhat arbitrary. Some species are readily assignable to
each genus; others don’t fit in either very well. One is faced with
the choice of establishing a series of poorly defined genera to accom-
modate the known species or of placing all of them in one. I believe
that the second alternative is the better one.

Genus Ischyropalpus LaFerte

Anthicus ( Ischyropalpus ) LaFerte 1848, 106, 141, 314 (Type-species:
Anthicus sericans Erichson 1834, designated by LaFerte on p. 106).
Pic 1911, 30. Blackwelder 1945, 433.

Ischyropalpus : Casey 1895, 656. Bonadona 1961, 150 (designates
Anthicus perplexus LaFerte 1848 as type-species; designation invalid
because of the original designation of sericans by LaFerte 1848.)
Lappus Casey 1895, 656 (Type-species Anthicus nitidulus LeConte

1851 NEW DESIGNATION — -NEW SYNONYMY.

Anthicus (Lappus): Pic 1911, 30. Leng 1920, 163. Blackwelder

1945, 433-

As here redefined, Ischyropalpus is a very isolated genus in the
family, with a number of features that are distinctive. The first of
these is the form of the mesosternum, as shown in Fig. 1. This
structure is slightly expanded, but not as greatly as in Acanthinus,
BauliuSj Euvacusus, Formicilla and Vacusus in the New World
fauna and some other groups in the Old World. The form of the
hind angles is unique. The sides extend obliquely laterally and
posteriorly to the level of the edges of the ventral portion of the
mesepisterna, meeting the hind margin abruptly and at an acute
angle well lateral to the middle coxae. The mesepisterna and mesepi-

1This study was supported by National Science Foundation Grant GB-427.
Agricultural Experiment Sta., Dept. Entomology, Journ. Art. (p 1068.

192

Psyche

[September

mera are not visible behind the hind margins of the mesosternum.
The sides of the mesosternum are flattened and meet the adjacent
flattened portion of the mesepisterna at a slight angle, only a fine
groove separating them. There is no fringe of setae along this groove.

Another feature that appears to be unique in the family is the
presence of a zone of tiny spines on the bases of the tibial spurs,
encircling the spurs. These are present in all of the species of Ischyro-
palpus examined in slide preparations. All other Anthicidae examined
have the spurs without spines or have the spines arranged in two
longitudinal rows on the surface nearest the tarsus.

Some other external features are apparently quite stable, although
not limited to the genus. The tactile setae are short and inconspicuous
and the elytral punctures are not arranged in rows. The genus
Anthicus , s. str., is similar in both these features, although some
species have both tactile and normal setae long.

The cf aedeagus is distinctive. The tegmen has the parameres
separate and sometimes very complex; these are asymmetrical in some
species. The apodeme (basal piece) is unusually broad, especially near
the attachment of the parameres. The penis is relatively simple, but
has an unusually long apodeme, which may extend beyond the end
of the apodeme of the tegmen in repose. The penis probably slides
freely in the tegmen, because it is sometimes found far extended in
dissected specimens. The internal sac is extremely complex and com-
pletely unlike this structure in any other genus examined. The entire
apical area has transverse ridges or thickenings and there is a pair of
plates about half-way along the sac. These have not been seen in any
other genus. The plates are absent in the known males of the Eryngii-
Group described below, but present in all other members of the
genus examined.

The Eryngii-Group also is different in the external feature most
easily seen in the rest of the members of the genus. This is the shape
of the last segment of the maxillary palpi, which is in the form of a
roughly isosceles triangle with the apex at the point of attachment.
Many species have this segment extremely broad. But a few, notably
I. sericeus Bonadona, have the sides of the triangle unequal and the
apex rather narrow. The members of the Eryngii-Group have the
segment narrowly securiform.

The typical species, not including the Eryngii-Group, have a rather
distinctive facies. The prothorax is quite deeply constricted laterally
in the basal third, with the sides rounded from there to the collar.
The disc is not grooved at the level of the constriction. The large
species that made up most of the genus Ischyropalpus as previously

1965]

Werner — Ischyropalpus

193

defined have the head and pronotum deeply sculptured, with the disc
somewhat flattened and the base of the head and sides of the pronotum
anterior to the constriction ending in a sharp margin that is lentic-
ular in profile.

Uniting Lappus with Ischyropalpus necessitates a rather large
number of recombinations. To this number are added some new
generic assignments and synonymy that have resulted from an examina-
tion of the types. Bonadona has recently treated most of the species
assignable to Ischyropalpus as previously defined (1961), and has
made the changes necessary for these species. The new combinations,
new synonymies, and the previous generic assignments of the species
involved are given below, followed by a list of the species known to
me, incorporating these changes.

The following are new combinations: i. Species described in
Anthicus: amplithorax (Pic), antic ef as ciatus (Pic), asphaltinus

(Champion) (= Anthicus distinctus Pic, Lappus thicaniformis
Werner, both new synonymy), aterrimus (Champion), baeri (Pic),
dilatipes (Champion), externenotatus (Pic), impressus (LaFerte),
laticeps (LaFerte), nigrofemoratus (Fairmaire and Germain), paral-
lels (Fairmaire and Germain), subtilissimus (Pic) (substitute name
for Lappus subtilis Casey, a junior secondary homonym in Anthicus) .
2. Species described in Anthicus but transferred to Lappus by Casey,
1895: bactrianus (Champion) (listed by Casey as backianus , a

typographical error), nitidulus (LeConte) (= Lappus alacer Casey,
asperulus Casey, canonicus Casey, cursor Casey, nubilatus Casey,
vigilans Casey, all new synonymy), obscurus (LaFerte), occiden-
talis (Champion), punctipennis (Champion), sturmi (LaFerte)
(substitute name for elegans LaFerte, a junior primary homonym in
Anthicus) , tumidicollis (Champion). 3. Species described in Anthicus
but transferred to Anthicus (Lappus) by Krekich, 1914: nitidus
(Boheman) ( — Anthicus atomarius Boheman). 4. Species described
in Anthicus (Ischyropalpus) : batesi (Pic), hemicyclius (Heberdey).
5. Species described in Anthicus (Lappus) : dormei (Pic), goyasensis
(Pic), postobscurus (Pic), proprius (Pic), sipolisi (Pic), vagema-
culatus (Heberdey). 6. Species described in Lappus: lividus (Casey)
(— Lappus bipartitus Casey, ornatellus Casey, solivagans Casey,
vividus Casey, all new synonymy), pinalicus (Casey), turgidicollis
(Casey) ( — Lappus animatus Casey, new synonymy).

Anthicus trigonocephalus LaFerte 1848, described in the subgenus
Ischyropalpus , is not included here. This species was described from
India on the basis of a specimen lacking maxillary palpi. LaFerte
placed it in a group separate from the South American species on the

194

Psyche

[September

basis of head shape. I have not seen a specimen of this species but
have never seen a member of the genus from the Old World. Pic
(1911) and Bonadona (1961) did not include it as a member of
Ischyropalpus and there is no reason to believe that it should be
retained in this genus.

Anthicus gibbithorax Pic, transferred to Lappus by Casey (1895)
from the description, does not belong to this genus. It was described
as from Texas but I have had an opportunity to examine the type.
It is undobutedly an erroneously labelled Old World species, and is
not assignable to Ischyropalpus.

List of Species — Group Names

[Names believed valid are in bold-face type]

adstrictus Krekich 1914 — Peru.
alacer (Casey) — see nitidulus.

albifasciatus (Pic) 1897 — probably South America.

var. argentinus (Pic) 1904 — Argentina,
alvarengai, new species — Brazil.
amplicollis (Boheman) — see sericans.
amplithorax (Pic) 1895 — Brazil.
animatus (Casey) — see turgidicollis.
anticefasciatus (Pic) 1910 — Argentina.
argentinus (Pic) — see albifasciatus.
asperulus (Casey) — see nitidulus.
asphaltinus (Champion) 1890 — - Mexico, Guatemala.
distinctus (Pic) 1897 — Mexico.
thicaniformis (Werner) 1956 — Mexico,
aterrimus (Champion) 1890 — Guatemala.
atomarius (Boheman) — see nitidus.
attenuatus Krekich 1913 — Argentina.
backianus (Casey) — see bactrianus.
bactrianus (Champion) 1890 — Mexico, Guatemala.

backianus (Casey) 1895 — typographical error,
baeri (Pic) 1902 — Peru,
batesi (Pic) 1914 — Colombia.
bipartitus (Casey) — see lividus.
boliviensis (Pic) 1909 — Bolivia.
bruchi (Pic) — see caesiosignatus.

caesiosignatus (Boheman) 1858 — probably Ecuador.
bruchi (Pic) 1904 — Argentina,
var. mendozanus (Pic) 1914 — Argentina.

1965]

W erner — Ischyropalpus

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canonicus (Casey) — see nitidulus.
catamarcanus (Pic) — see testaceoguttatus.
cursor (Casey) — see nitidulus.
curtisi (Solier) 1851 — Chile.

ssp. wittmeri Bonadona 1961 — Chile,
decoratus, new species — Brazil,
dentipes Bonadona 1961 — Argentina,
dilatipes (Champion) 1890 — Mexico.
distinctus (Pic) — see asphaltinus.
dormei (Pic) 1933 — Brazil.
elegans (LaFerte) — see sturmi.
eryngii, new species — Uruguay.
ester oensis (Pic) — see testaceoguttatus.
externenotatus (Pic) 1913 — Argentina,
freyi, new species — Venezuela,
goyasensis (Pic) 1904 — Brazil,
hemicyclius (Heberdey) 1937 — Costa Rica,
impressus (LaFerte) 1848 — Colombia,
interamnis (LaFerte) 1848 — Brazil,
jatahyensis (Pic) 1899 — Brazil,
jenseni Krekich 1913 — Argentina,
latereductus (Pic) 1913 — Chile,
laticeps (LaFerte) 1848 — Venezuela,
lividus (Casey) 1895 — U. S. A.
bipartitus (Casey) 1895 — U. S. A.
ornatellus (Casey) 1895 — U. S. A.
solivagans (Casey) 1895 — U. S. A.
vividus (Casey) 1895 — U. S. A.
maculosus (Fairmaire and Germain) i860 — Chile,
mapirianus (Pic) 1909 — Bolivia.

var. semirufescens (Pic) 1909 — Bolivia,
var. testaceitarsis (Pic) 1913 — Mexico.
mendozanus (Pic) — see caesiosignatus.
nigrofemoratus (Fairmaire and Germain) i860 — Chile,
nitidulus (LeConte) 1851 — U. S. A.
alacer (Casev) 1895 — U. S. A.
asperulus (Casey) 1895 — U. S. A.
canonicus (Casey) 1895 — U. S. A.
cursor (Casey) 1895 — U. S. A.
nubilatus (Casey) 1895 — - U. S. A.
vigilans (Casey) 1895 — U. S. A.
nitidus (Boheman) 1858 — probably Ecuador.

196

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atomarius (Boheman) 1858 — probably Ecuador.
nubilatus (Casey) — see nitidulus.
obscurus (LaFerte) 1848 — U. S. A.
occidentalis (Champion) 1890 — Mexico.
ornatellus (Casey) — see lividus.
parallelus (Solier) 1851 — Chile,
patagonicus (Pic) 1902 — Argentina,
perplexus (LaFerte) 1848 — Colombia,
pinalicus (Casey) 1895 — U. S. A.
planicollis (Fairmaire and Germain) — see sericans.
postobscurus (Pic) 1904 — Brazil,
propriiis (Pic) 1904 — Peru.

punctipennis (Champion) 1890 — Mexico to Venezuela,
puteifer, new species — Argentina,
quadriplagiatus (LaFerte) 1848 — Colombia.
semimfescens (Pic) — see mapirianus.
sericans (Erichson) 1834 — South America.

ajnplicollis (Boheman) 1858 — probably Ecuador.
planicollis (Fairmaire and Germain) i860 — Chile,
van trireductus (Pic) 1928 — Argentina,
sericeus Bonadona 1961 — Argentina,
sipolisi (Pic) 1933 — Brazil.
solivagans (Casey) — see lividus.

stur mi (LaFerte) 1848 — U. S. A. (substitute name).

elegans (LaFerte) 1848 — U. S. A.
subtilis (Casey) — see subtilissimus.
subtilissimus (Pic) 1896 — U. S. A. (substitute name).

subtilis (Casey) 1895 — U. S. A.
tenax (Pic) 1904 — Brazil.

var. tenuiculus (Pic) 1904 — Brazil.
tenuiculus (Pic) — see tenax.
testaceitarsis (Pic) — see mapirianus.

testaceoguttatus (Fairmaire and Germain) 1863 — Chile.
catamarcanus (Pic) 1926' — Argentina,
var. esteroensis (Pic) 1928 — Argentina.
thicaniformis (Werner) — see asphaltinus.
tibialis, new species — Venezuela,
titschacki (Pic) 1944 — Peru.
trireductus (Pic) — see sericans.
tumidicollis (Champion) 1890 — Panama,
turgidicollis (Casey) 1895 — U. S. A.
animatus (Casey) 1895 — U. S. A.

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vagemaculatus (Heberdey) 1938 — Brazil,
venezuelensis (Pic) 1900 — Venezuela.
vigilans (Casey) see nitidulus.
vividus (Casey) — see lividus.
wittmeri Bonadona — see curtisi.

Description of New Species
Ischyropalpus tibialis, new species
(Fig. 2)

2.48 - 3.02 mm, dark piceous, elytra slightly paler, legs and an-
tennae brown. Head and pronotum sparsely and finely punctured,
not sharply margined. Pubescence in weak postbasal transverse im-
pression of elytra not oblique or transverse. Hind tibiae of cf with
a tuft of long setae medially near apex.

Holotype cf : Length 2.82 mm. Head 0.49 mm long to the
distinct frontoclypeal suture, 0.58 wide across eyes, 0.49 just behind,
the base short and evenly rounded behind eyes, a line connecting the
hind margins of the eyes 0.17 mm from middle. Surface smooth and
shiny, the punctures small and 0.03 to 0.04 mm from center to center,
the intervals not curving down to them. Disc evenly curved, not
very convex, the clypeus on almost the same curve. Pubescence sparse
and inconspicuous, decumbent, almost appressed, 0.06 mm long,
slightly curved ; tactile setae erect, straight, fine, 0.08 mm. Last
segment of maxillary palpi 0.20 mm long on outer side, 0.15 on
inner, 0.17 across apex, the apical angles well-defined. Eyes large
and moderately prominent, 0.26 X 0.19 mm, separated by 0.41,
finely faceted, with curved setae ca. 0.0 1 mm long at intervals in
posterior half. Antennae with segment 1 slightly swollen in middle
and feebly pedunculate, segments 2-10 widest near apex and becoming
more triangular, 11 truncate at base, widest near middle, the sides
then curving to a blunt point. Measurements in 0.01 mm: 17/8,
10/5, 13/5, 12/5, 12/6, 12/6, 12/6, 12/7, 12/8, 10/8, 18/8.

Pronotum 0.64 mm long, 0.38 wide at base, 0.3 1 at the well-
! defined constriction 0.23 111m from base, 0.54 maximum at 0.44 mm
j from base, 0.23 at the short but well-defined collar. Basal impressed
line distinct. Sides almost evenly rounded from constriction to collar
| and disc evenly convex from basal impressed line to collar. Surface
as on head, the punctures slightly deeper and sparser. Elytra 1.69
mm long, 0.74 wide at the well-marked humeri, 0.90 maximum just
before middle, subparallel. Omoplates feebly elevated; postbasal
transverse impression feeble, with punctures no denser or deeper than

198

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on rest of surface, with setae that are directed backward. Suture
elevated in apical half. Surface shiny and smooth, with some very
sparse and fine punctulation and small punctures ca. 0.05 mm from
center to center, the intervals not curved down to punctures. Pubes-
cence decumbent, almost appressed, sparse, 0.10 mm long, incon-
spicuous; tactile setae suberect to erect, 0.08 mm.

Sternum 7 flattened, with an inconspicuous median rounded zone
of more erect, curved setae, all of which are directed backward.
Sternum 8 flattened, its apical margin very shallowly concave.
Tergum 8 with sides gradually convergent to apex, which is more
abruptly convergent with a short median point; 0.33 mm wide at
base of exposed portion. Femora gradually thickened to about apical
fourth. Front femur 0.54 X 0.18 mm, front tibia 0.42 X 0.07,
hind femur 0.64 X 0.20. All trochanters with a small tuft of setae
at outer apical portion. Front and middle tarsi slightly expanded,
with a denser pad than in ?. Hind tibiae flattened and slightly bowed
on surface facing tibia, with a conspicuous tuft of setae in apical
fourth of median side, the individual setae suberect for 0.08 mm,
then quite sharply curved toward apex, then almost straight for ca.
0.05 mm.

The combination of sparsely and finely punctured head, pronotum
and postbasal transverse impression of elytra, lack of oblique setae
in the impression, and presence of a tuft of long setae near the apex
of the hind tibiae in the cf , appears to be unique in the genus. Some
of the males have the tuft of setae on the hind tibiae shorter and
barely curved. Females have the legs unmodified.

Distribution: Known from northern Venezuela and the island of
Trinidad. Holotype cf : Maracay, Est. Aragua, VENEZUELA, 29
May 1942, en Astromelia (U. Venezuela — Maracay). Paratypes:
44 cf , 46 ?, same data (U. Ven. — Maracay and Werner). 4 cf,
9?> Maracay, 27 Jan. 1943, R. Araque, en flores Persea gratissima
(U. Ven. — Maracay and Werner). 2 cf, 3$, El Valle, [Est.
Aragua], Venezuela, 24 Sept. 1938, on Mimosa arenosa Poir. ; 3 9,
29 Mar. 1938, on Cordia cylindrostachya; 2 9, 14 Aug. 1939, on
Amaranthus duhius ; all C. H. Ballou (U. S. N. M.). 1 cf , La
Victoria, Est. Aragua, 26 Aug. i960, C. Bordon (Bordon). 1 cf,
San Antonio de los Altos, Est. Miranda, Venezuela, 30 Aug. 1959;
1 cf , 1300 m, 6 Sept. I959j both C. Bordon (Bordon). 1 9, Los
Angelinos (Los Teques), Venezuela, 27 Mar. 1959, C. Bordon
(Bordon). 3 cf , 5 9, Umg. Caracas, Venezuela, Jan. 1954, G. u. H.
Frey (Mus. G. Frey, Tutzing, Germany). 3 9, Sta. Lucia, Vene-
zuela, May 1922, L. R. Reynolds (F. Psota Coll., C. N. H. M.).

1965]

Werner — Ischyropalpus

199

1 §, San Esteban, nr. Puerto Cabello, Venezuela, 1-6 Dec. 1939,
P. J. Anduze (C. N. H. M.). 1 d\ Simla, N. TRINIDAD, W. I.,
1-9 May 1963; 2$, 24-30 Apr. 1963; all M. Emsley, light trap
(Werner) .

The Eryngii — Group

A small number of South American species forms a group that is
quite aberrant in the genus. The form of the last segment of the
maxillary palpi and of the prothorax are entirely different from those
of the typical members. The last segment of the maxillary palpi is
narrowly securiform rather than triangular. The prothorax is not
at all constricted laterally before the basal impressed line, being more
like that of Anthicus, s. str., but unusually enlarged in the cf of at
least one species.

The members of the group are assigned to Ischyropalpus on the
basis of the form of the mesosternum, the presence of a zone of
small spines around the basal portion of the tibial spurs (seen only
in puteifer ), and on the form of the tegmen and general features of
the internal sac of the <$ aedeagus in eryngii and puteifer, the only
species known from males. The paired plates of the middle of the
sac are absent. And the apodeme of the penis is extremely long.

The group is therefore very disinct within the genus. Whether
it is specialized or primitive is difficult to say. But it appears to be
much more closely related to normal species of Ischyropalpus than
to any other group, and is most conveniently treated as part of this
genus.

Key to Species of Eryngii — Group
1. Pubescence in basal half of elytra cinereous, darker, almost brown,
in apical half (as seen in side view with light from front).
Dark brown to black with bases of femora and antennae
slightly paler, occasionally paler with the region of the post-
basal transverse impression of the elytra rufescent. Setae
distinctly oblique between and behind the omoplates of the

elytra. Uruguay eryngii

Elytral pubescence of uniform color, not distinctly oblique in
omoplate region 2.

Ij 2. Elytra dull luteous, brown across base, in a postmedian crescent
and in a narrow strip connecting base and crescent, near and
parallel to suture. Elytral pubescence fine and moderately
dense. Punctures of pronotum slightly confluent longitudinally.

Est. Minas Gerais, Brazil decor atus

Elytra pale to dark but without dark markings 3.

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3. Piceous, the tibiae, tarsi, antennae except apex, labrum and mouth

parts pale. Tempora of head extremely reduced, the base almost
evenly and gently rounded behind the large eyes. Venezuela

freyi

Elytra brown to luteous. Tempora distinct 4.

4. Head black, prothorax and most of underside brown; elytra,

labrum, mouth parts, antennae and legs rufescent. Punctures
of forebody sparser but those on head separated by less than

half their diameters. 3.26 mm. Para, Brazil alvarengai

Brown to rufescent, the head not darker. Punctures of forebody
denser, those on head very narrowly separated but not con-
fluent. Prothorax unusually large, especially in cf, which has
a pit on the surface of the front and hind femora facing the
tibia. 3-53 - 4.24 mm. N. Argentina and Mato Grosso, Brazil
puteifer

Ischyropalpus eryngii, new species

(Fig. 3)

2.76 - 3.28 mm, piceous above, the elytra slightly brownish, the
appendages obscurely paler. Elytral pubescence moderately sparse,
appressed, that from base to posterior border of weak postbasal
transverse impression cinereous, the rest brown. The pale pubescence
is much more noticeable, but is no denser than the rest. Sides and
base of pronotum also with some cinereous pubescence, that on disc
brown. Two large females from Arro. Espinillo are paler, with the
elytra pale laterally in the postbasal transverse impression. These
have the sculpture and pubescence similar to the more typical speci-
mens, but one of them has the elytral pubescence uniform.

Holotype cf : Length 2.91 mm. Head subtruncate at base, 0.47
mm long, 0.59 wide across eyes, 0.45 behind, the tempora short, the
temporal angles distinct, and the base gently and evenly curved
between them. A line connecting the backs of the eyes is 0.15 mm
from the middle of the base. Disc slightly flattened, densely covered
with sharply defined flat-bottomed punctures, even on frons, the
punctures ca. 0.03 mm from center to center, the intervals narrow,
less than half the diameter of the punctures, shiny; the punctures
not at all confluent. Setae appressed, ca. 0.04 mm long, brown on
disc, pale across base; tactile setae erect, straight, ca. 0.04 mm.
Frontoclypeal suture barely indicated, the frons on same curve as
front and with the same sculpture. Last segment of maxillary palpi
narrowly securiform, 0.18 X 0.09 mm. Eyes large, 0.27 X 0.23
mm, prominent, moderately coarsely faceted, with slightly curved

1965]

Werner — Ischyropalpus

201

setae ca. o.oi mm long at intervals; separated by 0.38 mm on front
and covering most of sides. Antennae brown, with segment 1 1
obscurely paler, segments 2-10 widest near apex. Measurements in
0.01 mm: 14/7, 9/6, 12/6, 11/7, 12/7, 12/8, 10/8, 10/8, io/8,
10/10, 17/10. Segment 1 somewhat barrel-shaped, pedunculate, 2-10
widest near apex, becoming more triangular, 9-10 subtruncate at
base, 11 truncate at base, tapering to a blunt point from basal 2/5.
Setae and tactile setae rather shaggy.

Pronotum 0.64 mm long, 0.41 wide at base, 0.54 maximum, 0.26
at the well-defined collar. Basal impressed line distinct. Sides widest
at apical 2/5, curving evenly to collar and almost straight to basal
impressed line. Disc slightly flattened ; sculpture similar to head but
punctures slightly deeper, smaller and denser. Tactile setae longer,
ca. 0.05 mm. Elytra subparallel, 1.79 mm long, 0.73 wide at the
well-marked humeri, 0.90 maximum, slightly flattened but with suture
elevated behind the weak postbasal transverse impression ; omoplates
feebly elevated. Setae decumbent, straight, 0.05 mm long; tactile
setae suberect, slightly curved, 0.04 mm. The setae between and
just behind the omoplates are slightly oblique. Surface shiny, with
very fine punctulation and slight irregularities; punctures small but
sharply defined, ca. 0.03 mm from center to center, the intervals flat.

Sides of mesosternum and ventral portion of mesepisterna smooth
and shiny, with a few fine punctures and closely appressed setae.
Mesepimera apparently not visible along sides of mesepisterna. Mete-
pisterna and metasternum with fine punctures and feebly micro-
reticulate. Abdomen shiny, with fine punctures. Sternum 7 simple;
sternum 8 transversely excavated in apical third, with the apical
margin very shallowly concave. The dical excavation does not appear
to have any modification of the setae. Tergum 7 with a concave hind
margin. Tergum 8 flat, shiny, finely punctured, 0.26 mm wide, the
apical portion broadly and evenly rounded. Femora moderately heavy
beyond middle. Front femur 0.49 X 0.17 mm, front tibia 0.45
X 0.06, hind femur 0.61 X 0.18. Coxae all dark, trochanters and
rest of leg dark rufescent, the trochanters slightly paler. Front tarsi
somewhat expanded. Females are almost the same as the males, but
have the front tarsi narrower.

Distribution : Known only from Uruguay. Holotype cf : Maldo-
nado (Cerro Pelado), [uruguay], 25.XII.12, sobr z Eryngium sp.,
Coll. Tremoleras. Paratypes: 8 cf, 4?, same data. 1 cT, Maldonado
(Pan de Azucar), 15.XII.12, sobre Eryngium sp., Coll. Tremoleras.
7 cf, 5?, Minas — Arro. Espinillo, [uruguay], 10.XII.12, sobre
Eryngium sp., Coll. Tremoleras. Holotype and paratypes in Museo
de La Plata.

Psyche, 1965

Vol. 72, Plate 14

Werner — Ischyropalpus

1965]

W erner — Ischyropalpus

203

Ischyropalpus freyi, new species

Very similar to eryngii, differing, in the eyes being even larger, the
base of the head short and gently, evenly curved behind them, the
punctures, of the head especially, smaller and the intervals wider.
Elytral pubescence quite sparse, of uniform tan color.

Holotype Length 2.79 mm. Head 0.46 mm long to the extreme-
ly feebly indicated frontoclypeal suture, 0.54 to apex of clypeus, 0.56
wide across eyes, 0.46 just behind. Base very short, almost evenly
and gradually curved, the tempora very short and nearly obsolete ; a
line between the hind margins of the eyes is 0.13 mm from middle of
base. Disc slightly flattened. Surface smooth and shiny, with quite
small flat-bottomed punctures ca. 0.03 mm from center to center, the
intervals flat, and as wide as punctures in most cases. Setae appressed,
0.04 mm long, tactile setae erect, straight, 0.04 mm. Clypeus on same
curve as front and similarly sculptured. Last segment of maxillary
palpi narrowly securiform 0.15 X 0.08 mm. Eyes very large, promin-
ent, moderately coarsely faceted, separated by 0.36 mm and covering
most of sides of head, with curved setae ca. 0.01 mm long at the inter-
vals of the facets. Antennae similar to eryngii, bright rufescent at base,
gradually darker beyond segment 6. Measurements in 0.01 mm: 14/8,
8/5) 9/5, 8/6, 9/6, 9/6, 9/6, 8/8, 9/9, 8/9, 15/8. Pronotu-m similar
to eryngii, 0.64 mm long, 0.38 wide at base, 0.52 maximum and 0.22
at collar. Punctures denser than on head but of similar diameter,
separated by less than their diamaters in most cases, slightly oval longi-
tudinally but not at all confluent. Elytra also very similar to eryngii
except for the unicolorous pubescence, which is nearly longitudinal in
the region of the omoplates; length 1.69 mm, width at humeri 0.68,
0.83 maximum. Punctures ca. 0.04 mm from center to center; setae

EXPLANATION OF PLATE 14

Fig. 1. Ventral view of mesothorax and part of metathorax of Ischyropal-
pus caesiosignatus (Boheman).

Fig. 2. Ventral view of aedeagus of Isychyropalpus tibialis, new species,
paratype, Maracay, Venezuela.

Fig. 3. Ventral and left lateral views of aedeagus of Ischyropalpus
eryngii, new species, paratype, Maldonado, Uruguay.

Fig. 4. Ventral and left lateral views of aedeagus and of penis or median
lobe of Ischyropalpus puteifer , new species, Mojon de Fierro, Argentina.

Figs. 5-7. Details of Ischyropalpus puteifer, new species, holotype $. Fig.
5. Left maxillary palpus. Fig. 6. Left front femur in ventral view. Fig. 7.
Left hind femur in ventral view.

Fig. 8. Ischyropalpus puteifer new species, holotype $ (X 11). All
other figures at scale indicated.

204

Psyche

[September

0.05 mm long, tactile setae 0.04. Wings apparently functional. Meso-
sternum, mesepisterna and legs similar to eryngii. Front femur 0.44
X 0.15 mm, front tibia 0.41 X 0.07, hind femur ca. 0.51 X 0.16.

Holotype Rancho Grande, [nr.] Maracay, [Est. Aragua],
Venezuela, XI. i960, G. Frey, in Museum G. Frey, Tutzing, Ger-
many.

Ischyropalpus decoratus, new species

This species, known only from the female holotype, is the only one
in the group with a marked color pattern on the elytra. These are dull
luteous with the basal area and a crescentric marking behind the middle
on each elytron brown, the two connected by a narrow strip that
parallels the suture. The suture is pale in a strip 0.17 mm wide, be-
coming darker at the base but feebly indicated even there. The prono-
tal punctures are more or less confluent longitudinally and the elytral
pubescence is moderately dense, fine and appressed.

Holotype Length 2.88 mm. Brown, the head slighty darker, the
elytra as described above, the antennae, palpi and legs dull luteous also.
Head subquadrate, with distinct but short tempora, the base gently
curved between the distinct temporal angles. Length 0.49 mm, width
across eyes 0.59, behind 0.49. Eyes large, moderately prominent, 0.28
X 0.20 mm, separated by 0.40 on front, extending far down on sides
of head. A line connecting their hind margins is 0.15 mm from middle
of base. Facets moderately coarse, with curved setae at their intervals
ca. 0.02 mm long. Surface of head densely covered with sharply de-
fined flat-bottomed punctures ca. 0.03 mm from center to center,
separated by less than half their diameters, slightly sparser on anterior
part of frons; intervals smooth and shiny. Setae appressed, fine, ca.
0.03 mm long, tactile setae straight, erect, ca 0.01 mm. Frontoclypeal
suture feeble, the clypeus on same curve as front. Last segment of
maxillary palpi narrowly securiform, 0.14 X 0.08 mm. Antennae
moderately stout, with segments 2-10 thickest near apex, becoming
subtrianguler. Measurements in 0.0 1 mm: 15/6, 8/6, 8/5, 8/6, 8/6,
8/7, 8/8, 8/8, 8/8, 8/8, 13/8. Segment 1 subparallel, feebly pedun-
culate, 2-3 thickest near apex, 4-10 subtriangular, becoming heavier,
9-10 subtruncate at base, 11 truncate at base, the sides gently curved
to the blunt apex from near base. Setae and tactile setae rather shaggy.

Pronotum 0.67 mm long, 0.38 wide at base, 0.56 maximum and
0.26 at the short but well-defined collar. Sides widest at apical 2/5,
almost evenly curved to collar and almost straight back to the level
of the distinct basal impressed line. Disc almost evenly convex, slight-
ly concave behind middle. Punctures denser and smaller than on head,

1965]

Werner — Ischyropalpus

205

slightly oval and feebly confluent longitudinally. Pubescence similar to
head but tactile setae longer, ca. 0.04 mm. Scutellum pale. Elytra sub-
parallel, 1.73 mm long, 0.69 wide at the well-defined humeri, 0.90
maximum. Surface shiny but very feebly punctulate, the small punc-
tures 0.03 mm from center to center, slightly papillate, especially to-
ward base. Omoplates feebly elevated; suture feebly elevated behind
middle. Setae moderately dense, fine, appressed, 0.05 mm long; tac-
tile setae suberect, almost straight, 0.04 mm. The general color of the
elytra is dull luteous but the base from humeri to and including omo-
plates is brown, the suture here narrowly slightly paler. An obscure
brown strip extends from near the suture in this brown zone to a
crescentric marking on each elytron 0.90 mm from base and ca. 0.26
mm wide. The suture is pale for ca. 0.17 mm between the posterior
markings and the strips, the boundaries of this pale zone almost
straight. It continues obscurely into the basal area.

Mesosternum and ventral portion of mesepisterna smooth and shiny,
the former quite narrow laterally. Mesepimera very narrowly exposed
along edges of mesepisterna. Abdomen finely punctured and sparsely
pubescent, the first visible sternum shiny, the rest feebly microretic-
ulate. Legs, including front and middle coxae, dull luteous. Front
femur 0.46 X 0.15 mm, front tibia 0.41 X 0.05, hind femur 0.56 X
0.17.

Holotype Paracatu, Minas Gerais, [brazil], VII-1960 - 86/60,
Exp. Formosa Col., in Museu Nacional, Rio de Janeiro, Brazil.

Ischyropalpus alvarengai, new species

This species most closely resembles decoratus in general appearance,
but it lacks elytral markings. The punctures on the pronotum are not
confluent and the pubescence on the elytra is longer and coarser. If
the single known specimen is typically colored, the coloration is dis-
tinctive, as indicated in the key to the species of the group.

Holotype $: Length 3.26 mm. Head subquadrate, 0.52 mm long,
0.64 wide across eyes, 0.54 behind, with the tempora short but distinct,
the temporal angles well-marked and the base very gently and evenly
curved between them. The middle of the base is 0.17 mm from a line
connecting the hind margins of the eyes. Disc slightly flattened, evenly
covered with flat-bottomed punctures ca. 0.03 mm from center to
center, the intervals smooth and shiny, and less than half the diameter
of the punctures. The sculpture continues to the frontoclypeal suture,
which is fine but well marked, and onto the clypeus, which is on the
same curve as the frons. Setae appressed, ca. 0.05 mm long, tactile
setae erect, 0.04 mm. All of maxillae and labium rufescent. contrast-

206

Psyche

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ing with the dark head. Last segment of maxillary palpi narrowly
securiform, 0.15 X 0.08 mm. Eyes moderately large and prominent,
0.3 1 X 0.20 mm, moderately coarsely faceted, occupying much of side
of head, with curved setae ca. 0.02 mm long at the intervals of the
facets. Antennae of same general proportions as eryngii but the inter-
mediate segments stouter, 5-10 being subtriangular and gradually
stouter, and subtruncate at base. Measurements in 0.0 1 mm: 18/8,
8/6, 9/6, 9/6, 9/8, 9/8, 9/8, 9/9, 9/9, 9/8, 14/8.

Pronotum of form of decoratus, 0.76 mm long, 0.46 wide at base,
0.68 maximum and 0.28 at collar. Punctures denser than on head,
longitudinally oval, narrowly separated. A few of them are slightly
confluent longitudinally, but almost all are distinctly separated. The
disc is slightly flattened, but almost evenly convex. Elytra suparallel,
1.98 mm long, 0.79 mide at the well-marked humeri, 1.02 maximum,
somewhat flattened, with the omoplates and postbasal transverse im-
pression barely indicated and the suture not elevated. Surface as in
decoratus , the punctures fine and 0.04 mm from center to center;
setae appressed but not closely so, slightly curved, 0.05 mm long,
noticeably coarser and shaggier than in decoratus; tactile setae sub-
erect to erect, 0.04 mm. There is a narrow slightly darkened strip
along the suture, ca. 0.18 mm wide and most evident on the middle
third. The region of the omoplates is also slightly darkened, but
probably because the slightly translucent elytra reveal the top of the
thorax. Wings apparently of full length. Underside as in decoratus,
but mesepimera more distinctly visible. Legs rufescent, the coxae
darkened toward base, the hind coxae mostly dark. Front femur 0.49
X 0.18 mm, front tibia 0.44 X 0.08, hind femur 0.64 X 0.20.

Holotype Cachimbo, Para, brasil, VI- 1962, M. Alvarega,
in the collection of Col. Moacir Alvarenga, Rio de Janerio, Brazil.

Ischyropalpus puteifer, new species
(Figs. 4-8)

3.53-4.24 mm, brown with the appendages rufescent, to all rufes-
cent; subopaque, moderately slender and flattened. Prothorax,
especially of male, unusually large. Front and hind femora of male
with a pit on the surface facing the tibia.

Holotype <$ : Length 4.24 mm. Head 0.67 mm long, 0.72 wide
across eyes, 0.67 just behind, with short tempora, the base gently and
evenly rounded between the poorly defined temporal angles, moderately
retrosalient. Disc somewhat flattened, evenly covered with sharply
defined flat-bottomed punctures 0.03 mm from center to center even
on frons, the punctures only narrowly separated but not all con-

1965]

W erner — Ischyropalpus

207

fluent, the intervals ca. 1/2 the width of the punctures and shiny. The
surface appears subopaque because of the dense punctures. Setae
moderately dense, decumbent, 0.04 mm long, tactile setae suberect,
0.04 mm. Frontoclypeal suture distinct but not deep; surface of cly-
peus similar to frons. Labrum ogival, shiny. Last segment of maxil-
lary palpi narrowly securiform (Fig. 5), 0.20 X 0.09 mm. Eyes
moderately prominent, moderately coarsely faceted, 0.33 X 0.23 mm,
separated by 0.51, a line connecting their hind margins 0.26 mm from
middle of base. Antennae moderately slender, gradually thicker
apically, the segments distinctly separated. Measurements in 0.01 mm :
19/9, 9/6, 13/8, 12/8, 13/8, 10/9, 12/9, 12/9, 12/9, 12/9, 18/9.
Segment 1 parallel-sided, feebly pedunculate, 2-10 thickest near apex,
gradually more truncate at apex, 6-10 distinctly truncate at base,
subtriangular ; 11 truncate at base, tapering to a blunt point from
basal third.

Pronotum unusually large and broad, 1.13 mm long, 0.59 wide at
base, 0.84 at beginning of expanded portion 0.26 mm from base, 1.00
maximum, 0.36 at the short collar. The apical area behind the collar
is elevated above the collar, with the border of this elevated area
slightly concave. This anterior border meets the sides in an even but
rather sharp curve; the sides are then subparallel about half-way to
the base, then slightly convergent to another sharp curve, which ends
at the level of the distinct basal impressed line. Disc moderately
flattened. Surface similar to head but punctures smaller and ca. 0.02
mm from center to center anteriorly and laterally, almost as on head
in posterior half of disc. Lateral margins evenly rounded, but in a
curve that is about as sharp as the one that joins the front and side
margins. Elytra almost parallel, 2.44 mm long, 0.95 wide at the dis-
tinct humeri, 1.20 maximum, with the apices gently conjointly rounded
and narrowly rounded at suture. Disc almost evenly rounded in
transverse section ; postbasal transverse impression feebly indicated.
Surface with lightly papillate punctures ca. 0.04 mm from center to
center, appearing feebly shiny. Setae moderately dense, subappressed,
0.08 mm long, tactile setae subdecumbent, slightly curved, 0.04 mm.
Wings well-developed, probably functional.

Mesosternum shallowly punctured and pubesecent medially, shiny
and almost smooth laterally, the ventral portion of the mesepisterna
similar to the sides. Mesepimera narrowly visible along edges of ven-
tral portion of mesepisterna. Abdomen shiny, finely punctulate and
with setae similar to elytra. Sternum 7 not modified, truncate at apex.
Sternum 8 with a shallow transverse depression that is only very
sparsely punctulate with a few fine setae, deepest near apex and extend-

2o8

Psyche

[September

ing most of the way across the segment, which has a subtruncate apex.
Tergum 8 flattened, slightly concave longitudinally, 0.41 mm wide,
with the apex gently rounded, without any indication of a median
angulation. Surface shiny, finely punctulate and pubescent. Ventral
sculptured portion flat, with no indication of a median keel.

Femora moderately stout, widest at apical third. Front femur with
a deep, slightly margined, roughly circular pit, bearing a few erect
setae from the middle of its flat and apparently membranous bottom
(Fig. 6), on anterior portion of the side facing the tibia. Hind femur
with an elongate pit, which has only the sides elevated ( Fig. 7 ) . Mid-
dle femur without a pit. Front femur 0.64 X 0.23 mm, front tibia
0.56 X 0.10, hind femur 0.77 X 0.26.

The 9 has the pronotum less enlarged, with the sides only slightly
narrowed behind. The 9 from Colonia Benitez is 3.71 mm long. The
pronotum is 0.92 mm long, 0.56 wide at base, 0.67 at 0.26 from base,
0.77 maximum and 0.3 1 at collar. The front margin is only slightly
higher than the collar and slopes to it; and the disc is almost evenly
rounded in section. It is only slightly broader than the head, which is
0.69 mm wide across the eyes. If the elytral length of the holotype
were made equivalent, its pronotum would be 1.02 mm long and 0.89
mm in maximum width. The femora of the $ are slightly more slender
and completely lack the pits of the cf •

The holotype and most of the paratypes are all brown above, the
9 from Colonia Benitez rufescent with the elytra brown, and the
99 from Mojon de Fierro and Campo del Cielo all pale rufescent.

Holotype cf • Colonia Benitez, Terr. Chaco, Argentina, 12.XII.
1948, R. Golbach (Inst. Miguel Lillo). Paratypes: 1 9, same data
(Inst. Miguel Lillo). 1 cf, 1 9> Mojon de Fierro, Terr. Formosa,
Argentina, 25. XII. 1948, R. Golbach (Inst. Miguel Lillo). 1 9>
Campo del Cielo, [Prov. Santiago del Estero, Argentina], I.1934, J.
Bosq (Museo de La Plata). 1 cf , 9 9> Cuyaba, Matto Grosso, Brasi-
lien (Chicago Nat. Hist. Mus., from residual collection of Staudinger
and Bang-Haas).

References Cited

Blackwelder, R. E.

1945. Checklist of the coleopterous insects of Mexico, Central America,
the West Indies, ,and South America. Part 3. U. S. Nat. Mus.
Bull. 185 : i-iv, 343-550.

Bonadona, P.

1961. Revision des Ischyropalpus Laferte [Col. Anthicidae]. Rev.
frangaise Entomol. 28: 150-177.

1965]

W erner — Ischyropalpus

209

Casey, T. L.

1895. Coleopterological notices VI. Anthicini. Ann. New York Acad.
Sci. 8: 639-772.
von Krekich-Strassoldo, H.

1914. Beitriige zur Kenntnis der Anthiciden. Verh. Zool.-Bot. Ges.
Wien, 1914: 215-228.

DE LAFERTE-SENECTERE, F. T.

1848. Monographic des Anthicus et genres voisins, Coleopteres Heter-
omeres de la Tribu des Trachelides. De Sapia. Paris, xxiv “F
340 p., 16 pis.

Leng, C. W.

1920. Catalogue of the Coleoptera of America, North of Mexico. John
D. Sherman, Jr. Mount Vernon, N. Y. x + 470 p.

Pic, M.

1911. Coleopterorum Catalogus, pars 36, Anthicidae [vol. 17]. W.
Junk. Berlin. 102 p.

NOTES ON NEOTROPICAL TABANIDAE VI
A NEW SPECIES OF LEPISELAGA MACQ.
WITH REMARKS ON RELATED GENERA

By G. B. Fairchild

Gorgas Memorial Laboratory, Panama, R. de P.

The discovery by Dr. Vernon Lee of a new species of Lepiselaga
in Colombia, provides the opportunity to review the taxonomy of
the group and to make some suggestions as to the relationships of the
species to each other, as well as to other genera usually considered
to be related.

Lutz (1913) erected the subfamily Lepidoselaginae to contain
Lepiselaga , Selasoma , Himantostylus, and Stigmatophthalmus , the
last two described as new. Lutz accepted Osten Sacken’s invalid
emendation of Lepiselaga to Lepidoselaga , but used both inter-
changeably in this publication. He also stated that the subfamily
would contain other unnamed Australian and African species. Ender-
lein (1925) retained the group as a tribe Lepiselagini for 9 more or
less unrelated genera, but including Lepiselaga and Himantostylus ,
though not Selasoma or Stigmatophthalmus , which were placed in
the tribes T'abanini and Dichelacerini respectively. Krober (1934)
included Lepiselaga and Selasoma in the subfamily Lepiselaginae, but
placed Himantostylus in the Stenotabaninae and relegated Stigma-
tophthalmus to the position of a subgenus of Dicladocera in the sub-
family Tabanini. Fairchild (1942) placed Lepiselaga and Selasoma
in Tabaninae , tribe Lepiselagini. Finally Mackerras (1955) included
all the Neotropical Tabaninae with bare basicosta in one tribe,
Diachlorini.

As presently known, Selasoma, Stigmatophthalmus and Himanto-
stylus are monotypic, each containing but a single rather aberrant
species, and will be discussed first.

Selasoma Macquart

1838, Dipt. Exot. 1(2): 187. Type Tahanus tibialis Fab.

Krober (1934) included also T. nigrocoeruleum Rond. 1850 and
5. giganteum Lutz 1913. The latter is a species of Stibasoma, as
noted earlier (Fairchild 1961), while recent examination of the type
of nigrocoeruleum in Naples shows it to be the same as Dicladocera
acheronitens Krbb. 1931 (New synonymy). S. tibiale has a wide
range in the Neotropics, being recorded from Oaxaca, Mexico to
southern Brasil, but seems everywhere uncommon and local.

210

1965]

Fairchild — Neotropical T abanidae

21 1

The position of Selasoma , similar in its coloration and incrassate
tibiae to Lepiselaga, is dubious. It has generally been placed near
Lepiselaga (Lutz 1913, Enderlein 1925, Krober 1934) but there
are fundamental points of difference. The basicosta bears a fair
number of setae, the palpi are greatly inflated but not flattened, the
frons is narrow with a rugose linear callus nearly filling it and with-
out tubercle or ocelli at vertex, the labella are large and wholly
membranous, the first antennal segment is not elongate, the second
bears a dorsal tooth and the basal plate of third is wide and flattened.
The eyes are unpatterned in life, dull greenish black, while those of
Lepiselaga and Himantostylus are characteristically banded. Accord-
ing to Lutz (1913) the species is crepuscular and bites horses on the
belly. Records from Panama confirm this.

Stigtnatophthalmus Lutz

1913, Mem. Inst. Osw. Cruz, 5(2): 174, 184-185, PI. 13, fig. 23. Type
Stigmatophthalmus altivagus Lutz.

Stigmatophthalmus altivagus Lutz was placed by Lutz (1913)
with Selasoma and Lepiselaga, by Enderlein (1925) near Stibasoma ,
and by Krober as a subgenus of Dicladocera. It has, however, setose
basicosta, large membranous labella, and no vestiges of ocelli. The
wing pattern is somewhat like Selasoma , though more extensively
black, and the fore tibiae slightly incrassate. Except for the wing
color, there seem to be no characters that would exclude it from
Tabanus , much less allow placement in the Diachlorini. It certainly
does not seem in any way related to Lepiselaga or even Selasoma.
Lutz figures the species as with green eyes with a single dark band.
I have given a figure of head structures elsewhere (Fairchild 1961).
Nothing is recorded of its habits, and it seems to have been taken
only about Petropolis and in the mountains of Sta. Catarina in
southern Brasil.

Himantostylus Lutz
Fig. 1

1913, Mem. Inst. Osw. Cruz, 5 (2): 174, 183-184, PI. 13, fig. 22. Type H.
intermedins.

The genus and species were based on a single male from eastern
Peru. Stone (1934) later described the female from Bolivia. A long
series of females and a single male were secured in Darien, Panama,
in 1963, and there are other specimens collected by Luis E. Pena at
Quincemil, Dept. Cuzco, Peru, Oct. 1962 in the collection of Dr.
L. L. Pechuman. Panama specimens are darker, having the cubital
cell and anal area as dark as the basal cells, not paler as in the type
and other Peruvian material.

Psyche, 1965

Vol. 72, Plate 15

4. L. colombiana

Fairchild — Tabanidae

1965]

Fairchild — Neotropical T abanidae

213

Hhnantostylus Lutz, as suggested by Stone (1934) and Barretto
( 1949), is very similar to Lepiselaga but differs structurally in having
a strong tubercle at vertex with well marked vestiges of ocelli, a
quite different frontal callus, and in lacking the narrowed discal cell.
The labella are also without sclerotized strips, and the palpi less
highly modified. The palpi of male Himantostylus are porrect, oval,
inflated and shiny, those of Lepiselaga are decumbent and similar
to the female, though more slender and less flattened.

Many of the peculiarities and similarities of Lepiselaga and
Himantostylus may be due to what might be termed mimetic con-
vergence. All the species bear a strong resemblance in size, coloration,
and to some extent in actions, to stingless bees of the genus Trigona.
This resemblance is strongest in Hhnantostylus and the subgenus
Conoposelaga, less marked in L. crassipes. The latter at least is
preyed upon by certain solitary wasps, as noted by Bates over a
century ago (Bequaert 1926). Himantostylus intermedius is espe-
cially bee-like in its actions. In a jungle camp on the Rio Tacarcuna,
Darien Province, Panama, I collected a good series of this species,
and had an opportunity to observe its actions. The flies flew silently
close to the ground, clustering on my canvas and rubber boots and
seldom alighting above the ankles. They crawled busily over the
boots in the manner of stingless bees investigating an attractive sur-
face, but were unable to penetrate the thick material. When a hand
was placed on the boots, they crawled over the fingers, but made no
attempt to bite. L. crassipes also seems to prefer the lower part of
the body, and also sometimes crawls about in a bee-like manner, but
it bites viciously when a suitable site is found. Little is recorded
as to the habits of the other species. Lutz (1913) notes that the
original series of L. aberrans was taken at 3 p. m. and bit man and
horses avidly. The original specimens of L. albitarsis were collected
by D’Orbigny in 1827* His original journal is preserved in the

Explanation of Plate 15

Figures of heads, notopleural lobes and wing are to the same scale, The
wing measures 5.3 mm. Figures of antennae and palpi are also to the same
scale, but twice the magnification of heads.

Fig. 1 Himantostylus intermedius Lutz, a, palpus; b, dorsal view of noto-
pleural lobe ; c, antenna ; d, head, frontal view. Panama. Fig. 2 Lepiselaga
(Conoposelaga) albitarsis Macq. a, palpus; b, notopleural lobe; c, antenna.
Argentina. Fig. 3 Lepiselaga (Conoposelaga) aberrans Lutz, a, palpus; b,
notopleural lobe; c, antenna. Paratype. Fig. 4 Lepiselaga ( Conoposelaga )
colombiana n. sp. a, head; b, palpus; c, notopleural lobe; d, antenna; e,
wing. Holotype. Fig. 5 Lepiselaga (Lepiselaga) crassipes Fab. a, palpus;
b, notopleural lobe; c, antenna, Panama.

214

Psyche

[September

Museum d’Histoire Naturelle in Paris, and Dr. Tsacas kindly
secured for me a photocopy of the parts dealing with Diptera.
D’Orbigny records under number 9533-37, the number borne by the
types, that the species lives on the banks of the Parana along its
whole length, that its bites are very painful and that the local inhabi-
tants, who already suffer much from the attacks of mosquitoes and
a multitude of other insects, fear this species greatly, as it torments
them cruelly and returns ceaselessly to the attack. These remarks
indicate that albitarsis as well as aberrans have feeding habits similar
to crassipes, but different from Himantostylus.

In summary, I believe that Stigmatophthalmus and Selasoma are
both isolated and unrelated genera best placed in the tribe Tabanini.
They may well be relatively old elements whose connections have
disappeared. Stigmatophthalmus altivagus is a rather rare species,
seemingly confined to the higher altitudes of the coast range of south
eastern Brasil. Selasoma is widespread from Mexico to Brasil, but
apparently local in occurrence. Himantostylus I believe belongs in
the Diachlorini, but I think its resemblances to Lepiselaga are more
likely due to mimetic convergence rather than close relationship in a
phylogenetic sense. To me it appears closer to Diachlorus and Steno-
tabanus than to Lepiselaga , but it is in any event rather isolated.

I give below a key to the genera discussed here. All the species
are black or largely so, with basally black wings, rather stout and
chunky build, and tibiae more or less inflated.

Key to the genera

1. Discal cell (ist M2) of wings constricted, the vein forming its
posterior margin strongly arched upward, the cell entirely infus-

cated. Lepiselaga 2.

Discal cell normal, either wholly or partly hyaline, or with a
hyaline spot. 3*

2. Frons about twice as high as wide, parallel sided, the vertex with-
out bare area. Notopleural lobes not strongly protruding. Body

when undenuded clothed with greenish irridescent scales

(Lepiselaga)

Frons as wide or wider than high, widened below, the vertex
protuberant and shiny. Notopleural lobes strongly inflated and
protuberant. Body without greenish scales (Conoposelaga)

3. Basicosta bare. Small species less than 9 mm. long. Antennae

slender, the basal plate of third segment subcylindrical, about

twice length of style. Three distinct ocelli at vertex. Wing
basally black or dusky to ends of basal cells, beyond this wing

1965]

Fairchild — Neotropical T abanidae

215

glass clear with yellow veins and stigma FI im ant 0 stylus

Basicosta at least sparsely setose. Large species generally over
12 mm. Basal plate of antennae broad, or with a strong dorsal
angle. No ocelli. Wing more extensively black, the discal cell

at least half black. 4.

4. All tibiae greatly inflated. Palpi greatly inflated and shiny. Sub-
callus, face and cheeks largely shiny. Basal black of wing extend-
ing to middle of discal cell, with a small clear spot in first and
second basal cells near apex. Antennal plate broad and flattened,

much longer than style, obtusely angulate above. Selasojna

Fore tibiae slightly inflated, others normal. Palpi slender, polli-
nose. Subcallus, face and cheeks brown pollinose. Basal black
of wing extending beyond end of discal cell, the latter with a
small transverse oval hyaline fenestra beyond middle. Antennal
plate with a strong acute dorsal angle, almost a spine, the plate
but slightly longer than style. Stigmatophthalmus.

Lepiselaga Macquart

1838, Dipt. Exot., 1(1): 153. Krober, 1934, Rev. Ent. 4(2): 252. Fairchild
1942, Ann. Ent. Soc. Amer., 3 5 (3): 290. Type Tabanus lepidotus Wied.
{— eras sipes Fab.)

Hadrus Perty, 1833, Del. Anim. Artie., Brasil, p. 183 (nec Dejean 1833
Coleoptera) .

Lcpidoselaga Osten Sacken, 1876, Mem. Boston Soc. Nat. Hist., 2: 435.
Conoposelaga Barretto, 1949, An. Fac. Med. Univ. S. Paulo, 24: 87-88. Type
Lepiselaga aberrans Lutz.

The most recent definition of the genus is by Fairchild (1942).
To this may be added the condition of the discal cell, which is nar-
rowed in the middle by the strong anterior bending of vein M3 where
it forms the posterior border of the discal cell.

L. aberrans Lutz, (Fig. 3) for which Barretto erected the genus
Conoposelaga, differs from L. crassipes (Fig. 5) most notably in the
inflated notopleural lobes, inflated scutellum, broad frons with shiny
vertex, and longer first antennal segment. These characters are to
a considerable extent shared by L. albitarsis Macq. (Fig. 2) and the
new species described below, so that there is justification for Barretto’s
action. I prefer, however, in view of the obviously close relationship
of the four species, to retain Conoposelaga in a subgeneric sense.

Lepiselaga (Conoposelaga) colombiana n. sp.

Fig. 4

A small black species resembling L. albitarsis but with narrower
frons, longer third antennal segment, and abdomen black tomentose
dorsally.

216 Psyche [September

Female. Length 6 mm., of wing 5.5 mm. Eyes short and sparsely
pilose. Frons as figured, largely black and shiny, the pollinose areas
dark brown. Subcallus yellow brown, wholly shiny and rather
inflated. Frontoclypeus and genae wholly shiny black. Antennae and
palpi as figured. Proboscis short, about equalling head height, the
labella small and compact, at least partially sclerotized.

Mesonotum black, thinly dark pollinose with scattered brassy hairs,
the notopleural lobes globose, shiny, sparsely dark haired. Scutellum
black and shiny, somewhat inflated. Pleura and sternum dark
brown pollinose, sparsely black-haired. All legs, except white tarsi,
black and black-haired. All tibiae strongly incrassate. Halteres brown
with brown knob. Wing as figured, the costal cell and basal cells
dark yellowish brown, rest of pattern blackish.

Abdomen with first tergite velvety black with a small patch of
yellow at extreme sides. Second tergite velvety black in middle third,
lateral thirds shiny yellow. Third and fourth tergites largely velvety
black, the lateral yellow decreasing posteriorly. Remaining tergites
shiny black. The abdomen also bears scattered long yellow hairs,
denser laterally and posteriorly. Beneath the abdomen is wholly shiny,
yellow on first two or three segments, black on the remainder.

Holotype female, Barbascal, Intendencia de Meta, Colombia, 23
Sept. 1964, Vernon Lee coll. To be deposited in M.C.Z., Cambridge,
Mass.

Paratype female, same data. The paratype is slightly smaller and
with considerably less yellow on abdomen. The species can be sepa-
rated from related species by the characters given in the key.

Key to species of Lepiselaga

1. Basal plate of third antennal segment longer than first two an-

tennal segments. Black wing pattern with about 9 small round
clear spots. Frons about twice as high as wide, with a small
round basal callus. No bare area at vertex. Whole dorsum, in-
cluding frons, with greenish metalic scales

(Lepiselaga) crassipes Fab.

Basal plate shorter than first two antennal segments. Black wing
pattern with no more than 3 round clear spots. Frons as wide
as high or wider, the callus large and diffuse, the vertex exten-
sively bare. Body without greenish scales. .... (Conoposelaga) 2.

2. Tergites 2 to 4 of abdomen orange yellow, with broad transverse
orange pollinose bands. Dark wing pattern brownish, with a spur

reaching hind margin in fourth posterior cell (3rd M).

aberrans Lutz.

1965] Fairchild — Neotropical Tahanidae 2IJ

Abdomen at most yellowish horn colored at sides of anterior seg-
ments. Posterior spur of wing pattern fails to reach hind margin.

3-

3. Frons markedly wider than high, much wider below than at
vertex. Third antennal segment basally yellow, the plate much
less than twice length of style. Posterior spur of dark wing
fascia narrow, ending abruptly. Only first tergite velvety black,

the rest shiny. al bit arsis Macq.

Frons about as high as wide, slightly widened below. Third an-
tennal segment black, the plate twice length of style. Posterior
spur of wing fascia broad, fading out in fourth posterior cell
before wing margin. Tergites 1 to 4 extensively velvety black
dorsally. colombiana n. sp.

References

Barretto, M. P.

1949. Estudios sobre Tabanidas Brasileiros X. Conoposelaga n. gen.
An. Fac. Med. Univ. S. Paulo, 24: 87-93, PI. 1.

Bequaert, J.

1926. Tahanidae, in Med. Rep. Hamilton Rice 7th Exped. Amazon,
etc. Contr. no. IV Harvard Inst. Trop. Biol. Med. Chap. XV,
Insecta, pp. 214-235. Cambridge, Harvard Univ. Press.

Enderlein, G.

1925. Studien an blutsaugenden insekten 1. Grundlagen einer neuen
Systems den Tabaniden. Mitt. Zool. Mus. Berlin, 11 (2): 255-409.
Fairchild, G. B.

1942. Notes on Tahanidae from Panama. IX. The genera Stenotabanus
Lutz, Lepiselaga Macq. and related genera. Ann. Ent. Soc. Amer.,
35 (3) : 289-309, PI. 1.

1961. The Adolfo Lutz collection of Tahanidae. 1. The described genera
and species, condition of the collection, and selection of lectotypes.
Mem. Inst. Osw. Cruz, 59(2): 185-249, 2 plates.

Krober, O.

1934. Catologo dos Tahanidae da America do sul e Central, incluindo
o Mexico e as Antilhas. Rev. Ent., 4(2-3): 222-276, 291-333.

Lutz, A.

1913. Tabanidas do Brasil e alguns Estados visinhos. Mem. Inst. Osw-
Cruz, 5(2) : 142-191, PI. 12-13.

Mackerras, I. M.

1955. The classification and distribution of the Tahanidae. II. History:
Morphology: Classification: Subfamily Pangoniinae. Aust. J.

Zool., 3 (3) : 439-511.

Stone, A.

1934. Notes on Bolbodimyia and Himantostylus, with a new species.
Rev. Ent., 4(2) : 190-191.

SELF-BURYING BEHAVIOR IN THE
GENUS SICARIUS (ARANEAE, SICARIIDAE)1

By Jonathan Reiskind

Biological Laboratories, Harvard University

The family Sicariidae has only recently been recognized as a poly-
ph}detic taxon. Although it is difficult to establish natural limits,
Gertsch (1949) has more logically separated the group into the
Scytodidae, Diguetidae, Plectreuridae, Loxoscelidae, and Sicariidae
(s.s.), all of which had been subfamilies of the Sicariidae (Bonnet,
I945-I959). These five families are differentiated distinctly and
naturally by their ethological characters. The sole reason for their
being previously lumped is their common possession of simple geni-
talia. This character is automatically considered primitive and hence
these spiders were “representative of the ancestral stock.” Their
genitalia may, indeed, be relatively unchanged from the primitive
form but in many other characteristics these spiders are quite spe-
cialized and advanced and bear very little resemblance to the an-
cestral stock. One can readily talk of “primitive characters” but
rarely of “primitive, living organisms.”

Some of the highly specialized ethological characters of the
families mentioned are:

Scytodidae: Prey captured by spitting gummy substance from

chelicerae; egg cocoons carried; spiders sometimes live in web on
stones and rocks or in houses (Dabelow, 1958).

Diguetidae: Tubular cocoon constructed from silk and detritus
and suspended by web in vegetation over sheet web ; layers of egg
sacs in a cocoon which also acts as retreat for spider (Cazier & Mor-
tenson, 1962).

Plectreuridae: Low lying mesh web, associated with tubular re-
treat under rock or debris, in which spider sometimes hangs up-side-
down; eggs in loose cocoon (W. Eberhard, pers. comm.; Gertsch,
1958).

Loxoscelidae: Large, irregular sheet web (hackled band appear-
ance) ; eggs in loose sac.

1Research a by-product of National Institute of Health Grant No. AI-
01944 to Dr. H. W. Levi.

Manuscript received by the editor October 15, 1965.

2l8

1965] Reiskind — Genus Sicarius 219

Sicariidae: No web, unique ethological specializations outlined
below.

The family Sicariidae (s.s.) contains the single genus, Sicarius.
The genus has a predominantly south temperate distribution with
species also described from Peru, Chile, Argentina and Southern
Africa, with single species found in eastern Brazil, the Galapagos
Islands and Costa Rica. Merian (1913) states that this “distribu-
tion meridionale isolee est le resultat de la retraite de [ce genre], de
la zone tropical vers le Sud.” On biogeographic grounds Merian
considers Sicarius part of an ancient fauna.

The simple genitalia in the Sicariidae place them in the ecribellate
spider group known as the haplogynes. In regard to systematics,
“spiders lacking distinctive genitalia, in particular the Haplogynae,
tend to have been neglected rather than to have stimulated interest
in extra-genital characters” (Cooke, 1965). These extra-genital char-
acters must include aspects of the external morphology (as used in
Cooke’s study of Dysdera) as well as several non-morphological char-
acters— e.g., ecological and ethological ones.

In order for a character to be taxonomically useful, whether mor-
phological or not, it must be comparatively constant in the taxon
being studied.

Sicarius exhibits three behavioral traits that are of potential sys-
tematic value:

1. Stridulation - When disturbed the spider will rub the femur of
its pedipalp against ridges on the chelicerae, producing bee-like buzzing
(Simon, 1893 of the South African species). Simon suggests that the
buzzing frightens enemies.

2. Cocoon Construction - The structures built by an organism are
the physical summation of certain behavioral patterns. In spiders
these structures include webs and egg cocoons. As a hunting spider,
Sicarius does not spin webs but constructs egg cocoons that are char-
acteristically covered with an earthy coating. Simon (1899) utilized
this ethological character in noting the difference in the cocoons of
Sicarius hahni Karsch (S. W. Africa) and ? S. peruensis Keyser-
ling (Peru).

3. Self-burying - This phenomenon, found in two species of South
American Sicarius , is reported in this paper.

Two species of Sicarius collected by Dr. H. W. Levi in South
America were used in this study. Because of an inadequate knowl-
edge of the taxonomy of this genus I have designated by numbers
the two distinct species discussed in this paper, and have so labelled
the specimens and deposited them in the Museum of Comparative

220

Psyche

[September

Zoology. Sicarius sp. i was collected under a log in a pasture near
Santiago del Estero, Argentina, on April 2, 1965. Sicarius sp. 2 was
found under a rock at the base of a loma in the desert south of
Lima, Peru on February 7, 1965.

Detailed observations and timings were made by analyzing, frame
by frame, 16 mm. motion pictures (24fps) of the complete, self-bury-
ing activity. Several runs of each species were analyzed and twelve
frames from one take of Sicarius sp. 1 are presented in Plates 1 and 2.

The basic pattern of behavior observed was similar in both cases
though the species are distinct. This pattern is outlined in Table 1
and illustrated in Figures 1-12.

The first six steps (Table 1) are invariable, qualitatively, though
time duration of each action and period between actions varies a
great deal. In digging, the first two pairs of legs dig while the hind
pairs stabilize the spider (Fig. 1). Positioning of legs (Step 6) varies
from less than a second to four seconds.

Step 7 consists of the first pair of legs alternately throwing sand
over the body and exposed portions of the 4th legs (arrow in Fig. 6)
from 6-15 times (approx. 4 times/second). Tufts of hairs on abdo-
men (arrow in Fig. 2) aid in holding the sand-covering in place. In
Sicarius sp. 2 this action is split into two sections. First the throwing
of sand, then a pause of about 30-120 seconds, then a continuation of

Table 1. Self-burying in Sicarius sp. 1 (duration
in seconds, of each step within parentheses).

Step 1. Digs hole (1.0) Fig. 1

Step 2. Jumps into hole (0.2) Fig. 2
Step 3. Adjusts abdomen
(0.4)

Step 4. Adjusts 4th legs (0.6) Fig. 3
Step 5. Buries 4th legs partly

(0.2) Fig. 4

Step 6. Positions legs (2.0) Fig. 5
Step 7. 1st legs cover body

and 4th legs with sand
(1.8) Fig. 6

Step 8. Buries 1st legs partly

(0.6) Fig. 7

Step 9. 2nd left leg throws sand

forward (0.3) Fig. 8

Step 10. Buries 2nd left leg (0.5)
Step 11. 2nd right leg throws

sand forward (0.2) Fig. 9
Step 12. Buries 2nd right leg

(0.6) Fig. 10

Step 13. 3rd left leg throws sand

forward (0.2) Fig. 11
Step 14. Buries 3rd left leg (0.4)
Step 15. 3rd right leg throws
sand forward (0.2)

Step 16. Buries 3rd right leg

(0.4) Fig. 12

Explanation of Plate 16

Figs. 1-6. Self-burying of Sicarius sp. 1, steps 1, 2, 4-7 (see Table 1
and text for details).

Psyche, 1965

Vol. 72, Plate 16

Reiskind — Sicarius

Psyche, 1965

Vol. 72, Plate 17

Reiskind — Sicarius

1965]

Reiskind — Genus Sicarius

223

Step 7 or beginning of Step 8. This same behavior is also observed
in a more immature specimen of Sicarius sp. 2. In Sicarius sp. 1
there is usually no pause during Step 7 or between Steps 7 and 8.

The first legs are buried (Step 8), as are the second and third
legs, by forward, downward, jerky movements. Often the legs are
only partially buried, leaving the patella and part of the femur ex-
posed (arrows in Fig. 7).

Step 9 begins with either the right or left second leg (usually the
right). After throwing sand forward, covering the exposed part of
the first leg of its side, each leg buries itself (Steps 9-12).

The third leg actions (Steps 13-16) resemble those of the second
legs with either side beginning first. This usually completes the bury-
ing though there are often adjustments in the fourth legs, and some-
times fourth leg actions similar to the second and third legs.

The above description is of the complete, fixed behavior observed
in both species of Sicarius. But often the burying stops prematurely,
usually in response to some environmental difficulty — e.g., a cramped
position or an unsatisfactorily dug hole. At other times cause of
cessation before completion is unidentifiable.

To summarize, two species of Sicarius from South America show
basically similar, fixed self-burying behavior with only one, distinct,
qualitative difference (in Step 7). This fixed ethological character
appears to be of potential systematic value.

The author wishes to thank Dr. H. W. Levi for suggesting this
topic, collecting the organisms, and giving complete support in all
phases of this work; Dr. W. K. Weyrauch for bringing this specific
problem to notice: and both Drs. J. Abalos and P. Aquilar F. for
invaluable aid in the field work.

Explanation of Plate 17

Figs. 7-12. Self-burying of Sicarius sp. 1, steps 8, 9, 10-12, 16 (see Table
1 and text for details).

References

Bonnet, P.

1945-1959. Bibliographia Araneorum, Les Artisans de I’Imprimerie
Douladoure, Toulouse.

Cazier, M. A. and M. A. Mortensen

1962. Analysis of the habitat, web design, cocoon and egg sacs of the
tube weaving spider Diguetia canities (McCook). (Aranea, Di-
guetidae). Bull. So. Calif. Academy Sciences 61 (2) : 65-88.

224

Psyche

[September

Cooke, J. A. L.

1965. Systematic Aspects of the External Morphology of Dysdra cro-
cata and. Dysdera erythrina (Araneae, Dysderidae). Acta Zoo-
logica 4 6 : 41-65.

Dabelow, S.

1958. Zur Biologie der Leimschleuderspinne Scytodcs thoracica (La-
treille). Zool. Jb. Syst. 86: 85-126.

Gertsch, W J.

1949. American Spiders. D. Van Nostrand Company, Inc. Princeton,
New Jersey.

1958. The Spider Family Plectreuridae. Amer. Mus. Novitates No. 1920.

Merian, P.

1913. Les Araignees de la Terre de Feu et de la Patagonie. Rev. La
Plata 20 : 7-100.

Simon, E.

1893. Ann. Soc. Ent. France 62 Bull. ent. : CCXXIV-CCXXV.

1899. Note sur le cocon ovigere d’un sicarius du Perou. Bull. Soc. Ent.
France 1899 (19) : 367-368.

A SECOND AFRICAN SPECIES OF THE DACETINE
ANT GENUS CODIOMYRMEX 1

By Robert W. Taylor
Biological Laboratories, Harvard University.

Codiomyrmex Wheeler, sensu Brown 1948, includes 4 previously
described, worker-based species: C. thaxteri Wheeler, 1916 (Bull.
Mus. Comp. Z00L, 60 (8) : 327, Port of Spain, Trinidad) ; C.
loveridgei Brown, 1953 (Am. Midi. Nat., 50 (1): 21-23, Nyika
Plateau, Nyasaland) ; C. semicomptus Brown, 1949 (Breviora, 108:
9-1 1, Shipton’s Flat, Queensland, Australia); and C. flagellatus
Taylor, 1962 (Breviora 152: 7-9, Clump Point, Queensland,
Australia). Several others originally described in the genus have
been placed elsewhere by Brown (1948).

Codiomyrmex tetragnathus new species

Holotype. A unique worker taken in a Berlese funnel by a native
collector of the Museu do Dundo: ANGOLA: Dundo, Route Tur-
ismo (zb 70 02' S., 20° 51' E), Foret-galerie, R. Luachimo, 28-III-
1962 (ANG. 16888).

Type Deposition. Museu do Dundo, Lunda, Angola.

Description. The holotype has the following dimensions (for para-
meters of measurement and abbreviations see Brown, 1953) : TL c.
2.4 mm; HL 0.62 mm; HW 0.49 mm; Cl 79; Scape length (SL)
0.25 mm; ML 0.15 mm; MI 24; WL 0.60 mm.

Cephalic dorsum as shown in Figure 2 ; occipital lobes well devel-
oped ; short intervening transverse occipital border moderately arched,
narrowly carinate. Center of frons elevated, convex, its lateral
parts and dorsa of occipital lobes flat to feebly concave, extreme edges
forming a slightly raised flange, contours smoothly rounded through-
out. Clypeal disc flat, anterior portion sloping fairly abruptly to the
shallowly emarginate leading edge. Antennal scrobes strong, com-
pletely obscured in facial view, divided anteriorly by a narrow longi-
tudinal ridge; ventral border cariniform, ending anteroventrally in
a small tooth. Maximum ocular diameter 0.06 mm ; antennae as
usual for genus (Figure 1). Mandibles strongly convex; rising from
clypeus. Masticatory border bearing two sets of teeth: an apical

3Research supported U.S. National Science Foundation Grant No. GB 1634.

Manuscript received by the editor September 7 , 1965

225

226

Psyche

[September

series of 3 subequal denticles about 0.0 1 mm high, and a main series
of 5 strong, acute teeth of the type normally seen in Codiomyrmex.
Of the latter, the 3rd is largest (c. 0.06 mm high), 2nd and 4th
slightly smaller, 1st and 5th smallest (c. 0.05 and 0.04 mm high,
respectively). Apices of large teeth, especially the 5th, inclined
posteriorly. Denticles of apical series mounted on a process about
0.04 mm high and slightly wider at base, so that their apices are
aligned with those of the major teeth. Basal lamella arising imme-
diately behind posterior tooth, partly obscured by clypeus, apparently
triangular. Labrum transverse, distal border strongly biconvex with
a deep, acute median cleft.

Mesosomal profile as in Figure 1 ; dorsum transversely flat to feebly
concave, lateral margins defined by a fine carina. Sides almost with-
out sutures except the pro-mesonotal. Pronotum transversely mar-
ginate in front; 0.57 X as wide as head, 2.15 X as wide as propo-
deum. Humeral angles each with a low papilliform elevation. Pro-
mesonotal suture lacking on mesosomal dorsum, mesonotum and pro-
podeum separated by a fine transverse carina. Propodeal declivity
strongly concave; teeth only slightly divergent, their tips minutely
hooked upwards; infradental lamellae weak; spiracles simple, un-
flanged. Petiolar profile as in Figure 1 ; peduncle with a pair of
fine dorsolateral longitudinal carinae ; node rising to a narrow
transverse crest, its dorsum feebly convex in posterior view. Post-
petiolar disc elliptical, almost twice as wide as long. First gastric
segment 0.85 X as broad as long, 1.2 X as broad as deep; sides
submarginate basally ; basigastric costulae distinct, crowded, evenly
developed, extending back to mid-length of segment. Usual comple-
ment of areolate spongiform material on petiole and postpetiole;
posterodorsal bridges of both segments reduced to fine carina-like
vestiges; anteroventral pad lacking on gaster.

Mandibles shining, with a few scattered punctures; scapes finely
shagreened. Clypeus and anterior part of frons shining; remaining
cephalic dorsum and antennal scrohes less lucid, with coarse granular
shagreening. Mesosomal dorsum moderately shining, with effaced
shagreening overlain by vestigial fine longitudinal costulae on pro-
mesonotum, several median costulae moderately distinct. Sides of
mesosoma smooth to coarsely shagreened, generally shining, especially
on pronotum and propodeum. Propodeal declivity strongly shining.
Petiolar peduncle finely shagreened, node more coarsely so, its dorsum
with 2 fine longitudinal costulae near midline. Postpetiole finely longi-
tudinally costulate. First gastric tergum, behind basigastric costulae,
with a finely etched reticulate microsculpture. Legs shagreened.

1965]

Taylor — Genus Codiomyrmex

227

Figs. 1-2. Codiomyrmex tetragnathus n. sp. ; holotype worker: Fig. 1. Head,
mesosoma, petiole and postpetiole in lateral view. Fig. 2. Head in frontal
view.

Mandibles each with several hairs ventrally; distal edge of labrum
with a row of fine looped hairs, forming a thin tangle in its median
cleft. Body hairs few, elongate, 0.1-0.24 mm long, arranged in erect
bilateral pairs; completely lacking on head. Mesosoma, petiole and
postpetiole with 2, 1, and 4 pairs respectively, distributed as in
Figure 1. Petiolar and postpetiolar tergites also with a few fine
reclinate lateral hairs which arch back over their posterolateral fungi-
form masses. First gastric tergite with three basilateral pairs, and
1 pair on dorsum near its posterior margin.

Pubescence reduced, dense on antennae, very scattered on frons,
extremely scattered on first gastric tergite. Color very dark reddish-
brown, mandibles, antennae, legs and tip of gaster more yellowish.

Remarks. C. tetragnathus is easily distinguished from most other
members of its genus by many characters, including the head shape,
the peculiar mandibular dentition referred to in the specific name,
the structure of the propodeal teeth and infradental lamellae, the

228

Psyche

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lack of an anteroventral gastric fungiform mass, the coloration and
the extremely reduced pilosity. In the latter character this species
belies its generic name even more than does C. flagellatus.

The only other known African species, C. loveridgei , is close to
tetragnathus . It is smaller, with a straight transverse occipital bor-
der, a much more heavily sculptured head, and more dense pilosity.
The three apical mandibular teeth are reduced in size and fused
basally, much as in tetragnathus , and the anteroventral mass of spongi-
form material on the first gastric sternite is reduced to a diffuse
vestige. It is difficult to decide whether the dental heterogeniety of
these species is homologous with that seen in the related genus
Smithistruma , or whether it represents a convergently developed
specialization.

References

Brown, W. L., Jr.

1848. A preliminary generic revision of the higher Dacetini (Hymen-
optera-Formicidae) . Trans. Amcr. Ent. Soc., 74: 101-129.
1953. Revisionary studies in the ant tribe Dacetini. Amer. Midi.
Nat., 50: 1-137.

FIVE NEW SPECIES OF THE GENUS TMARUS
(ARANEAE, THOMISIDAE) FROM THE WEST INDIES

By Arthur M. Chickering
Museum of Comparative Zoology

The Genus Tmarus Simon, 1875, is widely distributed in Europe,
Asia, Africa, Australia and in the Americas. This is especially true
of South America. It appears to be abundant and highly diversified
into species in the Neotropical region. At the present time thirty
species are known from Panama alone. But, so far as I have been able
to determine, the genus has not been reported from the West Indies
until the present time. I was much interested, therefore, to find
several species of this genus in my collections made during the last
few years in Jamaica, Puerto Rico and Trinidad. These have now
been carefully studied with the result that I am obliged to recognize
five new species from these islands. Two of these species are known
from both sexes; two are known only from males and the remaining
one is known only from the female. These five species are described
in the following pages of this brief paper and may be listed as follows:
Tmarus crane ae sp. nov. ; T. farri sp. nov. ; T. insuet us sp. nov. ;
T. menotus sp. nov. ; T. vertumus sp. nov. The types will be deposited
in the Museum of Comparative Zoology at Harvard University.

Grant No. GB-1801 from the National Science Foundation made
it possible for me to spend seven months making collections of spiders
in the West Indies and Panama during the latter part of 1963 and
the first five months of 1964. This grant is also making it possible for
me to continue my studies at the Museum of Comparative Zoology
for a considerable period. A Guggenheim Fellowship made it possible
for me to collect in Jamaica, W. I. and in Panama in 1957 and 1958
and also to spend four months in the British Museum (Natural His-
tory). Dr. G. Owen Evans and Mr. D. J. Clark, Department of
Zoology (Arachnida), British Museum (N. H.) have recently
loaned me valuable specimens of the genus Tmarus to help me in my
studies. As I have frequently stated in my published papers, I am
deeply appreciative of the many privileges extended to me by the staff
of the Museum of Comparative Zoology, Harvard University, over a
period of many years. My studies could never have been carried on
as they have been continued without this encouragement. Special
acknowledgements should be extended to Dr. Ernst Mayr, Director;

229

230

Psyche

[September

Dr. P. J. Darlington, Jr., Alexander Agassiz Professor of Zoology;
Dr. Herbert W. Levi, Associate Curator of Arachnology; and Miss
Nelda Wright, Editor of Publications. These acknowledgements
should now also be extended to Dr. Frank M. Carpenter, Alexander
Agassiz Professor of Zoology and Editor of Psyche in which journal
several of my published papers have appeared.

Genus T?narus Simon, 1875

Tmarus craneae sp. nov.

Figures 1-3

This species is named after Miss Jocelyn Crane, Director, The
William Beebe Tropical Research Station, Simla, Arima Valiev,
Trinidad, W. I.

Male holotype. Total length 4.19 mm, including quite porrect
chelicerae and extended spinnerets. Carapace 1.52 mm long; 1.38
mm wide; about .53 mm tall, exclusive of the prominent lateral ocular
tubercles; with the usual long, slender spines; surface between the
spines smooth; with no median thoracic groove. Eyes: viewed from
above, posterior row strongly recurved, anterior row moderately so;
viewed from in front, anterior row straight, measured by centers.
Central ocular quadrangle wider behind than in front in ratio of
about 3:2; nearly as long as wide behind. Posterior row occupies
nearly four fifths of width of carapace at that level. Ratio of eyes
AME : ALE : PME : PLE = 5 : 10 : 6 : 10. AME separated
from one another by slightly more than twice their diameter, from
ALE by 2.6 times their diameter. PME separated from one another by
nearly three times their diameter, from PLE by 4.6 times their diam-
eter. Laterals separated from one another by 2.7 times their diameter.
Clypeus porrect in the usual manner; apparently with the usual long,
slender spines near ventral border (judged by scars) ; height, includ-
ing membranous ventral border, equal to about seven times the diam-
eter of AME. Chelicerae, maxillae, lip and sternum all apparently
typical of the genus. Legs: 1 = 243 in order of length; first and
second nearly twice as long as third and fourth; with many spines;
tarsal claws as usual; trichobothria observed on tibiae, metatarsi and
tarsi. Palp: patella short, unmodified; tibia with conspicuous

apophyses and features of the tarsal bulb as shown in Figures 1-2.
Abdomen : elongated and somewhat cylindrical ; without special mod-
ifications; with numerous long, conspicuous, slender, dorsal spines.
Color in alcohol : rather conspicuously colored ; carapace yellowish with
a narrow, red, ventral, marginal band; a dorsal, irregular, V-shaped

1965]

Chickering — Genus T mar us

231

Figs. 1-6. External anatomy of Tmarus. Figs. 1-2, T. craneae sp. nov.,
left palpal tibia and tarsus; vertral and retrolateral views, respectively.
Fig. 3, T. craneae sp. nov., another view of the ventral, tibial apopysis.
Fig. 4, T. farri sp. nov., left palpal tibia and tarsus; ventral view. Fig. 5,
T. farri sp. nov., left palpal tibia; retrolateral view. Fig. 6, T. farri sp.
nov., epigynum from below.

red mark extends from PLE and down the posterior declivity for
nearly half its decline; the clypeus bears a broad, somewhat U-shaped
red figure; each chelicera bears a red spot on its front surface; other
mouth parts generally yellowish; legs yellowish with many red dots
and larger spots and stripes; first and second femora and, to a lesser
extent, first and second patellae and tibiae, with many small, irregular,
reddish spots on the prolateral surfaces; abdomen with a mixture of
brown, white and red colors difficult to describe adequately; with a

232

Psyche

[Septembe

series of five irregular, white, transverse bars closely associated with
narrow red bars ; these separate the brown area into four more or less
rectangular blocks; lateral areas yellowish white with darker, oblique
bars; venter with a broad, very light brownish, median stripe flanked
on each side by ventrolateral, yellowish white areas. This species
seems to be closely related to T. aculeatus Chickering and others in
which the male palp has the conspicuously developed tibial apophyses.

Type locality. The holotype male is from Simla, Arima Valley,
Trinidad, W. I., April 18, 1964. Three immature specimens from
the same locality, April, 1964 probably belong to the same species.
The female is unknown.

Tmarus farri sp. nov.

Figures 4-6

The species is named after Dr. T. H. Farr, Science Museum, Insti-
tute of Jamaica, Kingston, Jamaica, W. I.

Male holotype. Total length from tip of porrect chelicerae to
posterior end of somewhat extended spinnerets 3.32 mm. Carapace
1. 1 6 mm long; almost exactly as wide as long; about .48 mm tall;
sharply constricted below posterior margin of ALE and then broadly
and regularly rounded to posterior border; with several long, slender
spines each arising from a small, chitinized tubercle ; without a median
thoracic groove or pit. Eyes: lateral ocular tubercles well developed;
viewed from above, posterior row definitely recurved, anterior row
gently so, exclusive of the very convex lenses; seen from in front,
anterior row straight, measured by centers. Central ocular quadrangle
wider behind than in front in ratio of about 13 : 8; wider behind
than long in ratio of about 26 : 23. Posterior row occupies about
10/13 of width of carapace at that level. Ratio of eyes AME : ALE :
PME : PLE = 3.5 : 8.5 : 5 : 7.5. AME separated from one an-
other by about three times their diameter, from ALE by a little more
than 3.7 times their diameter. PME separated from one another by
3.2 times their diameter, from PLE by 4.6 times their diameter.
Laterals separated by slightly more than twice the diameter of PLE.
Clypeus quite porrect; apparently with seven spines in an upturned
row but the lateral spines are small ; height equal to slightly more
than seven times the diameter of AME. Chelicerae, maxillae, and lip
all typical of males of the genus. Sternum broadly scutiform; slightly
convex; longer than wide in ratio of about 10 : 9; with many fine,
procurved bristles ; posterior end bluntly terminated opposite bases of
fourth coxae which are separated by about their width. Legs: 123 =

1965]

Chickering — Genus Tmarus

233

4 in order of length ; first and second more than twice as long as third
and fourth; numerous trichobothria observed on tibiae, metatarsi and
tarsi. Palp : reminiscent of the palp of T. ineptus O. P.-Cambridge
and T. intentus O. P.-Cambridge as these species are now under-
stood; essential features shown in Figures 4-6. Abdomen: with many
spines; about one third from base there is a pair of small, low, dorsal
tubercles each topped by a slender, stiff, somewhat recurved spine;
about two thirds from base there is a median, low tubercle correspond-
ing to the pronounced posterodorsal tubercle of the female; shortly
anterior to this reduced dorsal tubercle the abdomen is somewhat
widened ; otherwise apparently typical of males of the genus. Color in
alcohol : carapace with dorsal area yellowish in general but with small,
irregular, reddish brown spots and streaks extending from median
margins of tubercles bearing PLE to top of posterior declivity; clypeus
about the same ; lateral sides a mottled, darker, reddish brown ; central
part of posterior declivity yellowish with a central darker spot.
Chelicerae: front surface with yellowish margins, somewhat darker
centrally. Sternum yellowish with small, irregular, darker spots.
Maxillae and lip yellowish with variations. Legs: light brownish in
general ; with brownish, yellowish and reddish spots ; first two pairs of
femora conspicuously spotted with irregular brownish and whitish
spots; first two pairs of metatarsi and tarsi somewhat reddish. Abdo-
men : dorsum yellowish white with many irregular, small darker
spots; venter light brownish centrally, flanked by yellowish white
areas.

Female paratype. Total length, including somewhat porrect cheli-
cerae, 3.84 mm. Carapace 1.62 mm long; 1.56 mm wide; about .81
mm tall; with numerous spines each of which arises from a small,
chitinized tubercle ; with a series of grooves radiating over the lateral
regions from the dorsolateral areas. Eyes: lateral ocular tubercles
large; with a deep depression separating ALE from PLE; viewed
from above, posterior row recurved, anterior row slightly so, exclusive
of the very convex lenses of ALE ; viewed from in front, anterior row
slightly procurved, measured by centers. Central ocular quadrangle
wider behind than in front in ratio of about 19 : 12; wider behind
than long in ratio of 38 : 31. Posterior row occupies about 11/12 of
width of carapace at that level. Ratio of eyes AME : ALE : PME :
PLE = 4 : 10.5 : 7 : 9.5. AME separated from one another by
nearly four times their diameter, from ALE by a slightly greater dis-
tance. PME separated from one another by slightly more than 3.5
times their diameter, from PLE by about 4.7 times their diameter.

234

Psyche

[September

Laterals separated by slightly more than twice the diameter of ALE.
Clypeus quite porrect; with one erect spine beneath interval separat-
ing AME and two erect spines on each side near ventral border; height
nearly equal to eight times the diameter of AME. Mouth parts es-
sentially typical of females of the genus. Legs: 1243 in order of
length; spines are numerous and, in general, typical of females of
the genus ; trichobothria essentially as in male, also observed on palpal
tibiae. Abdomen: very spiny and irregular along dorsal region (prob-
ably associated with shrinkage following deposition of eggs) ; with
a well developed posterodorsal tubercle projecting upward at an
angle ; somewhat the widest in front of base of posterodorsal tubercle.
Epigynum : essential features shown in Figure 6 ; some variations
noted among paratypes. Color in alcohol : essentially as decribed for
male but all colors are more vivid than in that sex; first two pairs of
tarsi and metatarsi not reddish as in male but irregularly yellowish
brown.

Type locality. Male holotype and female paratype taken on north-
east slope of Long Mt., St. Andrew Parish, Jamaica, W. I., October
26th, 1957. Numerous paratypes of both sexes were taken with the
holotype and also in St. Catherine Parish, Hanover Parish, and on
other parts of St. Andrew Parish in 1954, 1957 and 1963.

Tmarus insuetus sp. nov.

Figures 7-8

The name of the species is a Latin adjective suggested by the
unusual type of male palp.

Male holotype. Total length 4.03 mm. Carapace 1.5 mm long;
1.5 mm wide; .63 mm tall; surface very finely granulate; with numer-
ous long, slender spines; posterior declivity gently arched, not steeply
slanted. Eyes : ocular tubercles less prominent than usual in the genus
but otherwise typical of the genus; viewed from above, posterior row
rather strongly recurved, anterior row moderately so; viewed
from in front anterior row slightly recurved, measured by
centers. Central ocular quadrangle wider behind than in front
in ratio of 1 1 : 8; longer than wide behind in ratio of 13 : 11.
Posterior row occupies about 6/7 of width of carapace at that level.
Ratio of eyes AME : ALE : PME : PLE = 4:10:5:9. AME
separated from one another by twice their diameter, from ALE by
2.75 times their diameter. PME separated from one another by 2.6
times their diameter, from PLE by nearly five times their diameter.
Laterals separated from one another by about 2.2 times the diameter

1965]

Chickering — Genus Tmarus

235

Figs. 7-11. External Anatomy of Tmarus. Fig. 7, T. insuetus sp. novv
left palpal patella, tibia and tarsus; ventral view. Fig. 8, T. insuetus sp.
nov., left palpal patella, tibia and tarsus; retrolateral view. Figs. 9-10, T.
menotus sp. nov., female abdomen: dorsal and left lateral view; respectively.
Fig. 11, T. menotus sp. nov., epigynum from below.

of ALE. Clypeus only moderately porrect ; with a row of long, slen-
der spines near ventral border; height, including membranous ventral
border, nearly equal to six times the diameter of AME. Chelicerae,
maxillae, lip and sternum all apparently typical of the genus. Legs:
1 — 234 in order of length; first two pairs very long and slender and
with numerous, long, slender spines; third and fourth less than half
as long as first and second and with less conspicuous spines; tarsal
claws and trichobothria apparently typical of the genus. Palp: com-

236

Psyche

[September

plicated ; patella with unusual apophysis ; tibia with a group of short
apophyses; tarsus with embolus encircling the bulb more than three
times (Figs. 7-8). Abdomen: cylindrical through anterior two thirds
and then much narrowed to posterior end; only a slight indication
of a median posterodorsal tubercle at about the beginning of the last
quarter. Color in alcohol : less colorful, in general, than usual in the
genus. Carapace: yellowish with a reddish streak around the medial
borders of the lateral ocular tubercles. Mouth parts and sternum
yellowish. Legs yellowish ; lacking the conspicuous markings character-
istic of many species in the genus. Abdomen : yellowish in general ;
dorsum with numerous white and reddish spots and transverse bands ;
venter with a broad, very light brown, central stripe flanked on each
side by whitish areas.

Type locality. The male holotype is from Simla, Arima Valley,
I rinidad, W. I., April 23, 1964. One male paratype is in the collec-
tion from the same locality, April 17, 1964. The female is unknown.

Tmarus menotus sp. nov.

Figures 9-1 1

The name of the species is an arbitrary combination of letters.

Female holotype. Total length 5.2 mm, including somewhat por-
rect chelicerae (length increased somewhat by stretching in preserva-
tion). Carapace 1.65 mm long; 1.54 mm wide; about .7 mm tall;
otherwise quite typical of females of the genus. Eyes : ocular tubercles
as usual in females ; viewed from above, posterior row moderately re-
curved, anterior row gently so, exclusive of the very convex lenses of
ALE ; viewed from in front, anterior row nearly straight, measured
by centers. Central ocular quadrangle wider behind than in front in
ratio of 19 : 12; wider behind than long in ratio of 19 : 15. Posterior
row occupies a little more than 4/5 of width of carapace at that level.
Ratio of eyes AME : ALE : PME : PLE = 5 : 12 : 7 : 10. AME
separated from one another by about three times their diameter, from
ALE by a slightly greater distance. PME separated from one an-
other by slightly more than 3.5 times their diameter, from PLE by
nearly 4.25 times their diameter. Laterals separated by a little more
than twice the diameter of PLE. Clypeus quite porrect; with a row
of seven slender spines; height nearly equal to seven times the diameter
of AME. Chelicerae, maxillae, lip and sternum all apparently typical
of females of the genus. Legs : 1 = 243 in order of length ; with
numerous spines; trichobothria observed on tibiae, metatarsi and tarsi;
tarsal claws as usual. Abdomen: short, broad and tall (Figs. 9-10) ;

1965]

Chickering — Genus Tmarus

237

Figs. 12-15. External Anatomy of Tmarus. Figs. 12-13, T. vertumus sp.
nov., left palpal tibia and tarsus; ventral and retrolateral views, respectively.
Fig. 14, T. vertumus sp. nov., female abdomen; dorsal view. Fig. 15, T .
vertumus sp. nov., epigynum from below.

with a short, obtuse, posterodorsal tubercle; spines short, inconspicu-
ous; other features essentially typical of females of the genus. Epigy-
num: essential features shown in Figure 11; obscurely distinctive;
unlike any other seen during my study of the genus in Panama and the

238

Psyche

[September

West Indies. Color in alcohol: carapace yellowish in general, with
a mottled appearance ; with many fine, brownish dots ; bases of largest
spines usually brownish ; posterior declivity a medium brown. Legs
and mouth parts yellowish in general; prolateral surfaces of first two
pairs of femora, patellae and tibiae very light yellowish with darker
spots. Abdomen: dorsum light yellowish in general but with many
fine, dark dots; posterolateral surfaces a medium brown; venter with
a narrow, median, light brown stripe flanked on each side by a very
light yellowish area.

Type locality. The female holotype is from St. Catherine Parish,
six miles east of May Pen, Jamaica, W. I., November 22, 1957.
There are no paratypes and the male is unknown.

Tmarus vertumus sp. nov.

Figures 12-15

The name of the species is an arbitrary combination of letters.

Male holotype. Total length, including somewhat porrect cheli-
cerae and extended spinnerets, 4.1 mm. Carapace 1.43 mm long; 1.41
mm wide ; about .65 mm tall ; ventral margin regularly rounded from
opposite interval between ALE and PLE to posterior border; other-
wise quite typical of males of the genus in the neotropical region.
Eyes: lateral ocular tubercles well developed; those bearing AME and
PME moderately so; viewed from above, posterior row rather
strongly recurved, anterior row gently so, exclusive of the strongly
convex lenses of ALE; viewed from in front, anterior row slightly
procurved, measured by centers. Central ocular quadrangle wider be-
hind than in front in ratio of about 3:2; wider behind than long in
ratio of about 6:5. Posterior row occupies nearly .8 of width of
carapace at that level. Ratio of eyes AME : ALE : PME : PLE =
4 : 9.5 : 6 : 9. AME separated from one another by about three
times their diameter, from ALE by 3.5 times their diameter. PME
separated from one another by slightly more than three times their
diameter, from PLE by 4.3 times their diameter. Laterals separated
by slightly more than twice the diameter of ALE. Clypeus : quite por-
rect; with the usual row of seven spines near ventral border; height
equal to seven times the diameter of AME. Chelicerae, maxillae, lip
and sternum all apparently typical of the genus. Legs: 123 — 4 in
order of length ; spines about as usual ; tarsal claws typical ; tricho-
bothria observed on tibiae, metatarsi and tarsi. Palp : essential features
shown in Figures 12-13; femur of moderate length; patella short, un-
modified ; tibia with two conspicuous apophyses ; tarsus with a hook
near center of bulb. Abdomen : first three fifths somewhat cylindrical

1965]

Chickering — Genus T marus

239

in form; not flattened; widened a little about 2/3 from base; a short
distance posterior to widened region there is a small, median, dorsal
tubercle corresponding to the more conspicuous tubercle in the fe-
male ; other features essentially typical of the genus. Color in alcohol :
carapace yellowish in general ; with many minute reddish dots and
brownish spots and streaks impossible to describe in detail. Legs:
yellowish with many small, reddish spots and streaks; first two pairs
of femora with the characteristic mottled prolateral and retrolateral
surfaces; third and fourth legs with more reddish spots than other
legs. Sternum yellowish with small, pinkish dots irregularly distrib-
uted near margin. Abdomen : dorsum pinkish in general ; a very
narrow, reddish band extends across the widened region ; irregular,
reddish streaks occur in the posterior third of the dorsum and an ir-
regular, narrow, reddish band separated in the middle occurs about
half way between the base and the widened region; lateral sides
yellowish with reddish dotst and streaks; venter very light brown
through the center flanked by whitish areas.

Female paratype. Total length, including porrect chelicerae and
somewhat extended spinnerets, 5.91 mm; total length from anterior
border of clypeus to posterior end of anal tubercle 5.4 mm. Cara-
pace 1.8 mm long; 1.65 mm wide; about .7 mm tall; with about
twelve long, slender spines around border of posterior declivity.
Eyes : curvature of rows essentially as in male. Central ocular quad-
rangle wider behind than in front in ratio of about 3:2; wider be-
hind than long in ratio of nearly 4:3. Ratio of eyes AME : ALE :
PME : PLE = 5 : 12 : 7 : 10. AME separated from one another
by a little more than three times their diameter; from ALE by
slightly more than 3.5 times their diameter. PME separated from
one another by about 3.5 times their diameter, from PLE by a little
more than 4.6 times their diameter. Laterals separated by 2.3 times
the diameter of PLE. Height of clypeus nearly equal to eight times
the diameter of AME. Chelicerae, maxillae, lip and sternum all typi-
cal of females of the genus. Legs: 123 — 4 in order of length; spines,
tarsal claws, palpal claws and trichobothria all typical of females of
the genus studied by the author. Abdomen : considerably more robust
than in male; much widened two thirds from base and then sharply
narrowed to posterior end ; posterodorsal tubercle somewhat more
prominent than in male (Fig. 14). Epigynum : obscurely distinctive;
unlike any other seen in the genus from the Neotropical region; es-
sential features shown in Figure 15; some variations noted among
paratypes. Color in alcohol: essentially like that of male.

Type locality. Male holotype from university farm east of campus,

240

Psyche

[September

Mayaguez, Puerto Rico, W. I., January 15, 1964; female paratype
from university farm north of campus, Mayaguez, January 17, 1964.
Several paratypes of both sexes were taken with the described male
and female types, at other localities in the same general region and
on Route 106, 5 km. east of Mayaguez during January and February,
1964.

Selected bibliography

Bonnet, Pierre

1959. Bibliographia Araneorum. Toulouse. Vol. 2 (5).

Cambridge, O. P.- and Cambridge, F. P.-

1889-1905. Arachnida-Araneida. Vols. I-II. In: Biologia Centrali-
Americana. Dulau & Co. London.

Chickering, A. M.

1950. The Spider Genus Tmarus (Thomisidae) in Panama. Bull. Mus.

Comp. Zool. at Harvard College, 103 (4), pp. 213-255, 4 pis.
1965. Panamanian Spiders of the Genus Tmarus (Araneae, Thomisidae).
Ibid. In press.

Roewer, C. Fr.

1954. Katalog der Araneae. Vol. 2, Pt. 1. Brussels.

A KEY TO NORTH AMERICAN STATIRA
(COLEOPTERA: LAGRIIDAE)

By Carl T. Parsons
Manchester Depot, Vermont

From time to time during the past several years I have studied
the Lagriidae with C. W. Leng’s excellent collection as a basis. To
this collection has been added valuable series especially from the
southwest and Mexico generously presented by Dr. Floyd Werner.
As indicated in the table of abbreviations below, various public and
private collections have been examined. I am deeply indebted to the
respective curators and individuals for extending this privilege.*

In the course of this study a few additions to our fauna turned
up and a few changes seem advisable. Much more material is needed
to clear up still pending problems. An account of Arthromacra will
follow.

Key to North American Statira

1. At least middle and hind tibiae distinctly sulcate along nearly

entire outer edge; disc of pronotum scabrous or rugulose .2
Tibiae on outer edge rounded or flat (rarely feebly sulcate
distally) as in dejecta, disc of pronotum variably punctate,
intervals between punctures usually alutaceous, rarely smooth

5

2. Setigerous punctures on first elytral interval not more than 6,

on 3rd not more than 8, on 5th not more than 5, on 7th not

more than 2, on 9th not more than 5 3

Setigerous punctures on first elytral interval not less than 10, on
3rd not less than 13, on 5th not less than 12, on 7th not less
than 10, on 9th not less than 11 4

3. Lateral margin of prothorax obliterated anteriorly, becoming

distinct posteriorly; pronotum finely scabrous, rich dark

^Specimens studied are referred to by abbreviations as follows:
(A.M.N.H.), American Museum of Natural History; (A.N.S.P.), Academy
of Natural Sciences Philadelphia; (C.A.F.), C. A. Frost collection in M.C.Z. ;
(C.T.P.), C. T. Parsons collection, Manchester Depot, Vermont; (C.U.),
Cornell University; (F.P.B.), Florida Plant Board, Gainesville; (H.C.F.),
H. C. Fall collection in M.C.Z. ; (H.F.H.), Henry F. Howden collection;
(111. N. H. S.), Illinois Natural History Survey; (M.C.Z.), Museum of Com-
parative Zoology; (U.S.N.M.), United States National Museum.

Manuscript received by the editor March 16, 1965.

241

242

Psyche

[Septembe

brown; labrum, base of antennae, and legs paler; subopaque;
in both sexes apical antennal segment equal to next 2^4 seg-
ments subnitida

Lateral margin of prothorax evanescent anteriorly but just dis-
cernible; pronotum densely, minutely, punctulate, rugulose,
and opaque; piceous, except prothorax, scutellum, legs, and
basal two antennal segments rufo-testaceous; male apical an-
tennal segment equal to next 3^4 segments color ata

4. Lateral margin of prothorax almost always obliterated anteriorly

but starting at the middle becoming distinct posteriorly, rarely
the margin continuing obsoletely to the anterior edge; margins
of sulcus on fore and middle tibiae similar; pronotum finely
rugulose and scabrous, much finer than eye facets; 1st elytral
interval with 11-16 setigerous punctures, 3rd with 14-21, 5th
with 12-20, 7th with 10-17 and 9th with 12-20 setigerous

punctures pluripunctata

Lateral margin of prothorax entire and distinct; either anterior
or posterior margin of sulcus on fore and mid tibiae raised
to form a blackish carina; pronotum moderately coarsely
scabrous, about as course as eye facets; 1st elytral interval
with 20-28, 3rd with 21-26, 5th with 22-26, 7th with 22-28,
and 9th with 22-35 setigerous punctures hirsuta

5. Elytra testaceous to rufous with brown to black markings; no

setigerous punctures on 5th elytral interval (except rarely 1

at apex) 6

Elytra unicolorous, usually dark (testaceous in teneral examples) ;
several setigerous punctures on 5th elytral interval 7

6. Male apical antennal segment equal to next 5 segments, in female

equal to next 3/4-4 segments; testaceous to rufous, elytra
slightly paler, with large scutellar spot and transverse fascia,
slightly behind middle and wider at sides than at middle,
brown or black; O-i setigerous puncture at base of 1st elytral
interval, 3rd elytral interval with 4-6 from base to apex, 7th

with 2 at base, 9th with 4 along apical fourth pulchella

Male apical antennal segment equal to next 4-4 1/5 segments,
in female equal to next 3 segments; rufo-testaceous, median
black spot on each elytron not reaching the suture; 3rd elytral
interval with 5-9 setigerous punctures, 7th with 2 at base,
9th with 3-4 along apical fourth nigromaculata

7- Setigerous punctures on 1st elytral interval 19-26, on 3rd interval
18-33, on 5th interval 17-23, on 7th interval 8-13, on 9th
interval 8-22 8

1965]

Persons — Statira

243

Setigerous punctures on 1st elytral interval 2-5, on 3rd interval
4-15, on 5th interval 4-1 1, on 7th interval 2-8, on 9th interval

4- i 1 9

8. Setigerous punctures on 1st elytral interval about 25, on 3rd

interval 26-33, on 5th interval 17-23, on 7th interval 8-13,
on 9th interval 8-17; setigerous punctures about as large as
strial punctures, pronotum opaque, male apical antennal seg-
ment equal to next 4% segments, in female equal to next

3S/4 ’> length 9-1 1 mm. opacicollis

Setigerous punctures on first elytral interval 19-25, on 3rd in-
terval 18-21, on 5th interval about 18, on 7th interval about
8, on 9th interval about 24; setigerous punctures at least
twice as large as strial punctures; pronotum sub-opaque; male
apical antennal segment equal to next 3%-4, in female equal
to next 3; length 12. 5-14 mm huachucae

9. Fore and middle tibiae rounded on outer edge; male apical an-

tennal segment equal to next 3%-7 segments, in female equal

to next 2 >4-4 segments 10

Fore and middle tibiae flat on outer edge (rarely feebly sulcate
distally) ; male apical antennal segment equal to next 2 >4-2%
segments, in female equal to next 2-2 >4 segments; disc of
pronotum very densely and finely punctate, the punctures less
than their diameters apart, the surface finely alutaceous; setig-
erous punctures on 1st elytral interval 4-5, on 3rd interval
3-6, on 5th interval 3-5, on 7th interval 1-3, on 9th interval

5- 6 defect a

10. Disc of pronotum with punctures on the average at least 3 times

their diameter apart; elytra brown to black (except bluish in

croceicollis T 1

Disc of pronotum with punctures separated by about their dia-
meter; rufous, legs testaceous, abdomen piceous, elytra black
with metallic bluish-green lustre; setigerous punctures on 3rd
elytral interval 5-7, on 5th interval 4-5, on 7th interval 2 at
base, on 9th interval about 6; male apical antennal segment
equal to next 5 >4-5% segments, in the female equal to next

3/4 segments ; length 9-10 mm liebecki

II. Setigerous punctures on 3rd elytral interval 7-15 12

Setigerous punctures on 3rd elytral interval 3-5 13

12. Length 10.5-12.5 mm.; male apical antennal segment equal to
next 3%-4 1/3 segments, in female equal to next 2 >4-3 seg-
ments; brown to piceous black, head darker; setigerous punc-
tures on 3rd elytral interval 10-15, on 5th interval about 11,

244

Psyche

[September

on 7th interval 5 on basal three-fifths, on 9th interval 8-1 1

rohusta

Length 7-9.5 mm.; male apical antennal segment equal to next
4^-5 segments, in female equal to next 3-3/2 segments; pro-
thorax and legs testaceous or rufous, head piceous, elytra
piceous or black, the elytra with a distinct bluish lustre;
setigerous punctures on 3rd elytral interval 7-14, on 5th in-
terval 7-12, on 7th interval either 2 at base or 6-8 all along,
on 9th interval 5-13 croceicollh

13. Setigerous punctures of elytra about the same size as the strial

punctures: larger, 8-11.5 mm 14

Setigerous punctures of elytra about twice as large as the strial
punctures; smaller, 6.5~8.8 mm. 15

14. Posterior margin of fifth ventral segment produced around a

distinct fovea; male apical antennal segment equal to next
4 2/3-5 segments, in the female equal to next 3 2/3-3 3/4 seg-
ments; color paler, tending to brown rather than piceous,

elytra tending to brown especially along sutural margins

dolera n. sp.

Posterior margin of fifth ventral segment simple, without fovea;
male apical antennal segment equal to next 6/2-7 segments,
in the female equal to the next 4 segments or slightly less;
color darker, tending to piceous black, elytra sometimes slightly

more pale and with very narrow brown sutural margins

basalts

15. Pronotum brownish piceous to piceous black usually concolorous

with elytra; male apical antennal segment equal to next 5/2
segments, in the female equal to next 3 1/3 segments, range
Vermont to Wisconsin south to North Carolina and Tennessee

gag atm a gagatina

Pronotum testaceous to brownish rufous, head and elytra brown-
ish piceous to piceous black; male apical antennal segment
equal to next 5/2 -6/2 segments, in the female equal to next
31/3 segments; range more southern, New York City to
Alabama, also Michigan gagatina resplendens

Statira snbnitida Leconte

Statira subnitida Lee., 1866, Smithsonian Misc. Coll., Wash., 167: 141-142.

Types: lectotype cf (not 9 as Horn stated) no. 4749 and 1$ cotype

collected by John Xantus at Cape San Lucas, Baja California, in
M.C.Z.

Range: Known only from Baja Calif ronia, Mexico, from the two

1965]

Parsons — Statira

245

types and female from Santa Rosa (H.C.F. in M.C.Z.). In the
Horn collection (A.N.S.P.) are three specimens identified as sub-
nitida in Horn’s handwriting. The first from San Jose del Cabo is
colorata and the other two are dejecta from “Ariz”.

Statira colorata Fall

Statira colorata Fall, 1909, Canadian Entomologist, 41: 165-166.

Types: holotype no. 24613 collected by Charles Fuchs at San
Jose del Cabo, Baja California, Mexico (H.C.F.) With identical
data are two males in the Casey collection (U.S.N.M.) and one male
in the Horn collection (A.N.S.P.)

Statira pluripunctata Horn

Statira pluripunctata Horn, 1888, Trans. Amer. Ent. Soc. 15: 29.

Statira sulcicrus Champion, 1889, Biol. Centr.-Amer. Coleop. 4(2): 51-52.
NEW SYNONYMY.

Types: of pluripunctata lectotype $ no. 8018 from Arizona

(A.N.S.P.) and 1 9 evidently a cotype in the Leconte coll. (M.C.Z.)
Of sulcicrus 2 cT cT and 2 $9 cotypes collected by Hoge at Chilpan-
cingo, Guerro, Mexico in the British Museum.

In the United States this common species is most nearly allied to
hirsuta. But in must be nearer pucblensis Champion (not seen),
which is said to be more stout, antennae less slender, hairs on femora
longer, and pronotum densely, finely, and irregularly punctate and
wrinkled, with prothorax more distinctly margined. Champion’s
sulcicrus is made a synonym from description only.

G. H. Nelson has collected in Arizona two examples which do
not key out. A male taken on Ephedra trifurca, Aug. 28, 1954, at
Portal has setigenous punctures on base only of first elytral interval
and half the normal number of punctures on the other intervals. The
other specimen, collected at light in Sabino Canyon, Santa Catalin
Mts. Aug. 25, 1959, has typical male terminalia but only 4 setigerous
punctures on first, 10 punctures on third, and 7 punctures on fifth
elytral interval.

Range: This species occurs from June 20 to Sept. 20, chiefly late
July to earlv August in southern Utah: state label (C.T.P.), St.
George (C.U., A.M.N.H.F South Creek, Beaver Co. (C.U.)
Texas: Alpine (C.U., H.C.F.), 2 and 15 miles west of Fort Davis
(H.F.H.) New Mexico: La Cueva. Organ Mts. (H.C.F.); Ari-
zona: Cochise Co., Santa Cruz Co., Pima Co., Pinal Co., Gila Co.,
Maricopa Co., Yavapai Co.; Mexico: Sonora (A.M.N.H.), Chihua-
hua (A.M.N.H., C.T.P.) Durango (A.M.N.H., C.T.P.), Guerrero
(B.M.)

246

Psyche

[September

Habitat: It has been collected on cotton, Datura meteloides , and
from leaf axils of Yucca with dead flowers in them. Werner and
Nutting have taken it at light in the following plant associations in
Arizona (C.T.P.) : Larrea , mesquite-cholla, chaparral, sycamore-
oak-ash, and mesquite-desert grassland.

Statira hirsuta Champion

Statira hirsuta Champion, 1889, Biol. Centr.-Amer. Coleop. 4(2) : 50-51.
Statira simulans Schaeffer, 1905, Journ. New York Ent. Soc. 13: 180. new
SYNONYMY.

Types: of hirsuta cotypes from “Mexico (coll. F. Bates), Jalapa,
Iguala in Guerrero, Tapachula in Chiapas (Hoge) ; Nicaragua,
Chontales (Belt) all in the British Museum. Jalapa is here designated
the type locality since a series, of which two are in the A.M.N.H.,
was collected there. Of simulans from Brownsville, Texas
(U.S.N.M.).

Range: In the United States hirsuta occurs from June 7 to August
8 in southeastern Texas at Brownsville, usually at the Esperanza
Ranch. Also in the authors’s collection is a specimen taken by Henry
Wenzel in Dimmit Co., 250 miles northwest of Brownsville. The
species extends through Mexico: (May 24 to August 22) Nuevo

Leon, Tamaulipas, Vera Cruz, Mexico City, Guerrero, Chiapas to
Nicaragua.

Habitat: F. Werner and W. Nutting collected hirsuta at light in
desert shrub in Nuevo Leon and at light in mesophytic forest in
Tamaulipas.

Statira pulchella Maklin

Statira pulchella Maklin, 1863, Act. Soc. Sci. Fenn. 7: 589. Reprinted April
13, 1863, pp. 101-2. Figured by Champion, 1889, Biol. Centr.-Amer.
pi. 2, fig. 8.

Type: from Mexico presumably in the Zool. Mus. in Helsinki.
Range : This species occurs in the United States in extreme south-
eastern Texas at Brownsville where J. N. Knull collected a series
on June 1-8, 1934 (H.C.F.) In Mexico it has been taken on the
Atlantic slope in Vera Cruz: Jalapa (B.M., A.M.N.H.), Cordova
(B.M.), Atoyac (B.M.) and in eastern San Luis Potosi: Tama-
sunchale (C.T.P.)

Habitat: F. Werner and W. Nutting took the Tamasunchale speci-
men at light in open river bottom on May 30.

Statira nigromaculata Champion

Statira nigromaculata Champion, 1889, Biol. Centr.-Amer. Coleop. 4(2):
33-34, pi. 2, fig. 9.

1965]

Parsons — Statira

247

Types: from Mexico: Jalapa in Vera Cruz and Yolos in Oxaca;
Guatemala: San Geronimo in the British Museum.

Range: This rare species occurs in southeastern Arizona: Aug. 6,
1948, Geronimo, Graham Co. (C.T.P.) ; July 1 - 1 5 , 1923, Babo-
quivari Mts. (H.C.F.) ; July 18, 1948, Sonoita River, Patagonia,
Santa Cruz Co. (A.M.N.H.) ; at light July 7, 1957 and at light
July 26, 1948, Sabino Canyon, Santa Catalina Mts. (C.T.P.,
C.A.F.) ; Texas: July 20, 1956, 2 miles west of Ft. Davis (H.F.H.,
G.N., C.T.P.) ; July 26, 1956, Limpia Canyon near Ft. Davis
(C.U.) ; Mexico: July 14 and 15, 1947, Camargo and 15 miles east
of Parrah in Chihuahua (A.M.N.H.) ; also the above type localities.
Habitat: H. & A. Howden took a series at Ft. Davis on willow.
Werner and Nutting took a series at light in the sycamore-oak-
mesquite association in Sabino Canyon and the Geronaimo, Arizona
specimen in willow-mesquite-cottonwood association.

Statira opacicollis Horn

Statira opacicollis Horn, 1888, Trans. Amer. Ent. Soc. 15: 30.

Types: lectotype cf no. 8017 collected by Morrison in Arizona
(A.N.S.P.). Two evident cotypes are in the Leconte coll. (M.C.Z.)
Range : This rare species is known definitely from southeastern

Arizona in Gila Co.: Sierra Ancha Mts. (C.U.) and Cochise Co.:
Chiricahua Mts. A series of 67 specimens were taken by M. Statham
May 3-19, 1956 at 5400 ft. five miles west of Portal, Cochise Co.,
Arizona (A.M.N.H., C.T.P., C.A.F.)

Statira huachucae Schaeffer

Statira huachucae Schaeffer, 1905, Science Bull, of the Museum of the
Brooklyn Inst, of Arts and Sciences 1(7) : 176.

Types: cotype J1 and cotype $ both no. 42558 (U.S.N.M.), both
labelled “type” in Schaeffer’s writing. They were collected by
Charles Schaeffer June 13, 14 in the Huachuca Mts. Arizona.
Range: This rare species is known from southeastern Arizona: July
14, 1952, Madera Canyon, Santa Rita Mts. in Pima Co., George
Bradt (A.M.N.H.) ; July, Palmerlee, Cochise Co. (C.T.P.) ; and
the following records from the Huachuca Mts.: July 1905 (C.T.P.) ;
July 14, 1928, 6000 ft., A. A. Nichol (C.U.) ; July 14, 1928, 6000
ft., A. A. Nichol (C.U.) ; June 24 and July 13 (C.U.).

Champion (1889, p. 30) described Statira alternans based on a
unique male from Tepansacualco, Mexico. It is possible that hua-
chucae and alternans are identical. A specimen that is questionably
alternans was taken June 4, 1948 at 8000 ft. at Gaborachie, Chihua-
hua, Mexico by George Bradt (A.M.N.H.).

248

Psyche

[September

Static a dejecta Schaeffer

Statira dejecta Schaeffer, 1905, Science Bull., Mus. Brooklyn Inst, of Arts
and Sciences. 1(7): 175.

Type: Holotype no. 42557 Palmerlee, Cochise Co., Arizona

(U.S.N.M.)

Range : This common species occurs in southeastern Arizona and
adjoining New Mexico from July 5 to September 3, preponderantly
during the latter half of July. In Arizona it extends from extreme
southern Navajo Co., Gila Co., Pima Co., Santa Cruz Co., to
Cochise Co., and across into Hidalgo Co., New Mexico.

Habitat: Werner and Nutting have taken it at light in the follow-
ing plant associations in Arizona (C.T.P.) : cholla-mesquite, chapar-
rel, oak-hackberry, sycamore-oak-mesquite, and rich mesophytic valley
in pine zone.

Statira liebecki Leng

Statira liebecki Leng, 1923, Journ. New York Ent. Soc. 31: 185-187.

Types: holotype cf from Enterprise, Florida and allotype $ collected
by H. P. Loding May 15, 1919 at Spring Hill, Alabama, both in
author’s collection (ex coll. C. W. Leng).

Range: This very rare spring species is known from the following
records in addition to the above mentioned types: two male para-
types no. 26543 from Enterprise, Florida (H.C.F. ex Liebeck coll.) ;
one male collected by H. P. Loding at Orchard, Alabama (H.C.F.) ;
one female paratype, April 24, Crescent City, Florida (A.M.N.H.)
and one paratype collected by W. T. Davis, April 29 at South Bay,
Lake Okeechobee, Florida in Staten Island Inst, of Arts and Sci. ;
one male collected and presented by H. V. Weems, Jr. at Highland
Hammock State Park, Florida on March 20. (C.T.P.)

This species appears to be closely related to or possibly identical
with var. a of Statira mexicana Champion which occurs on the
eastern slope of Mexico.

Statira robusta Schaeffer

Statira robusta Schaeffer, 1905, Journ. New York Ent. Soc. 13: 180.

Type: holotype $ no. 42556 collected by Charles Palm at Globe,
Arizona (U.S.N.M.)

Range: This species occurs May to August in Colorado: Colorado
Springs (111. N. H. S.) ; Arizona: Yavapai Co.; Maricopa Co., Gila
Co., Cochise Co.; New Mexico: San Miguel Co., Otero Co.; Texas:
Brewster Co.

Habitat: Werner and Nutting collected robusta on July 9 at light

1965]

Parsons — Statira

249

in the pinon-juniper-oak association in the Chisos Mts. of Texas

(C.T.P.) •

Evidently robusta is closely related to or possibly identical with
tuberosa which Champion described, p. 31, from one female from
Jalapa, Mexico. So far no Mexican examples have been seen.

Statira croceicollis Maklin

Statira croceicollis Maklin, 1863, Act. Soc. Sci. Fenn. 7: 594. ReprintedApril
13, 1863, p. 106.

Type: collected by Motschulsky at Mobile, Alabama; presumably
in the Zool. Mus. in Helsinki.

Range: This early spring species has been collected during March 9
to April 24 most often in peninsular Florida: Dunedin (H.C.F.,
C.U.), Enterprise (C.T.P., H.C.F.), Jupiter (M.C.Z., 111.

N.H.S.), Kissimmee (A.M.N.H.), Lake Worth (C.U.), Biscayne
(A.N.S.P.), Miami (CT.P.), Paradise Key (H.C.F., C.A.F.), Key
Largo (A.M.N.H.) There are two specimens from Georgia: St.
Simons Island (C.T.P.) , “Geo.” (H.C.F.) ; two specimens from
Mississippi: Ocean Springs (C.U.), Lucedale (C.U.) ; and a female
collected by A. Nicolay on June 23, 1933 at Chesapeake Beach,
Maryland (H.C.F.)

Statira dolera new species
fig. 1.

Having the appearance of a teneral basalts but a little larger and
distinctly larger than gagatina. The color tends to be more pale
than basalts with the sutural margins of the elytra more pale than
the rest of the elytra ; shining, piceous, with the head darker and the
elytra usually more pale, legs testaceous to brown with a tendency
for the apices of the femora to be darker. Antennae and anterior
part of head rufous. Vertex narrower than in gagatina, densely
rather coarsely punctate, much more coarsely punctate than prono-
tum but a little more finely punctate than elytral striae. Eleventh
antennal segment as long as 10th, 9th, 8th, 7th, and Y\. of 6th to
all of 6th segments combined in the male and as long as the 10th,
9th, 8th, and 2/3-3/ 4. of 7th segments combined in the female.
Prothorax of holotype with width/length as 1/.90, in the allotype as
1 /-9 3 i anterior margin truncate or very feebly emarginate, sides
more rounded than in gagatina or basalis, posterior margin truncate,
lateral line distinct and entire, disc very finely and sparsely punctate,
the punctures about 6X their diameters apart, the intervals very
finely granular. In the allotype and some paratypes the punctures

Psyche, 1965

Vol. 72, Plate 18

7

8

9

n

Parsons — Statira

1965]

Parsons — Statira

251

may be variably obsolete and the intervals almost smooth. Scutellum
distinctly or obsoletely granular and distinctly or obsoletely very
finely punctate.

Elytra striate with closely placed punctures along the striae, the
intervals moderately convex and with the setigerous punctures the
same size as the strial punctures. In presenting the numbers of
setigerous punctures along the elytral intervals the first figure is of
the holotype, the second where present of the allotype, and figures in
parentheses represent variations in the paratypes. First elytral interval
with 1 setigerous puncture at base and 2 (3) at apex, 3rd interval
with 6, 5 (5-8) setigerous punctures placed all along, the 5th interval
with 6, 4 (4-6), 7th interval with 2, 2 at base (there may also be
1 at apex), 9th interval with 6, 7 ( 5- 1 1 ) along apical half. Femora
glabrous, tibiae rounded on outer edge. In both sexes the posterior
margin of the fifth ventral segment is produced around a distinct
fovea, a character lacking in our other species. Male terminalia as
figured. Holotype: length 10.7 mm., width of prothorax 1.95 mm.,
width of elytra at humeri 2.9 mm.; allotype: length 9.3 mm., width
of prothorax 1.6 mm., width of elytra at humeri 2.5 mm. For this
species the dimensions of the holotype are typical whereas dimensions
of the allotype are minimum.

Range: All the specimens were taken from February 19 to May 1-12
in Florida as follows. Holotype cf Gainsville, H. L. Dozier (C.T.P.
ex C. W. Leng) ; allotype 9 April 1, 1947, Lake Placid, J. W.
Green ( C.T.P. ). The following are designated paratypes. 19 April,
Enterprise (C.T.P.) ; i$ Key West (C.T.P.) ; 2 9 9? 1 cf Florida
(C.T.P., H.C.F.) ; 1 cf March 5, 1939, Edgewater, C. A. Frost
(C.A.F.); 2d* cf April 6, 1929 Dunedin, W. S. Blatchley (C.U.,
C.A.F.), 9 washup Feb. 26, 1939 Coronado Beach, C. A. Frost
(C.A.F.) ; 1 cf Feb. 19, Crescent City, Hubbard & Schwarz

(U.S.N.M.) ; 1 cf Florida, Hubbard & Schwarz (U.S.N.M.) ; 1 cf
March 1, Haulover, Hubbard & Schwarz (U.S.N.M.) ; i9 March
5, 1948, Gainsville, L. A. Hetrick (Florida Plant Board); Florida,
A. T. Slosson (A.M.N.H.); 1 cf Lake Worth, A. T. Slosson
(A.M.N.H.) ; 2 Ormond, A. T. Slosson (A.M.N.H.) (M.C.Z.) ;
1 cf Orange Grove, O. Seifert (C.T.P.) ; 19 March 20, 1955 and
3 cfcf, 48 99 March 23-30, 1954, Oneco, Manatee Co. (C.U.,

Explanation of Plate 18

Figs. 1-12. Lateral and dorsal views of the terminalia of Statira. 1.
dolera n. sp. 2. basalis. 3. gagatina. \.gagatina resplendens. 5. species “S.
Cal.” 6. liebecki. 7. huachucae. 8. robusta. 9. opacicollis. 10. pluripunctata.
11. colorata. 12. simulans.

252

Psyche

[September

C.T.P.); i May 1-12, 1955, Welaka (C.U.) ; 2 Florida (111.
N.H.S.) ; March 6, 1927, Micanopy (C.U.) ; March 6, 1945, Lake
Placid (C.U.) ; 4 at light March 11, 1956, Titusville (H.F.H.);
13 Georgetown, March, April (M.C.Z.); 1$ Sebastian (M.C.Z.);
1 cf 1? Waccassa River, Gulf Hammock, Levy Co. March 20, 1954

(c.u.)

Relationships: This species is most nearly related to basalis (Fig. 2),
but averages larger, paler, and has shorter apical antennal segments,
and distinctive hypopygidium and male terminalia (Fig. 1). Also
the pronotum is usually more finely and sparsely punctate and the
elytral intervals usually more convex.

Statira basalis Horn

Statira basalis Horn, 1888, Trans. Amer. Ent. Soc. 15: 31-32.

Type: lectotype no. 8016 from Florida in Acad. Nat. Sci. Phila-
delphia.

Range: This species occurs from February 15 to June, chiefly during
April from North Carolina: Lake Mattamuskeet, Wilmington;

South Carolina: Camden, Sampit; Georgia: Atlanta, Clarke Co.;
to northern Florida: St. Johns Co., Rock Bluff, Osceola Nat. Forest;
west through Alabama: Washington Co., Hazen, Florala, Mobile;
Mississippi: Lafayette Co., Oxford, Richton, Lucedale; Louisiana:
Vowell’s Mill; Missouri: Poplar Bluff; Arkansas: South West,
Cove Lake near Paris, Mt. Magazine; to Texas: Dallas.

Habitat: At Hazen, Alabama L. B. Woodruff beat basalis from oak
and Crataegus blossoms in April.

Statira gagatina gagatina Melsheimer

Fig. 3

Statyra gagatina Melsh., 1846, Proc. Acad. Nat. Sci. Philadelphia 2: 311.
Statyra resplendens var. fusca Melsh. ibid, new synonymy.

Types: of gagatina lectotype from Pennsylvania in Melsheimer col-
lection (M.C.Z.) The lower of two specimens on one pin is desig-
nated the lectotype. What Leconte chose as type and labelled “S.
gagatina ! Mels.” is basalis Horn. Of var. fusca holotype labelled
“var. fusca Melsh.” in Melsheimer’s handwriting in the Melsheimer
collection (M.C.Z.)

The variability of gagatina has led to confusion which cannot yet
be cleared up. The color varies from pale brown to black, the paler
specimens being teneral. As Leng points out (Journ. N.Y. Ent. Soc.
31: 186-7) gagatina means “like asphalt or jet”. Melsheimer

described gagatina as “black, tinged with bluish, glossy”. But
specimens in his collection are brown to piceous. My notes on the

1965]

Parsons — Statira

253

lectotype read “dark brown”. Melsheimer described fusca as “uni-
formly yellowish-brown”. It is just like teneral northern examples
of gagatina. There is an occasional tendency for the pronotum to be
paler than the head and elytra as in resplendens. The more extreme
variations are discussed under resplendens and “sp. near gagatina A
Range: This species occurs, March 29 to July 25 chiefly in May,
from southern Vermont: Bennington Co.; Massachusetts: Woods
Hole, Marion; to North Carolina: Bell Island; Tennessee: Bristol,
Knoxville; Arkansas: “Ark.” (111. N. H. S.) a doubtful dealer’s label;
Iowa and Wisconsin.

Habitat: Melsheimer collected gagatina in June on leaves of black
oak and service berry. It has also been taken by beating apple,
hickory, sweet gum, wild cherry, and Pinus virginiana.

Statira gagatina resplendens Melsheimer
Fig. 4

Statyra resplendens Melsh., 1846, Proc. Acad. Nat. Sci. Philadephia 2: 311.
Type: described from Pennsylvania. Leconte has stated that the
unique types of Melsheimer are in his collection. Some of these were
subsequently returned to the Melsheimer collection. In the Leconte
coll. (M.C.Z.) is a specimen labelled “S. resplendens! Mels.” which
means Leconte considered it the type. Unfortunately the locality
label is an orange disc which indicates “southern states”.

This name is questionable. It stands for gagatina in the more
southern part of its range with a more or less rufous pronotum and
last antennal segment a little longer. The most distinctive specimen
seen is a male, July 4, 1925, Oakland Co., Michigan in the G. H.
Nelson collection. This specimen has the last antennal segment equal
to the next 6^/2 segments. Its slightly different terminalia is figured.
Only very few specimens, ranging from New York City to Alabama,
have been seen.

Statira sp. near gagatina
Fig. 5

In the collection of the Illinois Natural History Survey (Andreas
Bolter coll.) is a male labelled “S. Cal.” It is evidently closely
related to gagatina but has a shorter last antennal segment, fewer
setigerous punctures, and slightly deeper terminalia. Also the prono-
tum is more sparsely and finely punctate than in gagatina. The
locality label is suspiciously like that of an unreliable dealer’s.

The color, probably teneral, is testaceous with head brown between
the eyes. Last antennal segment as long as next 4V2 segments. All
tibiae rounded on outer edge; all femora glabrous. Pronotal margins

254

Psyche

[September

distinct and entire. Last ventral segment simple. Disc of pronotum
with five punctures averaging about 4 X their diameters apart, sur-
face finely granular. Setigerous elytral punctures about twice as
large as strial punctures. Setigerous punctures as follows. Left
elytron: 1st interval 1 at apex, 3rd interval 2 at apical fourth; 5th
interval with 1 at base and 1 at apical fifth; 7th interval 2 at base;
9th interval 5 along apical fourth. Right elytron: 1st interval 1 at
base, 3rd interval 1 at apical fifth, 5th interval 1 at basal fourth and
I at apical fifth; 7th interval 2 at base, 9th interval 5 along apical
fifth. Elytral punctures, striae, and convexity of intervals as in
gagatina. Length 7.5 mm.

THE GENUS NEOHERMES (MEGALOPTERA:
CORYDAL1IDAE)*

By Oliver S. Flint, Jr.

Smithsonian Institution, Washington, D. C.

Neohermes was erected in 1908 by Nathan Banks for several
American species then placed in Chauliodes. The only comprehensive
treatment of the genus is that of Weele (1910), who in his mono-
graphic revision of the Megaloptera synonymized all the names
available for the North American members of the genus. Recently,
I happened to compare the genitalia of eastern and western specimens
of Neohermes and it was immediately obvious that they were differ-
ent. After careful study of much more material I am able to recognize
five species in the genus.

I wish to thank the following who generously loaned specimens
and provided other valuable information: Mr. D. E. Kimmins,

British Museum (Natural History), London, England; Dr. Ellis
G. MacLeod, Harvard University, Cambridge, Massachusetts; Dr.
J. G. Franclemont and Dr. L. L. Pechuman, Cornell University,
Ithaca, New York; Dr. Lewis P. Kelsey, University of Delaware,
Newark, Delaware; Dr. Paul H. Arnaud, Jr., California Academy
of Sciences, San Francisco, California; and Dr. Jerry A. Powell.
University of California, Berkeley, California.

Neohermes Banks

Banks, 1908, Proc. Ent. Soc. Wash. 10: 29. Weele, 1909. Notes Leyden Mus.
30: 258; 1910, Coll. Zool. Selys 5 (1): 52. Munroe, 1951, Canad. Ent. 83:
33-35 ; 1953, Canad. Ent. 85: 190-192. Kimmins, 1954, Bull. Brit. Mus.,
Ent. 3 : 418-419. Chandler, 1956, Aquatic Ins. Calif. : 232.

Male with moniliform antennae, about three-fourths as long as
fore wings, each segment bearing a whorl of bristly setae; antennae
of female subserrate, about one-half as long as forewings. Forewing
with R3 and R4 fused for nearly half of their length, and generally
with a crossvein beyond their separation ; anterior branch of 2A united
with 1 A for a short distance. Hind wing with first branch of M
forked apically. Wing membrane slightly grayish ; forewing and
costal margin and apex of hindwing densely marked with black spots
set at right angles to the veins ; forewing with a large black mark at
the level of m-cu. Male genitalia with anal plate cylindrical, apex

^'Manuscript received by the editor June 23, 1965

255

256

Psyche

[September

Fig. 1. Wings of N eohermes (after Chandler 1956).

often decurved, with an apicomesal patch of short, black setae; ninth
sternum forming a broad scoop; aedeagus flat, tip bilobed, with a
central ridge. Female genitalia with anal plate triangular, gona-
pophysis lateralis may bear an apical papilla.

Type-species: Chauliodes filicomis Banks (original designation).

The genus N eohermes ", known only from North America, is closely
related to Protochauliodes Weele which is known from Western
North America, Chile, and Australia. The males of N eohermes are
easily recognized by their long hairy antennae (similar to a bottle-
brush). The females of the two genera sometimes are very difficult
to separate; the presence of a crossvein in cell R3 of the forewing is
generally reliable, but it is sometimes lacking in N eohermes or present
in Protochauliodes; the gonapophyses laterales in the western species
of N eohermes lack an apical papilla (present in the eastern ones)
which is present in the Protochauliodes species in the same region.

There is some variation in the shapes of the anal plates and aedeagi
in all species. The three eastern species recognized probably do not
have significantly differently shaped anal plates but do seem to have
quite different aedeagi. It is possible that additional material from
more localities in the southeastern United States will show inter-
gradation in the shape of the aedeagus.

N eo h erm es filico rn is (Banks)

Figures 2-4, 9

Chauliodes filicomis Banks, 1903, Proc. Ent. Soc. Wash. 5: 238.

N eohermes filicomis (Banks): Banks, 1908, Proc. Ent. Soc. Wash. 10: 29.
Weele 1910, Coll. Zool. Selys 5 (1) : 52-54. Van Dyke, 19+4, Pan-Pacific Ent.
20: 110.

1965]

Flint — Neohermes

257

The male t3^pe of the species is present at the Museum of Compara-
tive Zoology at Harvard University. In addition to the type many
more specimens of the species from New Mexico, Arizona, and Cali-
fornia have been studied. Although this species and calif ornicus are
both found in California, they occupy different ranges within the state.

The male is easily separated from the closely related N. calif ornicus
by the shape of the anal plate which is not bifurcated but produced
apicoventrally. The females of the two species are difficult to
separate although the anal plate is usually not produced into a point
in filicornis.

Male genitalia. — Ninth tergum large, sternum scoop-like. Anal
plate with apicoventral angle produced, bearing mesally many short
black setae in this region; apicodorsal angle rarely slightly produced.
Aedeagus scoop-shaped, apex produced into a pair of submesal lobes,
from which run rather straight ridges to the base; lateral margin
evenly convex.

Female genitalia. — Anal plate triangular, with apical point barely
developed. Gonapophysis lateralis without papilla.

Length of forewing. — male (70 examples) 29 to 50 mm., ave.
39.0; female (25 examples) 37 to 54 mm., ave. 45.8.

Holotype, male — Arizona, Jerome, June 24, 1902, Oslar. Mu-
seum of Comparative Zoology.

Distribution. — new Mexico: Silver City (July). Arizona:

Chiricahua Mts. (June, July, May) ; Huachuacha Mts. (June) ;
Santa Rita Mts. (June); Baboquivari Mts.; Sta. Catalina Mts.
(June); Coconino Co. (July); Garcia; Palmerlee (July, Aug.,
Oct.). California: San Diego Co. (May, June); Orange Co.
(July) ; Riverside Co. (July) ; Los Angeles Co. (June, July, Aug.) ;
Ventura Co. (Aug.) ; San Luis Obispo (July, Aug.) ; Santa Clara
Co. (June) ; Alameda Co. (June) ; Contra Costa Co. (Sept.) ;
Marin Co. (July) ; Sonoma Co. (Feb., July) ; Mendocino Co.
(Aug.) ; Lake Co. (Aug.).

N eoherines calif ornicus (Walker)

Figures 5-8

Chauliodes calif ornicus Walker, 1853, Neur. Brit. Mus. 2: 199. Hagen,
1861, Syn. Neur. N. Amer. : 190. MacLachlan, 1867, Jour. Linn. Soc. Zool.,
9: 259; 1869, Ann. & Mag. Nat. Hist. 4(4): 40. Banks, 1892, Trans. Amer.
Ent. Soc. 19: 357; 1907, Trans. Amer. Ent. Soc. 33: 21. Davis, 1903, Bull.
N. Y. State Mus. 68: 463.

Neohermes californicus (Walker): Banks, 1908, Proc. Ent. Soc. Wash. 10:
29. Weele, 1910, Coll. Zool. Selys 5(1): 53. Caudell, 1933, Pan-Pac. Ent.
9: 125. Van Dyke, 1944, Pan-Pac. Ent. 20: 110. Munroe, 1951, Canad. Ent.
83: 33. Chandler, 1956, Aquat. Ins. Cal. : 232.

Psyche, 1965

Vol. 72, Plate 19

Flint — Nf.ohermes

1965]

Flint — Neohermes

259

Examples of this species from California and Nevada have been
studied. D. E. Kimmins kindly prepared the accompanying figure of
the female genitalia from the holotype located in the British Museum.
He further states that there is a second specimen (male) which was
apparently the one used by Weele for his figure of the genitalia, but
that it is not a type candidate.

The species is closest to N. filicornis from which it is easily sepa-
rated in the male sex by the forked anal plate, and in the female
by the more pointed anal plate.

Male genitalia — Anal plate decurved for apical half, with tip
bifid; mesal surface of ventral lobe with short, black setae. Aedeagus
scoop-like, apex produced into a pair of submesal lobes, which are
developed into slightly sinuous ridges running to the base; lateral
margin convex.

Female genitalia — Anal plate triangular, apex produced into a
distinct point. Gonapophysis lateralis without papilla.

Length of forewing. — male (20 examples) 34 to 47 mm., ave.
39.9 mm.; female (21 examples) 36 to 48 mm., ave. 42.9 mm.

Distribution. — California: Modoc Co. (July); Lassen Co.;

Siskiyou Co. (June) ; Shasta Co. (July) ; Tehama Co. (April, May) ;
Plumas Co.; Sierra Co.; (Aug.) ; Mendocino Co. (June) ; El Dorado
Co. (July); Sacramento Co. (June), Tuolumne Co. (June, July,
Aug.) ; Mariposa Co. (June) ; Fresno Co. (June) ; Tulare Co.
(June, July). Nevada: Reno.

Neohermes angusticollis (Hagen)

Figures 10-13

Chauliodes angusticollis Hagen, 1861, Syn. Heur. N. Amer. : 191. Mac-
Lachlan, 1869, Ann. & Mag. Nat. Hist. 4 (4) : 40. Banks, 1892, Trans. Amer.
Ent. Soc. 19: 357; 1907, Trans. Amer. Ent. Soc. 33: 20. Davis, 1903, Bull.
N. Y. State Mus. 68 : 462.

Neohermes angusticollis (Hagen): Banks, 1908, Proc. Ent. Soc. Wash.
10: 29. Weele, 1910, Coll. Zool. Selys 5 (1): 52.

This species, as defined herein, is known only from Georgia. There
are no types of this species in the Hagen collection at the Museum
of Comparative Zoology, nor are any known to me elsewhere. There-
fore I am taking the following action which preserves all names. The

Explanation of Plate 19

Neohermes filicornis (Bks.). Fig. 2, $ genitalia, lateral. Figs. 3 and 4,
aedeagus and anal plate, dorsal. N. calif ornicus (Walk.). Fig. 5, $ geni-
talia, lateral. Figs. 6 and 7, aedeagus and anal plate, dorsal. Fig. 8, $
genitalia, lateral. N. filicornis (Bks.). Fig. 9, $ genitalia, lateral. N. angu-
sticollis (Hag.). Fig. 10, $ genitalia, lateral.

26o

Psyche

[September

type locality is restricted to the first mentioned state, Georgia, and a
neotj^pe is designated below.

The three eastern species are closely related; indeed I am unable
to find any differences between the females. The aedeagus of angusti-
collis differs from that of the other species in its deeply and narrowly
divided tip and the shallow basolateral excisions.

Male genitalia — Anal plate tubular, slightly constricted at mid-
length; mesal surface of tip with short black setae. Aedeagus scoop-
like, tip divided by a deep, narrow, mesal slit; basolateral excision
shallow, extending only about a fourth of the length of aedeagus.

Female genitalia. — Anal plate with apex produced into a short
point. Apex of gonapophysis lateralis with a short, dark appendage,
above which is a semicircular excision and a short protuberence.

Length of forewing. — Male (2 examples) 28 mm.; female (2
examples) 33 and 35 mm.

Neotype, male. — Atlanta, Georgia, 6-11-39, P. W. Fattig.
USNM type No. 68040.

Distribution — Georgia: Atlanta, 19 June 1946, P. W. Fattig,
I cf i§f; Emerson, 14 June 1940, P. W. Fattig, 1$.

Neohermes matheri Flint, new species
Figures 17-19

The species is only known from a male and 2 females, all from
Mississippi.

The aedeagus of matheri is considerably thicker than that of the
other species, the posterior margin is strongly upturned with the tip
bifid and the halves divergent, and there is a strong lateral process.

Male genitalia. — Anal plate, elongate, tubular, slightly constricted
at midlength ; mesal face with short, black setae. Aedeagus not turned
up basally, central ridge high; tip slightly divided, halves divergent;
basolateral excision about a third the length of aedeagus, with lateral
point strong and upturned.

Female genitalia. — Inseparable from that of N. angusticollis.

Length of forewing. — Male and female, each 29 mm.

Holotype, male. — Clinton, Hinds Co., Mississippi, 12 June, i960,
collection of Bryant Mather, USNM type No. 68041.

Explanation of Plate 20

Neohermes angusticollis (Hag.). Fig. 11, $ genitalia, lateral. Figs. 12
and 13, aedeagus and anal plate, dorsal. N. concolor (Dav.). Fig. 14, $
genitalia, lateral. Figs. 15 and 16, aedeagus and anal plate, dorsal. N.
matheri Flint. Fig. 17, $ genitalia, lateral. Fig. 18 and 19, aedeagus and
anal plate, dorsal.

Psyche, 1965

Vol. 72, Plate 20

Flint

Neohermes

262

Psyche

[September

Paratypes, female. — Hattiesburg, Forrest Co., Mississippi, 31
May 1964. R. & B. Taylor, 2$.

Neohermes concolor (Davis)

Figures 14-16

Chauliod.es concolor Davis, 1903, Bull. N. Y. State Mus. 68: 462. Banks,
1907. Trans. Amer. Ent. Soc. 38: 21.

Neohermes concolor (Davis): Weele, 1910, Coll. Zool. Selys 5 (1): 53.

There are six syn types of this species at Cornell University and
one at the Museum of Comparative Zoology. All, however, are
females, of which one in the Cornell Collection is designated lecto-
type. In addition to these examples, I have studied specimens from
New York, Massachusetts, Delaware, New Jersey, Pennsylvania,
Maryland, District of Columbia, Virginia, North Carolina, Ken-
tucky, Missouri, and Arkansas.

The aedeagus of this species is rounded apically with a shallow,
mesal excision, and the basolateral excisions are rather deep and wide.

Male genitalia. — Anal plate tubular, scarcely constricted, but
some variation in contours among examples; mesal face with short,
black setae. Aedeagus scoop-like, slightly turned up basally; tip with
a slight mesal excision, basolateral excision deep and about one-third
length of aedeagus; central ridge well developed.

Female genitalia. — Inseparable from that of N. angusticollis.

Length of forewing. — Male (13 examples) 26 to 32 mm., ave.
28.2 mm.; female (18 examples) 30 to 39 mm., ave. 35.4 mm.

Lectotype, female (here designated). — “Cornell U. No. 815 sub.”
“Lectotype Cornell U. No. 4269” “Chauliodes concolor type” “Lecto-
type Chauliodes concolor Davis By Flint 1965”.

Distribution. — Massachusetts: Southbridge (July). new

York: Wayne Co. (July); Ithaca (July, Aug.); Binghamton

(July). DELAWARE: Porters; Newark (July), new jersey:

Middlesex Co. (July). Pennsylvania: State College (July).

Maryland: Dorchester Co. (July); Montgomery Co. (July) ;

Laurel (June), district of Columbia: Washington (June). Vir-
ginia: Falls Church (May, June, July); Blacksburg (June).

NORTH CAROLINA: Durham (June). Kentucky: Mammoth Cave
Nat. Pk. (June). Missouri: Willard (June). Arkansas: Imbo-
den; Hot Springs Nat. Pk. (June).

Literature Cited

Banks, Nathan

1892. A synopsis, catalogue, and bibliography of
insects of temperate North America. Trans.
19: 327-373.

the neuropteroid
Amer. Ent. Soc.,

1965]

Flint — Neohertnes

263

1903. Neuropteroid insects from Arizona. Proc. Ent. Soc. Wash.,
5:237-245.

1907. Catalogue of the neuropteroid insects (except Odonata) of the
United States. Trans. Amer. Ent. Soc., 33: 53 pp.

1908. On the classification of the Corydalinae, with description of a
new species. Proc. Ent. Soc. W^ash., 10: 27-30.

Caudell, A. N.

1933. Neohermes infuscatus, a new Sialid from California. Pan-Pac.
Ent, 9: 125-126.

Chandler, H. P.

1956. Megaloptera, in Aquatic insects of California. Pages 229-233.
Berkeley, California.

Davis, K. C.

1903. Sialididae of North and South America. N. Y. State Mus. Bull.
68: 442-487.

Hagen, Herman

1861. Synopsis of the Neuroptera of North America. Smiths. Misc.
Coll., 4: 1-347.

Kimmins, D. E.

1954. A new genus and some new species of the Chauliodini (Mega-
loptera). Bull. Brit. Mus., Ent. 3: 417-444.

McLachlan, R. L.

1868. A revision of the “List of the specimens of the Neuropterous
insects in the collection of the British Museum. Part II, 1853.
By F. Walker” as far as the end of the genus Myrmeleon, pp.
193-410. Jour. Linn. Soc. Lond. Zool. 9: 258-281.

1869. Considerations on the neuropterous genus Chauliodes and its allies,
with notes and descriptions. Ann. & Mag. Nat. Hist., 4(4) : 35-46.

Munroe, E. G.

1951. The identity and generic position of Chauliodes disjunctus Walker
(Megaloptera: Corydalidae) . Canad. Ent., 83: 33-35.

1953. Chauliodes disjunctus Walker: a correction, with descriptions of
a new species and a new genus (Megaloptera: Corydalidae).

Canad. Ent., 85: 190-192.

Van Dyke, Edwin C.

1944. A new dobsonfly (Megaloptera) from California. Pan-Pac. Ent.,
20: 110.

Walker, F. M.

1853. List of the specimens of neuropterous insects in the collection of
the British Museum. Part II: 193-475.

Weele, H. W. van der

1909. New genera and species of Megaloptera Latr. Notes Leyden
Mus., 30: 249-264.

1910. Megaloptera, in Collections zoologiques du Baron Edm. de Selys
Longchamps. Fasc. 5 ( 1) :l-93. Bruxelles.

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PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol. 72 December, 1965 No. 4

CONTENTS

Further Studies on Epyrini (Hymenoptera, Bethylidae).

Howard E. Evans 265

The Taxonomic Problem of Sexual Dimorphism in Spiders and a
Synonymy in Myrmccotypus (Araneae, Clubionidae) .

Jonathan Reiskind 279

The Habits and Distribution of Macromischa subditiva Wheeler
(Hymenoptera: Formicidae). tVilliam S. Creighton 282

Notes on the Indo-Australian Ants of Genus Simpone Forel (Hymen-
optera, Formicidae). Robert IV. Taylor 287

A New Species of the Spider Genus Xysticus (Araneae: Thomisidae)
from Arizona. J. H. Redner and C. D. Dondale 291

The Types of Proctotrupoidea (Hymenoptera) in the Charles T. Brues
Collection at the Museum of Comparative Zoology. Lubomir Masner 295

The Habitat of Platypatrobus Darlington (Coleoptera: Carabidae).

Henri Goulet 305

The Genus Zomosemata , with Notes on the Cytology of Two Species
(Diptera — Tephritidae) . Guy L. Bush

Author and Subject Index for Volume 72

307

325

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1965-66

President IL Reich ardt, Harvard University

Vice-President S. Vogel, Harvard University

Secretary C. C. Porter, Harvard University

Treasurer F. M. Carpenter, Harvard University

Executive Committee W. L. Brown, Jr., Cornell University

J. Reiskind, Harvard University

EDITORIAL BOARD OF PSYCHE

F. M. Carpenter (Editor), Professor of Entomology , and Alexander
Agassiz Professor of Zoology , Harvard University

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University; Associate in Entomology , Museum of Comparative
Zoology

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parative Zoology

H. E. Evans, Curator of Insects, Museum of Comparative Zoology

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PSYCHE

Vol. 72

December, 1965

No. 4

FURTHER STUDIES ON NEOTROPICAL EPYRINI
(HYMENOPTERA, BETH YLIDAE ) 1

By Howard E. Evans
Museum of Comparative Zoology

This paper is a supplement to two recent papers published in the
Bulletin of the Museum of Comparative Zoology.2 Although these
papers were several years in the making, as so often happens they had
scarcely appeared when a considerable amount of new material came
into my hands. The present paper is concerned with range extensions,
notes on variation, and new species in the genera Rhabdepyris , Bakeri-
ella, Calyozina, and Epyris. The abbreviations employed are the
same as those used in my two previous papers, but for the sake of
ready reference the abbreviations are listed again at the conclusion
of this paper.

Rhabdepyris (Rhabdepyris) muesebecki Evans, 1965
This species was recorded from the United States only from two
specimens intercepted at quarantine in Texas. However, it is ap-
parently established in Texas, as I took a male while sweeping herba-
ceous vegetation at the north edge of the city of Brownsville on
June 29, 1965. I also have a female before me from Mera, Ecuador,
collected January 26, 1923, by F. X. Williams [coll. Bishop Mus.,
Honolulu]. This is within the known geographic range of this
species, but the specimen is of interest because of its large size ( LFW
2.1 mm). It keys out well in my revision if one makes allowance
for greater variation in size than previously indicated.

Research and publication supported by a grant from the National Science
Foundation, No. GB-1544. Manuscript received by the editor December 7 y
1965.

21964. A synopsis of the American Bethylidae (Hymenoptera, Aculeata).
Bull. Mus. Comp. Zool., 132: 1-222.

1965. A revision of the genus Rhabdepyris in the Americas (Hymenop-
tera, Bethylidae). Bull. Mus. Comp. Zool., 133:67-151.

265

266

Psyche

[December

Rhabdepyris ( Rhabdepyris ) nigriscapus Evans, 1965

This species was described from a single female from the province
of Jujuy, Argentina, unfortunately indicated as a “c?” in the orig-
inal description, although stated to be a female two lines later. Mr.
Fritz Plaumann has recently sent me a second female, this one from
Nova Teutonia, Santa Catarina, Brazil, collected in March, 1965.
I his specimen agrees closely with the type in color and most struc-
tural characters, but it is considerably larger (LFW 2.1 mm) and
the front femora measure only 2.65 X as long as their maximum
width. The head is also slightly wider (WH 1.02 X LH) and the
front narrower (WF 1.02 X HE) ; the ocellar triangle is broad,
OOL only .77 X WOT. Thus this specimen bridges some of the
gaps supposed to separate nigriscapus and minutulus. The latter
species may still be separable on the basis of its extremely small size,
more compact antennae, pale scape, and somewhat more robust front
femora; but only the accumulation of much more material of this
complex will permit final clarification of specific characters.

Rhabdepyris (Rhabdepyris) multilineatus n. sp.

This species is of unusual interest because the pronotum lacks a
foveolate groove paralleling its posterior margin; actually such a
groove is partially developed in one of the two available males, but
absent in the other male and in all six females. I used the presence
of such a groove as a subgeneric character in my 1965 revision, but
evidently it will not hold. Otherwise multilineatus is a reasonably
typical member of this subgenus, being related to those species in
which the front angle of the ocellar triangle of the female is less than
a right angle. In the male the ocellar triangle is somewhat broader,
so that the males will run to gracilis in my key (as they also will
with respect to head shape). Actually the males are readily separable
from those of both gracilis and muesebecki on the basis of the pale
and much more compact antennae.

This species is also of special interest since the type series has as-
sociated with it the first host data for any member of this genus.
Pinned with the type series is an adult and a larva of the dermestid
Cryptorhopalum septemsignatum Sharp [det. J. M. Kingsolver] ; the
word “notes” is written on the label in the handwriting of the col-
lector, F. X. Williams. Dr. Williams writes that he can no longer
find these notes, but he remembers collecting some empty cocoons
under loose bark along the edge of a forest; he believes that the
Rhabdepyris adults were reared from larvae which were feeding on

1965]

Evans — Epyrini

267

dermestid larvae breeding in the refuse in the cocoons. Perhaps this
should be considered a doubtful host record, since the notes are not
extant. However, it seems a very logical association, since dermestids
are the hosts of several species of Laelius. As I pointed out in 1965,
the morphological gap between Laelius and Rhabdepyris sensu stricto
is a very small one.

Holotype. — Ecuador: Milagro, December 1922 (F. X. Wil-
liams) [Bishop Mus., Honolulu].

Description of female type. — Length 3.1 mm; LFW 2.4 mm.
Black; mandibles and antennae rufo-testaceous except flagellum
weakly infuscated on upper surface; coxae and femora dark brown,
legs otherwise bright rufo-testaceous; wings hyaline, veins and stigma
amber. Clypeus sharply, obtusely angulate; third antennal segment
about as long as thick. Head slightly higher than wide, WH .97 X
LH ; vertex produced above eye tops by a distance equal to about half
the eye height; front of moderate breadth, WF .59 X WH, 1.17 X
HE ; front angle of ocellar triangle very slightly less than a right
angle, OOL 1,25 X WOT. Front wholly alutaceous, with shallow,
widely spaced punctures. Pronotum alutaceous but somewhat shining,
strongly punctate, its posterior margin depressed but not paralleled
by a groove. Scutellar groove a slightly arcuate band of nearly equal
width throughout. Propodeal disc 1.2 X as wide as long, with five
strong discal carinae plus strong lateral and sublateral carinae, all
nine carinae reaching the posterior margin or nearly so. Front femora
2.5 X as long as their maximum width.

Allotype. — cf, Ecuador: same data as type [Bishop Mus.,
Honolulu].

Description of male allotype. — Length 2.8 mm; LFW 1.9 mm.
Coloration as described for female. First four antennal segments in
a ratio of about 17:7:7:8, segments three and eleven each about 1.2
X as long as thick. WH/LH = 1.0; WF .63 X WH, 1.36 X
HE; ocellar triangle rather broad, front angle about a right angle;
OOL 1. 16 X WOT. Front alutaceous, somewhat shining, punc-
tures inconspicuous. Pronotum and scutellar groove as described for
female; propodeal disc 1.2 X as wide as long, carinate much as in
the female; side-pieces of propodeum strongly beaded.

Paratypes. — 5 99 j 1 cf , Ecuador : same data as type except 4
99 dated December 4, 1922 [Bishop Mus., Honolulu; MCZ].

Variation. — - The females show only slight variation in size
(LFW 2. 2-2. 4 mm) and only minor variation in color and sculpture;
WH varies from .96 to 1.0 X LH, WF from 1.15 to 1.20 X HE,

268

Psyche

[December

OOL from 1.2 to 1.3 X WOT. The male paratype is slightly
smaller than the allotype (LFW 1.8 mm) and is very similar except
that the posterior margin of the pronotum is preceded by a row of
shallow foveae.

Rhabdepyris (Rhabdepyris) gracilis Evans, 1965

I described this species from a female from Sacramento, California,
and a male from near Durango, Mexico, remarking that the associa-
tion seemed probable in spite of the distance between these two lo-
calities. I have since seen a male from Danville, Contra Costa Co.,
California (F. X. Williams, Aug. 8, 1948) [CAS] agreeing so
closely with the allotype from Durango that I feel confident that my
association of the sexes was correct. This specimen is identical to
the allotype in size, color, and most standard measurements, but
OOL is only 1.15 X WOT. A female collected by F. X. Williams
at Danville on May 29, 1952 [CAS] probably also belongs here,
but the groove paralleling the posterior margin of the pronotum is
virtually absent and OOL is only 1.10 X WOT; in size, color,
and other standard measurements it is similar to the Sacramento type.

Rhabdepyris ( Rhabdepyris ) nigripilosus (Ashmead),
new combination

Mesitius nigripilosus Ashmead, 1895, Proc. Calif. Acad. Sci., (2) 5: 539.

[Type: 9, MEXICO: NAYARIT: Tepic (CAS)].

Epyris nigripilosus Kieffer, 1914, Das Tierreich, 41: 345. — Evans, 1964,
Bull. Mus. Comp. Zool. Harvard, 132: 108.

Rhabdepyris (Rhabdepyris) huachucae Evans, 1965, Bull. Mus. Comp. Zool.
Harvard, 133 : 73. [Type: $, ARIZONA: Ramsey Canyon, Huachuca
Mts., March 22, 1956 (Werner & Butler) (MCZ)]. New synonymy
Through the courtesy of Mr. Hugh B. Leech, I have recently had
an opportunity to study the type of Ashmead’s nigripilosus , and I
find that Kieffer and I were incorrect in assuming it to be an Epyris.
Beyond much question it is a second specimen of the species I de-
scribed from southeastern Arizona as huachucae. The type of nigri-
pilos'us differs from that of huachucae in the following particulars:
slightly larger (LFW 2.6 mm) ; front femora with only a small
brownish blotch toward base, otherwise testaceous, middle femora
wholly testaceous; head more nearly circular, WH .98 X LH ; front
very narrow, WF and HE subequal, the punctures very slightly
stronger and more closely spaced; OOL 1.35 X WOl ; propodeal
disc somewhat broader, measuring 1.35 X as wide as its median
length. This is a relatively large and distinctive species of the sub-
genus.

1965]

Evans — Epyrini

269

Rhabdepyris (Trichotepyris) hirticulus new name

Rhabdepyris ( Trichotepyris ) nigropilosus Evans, 1965, Bull. Mus. Comp.
Zool. Harvard, 133: 84. Preoccupied by Ashmead, 1895.

Although Ashmead called his species nigripilosus (see preceding
species), according to the International Code of Zoological Nomen-
clature, article 58, compound words differing by only a connecting
vowel are to be considered homonyms. A new name is therefore
proposed for this species, which is known from several females from
Panama and Brazil.

Rhabdepyris (Trichotepyris) apache Evans, 1965

I described this large and striking species from three females from
Arizona and a male from Sonora, Mexico. It now appears that it is
widely distributed and locally not uncommon in western Mexico. I
collected 1 1 99 and 1 c? from the tops of young Ambrosia plants,
apparently containing honeydew, on the outskirts of Guadalajara,
Jalisco, Mexico, July 17-28, 1965. I have also seen 1 c? from 8
miles SE of Elota, Sinaloa, Mexico, collected May 19, 1962, by
L. A. Stange [UCD] and 1 c? from Maria Madre Isl., Tres Marias,
Nayarit, Mexico, collected May 22, 1925 (H. H. Keifer) [CAS].
In the original description I noted that the pronotal disc of the male
is sharply declivous anteriorly and laterally. In all three of the males
listed above, the disc is actually subcarinate anteriorly and along the
anterior half of the lateral margins. Also, in some of the females
from Guadalajara, one can detect a faint, irregular carina along the
anterior margin of the pronotal disc. I have already suggested that
Anisepyris and Trichotepyris are closely related by way of the mega-
cephalus species group of the latter subgenus. It would appear that
apache provides an almost perfect intermediate between these two
taxa. For the present I shall, however, leave it in Rhabdepyris
(Trichotepyris) , since on the whole it seems to fit best in the mega-
cephalus group of that subgenus.

Rhabdepyris (Trichotepyris) fortunatus Evans, 1965

This species was based on a single female from Costa Rica. I
have before me two females which resemble the type very closely,
one from Bucay, Ecuador, Oct. 4, 1922, and one from Blairmont,
British Guiana, Oct. 1923 (both collected by F. X. Williams)
[Bishop Mus., Honolulu]. Both are slightly larger than the type
(LFW 2. 6-2. 9 mm) and have the ocellar triangle slightly more re-

270

Psyche

[December

mote from the eye margins (OOL 1.3- 1.4 X WOT). The very
short third antennal segment is characteristic of this species.

Rhabdepyris (Chlorepyris) virescens Evans, 1965

This large, well characterized species was known to range through-
out tropical South America and into Panama. Apparently it also
ranges throughout the remainder of tropical Central America and
Mexico, for I took a very typical female of this species while sweep-
ing vegetation just behind the beach at Playa Matanchen, near San
Bias, Nayarit, Mexico, on August 7, 1965. As might be expected,
this specimen is more similar to material from Panama than to the
type (from southern Brazil) ; the apical third of the abdomen is
rufous, the femora are wholly rufous, and the front is relatively nar-
row (WF 1. 10 X HE). LFW measures 4.0 mm; WH/LH =
1.05; the front femora measure 2.05 X as long as wide.

Bakeriella inconspicua Evans, 1964

On July 6, 1965, I collected two males of this species on banana
leaves at Tamazunchale, San Luis Potosi, Mexico, thus extending the
known range of the species several hundred miles northwestward.
These males are small (LFW 2. 1-2.3 mm) and agree with males
from Tabasco in having the scape slightly infuscated and the front
and thoracic dorsum rather strongly shining. The female of this
species is still unknown.

Bakeriella brasiliana Evans, 1964

This species was described from a single female from Minas Gerais,
Brazil. I have seen one additional female from a place far distant
from the type locality but so similar to the type that its conspecificity
can scarcely be questioned. This specimen was taken on July 14,
1918, at the Limon Plantation, Chagres R., Panama (sweeping
around cornfield, H. Dietz & J. Zetck) [USNM]. This specimen
is of the same size as the type and of similar coloration except that
the apex of the abdomen is dark brown rather than ferruginous;
standard measurements are very much like those of the type except
that OOL is 1.7 X WOT, the propodeal disc 1.1 X as wide as
long. It is conceivable that brasiliana represents the otherwise un-
known female sex of depressa Kieffer or flavicornis Kieffer, though
by analogy with olmeca Evans and with subcarinata n. sp. I would
expect the females of depressa and flavicornis to have five-toothed

271

1965]

Evans — Epyrini

mandibles and at least some evidence of a median carina on the prono-
tum.

Bakeriella subcarinata n. sp.

This is the fourth known species in which the male possesses a
vertical carina on the temples and a median carina on the pronotum ;
it is the second of these four species in which the female is known (or
at least tentatively associated with the male) . In this case the female
lacks a carina on the temples (as in olmeca Evans) but possesses a
weak median carina on the pronotum (in contrast to olmeca). The
female will run to inca Evans in my 1964 key, but it differs in
several respects, most conspicuously in having 5-toothed mandibles and
in having the median pronotal carina very weak and not set in a
depression. The male runs to olmeca in my key, but differs in having
well formed lateral ridges on the pronotum and in having the third
antennal segment only 1.3 X as long as thick.

Allotype. — PERU: Quincemil, Dept. Cuzco, 750 meters, Nov.
5-16, 1962 (L. Pena) [MCZ].

Description of female allotype. — Length 5.0 mm; LFW 4.3 mm.
Black; mandibles testaceous except black at base, the teeth rufous;
antenna pale ferruginous, scape and outer flagellar segments strongly
infuscated on upper surface; legs black except tibiae and tarsi dull
ferruginous; wings lightly tinged with yellowish brown. Mandibles
with a broad, oblique apical margin which bears five teeth, the basal
three teeth very much smaller than the two large apical teeth. Clypeus
very short, broadly subangulate. Eyes weakly hairy; antennal scrobes
not margined; third antennal segment about 1.5 X as long as thick.
WH .95 X LH ; eyes converging below, WF 1.20 X HE; OOL
1.5 X WOT. Vertex broadly rounded off a considerable distance
above eye tops; temples fairly broad but ecarinate. Front shining,
weakly alutaceous, with strong punctures which are separated by
1-3 X their own diameters. Pronotal disc somewhat more strongly
alutaceous, also with strong punctures; anterior, transverse carina
straight; median carina weak, alutaceous, not set in a depression;
posterior margin with a polished ridge; sides of disc subcarinate in
front; side-pieces rugose, especially below. Mesoscutum polished
anteriorly, posterior half alutaceous, punctate, slightly depressed ;
scutellar pits large, transverse, separated by a thin septum. Propodeal
disc 1. 1 5 X as wide as long, transversely striate between the three
discal carinae, otherwise with only weak sculpturing. Front femora
measuring 2.4 X as long as wide.

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Psyche

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Holotype. — cf , PERU : Tingo Maria, Dept. Huanuco, 620 meters,
Oct. 5-12, 1964 (C. C. Porter) [MCZ].

Description of male type. — Length 4.0 mm; LFW 2.6 mm.
Black; mandibles testaceous, except basal third black, teeth rufous;
scape black except apical fourth, as well as all of following three
segments, testaceous, remainder of antennae mostly fuscous; legs
black except tarsi straw-colored, tibiae only partially infuscated;
wings lightly tinged with yellowish brown. Mandibles with five
sharp teeth ; clypeus obtusely angulate ; scrobes carinate ; eyes weakly
hairy. First four antennal segments in a ratio of about 20:8:9:10,
segment three about 1.3 X as long as thick, segment eleven 1.6 X as
long as thick. WH 1.08 X LH; eyes converging strongly below,
WF 1. 1 5 X HE; OOL 1.4 X WOT. Temples with a carina
which starts at the vertex but extends only to about the middle of the
eyes. Front alutaceous, moderately shining, obscurely punctate.
Pronotal disc alutaceous, even over the crest of the median carina,
which is stronger than in the female; anterior, transverse carina
angled backward only very weakly at the midline; sides of disc sub-
carinate anteriorly, posterior margin with a polished elevation pre-
ceded by a row of small foveae. Mesoscutum strongly alutaceous and
slightly depressed on the posterior half ; scutellar pits large, separated
by a thin septum medially. Propodeal disc 1.1 X as wide as long,
disc transversely striate between the three discal carinae; posterior
angles dentate in front of the strongly developed foveae.

Paratypes. — 3 cf cf , PERU: Monson Valley, Tingo Maria, Oct.
10, Nov. 29, 1954 (E. I. Schlinger and E. S. Ross) [CAS].

Variation. — The three male paratypes vary slightly in size (LFW
2. 4-2. 8 mm) but resemble the type very closely in color and standard
measurements. In all three specimens the carina on the temples ter-
minates near the middle of the eyes; in all three the median carina
of the pronotum is slightly weaker than in the type, although more
distinct than in the female allotype.

Bakeriella inca Evans, 1964

This species was described from a single female from Puerto Ber-
mudez, Rio Pichis, Peru. I have recently studied 1 9 and 2 cf cf of
this species collected by E. I. Schlinger and E. S. Ross in the Monson
Valley, Tingo Maria, Peru, Sept.-Dee. 1954 [CAS]. The female
is slightly larger than the type (LFW 3.3 mm) and the head is as
wide as high, but there are no other differences worthy of note. The
males lack a carina on the temples, have a strong median carina on

1965]

Evans — Epyrini

273

the pronotum which is set in a depression, and have hairy eyes; thus
they key out readily in my key. In these males LFW is 2. 1-2.6 mm;
WH is 1. 02-1. 05 X LH, 1 .25-1.30 X HE, OOL about 1.5 X
WOT. There are no important differences in color or sculpturing
as compared to the female except that the front is only obscurely
punctate and the antennal scrobes are carinate (as is usual in males
of this genus).

Bakeriella rossi Evans, 1964

The collecting of Schlinger and Ross in the Monson Valley, l ingo
Maria, Peru, in September and October 1954 also turned up 1 $
and 3 cf cf of this species, previously known from 3 cf cf from Co-
lombia. The female will run to couplet 8 of my key; the 5-toothed
mandibles at once separate it from brasiliana , the very weakly arched,
non-angulate transverse pronotal carina from olmeca , the fuscous
coxae and femora and much larger size from floridana. The female
is 5.5 mm long, LFW 3.9 mm; the body is black except that the
apical fourth of the abdomen is suffused with dull ferruginous. The
clypeus is sharply, obtusely angular; the eyes are hairy; the vertex is
broadly rounded off far above the eye tops, WH being only .9 X
LH; WF is 1.22 X HE, OOL 1.8 X WOT, the propodeal disc
1.2 X as wide as long and bearing the usual three carinae.

Bakeriella depressa Kieffer

Since treating this species in 1964, I have seen one additional male,
from Tena, Ecuador, collected February 28, 1923, by F. X. Williams
[Bishop Mus., Honolulu], It is the largest specimen I have seen
(LFW 3.0 mm) but presents no unusual structural features. The
antennae of this species are consistently more elongate than in flavi-
cornis Kieffer, segments three and eleven being 1.35- 1.5 X as long as
thick as compared to 1.1-1.3 X as long as thick in flavicornis. Also,
the antennae of most specimens are decidedly dusky apically, while in
flavicornis they are wholly light yellowish brown. In addition to the
type of the latter species I have seen four topotypic paratypes in the
collection of Cornell University.

Calyozina mexicana n. sp.

This remarkable species seems to be closer to the type species,
ramicornis Enderlein, from Formosa, than to the two neotropical
species I assigned here in 1964. In particular, there are ten long
branches on the antennae as opposed to only seven or eight short

274

Psyche

[December

branches in neotropica and amazonica. This species also differs from
the latter two species in having 5-toothed mandibles, dentate claws,
a groove connecting the two scutellar pits, and some unusual sculptur-
ing on the sides of the thorax and propodeum. It is possible that
mexicana should not be regarded as congeneric with neotropica and
amazonica , but considering the fact that all three species are known
from only one specimen ( in each case a male ) , as well as the many
problems of generic classification in this section of the Epyrini, it
seems best to be conservative.

Holotype . — cT , MEXICO: 23 mi. S Matias Romero, Oaxaca, Aug.
14, 1963 (F. D. Parker & L. A. Stange) [MCZ].

Description of male type. — Length 6.2 mm; LFW 3.8 mm. Black,
head and thoracic dorsum with a faint bluish cast, last two abdominal
segments rufous; mandibles rufo-testaceous; scape black, flagellum
dull ferruginous, the branches dusky; coxae black, legs otherwise
wholly bright rufo-testaceous; wings lightly tinged with brown.
Mandibles with a large apical tooth and four small teeth in a slightly
oblique series; clypeus broadly angulate, its median carina nearly
straight in profile. Antennae strongly pectinate; segment three con-
siderably shorter than two, much wider than long, bearing a rather
short process; segments 4-12 each bearing a slender process slightly
longer than the segment proper (segments 4 and 12) up to about
3 X as long as the segment proper (segments 7-10) ; segment 13
elongate, clavate; segments 2-5 each bearing long, erect setae, fol-
lowing segments with shorter setae but with long setae on the proc-
esses, these setae much longer than the width of the processes. Head
unusually broad, WFI 1.25 X LH ; front broad, WF .70 X WH,
1.50 X HE; vertex broadly rounded off a short distance above eye
tops; ocelli in a broad triangle, WOT slightly exceeding OOL.
Front polished, non-alutaceous, with small, widely spaced punctures.
Thoracic dorsum slightly alutaceous, duller than front; pronotal disc
sloping gradually toward the front and its sides approaching to a
rounded anterior margin, the disc not at all prominent antero-lat-
erally; scutellar pits large, subcircular, separated by slightly less than
their own diameters, connected by a very slender but fairly deep
groove. Propodeal disc 1.15 X, as wide as its median length, with
three strong discal carinae between which it is transversely ridged;
laterad of the discal carinae the surface is smoother, but there is a
low longitudinal ridge between the lateral discal and strong sub-
lateral carinae; propodeal side-pieces polished, with several prominent
longitudinal ridges which are continuous with similar ridges on the

1965]

Evans — Epyrini

275

mesopleura and which form (more especially on the mesopleura) two
concentric ellipses. Middle and hind tibiae somewhat spinose; claws
with a short, erect tooth. Thoracic venter rather flat, the middle
and hind coxae rather widely separated on the venter.

Epyris cubanus n. sp.

This is the first Epyris known from Cuba, and only the third
species reported from the West Indies. It is one of few species of
Epyris of metallic coloration.

Holotype. — CUBA: Rio Cauto, Oriente Prov., April 1928
(C. F. Stahl) [USNM].

Description of female type. — Length 3.0 mm; LFW 2.0 mm.
Head and thorax dark olive-green, shining; pronotal collar fer-
ruginous; propodeum black; abdomen dark reddish brown, fading to
light brown laterally and apically; mandibles testaceous, black at
extreme base ; antennae testaceous except most of scape and upper side
of flagellum dark brown; coxae fuscous, front and hind femora dark
brown, legs otherwise testaceous; wings hyaline. Mandibles bi-
dentate; clypeus obtusely angulate, rounded at the midline, elevated
medially but without a distinct carina. Antennal segment three
wider than long, segment eleven 1.4 X as long as wide. Head
elongate, WH .89 X LH ; WF .62 X WH, 1.20 X HE; vertex
passing straight across a considerable distance above eye tops, the
ocelli in a broad triangle far above eye tops; OOL 1.33 X WOT.
Front polished, very weakly alutaceous, with small, widely spaced
punctures. Thoracic dorsum also weakly alutaceous, obscurely punc-
tate; pronotal disc nearly twice as long as mesoscutum, with smooth
contours; notauli linear, except very slightly widened near the pos-
terior margin ; scutellar pits small, oblique, longer than wide, the two
pits separated by about 3 X their own greatest diameter. Propodeal
disc 1.2 X as wide as its median length, with a complete median
carina plus several short carinae in the basal triangle, but otherwise
with only weak surface sculpturing; posterior angles subfoveolate ;
side-pieces shining, somewhat alutaceous. Mesopleura weakly aluta-
ceous. Mesopleura weakly alutaceous, with a small, deep pit above
and a longitudinal ridge below. Front femora strongly incrassate,
measuring only 1.8 X as long as wide; middle tibiae very weakly
spinose; claws dentate. Abdomen robust, subfusiform, not at all
depressed apically.

Epyris depressigaster n. sp.

This species is of interest not only because of its brilliant colora-

276

Psyche

[December

tion, but also because the apical portion of the abdomen is strongly
depressed, suggesting some of the apterous Pristocerini. The head
is less elongate and flattened than is usual in this genus.

Holotype. — brazil: Nova Teutonia, Santa Catarina, July 17,
1937 (F. Plaumann) [BMNH].

Description of female type. — Length 6.0 mm; LFW 3.5 mm.
Head and thorax dark blue-green; propodeum black; abdomen shining
black except apical third bright ferruginous; mandibles and antennae
wholly rufo-testaceous ; legs testaceous except all coxae fuscous, hind
femora medium brown ; wings subhyaline. Mandibles bidentate, the
outer tooth sharp, the inner tooth broad and blunt. Clypeus broadly
angulate, the median carina nearly straight in profile. Antennae
compact, segments three and eleven each slightly longer than wide.
Head only slightly higher than wide, WH .97 X LH ; eyes large,
prominent, protruding; front convex, with a linear impression on the
lower fourth. WF subequal to HE ; ocelli in about a right triangle,
OOL 1. 1 X WOT; bottom of anterior ocelli on a line with the
eye tops, the vertex broadly rounded off a distance above the eye tops
equal to only about one third X HE. Front strongly alutaceous be-
low, the punctures separated by little more than their own diameters;
at the level of the ocelli the front is less strongly alutaceous and the
punctures more widely spaced. Thoracic dorsum weakly alutaceous,
wholly covered with small punctures except the scutellar disc im-
punctate; pronotal disc about 1.6 X as long as mesoscutum; scutellar
pits oblique, elongate, separated by 1.3 X their own length. Pro-
podeal disc about as long as wide, with three strong, complete discal
carinae plus lateral carinae, between these five carinae with weak,
irregular transverse striae; posterior angles weakly foveolate. Meso-
pleura pleura shining, with small punctures; upper fovea small but
well defined, lower fovea large, broadly open above. Front femora
slender, 2.3 X as long as wide; middle tibiae with dense, short
spines above; claws with a strong, erect tooth, the outer ray curved
so as to be nearly parallel to the tooth. Abdomen depressed through-
out, but more especially so on the apical half, the last segment very
thin in lateral view.

Epyris crassipes n. sp.

This distinctive species appears to be one of the commoner elements
in the Epyris fauna of southeastern Brazil. It is a glossy, black species
having unusually robust femora in both sexes and having the pro-
podeal sculpturing largely restricted to a basal triangle.

1965]

Evans — Epyrini

277

Allotype. — 9, brazil: Nova Teutonia, Santa Catarina, Feb. 1,
1963 (F. Plaumann) [MCZ].

Description of female allotype. — Length 4.2 mm; LFW 3.0 mm.
Black, shining, the extreme tip of the abdomen suffused with brown;
mandibles ferruginous, black at extreme base; scape dark brown
above, paler beneath, flagellum testaceous, weakly suffused with brown
on upper surface ; legs fuscous except front and middle tibiae and all
tarsi testaceous; wings subhyaline. Mandibles slender, with a strong
apical tooth, basad of this with a truncate cutting edge. Clypeus
with a protuberant, narrowly rounded median lobe. Third antennal
segment about as long as wide, segment eleven 1.5 X as long as
wide. WH .90 X LH ; eyes convergent below, WF 1.08 X HE;
vertex passing straight across far above eye tops (except slightly gib-
bous just behind the ocellar triangle) ; ocelli in a broad triangle,
OOL 1.4 X WOT; anterior ocellus far above eye tops. Front
strongly shining, obscurely alutaceous, with small, widely spaced
punctures. Thoracic dorsum more evidently alutaceous than head ;
pronotal disc 1.7 X as long as mesoscutum, with distinct, somewhat
irregularly spaced punctures; mesoscutum with a few weak punc-
tures; scutellar pits oblique, about twice as long as wide, separated
by twice their own length. Propodeal disc 1.3 X as wide as long,
with complete median and lateral carinae, otherwise shining and with
weak sculpturing except in a large, median, basal triangle, where
there are several irregular longitudinal carinae and transverse ridges.
Mesopleura shining, foveae not defined, but with a ridge which
passes above the pit and then arches downward to form what would
usually be the lower margin of the lower fovea. Front femora very
broad, measuring twice as long as their greatest width, the middle
femora even more robust, 1.7 X as long as wide; middle tibiae with
long spines above and below; claws weakly dentate. Abdomen
robust, not depressed.

Holotype. — cf, brazil: same data as allotype except collected
in June, 1952 [MCZ].

Description of male type. — Length 4.0 mm; LFW 3.1 mm.
Black, shining; mandibles testaceous except black at base, rufous at
apex; antennae wholly rufo-testaceous, the flagellum a bit dusky,
especially apically; coxae and femora dark brown, legs otherwise
light brown; wings subhyaline. Mandibles very slender, the apex
acute, simple. Clypeus with a protuberant, subangulate median lobe.
First four antennal segments in a ratio of about 22:10:11:16, seg-
ment three 1.3 X as long as thick, segment eleven 2.1 X as long

278

Psyche

[December

as thick. WH .98 X LH ; eyes convergent below, WF 1.1 X HE;
ocelli in a compact triangle well above eye tops, OOL 1.4 X WOT.
Vertex passing straight across a distance above eye tops equal to
about .6 X HE. Front strongly polished, with small punctures which
are separated by several times their own diameters at the level of the
ocelli, but much more closely spaced below. Thoracic dorsum also
strongly polished, the mesoscutum, however, weakly alutaceous; pro-
notal disc more than twice as long as mesoscutum. Scutellar pits and
features of propodeum and mesopleura as described for female; pro-
podeal disc measuring 1.4 X as wide as long, its posterior angles
foveolate. Legs robust, front femora measuring only 2.2 X as long
as their greatest width ; claws weakly dentate.

Pnratypes. — 1 4 c? cf , brazil : same data as type and allo-

type except various dates (July 1937, May 1938, June 1962, Nov.
1963) [MCZ, BMNH].

Variation. — The female paratype is of the same size as the allo-
type and shows no important differences in color, sculpture, or
standard measurements. The four male paratypes vary in length
from 3.2 to 4.5 mm, LFW 2.7-3. 1 mm; WF varies from 1.00-1.15
X HE. One of these specimens has an unusually broad ocellar
triangle, and in this specimen OOL is only 1.22 X WOT; in this
specimen the propodeal disc is also shorter than usual, measuring 1.6
X as wide as its median length.

ALPHABETICAL LIST OF ABBREVIATIONS USED IN TEXT

Structures

HE: height of eye (maximum, lateral view)

LFW : length of fore wing

LH : length of head (apical margin of clypeus to median vertex

crest)

OOL: ocello-ocular line (minimum distance from eye to lateral

ocellus)

WF : width of front (measured at its minimum point)

WH: width of head (maximum, including eyes)

WOT: width of ocellar triangle (including lateral ocelli)

Institutions

BMNH: British Museum (Natural History), London

CAS: California Academy of Sciences, San Francisco

CU : Cornell University, Ithaca, N.Y.

MCZ : Museum of Comparative Zoology, Cambridge, Mass.

UCD : University of California, Davis

USNM: United States National Museum, Washington

THE TAXONOMIC PROBLEM OF SEXUAL
DIMORPHISM IN SPIDERS AND A SYNONYMY IN
MYRMECO TYPUS (ARANEAE, CLUBIONIDAE)'1

By Jonathan Reiskind
Biological Laboratories, Harvard University

With the major exception of many of the vertebrate groups, some
mollusks, and butterflies, much of the current evolutionary work in
zoology requires, concurrently, research at the “alpha” level of tax-
onomy in which species are characterized and named (Mayr, Linsley
and Usinger, 1953). While this appears to be just a continuation of
the work of Linnaeus, Clerck, and others it is by no means an old-
fashioned, unchanging endeavor. With the advent of Darwin’s theory
and, later, the new systematics of the 20th century alpha tax-
onomy has continued to incorporate the most recent advances in
evolutionary biology. Behavioral, ecological, distributional, physiologi-
cal and biochemical characteristics must be utilized in this “lowest
level” of taxonomy in addition to the traditional morphology. I11
this way the typological and morphological result of overlooking two
sibling species or splitting a single species into two species can be
avoided. An error of the latter type is often a result of sexual dimor-
phism.

In non-hermaphroditic, sexually reproducing organisms there
usually exists some sort of morphological sexual dimorphism. Some-
times this is limited to the sexual structures themselves, but more
often it is extended to secondary sexual characters. In spiders there
are three types of secondary characters. Type one is intimately in-
volved in the physical act of copulation and is represented by the
pedipalps of males which are modified into organs for the transfer-
ence of seminal fluid to the female during mating. Type two is as-
sociated with the courtship before mating which includes the bizarre
and colorful structures that are observed in the Salticidae (especially
the males) as well as the size differences in certain argiopoid groups.
All structures resulting from sexual selection or reproductive re-
quirements fall into this second type. Type three is unrelated to the
procreative process and includes size, color and shape differences
whose origins are either due to non-sexual selection or possible pleio-
tropic effects.

’Research a by-product of National Institutes of Health Grant No. AI-01944
to Dr. H. W. Levi, and of an NSF Graduate Fellowship.

Manuscript received by the editor January 28, 1966.

279

280

Psyche

[December

The problem that results from this dimorphism is that of properly
pairing the male and female of the same species. With the rare ex-
ception of finding two spiders in copula or, in some Araneidae for
example, finding the apical tip of the embolus broken off inside the
seminal receptacle of the female, this problem must be solved indirect-
ly. This is the point at which a taxonomist’s intuition or reason is
required. In addition to the comparison of the two adult sexes the
use of the immature forms may now come in use. Many araneologists
consider pre-adult forms (i.e. those lacking male palpi or epigyna) to
be taxonomically useless. But the external morphology of the im-
matures will be less affected by the strong sexual dimorphism found in
many adults and therefore exhibit the more stable characters in the
species.

In a recent article (Unzicker, 1965) two new species of Myrme-
cotypus were described. Each one was described from a single speci-
men: M. rettenmeyeri Unzicker from a male, and M. discreta

Fig. 1. Lateral view of Myrmceotypus rettenmeyeri Unzicker.
Fig. 2. Lateral view of Camponotus serieeiventris (Guerin).

1965]

Reiskind — Dimorphism in Spiders

281

Unzicker from a female. Both species were collected at the same
locality, in the same way, within three days of one another. The two
spiders are almost identical with the obvious exception of sexual
structures. Hence it is strongly suggested that M. discreta is the
synonym of M. rettenmeyeri. In addition, examination of an im-
mature specimen of this species reveals a basic external morphology
and color pattern that is in common with both adults. This species
does not exhibit as much sexual dimorphism as in many other mem-
bers of the Micariinae where the immature forms are even more
useful.

Often a species can be properly delimited only after some contact
with the living form in its natural habitat. Both M. rettenmeyeri and
J\l. discreta have a strange “longitudinal band of black hairs extend-
ing along midline of cephalothorax from dorsal depression to base of
constriction at posterior end” (Unzicker, 1965) (see Fig. 1). From
observations and collections made by Dr. A. M. Chickering at the
type locality in Panama it appears that in both sexes the band of hair
enhances the resemblance to the ant Camponotus sericeiventris
(Guerin) with which they share the same habitat. The crests of
black hairs correspond to the solid longitudinal keel-like dorsal ex-
tensions of the posterior sections of the thorax of C. sericeiventris
(Fig. 2). The chances that two spiders in the same genus imitate
the same ant at the same place and time are ecologically improbable.
This is strong additional evidence supporting the synonymy of two
species.

To summarize, “alpha taxonomy” is presently a dynamic science
incorporating many new techniques. Sexual dimorphism presents the
problem of inadvertant splitting of species which can often be
avoided by the use of careful comparisons as well as immature stages
and field observations. Using these methods Myrmecotypus discreta
is found to be the female form of M . rettenmeyeri. As first reviser
I select M. rettenmeyeri as the valid name.

I wish to thank Dr. Carl Rettenmeyer for the loan of the types
of the spiders cited above.

References

Mayr, E., E. G. Linsley, and R. L. Usinger.

1953. Methods and Principles of Systematic Zoology. McGraw-Hill,
New York, 336 pp.

Unzicker, J. D.

1965. Two New Species of the Genus Myrmecotypus from Central
America (Clubionidae : Araneae). Journ. Kansas Ent. Soc. 38
(3) :2 5 3 -2 57.

THE HABITS AND DISTRIBUTION OF
MACROMISCHA SUBDITIVA WHEELER
(HYMENOPTERA: FORMICIDAE)

By William S. Creighton'1
Department of Biology, City College New York

Before discussing Macromischa subditiva I wish to review some
older observations on the habits of this beautiful genus. The nests
of Macromischa are seldom easy to find. The one exception to this
rule appears to be M. sallei Guerin, whose abundant and conspicious
carton nests, built around the twigs of bushes, are certain to attract
attention. It thus happens that M. sallei is the only species in the
genus for which adequate field data exist. In 1913 W. M. Mann
studied hundreds of nests of sallei in Haiti (1). The uniformity
of these nests was remarkable; they were invariably constructed of
carton and invariably suspended from the branches of bushes or
small trees. While Salle’s San Domingan nests had come from bushes
growing on marshy plains (2) the Haitian nests occurred on rocky,
well-drained ridges, hence there was no reason to suppose that the
latter nests had been placed in bushes to avoid water-soaked soil. In
short, the nesting habits of M. sallei appeared to be not only arboreal
but also those of a non-adaptable arboreal.

This circumstance strongly influenced W. M. Wheeler in the
nidification list which he published in his 1931 study of Macro-
mischa (3). At that time Wheeler had nesting data for 28 species
of Macromischa and he knew that there is a wide range of nesting
response within the genus. His list carries eight nesting categories
and, while some of these are rather similar, the range extends from
strictly arboreal to strictly terrestrial species. This list is invaluable
to anyone who is studying Macromischa in the field and, since
Wheeler was trying to show no more than the variable nesting habits
of different species of Macromischa, he amply accomplished his pur-
pose. The objectionable feature of the nidification list is its failure
to give the number of nests on which the inclusions are made. To
secure this information it is necessary to consult the authorities
whose names are carried after the names of the species whose nests
they observed. When this is done it is clear that 17 of the 28
species listed were known from a single nest and hence could appear

Tmeritus Professor, Department of Biology, City College.

Manuscript received by the editor October 3, 1965

282

1965]

Creighton — Macromischa subditiva

283

in only one nesting category. Nor was the situation much better
with most of the 1 1 other species. Seven of these were known from
two colonies each and two from three colonies each. Thus there
were only three species ( flavitarsis 5 colonies; wheeleri 8 colonies,
squamifera 15 colonies) which might be said to furnish more than
a suggestion of nest preferences. What Wheeler did with these last
two species is astonishing. He knew that both wheeleri and squamata
have flexible nesting habits. In 1920 W. M. Mann published obser-
vations that leave no doubt on this score and I had later amplified
Mann’s data in a personal communication to Dr. Wheeler (4).
Yet both wheeleri and squamata each appeared in a single category
in the nidification list. Thus, although Wheeler saw clearly that
nesting responses vary widely within the genus Macromischa, he
failed to appreciate that the nesting response of the individual species
may also be variable. On the basis of present data it is impossible
to say what percentage of species in the genus possess flexible nesting
habits but, if further progress is to be made with the habits of Macro-
mischa, it is imperative to recognize that some of the species, among
them subditiva, behave in this fashion.

Remarkably few records of any kind have been published for M.
subditiva since Wheeler described this species in 1903 (5). In 1912
Mitchell and Pierce provided a two-line habit note on specimens
taken in Victoria County, Texas which repeated the observations
carried in Wheeler’s original description (6). When M. R. Smith
monographed our species in 1939 he gave no new data on habits
and added only one new locality record ( 7 ) . Apparently there are
no other published records for subditiva, although M. R. Smith
stated in a paper published in 1947, that the species occurs in Lou-
isiana (8). This reference is enigmatic, since no locality was cited
and since repeated surveys in the area between Austin and the
Louisiana border have failed to turn up subtiva in eastern Texas.
The record is not included in the list presented in this paper.

From what has already been said it should be obvious that it is
important to distinguish between records based on strays and records
where the nest was found. I have, therefore, divided the records
into two groups, the first based on strays (Table I), the second on
nests taken (Table II).

In six of the above colonies a single female was present. It is
impossible to say whether this was true of the seventh nest (Wim-
berley colony) for part of this colony was scattered when the crevice
in which it was living was forced open. In addition to the female

284

Psyche

[December

Table I

Records for

M. subditiva based upon strays

Station

Collector

Where taken

TEXAS :

Austin (Travis Co.)

W. M. Wheeler

On leaves of bushes

New Braunsfels

W. M. Wheeler

Dead limbs on ground

(Comal Co.)

Harlingen

R. A. Vickery

Not stated

(Cameron Co.)

10 miles west of

W. S. Creighton

Cavities in dead

Boca Chica

mesquite limb

(Cameron Co.)

Fowlerton

W. S. Creighton

Cavities in dead

(La Salle Co.)

mesquite limb

Delta Lake

W. S. Creighton

On willow bark

(Hidalgo Co.)

Table II

Records for M. subditiva based

upon nests

Station

Collector

Nest site

TEXAS :

Austin (Travis Co.)

W. M. Wheeler

In willow bark
( Salix sp.)

Victoria

(Victoria Co.)

J. D. Mitchell

Under willow bark
( Salix sp.)

30 miles NE of

W. S. Creighton

Burrows in live-oak

Raymondsville

limb ( Q . virginiana )

(Kenedy Co.)

2 miles west of

W. S. Creighton

Crevice in limestone

Wimberley (Hays Co.)

ledge.

Bentsen State Park

W. S. Creighton

Under hackberry bark

(Hidalgo Co.)

( Celtis laevissima )

La Feria

W. S. Creighton

Abandoned termite

(Cameron Co.)

burrows in partly
buried plank

NUEVO LEON:

Chipinque Mesa

W. S. Creighton

Burrows in live-oak

(Monterrey)

limb ( Q . fusiformis)

1965] Creighton — Macrotnischa subditiva 285

the largest colony contained 145 workers, the smallest one only
twelve workers. It appears, therefore, that the colonies of subditiva
are always small and seldom, if ever, pleometrotic.

Two captive colonies were established in February 1965 and it
was soon apparent that subditiva is a very easy ant to maintain in
artificial nests. It appears to be omnivorous, since the captive colonies
rarely refused anything edible, but it has a strong preference for
insect food. The captive colonies ate termites, fruit flies, house
flies, crane flies, mosquitos, dermestid beetles, crickets, caterpillars
and various moths and butterflies. With the exception of adult
sawflies and stoneflies, which they plainly disliked, although they
ate them, they accepted this varied insect diet without hesitation.
They are one of the few ants which the writer has studied that
would eat cut-worms. Their favorite food appeared to be the
larvae of buprestid bettles. The ant larvae were mostly fed by
regurgitation but, on occasion, bits of insect tissue were placed in
their jaws.

In the captive colonies the rate of egg-laying averaged out to
slightly less than three eggs per day. The eggs hatched into larvae
in about 30 days and these transformed into pupae in about 23 days.
The pupal period was about 19 days. The pupae darken extensively
after 14 days and at emergence are so deeply colored that there is
no callow period in the strict sense of the term. These newly
emerged adults can be told from their older nestmates but this is
by no means easy for the color difference is slight and largely con-
fined to the lower surface of the body. It is usually easier to tell
a “callow” by its actions for, during the first two or three days
after emergence they take little part in the nest activities. The
pupal exuvium is pulled off in long strips by the workers. Two or
three will often work together at stripping off the exuvium, which
they apparently eat. During the stripping the emerging imago often
assists the process by bending its body from side to side.

It appears that subditiva produces and matures brood throughout
the year. The writer has taken nests of subditiva from the middle
of October to the middle of March and these have invariably con-
tained brood. With the exception of two male pupae this brood has
been free of sexual forms, hence it seems likely that under ordinary
conditions only worker brood is produced during the winter months.
It may be added that subditiva has no trouble bringing brood through
in artificial nests. During the time that the captive colonies were
under observation the population of one of them more than tripled.

286

Psyche

[December

Literature Cited

1. Wheeler. W. M. & Mann, W. M., Bull. Amer. Mus. Nat. Hist. 33: 35

2. Guerin-Meneville, F. E., Rev. Mag. Zool. Paris, 2nd Ser. 4: 73 (1852).

(1914).

3. Wheeler, W. M., Bull. Mus. Comp. Zool. 72 (1): 5 (1931).

4. Mann, W. M., Bull. Amer. Mus. Nat. Hist. 42: 413, 423 (1920).

5. Wheeler, W. M., Psyche, 10: 99 (1903).

6. Mitchell, J. D. & Pierce, W. D., Proc. Ent. Soc. Wash. 14: 73 (1912).

7. Smith, M. R., Ann. Soc. Ent. Amer. 32, (3) : 506 (1939).

8. Smith, M. R., Amer. Mid. Naturalist, 37, (3) : 575 (1947).

NOTES ON THE INDO-AUSTRALIAN ANTS OF GENUS
SIMOPONE FOREL ( HYMENOPTERA-FORMICIDAE)1

By Robert W. Taylor
Biological Laboratories, Harvard University

I recently discussed a Philippine specimen of Simopone which was
tentatively identified as S. bakeri Menozzi (Taylor, 1965: 6). Sub-
sequent examination of the bakeri holotype has shown the Philippine
specimen to be representative of a new species, which is described be-
low, along with diagnostic notes on the bakeri type. Three species of
Simopone are now known from the Indo-Australian area. All are
represented only by their worker holotypes. They are similar in
habitus, and constitute a compact and undoubtedly closely related
complex within this predominantly African cerapachyine genus. The
generic diagnosis presented earlier (Taylor, loc. cit.) needs no mod-
ification at this point.

Simopone chapmani sp. n.

Type locality. Philippine islands: Negros: Horns of Negros,
3600 ft. The unique worker holotype was collected, presumably in
rain forest, by the late Dr. James W. Chapman, for whom this
species is named.

Type deposition. Museum of Comparative Zoology, Harvard Uni-
versity (Type No. 31 199).

Description. Dimensions (in mm) : aggregate total length (TL)
ca. 5.0; head length at midline (HL) 0.91; maximum head width
(HW) 0.68; cephalic index (HW X 100/HL) (Cl) 75; maxi-
mum eye diameter 0.30; ocular index (eye diameter X 100/HW)
(OI) 44; minimum interocular distance 0.34 (1.14 X eye diam-
eter) ; PW 0.55; WL 1.2; petiolar node length at midline 0.68;
maximum petiolar node width 0.56; postpetiolar length at midline
0.64; maximum postpetiolar width 0.61. Generally similar to the
New Guinean S. gressitti Taylor, and agreeing with its original de-
scription (Taylor, 1965, Breviora 22 1 : 3), except in the following
characters :

Research supported by U. S. National Science Foundation Grant No. GB
1634.

The Simopone bakeri Menozzi holotype, discussed here, was generously
loaned for study by Prof. Dr. Guido Grandi, Istituto di Entomologia.
Universita degli studi, Bologna, Italy.

Manuscript received by the editor January 19, 1966.

287

288

Psyche

[December

1. Smaller size, with narrower head and proportionately large
eyes, which are less widely separated relative to their maximum
diameter; postpetiole slightly longer than broad in dorsal view
(length 1.05 X width). (The relevant dimensions of gressitti are:
TL ca. 6.4 mm; Cl 81; OI 38; eyes separated by 1.53 X their
maximum diameter; postpetiole length 0.70 mm, width 0.75 mm —
length 0.93 X width.)

2. Transverse occipital carina lacking distinct ribs along its an-
terior edge.

3. Lateral suturation of mesosoma complete, but less distinct.
Transverse ribbing vestigial on dorsal promesonotal suture and lack-
ing on dorsal mesometanotal suture.

4. Transverse anterior petiolar carina without ribs.

5. Sides of postpetiole, in dorsal view, almost parallel, very
feebly divergent posteriorly.

6. Body entirely smooth and shining, except for some vestigial
fine longitudinal striae on frontal carinae, and scaly microsculpture on
sides of pronotum and propodeum, and on metepisternal areas.

7. Post-cephalic ground pilosity more abundant. Very line short
reclinate to suberect hairs scattered on dorsum of mesosoma,
most abundant on sides of propodeal dorsum. Similar pilosity mod-
erately abundant on dorsa of petiole and postpetiole. Single, more
erect, but barely longer hairs at midlength of each frontal carina and
above each eye, but not elsewhere. No subpetiolar “brush” developed;
only 2 hairs break the subpetiolar profile.

The mandibular dentition is worn but was probably originally
like that of bakeri , as described by Menozzi. The legs are damaged;
so details of the tibial and tarsal armament are not known.

Relationships. See below under S. bakeri.

Sunopone bakeri Menozzi

Simopone bakeri Menozzi, 1926, Atti Soc. Nat. Mat. Modena, (6)

4: 92 (1925), worker. Type locality: Singapore.

The notes below are based on the unique holotype, which has the
following dimensions (in mm, see preceding description for explana-
tion of symbols) : TL ca. 5.6; HL 1.00; HW 0.65; Cl 65; maxi-
mum eye diameter 0.32; OI 49; minimum interocular distance 0.32
(equal to eye diameter) ; PW 0.58; WL 1.36; petiolar node length
at midline 0.72 ; maximum petiolar node width 0.62 ; postpetiolar
length at midline 0.73; maximum width of postpetiole 0.65. Close
to chapmani and gressitti, and agreeing with the original description
of the latter, except in the following characters:

1965]

Taylor — Genus Simpone

289

1. Smaller size (about as in chapmani) , with much narrower
head and proportionately much larger eyes, which are less widely
separated relative to their maximum diameter; postpetiole distinctly
longer than broad in dorsal view (length 1.12 X width).

2. Anterior clypeal border strongly biconcave in frontal view,
with a distinct median anterior tooth.

3. Transverse occipital and anterior petiolar carinae as in chap-
mani.

4. Lateral mesosomal suturation weaker, as in chapmani. Dorsal
promesonotal suture distinctly transversely ribbed ; mesometanotal
suture lacking on mesosomal dorsum.

5. Sides of postpetiole as in chapmani , almost parallel in dorsal
view.

6. Sculpturation generally as in chapmani , but scaly microsculp-
ture lacking on sides of pronotum, which are smooth and shining.

7. Vestiture much more abundant than in either gressitti or
chapmani. Moderately long reclinate to sub-erect hairs fairly abun-
dant on head, those at center of frons shortest and least abundant,
those along frontal carinae more erect, a single more elongate hair
above each eye. Dorsa of mesosoma, petiole and postpetiole with
similar, but somewhat longer pilosity, least abundant on disc of prono-
tum, best developed on propodeum and petiole. Single longer hairs
on pronotal humeri, on petiole slightly behind its anterior corners,
and dorso-laterally at about mid-length of postpetiole. Anterior
gastric pilosity slightly less abundant and shorter than that of nodes,
several more elongate hairs laterally, apex surrounded by long arched
hairs. The subpetiolar process is partly obscured by glue; it appears
that about 10-12 long hairs normally break its ventral profile; they
do not form a regular brush-like series as in gressitti.

Relationships. S. bakeri is more similar, and probably more closely
related to chapmani than to gressitti. It is easily distinguished from
chapmani by the cephalic and ocular proportions, the structure of the
anterior clypeal border, and the more abundant vestiture. These
three species form an apparently morphoclinal sequence (bakeri — >
chap??iani—> gressitti) in various characters, including decrease in
relative size of the eyes (i.e. OI), and increase in relative head width
(i.e. Cl), in the spacing of the eyes relative to their maximum diam-
eter, and in the width of the postpetiolar dorsum relative to its length.
Reduction and specialization of the pilosity also follow this sequence.
S. bakeri appears to have the most primitive habitus and probably
most resembles the ancestral Indo-Australian stock, except perhaps
for its possession of a median clypeal tooth, and the lack of a dorsal
mesometanotal suture.

290

Psyche

[December

Reference

Taylor, R. W.

1965. New Melanesian ants of the genera Simopone and A7nblyopone
(Hymenoptera-Formicidae) of Zoogeographic significance. Bre-
viora, 221: 1-11 (pp. 1-6).

A NEW SPECIES OF THE SPIDER GENUS XYSTICUS
(ARANEAE: THOMISIDAE) FROM ARIZONA*

By J. H. Redner and C. D. Dondale
Research Institute, Research Branch, Canada Department of
Agriculture, Belleville, Ontario

The North American crab spiders are now comparatively well
known taxonomically. This is particularly true of the species in the
large genus Xysticus , which has been twice revised continentally by
Gertsch (1939, 1953) and treated in more regional works by Buckle
and Redner (1964), Schick (1965), and Turnbull, et al. (1965).
It seems probable that any additional new forms that will be dis-
covered will be from remote parts of the continent or in sibling re-
lationship with known species. Several species have, however, been
described only from one sex.

The purpose of this paper is to describe a distinctive new species
of Xysticus from the mountainous parts of Arizona. Its structure
clearly places it in the locuples group of the apophysate division of
the genus, and its range suggests it to be an inland endemic of the
southwestern United' States.

Xysticus humilis sp. n.

(figs. 1-5)

Male: Total length 3.45 and 3.16 mm.; carapace 1 .88 and 1.68
mm. long and 1.82 and 1.59 mm. wide; femur II 1.86 and 1.68 mm.
long (measurements made, respectively, on the holotype and a para-
type). Carapace low and smoothly rounded from side to side, being
little higher at level of legs II than at level of posterior eye row;
front nearly vertical ; surface with thin coat of short, stiff, recumbent
setae; eye area and front set with several spiniform setae; orange-
yellow in color, with indistinct pale V in front of dorsal groove,
which is uncolored, and with thin black line at side margins. Both
rows of eyes recurved; ocular quadrangle wider than long, slightly
wider behind than in front; laterals larger than medians; posterior
medians only slightly closer to each other than to posterior laterals.
Legs pale yellowish, I and II slightly darker than III and IV and
with orange-brown tibiae; femur I with 3-4 weak prolateral spini-
forms, 1-2 dorsals, o retrolaterals; tibia I with 0-3 weak dorsal spini-
forms, o prolaterals, 4 pairs of ventrals, o retrolaterals; tarsal claws

* Manuscript received by the editor January 3, 1966.

291

Psyche, 1965

Vol. 72, Plate 21

Redner and Dondale

Xysticus

1965]

Redner and Dondale — Xysticus

293

with 5 teeth. Abdominal dorsum pale yellowish to off-white, with
pattern composed of thin lateral black lines on anterior half and 4-5
similar transverse lines posteriorly.

Tibia of palpus with ventral apophysis flattened and bladelike, and
bearing small basal lobe ; distal tegular apophysis flattened ; embolus
not thickened at tip (Figs. 2, 4).

Female: Total length (allotype) 5.21 mm.; carapace 2.61 mm.
long and 2.46 mm. wide; femur II 2.32 mm. long. Structure and
color essentially as in male, but femora I and II concolorous with other
segments; abdominal pattern and carapace setation as in Figure 1.
Epigynum with deep atrium and paired atrial sclerites as in Figure
3; spermathecae as in Figure 5, each copulatory tube arising pos-
teriorly and forming convoluted mass dorsal to anterior end of
spermatheca.

Range: Arizona.

Type Locality: Holotype male and one paratype male from Portal,
Cochise County, Arizona, July 19, 1964, J. A. Woods and V. Roth
collectors. Deposited in the American Museum of Natural History,
New York.

Other Locality: Allotype female from 2 miles north of Rodeo,
Arizona, June 12, 1957, Statham and Plimton collectors. Deposited
in the American Museum of Natural History, New York.

Diagnosis: X. kumilis most resembles X. bradti Gertsch, which
is known only from males taken in the State of Chihuahua, and X .
texanus Banks, a better-known species, from Nuevo Leon, Texas,
Arizona, Colorado, and the southeastern United States. Both X.
bradti and X . texanus are relatively small in size and weakly setaceous
in carapace and legs. X. kumilis is distinct from these and all other
known species of Xysticus in its low, pale, smoothly-convex carapace,
in its abdominal pattern of thin black lines, and in details of the ex-
ternal genitalia. The male palpus has a stout basal tegular apophysis
with a small irregularity midway along its basal margin (Fig. 4),
whereas in bradti and texanus this structure is slender and smoothly
tapered. The distal tegular apophysis further differs from that of
texanus in being flat and rounded in outline rather than slender and
“heeled”, while the epigynal atrium is nearly circular instead of
broadened, and the atrial sclerites are approximately ovoid instead
of elongate and slender.

Explanation of Plate 21

Figs. 1-5. Xysticus kumilis sp. n. 1. Dorsal view of female. 2, 4. Male
palpi. 3, 5. Female epigynum and spermathecae.

294

Psyche

[December

Acknowledgments

The specimens described in this paper were lent to us by Dr. W.
J. Gertsch, American Museum of Natural History, New York, to
whom we are sincerely thankful.

Literature Cited
Buckle, D. J. and J. H. Redner

1964. The nearctic species of the Xysticus labradorensis subgroup
(Araneae: Thomisidae). Canad. Ent., 96: 1138-1142.

Gertsch, W. J.

1939. A revision of the typical crab-spiders (Misumeninae) of Amer-
ica north of Mexico. Bull. Amer, Mus. Nat. Hist., 76: 277-442.
1953. The spider genera Xysticus, Coriarachne , and Oxyptila (Thom-
isidae, Misumeninae) in North America. Bull. Amer. Mus. Nat.
Hist., 102: 413-482.

Schick, R. X.

1965. The crab spiders of California (Araneida, Thomisidae). Bull.
Amer. Mus. Nat. Hist., 129: 1-180.

Turnbull. A. L., C. D. Dondale and J. H. Redner

1965. The spider genus Xysticus C. L. Koch (Araneae: Thomisidae)
in Canada. Canad. Ent., 97: 1233-1280.

THE TYPES OF PROCTOTRUPOIDEA
(HYMENOPTERA) IN THE CHARLES T. BRUES
COLLECTION AT THE MUSEUM OF COMPARATIVE

ZOOLOGY

By Lubomir Masner*

Institute of Entomology, Czechoslovak Academy of Sciences, Prague

During a two week visit to the Museum of Comparative Zoology
in June, 1964, I examined the types of 60 species of Proctotrupoidea
described by Charles T. Brues. These belonged to the following
families: 21 Proctotrupidae, 7 Diapriidae, 13 Scelionidae, 15 Platy-
gasteridae, and 4 Ceraphronidae. In the present report a number
of lectotypes are selected, some new combinations are suggested, and
an account of the state of preservation of all types listed is given.
The names and synonymy used in this paper are those proposed by
Masner and Muesebeck (Supplement 2, Hymenoptera of America
north of Mexico. United States Department of Agriculture,
in press). The taxonomic categories subfamily and tribe are not used
in the present paper; the genera and species are arranged alpha-
betically throughout the families. Under each species are listed the
various labels on the type specimen, from top to bottom, each “label”
being separated by a semicolon. Specimens are glued to card points
except when specifically mentioned as pinned (on minuten nadeln).

Family PROCTOTRUPIDAE

Codr us Panzer, 1801
Codrus angusticeps (Brues)

Proctotrypes angusticeps Brues, 1910. Wis. Nat. Hist. Soc. Bui. 7: 112.
Mt. Constitution, Jul. 09, Wash.; Type; MCZ type label; Exallonyx
angusticeps Brues

Holotype female No. 31002, well preserved. Unique.

Codrus ashmeadi (Brues)

Exallonyx ashmeadi Brues, 1919. N. Y. Ent. Soc. Jour. 27: 10, 13.

*Postdoctorate Fellow, National Research Council of Canada, Research
Institute, Canada Department of Agriculture, Belleville, Ont., Canada.

Thanks are extended to the staff of the Museum of Comparative Zoology,
particularly to Professor E. Mayr, Dr. H. E. Evans, and Professor P. J.
Darlington, Jr. Thanks are also due to the National Research Council of
Canada for support during completion of this paper.

Manuscript received by the editor January 16, 1966.

295

296

Psyche

[December

Machias, Me., VII-20-09, C. W. ; MCZ type label; Exallonyx ashmeadi
Brues

Holotype male No. 31006, well preserved. Three paratype males.
Codrus carinatus (Brues)

Exallonyx carinatus Brues, 1919. N. Y. Ent. Soc. Jour. 27: 11, 15.

Oroville, Wash., May 1, 1912; Type; MCZ type label; Exallonyx cari-
natus Brues

Holotype male No. 30993, well preserved. Unique.

Codrus fuscicornis (Brues)

Exallonyx fuscicornis Brues 1919. N. Y. Ent. Soc. Jour. 27: 10, 12.
Woods Hole, Mass.; Type; MCZ type label; Exallonyx fuscicornis Brue^
Holotype male No. 31005, well preserved. Unique.

Codrus grandis (Brues)

Exallonyx grandis Brues, 1919. N. Y. Ent. Soc. Jour. 27: 11, 17.

Ramsey, N. J. ; VIII-22-9; Type; MCZ type label; Exallonyx grandis
Brues

Holotype female No. 3 1010, well preserved. Unique.

Codrus ohscuripes (Brues)

Proctotrypes ohscuripes Brues, 1910. Wis. Nat. Hist. Soc. Bui. 7: 112, 114.
Mt. Constitution, Jul. 09, Wash.; Type; MCZ type label; Exallonyx oh-
scuripes Brues

Holotype male No. 31007, well preserved. One paratype male +
documentary specimens.

Codrus pallidicornis (Brues)

Exallonyx pallidicornis Brues, 1919. N. Y. Ent. Soc. Jour. 27: 11, 14.
Putnam Ct., 12 July 1905, HL Yiereck ; Type; MCZ type label; Exallonyx
pallidicornis Brues

Holotype male No. 30994, well preserved. Two paratype males.
Codrus parvulus (Brues)

Exallonyx parvulus Brues, 1919. N. Y. Ent. Soc. Jour. 27: 11, 16.
Oroville, Wash., I-IV, Melander; Type; MCZ type label; Exallonyx
parvulus Brues

Holotype male No. 30992, left antenna after 9th segment off.
Eleven paratype males.

Codrus placidus (Brues)

Proctotrypes placidus Brues, 1910. Wis. Nat. Hist. Soc. Bui. 7: 112, 113.
Mt. Constitution, Jul. 31-8, Wash.; Type; MCZ type label; Exallonyx
placidus Brues

Holotype male No. 31004, well preserved. Unique.

Codrus pleuralis (Brues)

Exallonyx pleuralis Brues, 1919. N. Y. Ent. Soc. Jour. 27: 10, 14.

Type; MCZ type label; Exallonyx pleuralis Brues
Holotype male No. 30991, well preserved. Two paratype males.
Codrus serricornis (Brues)

Proctotrypes serricornis Brues, 1910. Wis. Nat. Hist. Soc. Bui. 7: 112, 115.
Puget Sound, Wash., Jul. 08; Type; MCZ type label; Exallonyx serri-
cornis Brues

1965]

Masner — Proctotrupoidea

297

Holotype male No. 31009, right antenna after 6th segment and
left antenna after 3rd segment missing. Two paratype males.

Codrus similis (Brues)

Exallonyx similis Brues, 1919. N. Y. Ent. Soc. Jour. 27: 10, 11.

Tacoma, Wash., 28-VIII-ll; Type; MCZ type label; Exallonyx similis
Brues

Holotype female No. 31003. Four paratype females.

Codrus simplicior (Brues)

Proctotrypes simplicior Brues, 1910. Wis. Nat. Hist. Soc. Bui. 7: 112, 116.

Puget Sound, Wash., Jul. OS; Type; MCZ type label; Exallonyx simplicior
Brues

Holotype male No. 30990, right antenna after 7th segment missing,
left antenna and wings missing, legs and gaster glued separately to
card point. Unique ( !).

Cryptoserpkus Kieffer, 1907
Cryptoserphus melanderi Brues

Cryptoserpkus melanderi Brues, 1919. N. Y. Ent. Soc. Jour. 27: 8.

Pullman, Wash., V-18-09; Type; MCZ type label; Cryptoserphus melan-
deri Brues

Holotype male No. 31000, well preserved. Two paratype males.
Cryptoserphus occidentals Brues

Cryptoserphus occidentalis Brues, 1919. N. Y. Ent. Soc. Jour. 27: 7.

Chatcolet Lake, Idaho, VIII-1915, A. L. Melander; Type; MCZ type
label; Cryptoserphus occidentalis Brues

Holotype female No. 30999, left antenna missing after 2nd seg-
ment. Four paratype females.

Nothoserphus Brues, 1940
Nothoserphus mirabilis Brues

Notroserphus mirabilis Brues, 1940. Amer. Acad. Arts Sci. Proc. 73: 263.

Taihorin, Formosa, H. Sauter, 1911; 7. VII; MCZ type label; Nothoser-
phus mirabilis Brues

Holotype male No. 29789, left antenna broken after 3rd segment,
but remainder glued to mount; right antenna missing after 9th seg-
ment (pinned). One paratype male.

Phaenoserphus Kieffer, 1908
Phaenoserphus longipes Brues

Phaenoserphus longipes Brues, 1919. N. Y. Ent. Soc. Jour. 27: 9.

Almota, Wash., 24-VI, Melander; Type; MCZ type label; Phaenoserphus
longipes Brues

Holotype male No. 31001, well preserved. Unique.

Proctotrupes Latreille, 1796
Proctotrupes cocker elli (Brues)

Serphus cockerelli Brues, 1919. N. Y. Ent. Soc. Jour. 27: 3, 4.

Eldora, Colo., Aug. 18 (Cockerell); Type; MCZ type label; Serphus
cockerelli Brues

Holotype male No. 30997. well preserved (pinned). Unique.

298

Psyche

[December

Proctotrupes debilis (Brues)

Serphus debilis Brues, 1919. N. Y. Ent. Soc. Jour. 27: 3, 5.

Wawai, Wash., 20-V-ll; Type; MCZ type label; Serphus debilis Brues

Holotype male No. 30998, well preserved. Unique.

Proctotrupes sequoiarurn (Brues)

Serphus sequoiarurn Brues, 1919. N. Y. Ent. Soc. Jour. 27: 3, 4.

Muir Woods, VIII-7-15, Cal., A. L. Melander ; Type; MCZ type label;
Serphus sequoiarurn Brues

Holotype male No. 30996, left antenna after 8th segment off, left
front leg after coxa missing. Unique.

Proctotrupes zabriskiei (Brues)

Serphus zabriskiei Brues, 1919. N. Y. Ent. Soc. Jour. 27: 2, 3.

Rochester, N. Y., J. L. Zabriskie, 10. VI. 1905; Type; MCZ type label;
Serphus zabriskiei Brues

Holotype female No. 30995, left antenna after 8th segment off.
Unique.

Family DIAPRIIDAE

Auxopaedeutes Brues, 1903
Auxopaedeutes lyriformis Brues

Auxopaedeutes lyriformis Brues, 1910. Wis. Nat. Hist. Soc. Bui. 8: 82.
Forest Hills, Mass., V-16-1910; MCZ type label; Auxopaedeutes lyri-
formis Brues Type

Holotype female No. 31027, well preserved. Unique.

Entomacis Foerster, 1856
Entomacis ambigua (Brues)

Hemilexodes ambigua Brues, 1903. Amer. Ent. Soc. Trans. 29: 126.

Nest Stcnamma fulvum piceum , Colebrook, 8.25.01; Type; MCZ type
label; lectotype selected by L. Masner, 1964; Hemilexodes ambigua Brues
Lectotype female No. 3 1021, well preserved. Two additional fe-
males ( paralectotypes ) .

Hoplopria Ashmead, 1893
Ploplopria grandis Brues
Hoplopria grandis Brues, 1915. Psyche 22: 11.

Porto Velho, Rio Madeira, Brazil, Mann & Baker; Type; MCZ type
label; Hoplopria grandis Brues $ type

Holotype female No. 31022, well preserved. Unique.

Psilus Panzer, 1801
Psilus debilis (Brues) comb. n.

Galesus debilis Brues, 1915. Psyche 22: 12.

Independence, Parahyba, Brazil, Mann & Heath; Type; MCZ type label;
Galesus debilis Brues $ type

Holotype female No. 31025, well preserved. Unique.

Trichopria Ashmead, 1893
Trichopria acutiventris (Brues)

Phaenopria acutiventris Brues, 1902. Amer. Nat. 36: 372.

1965]

Masner — Proctotrupoidea

299

Austin, Tex., 3.01; Type; MCZ type label; in nest Ec'iton caecum;
Phaenopria acutiventris Brues

Holotype female No. 31026, well preserved. Unique.

Xanthopria Brues, 1915
X anthopria nitida Brues
Xanthopria nitida Brues, 1915. Psyche 22: 11.

Independencia, Parahyba, Brazil, Mann & Heath; Type; MCZ type
label; Xanthopria nitida Brues $ type

Holotype female No. 31024, well preserved. Two paratype fe-
males ( ! ) .

Xanthopria opaca Brues

Xanthopria opaca Brues, 1915. Psyche 22: 10.

Independencia, Parahyba, Brazil, Mann & Heath; Type; MCZ type label;
Xanthopria opaca Brues $ type

Holotype female No. 31023, well preserved. Two paratype fe-
males.

Family SCELIONIDAE

Calotelea Westwood, 1837
Calotelea parvipennis Melander et Brues
Caloteleia (!) parvipennis Melander et Brues, 1903. Biol. Bui. 5: 22.
Woods Hole, Mass.; Halictus ; Type; MCZ type label; Caloteleia parvi-
pennis Melander et Brues type

Holotype female No. 3 1014, right antenna after 7th segment and
left antenna after 6th segment off. Unique (!).

Gryon Haliday, 1833
Gryon robustus (Brues)

Hadronotus robustus Brues, 1907. Wis. Nat. Hist. Soc. Bui. 5: 156.

Austin, Tex.; Type; MCZ type label; Hadronotus robustus Brues

Holotype female No. 3 1017, well preserved. Unique.

Lepidoscelio Kieffer, 1905
Lepidoscelio viatrix Brues

Lepidoscelio viatrix Brues, 1917. Nat. Acad. Sci. Proc. 3: 139.

MCZ type label ; Lepidoscelio viatrix Brues, type
Holotype female No. 31018. .Two paratype females (“On an
Orthacris sp. det Uvarov”).

Macroteleia Westwood, 1835
Macroteleia caerulea (Brues) comb. n.

Cacellus caeruleus Brues, 1918. Mus. Comp. Zool. Harv. Bui. 62: 122.
Graciosa Bay, Santa Cruz Is., W. M. Mann ’16; MCZ type label; Cacellus
caeruleus Brues

Holotype female No. 9070, well preserved. Unique.

Classified in Macroteleia with some doubts.

Macroteleia herbigrada Brues

Macroteleia herbigrada Brues, 1915. Psyche 22: 7.

3oo

Psyche

[December

Independencia, Parahyba, Brazil, Mann and Heath; Type; MCZ type
label ; Macroteleia herbigrada Brues, type $

Holotype male No. 31016, right antenna after 6th segment off.
Unique.

Macroteleia pallipes (Brues) comb. n.

Hoploieleia pallipes Brues, 1915. Psyche 22: 8.

Para, Brazil, W. M. Mann; Type; MCZ type label; Hoploieleia pallipes
Brues, type $

Holotype male No. 3 1013, well preserved. Unique.

Classified in Macroteleia Westwood with some doubts.

Paridris Kieffer, 1908
Paridris nigricornis (Brues)

Idris nigricornis Brues, 1903. Amer. Ent. Soc. Trans. 29: 125.

Mixed nest Myr. Lep., Colebrook, 9-10-01; MCZ type label; lectotype
selected by L. Masner, 1964; Idris nigricornis Brues, types (!).

Lectotype male No. 31016, left antenna after 8th segment off.
Two paralectotype males.

Scelio Latreille, 1805
Scelio nitens Brues

Scelio nitens Brues, 1906. Wis. Nat. Hist. Soc. Bui. 4: 110.

Type; Algoa Bay, Capland, 29.11.96, Dr. H. Brauns; MCZ type label;
lectotype selected by L. Masner, 1964; Scelio nitens Brues

Lectotype female No. 3 1019, well preserved. Two paralectotype
females (!) ( ? conspecific with lectotype).

Scelio pulchripennis Brues

Scelio pulchripennis Brues, 1906. Wis. Nat. Hist. Soc. Bui. 4: 109.

Type: Algoa Bay, Capland, 22.1.96, Dr. Brauns; MCZ type label; Scelio
pulchripennis Brues

Holotype female No. 31020, well preserved. Unique (!).

Scelio setiger Brues

Scelio setiger Brues, 1918. Mus. Comp. Zool. Harv. Bui. 62: 121.

Ugi, British Solomon Is., W. M. Mann; MCZ type label; Scelio setiger
Brues

Holotype female No. 9069, well preserved.

Telenomus Haliday, 1833
Telenomus fiskei Brues

Telenomus fiskei Brues, 1910. Psyche 17: 106.

Machias, Me., Aug. 20, 1909; Type; MCZ type label; Telenomus fiskei
Brues

Holotype female No. 3101 1, right antenna missing. Unique (!).
Telenomus heracleicola Brues

T elenomus heracleicola Brues, 1906. Wis. Nat. Hist. Soc. Bui. 45: 143, 146.

Wash. Exp. Sta. No. 026; Type; MCZ type label; lectotype selected by
L. Masner, 1964; Telenomus heracleicola Brues

Lectotype female No. 31012, well preserved (pinned). One para-
lectotype female (!).

1965]

Masner — Proctotrupoidea

301

T'rimorus Foerster, 1856
Trimorus fortis (Brues) comb. n.

Hoplogryon forth Brues, 1915. Psyche 22: 6.

Independencia, Parahyba, Brazil, Mann & Heath ; Type ; MCZ type label ;
Hoplogryon forth Brues $ type

Holotype male No. 30987, well preserved. Unique.

Family PLATYGASTERIDAE

Fidiobia Ashmead, 1895
Fidiobia tritici (Brues)

Rosneta tritici Brues, 1908 Wis. Nat. Hist. Soc. Bui. 6: 157.

Middleport, N Y., 1908; Type; MCZ type label; Rosneta tritici Brues

Holotype female No. 300, missing from the card point (fore wing
only left). Unique.

Gastrotrypes Brues, 1922

Gastrotrypes caudatus Brues

Gastrotrypes caudatus Brues, 1922. Amer. Acad. Arts Sci. Proc. 57: 272.

Forest Hills, Mass., 5,1916; MCZ type label; lectotype selected by L.
Masner, 1964; Gastrotrypes caudatus Brues, types (!)

Lectotype female No. 3 1041, well preserved. 16 paralectotype
females.

Gastrotrypes spatulatus Brues

Gastrotrypes spatulatus Brues, 1922. Amer. Acad. Arts Sci. Proc. 57: 271.

Kartabo, B. G., Jul. Aug. 1920, W. M. Wheeler; MCZ type label; lecto-
type selected by L. Masner, 1964; Gastrotrypes spatulatus Brues

Lectotype female No. 31040, well preserved. Two paralectotype
females.

Isostasius Foerster, 1856
Isostasius crassus Brues

Isostasius crassus Brues, 1922. Amer. Acad. Arts Sci. Proc. 57: 277.

Kartabo. B. G., Jul. Aug. 1920, W. M. Wheeler; MCZ type label; Iso-
stasius crassus Brues, type

Holotype female No. 31030, well preserved. Two paratype fe-
males.

Platygaster Latreille, 1809

Platy g aster anor mis (Brues)

Aneuron anormis Brues, 1910. Wis. Nat. Hist. Soc. Bui. 8: 49.

Albany, N. Y., IV-18, 1908; MCZ type label; lectotype selected by L. Mas-
ner, 1964; Aneuron anormis Brues

Lectotype male No. 31039, left antenna after 4th segment missing.
Four paralectotype males and one paralectotype female.

Platygaster carinifrons (Brues) comb. n.

Polygnotus carinifrons Brues, 1910. Broteria 9: 155.

MCZ type label; lectotype selected by L. Masner, 1964; Polygnotus carini-
frons Brues

302

Psyche

[December

Lectotype male ( ! ) No. 31034, well preserved. Three paralecto-
type males but no female.

Platyg aster coronatus (Brues) comb. n.

Polygnotus coronatus Brues, 1910. Broteria 9: 154.

MCZ type label; lectotype selected by L. Masner, 1964; Polygnotus cor-
onatus Brues

Lectotype female No. 31035, well preserved. Three paralectotype
males.

Platygaster latescens (Brues) comb. n.

Polygnotus latescens Brues, 1910. Broteria 9: 156.

MCZ type label; lectotype selected by L. Masner, 1964; Polygnotus
latescens Brues

Lectotype female No. 31036, well preserved. Three paralectotype
females.

Platygaster mediocris (Brues) comb. n.

Polygnotus mediocris Brues, 1910. Broteria 9: 157.

MCZ type label; lectotype selected by L. Masner, 1964; Polygnotus medio-
cris Brues

Lectotype female No. 31037, well preserved. Three paralectotype
males.

Platygaster novehoracensis (Brues)

Eritrissomerus novehoracensis Brues, 1910. Wis. Nat. Hist. Soc. Bui. 8: 48.

Albany, N. Y., IV. 5, 1907; Type; MCZ type label; lectotype selected by
L. Masner, 1964; Eritrissomerus novehoracensis Brues, types (!)

Lectotype male No. 31038, well preserved. Two paralectotype
males.

Platygaster simplex (Brues) comb. n.

Polygnotus simplex Brues, 1922. Amer. Acad. Arts Sci. Proc. 57: 257.

#b; MCZ type label; lectotype selected by L. Masner, 1964; Polygnotus
simplex Brues

Lectotype female No. 31032, well preserved. Two paralectotype
females ( !) .

Platygaster tubulosa Brues

Platygaster tubulosa Brues, 1922. Amer. Acad. Arts Sci. Proc. 57: 276.

MCZ type label; Platygaster tubulosa Brues, type

Holotype female No. 31033, well preserved. Two paratype fe-
males.

Sceliotrachelus Brues, 1908

Sceliotrachelus braunsi Brues

Sceliotrachelus braunsi Brues, 1908. Genera Insectorum, fasc. 80: 13.

Algoa Bay, Capland, Dr. Brauns; MCZ type label; Sceliotrachelus braunsi
Brues, type

Holotype male ( !) No. 30989, right antenna after 7th segment
off (pinned). One paratype male.

1965]

Masner — Proctotrupoidea

303

Synopeas Foerster, 1856
Synopeas minor (Brues)

Polymecus (Dolichotrypes) minor Brues, 1922. Amer. Acad. Arts Sci.
Proc. 57: 268.

Kartabo, B. G., Jul. Aug. 1920, W. M. Wheeler; MCZ type label; lecto-
type selected by L. Masner, 1964; Polymecus (Dolichotrypes) minor
Brues $ $

Lectotype female No. 31028, well preserved. Two paralectotype
females ( ! ) .

Trichacis Foerster, 1856
Trichacis meridionalis (Brues) comb. n.

Xestonotus meridionalis Brues, 1910. Broteria 9: 150.

MCZ type label; lectotype selected by L. Masner, 1964; Xestonotus meri-
dionalis Brues; Trichacis (Fouts)

Lectotype female No. 3 1031, well preserved. One paralectotype
female ( ! ) .

Family CERAPHRONIDAE

Ceraphron Jurine,. 1807
Ceraphron croceipes Brues
Ceraphron croceipes Brues, 1902. Amer. Nat. 36: 369.

Type; in nest E. coecum; MCZ type label; Ceraphron croceipes Brues

Elolotype female No. 30984, well preserved. Unique.

Conostigmus Dahlbom, 1858
Conostigmus orcasensis (Brues)

Megaspilus orcasensis Brues, 1910. Wis. Nat. Hist. Soc. Bui. 7: 118
Puget Sound, Wash., Jul. 08; MCZ type label; Megaspilus orcasensis
Brues, $ type

Holotype male No. 30985, well preserved. Unique.

Ecitonetes Brues, 1902
Ecitonetes suhapterus Brues
Ecitonetes suhapterus Brues, 1902. Amer. Nat. 36: 370.

Type; Austin, Tex.; MCZ type label; Ecitonetes suhapterus Brues
Holotype female No. 30983, thorax slightly cracked. Unique.
Lygocerus Foerster, 1856
Lygocerus constrictus Brues

Lygocerus constrictus Brues, 1910. Wis. Nat. Hist. Soc. Bui. 7: 119.

Mt. Constitution, Jul. 09, Wash.; MCZ type label; Lygocerus constrictus
Brues, type

Holotype male No. 30986, well preserved. Two paratype males.

304

Psyche

[December

List of the types of Proctotrupoidea

acutiventris, Trichopria, 298
ambigua, Entomacis, 298
angusticeps, Codrus, 295
anormis, Platygaster, 301
ashmeadi, Codrus, 295
braunsi, Sceliotrachelus, 302
caerulea, Macroteleia, 299
carinatus, Codrus, 296
carinifrons, Platygaster, 301
caudatus, Gastrotrypes, 301
cockerelli, Proctotrupes, 297
constrictus, Lygocerus, 303
coronatus, Platygaster, 302
crassus, Isostasius, 301
croceipes, Ceraphron, 303
debilis, Proctotrupes, 298
debilis, Psilus, 298
fiskei, Telenomus, 300
fortis, Trimorus, 301
fuscicornis, Codrus, 296
grandis, Codrus, 296
grandis, Hoplopria, 298
heracleicola, Telenomus, 300
herbigrada, Macroteleia, 299
latescens, Platygaster, 302
longipes, Phaenoserphus, 297
lyriformis, Auxopaedeutes, 298
mediocris, Platygaster, 302
melanderi, Cryptoserphus, 297
meridionalis, Trichacis, 303

minor, Synopeas, 303
mirabilis, Nothoserphus, 297
nigricornis, Paridris, 300
nitens, Scelio, 300
nitida, Xanthopria, 299
noveboracensis, Platygaster, 302
occidentalis, Cryptoserphus, 297
obscuripes, Codrus, 296
opaca, Xanthopria, 299
orcasensis, Conostigmus, 303
pallidicornis, Codrus, 296
pallipes, Macroteleia, 300
parvipennis, Calotelea, 299
parvulus, Codrus, 296
placidus, Codrus, 296
pleuralis, Codrus, 296
pulchripennis, Scelio, 300
robustus, Gryon, 299
serricornis, Codrus, 296
sequoiarum, Procotrupes, 298
setiger, Scelio, 300
similis, Codrus, 297
simplex, Platygaster, 302
simplicior, Codrus, 297
spatulatus, Gastrotrypes, 301
subapterus, Ecitonetes, 303
tritici, Fidiobia, 301
tubulosa, Platygaster, 302
viatrix, Lepidoscelio, 299
zabriskiei, Proctotrupes, 298

THE HABITAT OF PLA TYPA T'ROBUS DARLINGTON
(COLEOPTERA: CARABIDAE)

By Henri Goulet

College Bourget, Rigaud, Quebec, Canada.

Since its description in 1938, Platypatrobus lacustris Darlington has
remained a great mystery. Despite intensive searches both in Canada
and the United States, only six examples had been found. But in
1965 the mystery seemingly was solved, for 51 specimens were cap-
tured by the author and his companions during three visits to Meach
Lake, Que., in Gatineau Park, near Ottawa.

Platypatrobus lacustris was described as a new genus and a new
species from a single female taken by Hubbard and Schwarz at
Batchawana Bay, Ontario (north of Sault Sainte Marie) on August
15, probably in 1876 or 1877. The species was known only from the
type until Dr. C. H. Lindroth and Dr. P. J. Darlington discovered
in the C. H. Frost collection a male taken at light by Dr. A. E.
Brower at Sinclair, in northernmost Maine, on July 30, probably in
the 1950’s. Br. A. Robert found another male at light in Mont
Tremblant Park, Que., on July 1961. On July 28, 1963, two
specimens were captured at light by Mr. and Mrs. R. T. Bell at
Stowe, in northern Vermont. Finally Mr. J. Larson found one
example on Simpson Island, in Great Slave Lake, N.W.T., Canada,
on July 18, 1964; and found another example at light on July 28 at
Lethbridge, Alberta, in 1965.

The rarity of Platypatrobus indicates a special habitat. Dr. Dar-
lington noted that, unlike species of allied genera, specimens of Platy-
patrobus frequently bore mites and he suspected that these might offer
a clue to the habitat of the beetle. Dr. E. Lindquist indentified the
single mite carried by Mr. Larson’s specimen as Protodinychus sp.
and pointed out that Protodinychus occurred commonly in beaver
houses in Algonquin Provincial Park, Ontario (collector: Dr. M.
Wood).

At Meach Lake, Que., about 15 miles northwest of Ottawa,
Ontario, the author discovered an abandoned beaver pond. The water
level was very low and the beaver house was largely surrounded by
very wet soil. On June 26, collecting very close to this house pro-
duced one teneral specimen of Platypatrobus ; 40 specimens, including
several that were teneral, were found on July 6, and 10 more on
July 10. None were found on August 6, when the soil was less wet;

305

306

Psyche

[December

and none were found on any date around a nearby beaver house in a
drier situation. The beetles occurred beneath embedded logs and
under loose bark of old stumps, sometimes several inches beneath the
surface of the extremely wet soil which was exposed to the sun and
which contained much organic matter and debris. Numerous scarce
or local Carabidae that occur in the same general habitat are Elaphrus
olivaceus LeConte, Bembidion versutum LeConte, and Agonum
cincticolle (Say). We noted other beetles there that bore the mite
Protodinychus : Leptinellus validus (Horn), Agonum melanarium
(Dejean), and Quedius sp. (those specimens were found around the
beaver house only).

Literature Cited

Bell, R. T. and J. R. Bell

1964. Platypatrobus lacustris Darlington in Vermont. Proc. Ent. Soc.
Wash. 66 : 100.

Darlington, P. J.

1938. The American Patrobini. Ent. Amer. 18: 135-183.

Lindroth, C. H.

1962. The Male of Platypatrobus lacustris Darlington. Psyche 69:
7-10.

Robert, A.

1963. Une premiere station du Platypatrobus lacustris Darlington dans
le Quebec. Ann. Soc. Ent. Quebec 8: 58-59.

THE GENUS ZONOSEMATA , WITH NOTES ON THE
CYTOLOGY OF TWO SPECIES
( DIPTERA — TEPHRITIDAE) 1

By Guy L. Bush

Department of Biology, Harvard University

The genus Zonosemata (Trypetinae, Trypetini) was established
by Benjamin (1934) to include the type, Trypeta electa Say, and
Zonosema vittigera Coquillett. The latter species he considered as
possibly only a western race of electa. Both are recognized as distinct
in this revision and two new species from Mexico and one from
Jamaica are described.

Zonosemata is closely related to the Holarctic and Neotropical
Rhagoletis Loew (including Zonosema Loew), the monotypic
Nearctic Rhagoletoides Foote, the Neotropical Rhagoletotrypeta
Aczel, and the Palearctic and Indian Carpomyia Rondani2. It shows
particularly close affinities in habitus to certain Mexican and Neo-
tropical Rhagoletis , such as R. striatella, R. ferrugineus, R. ly co-
per sella, R. ochraspis, and others, some of which infest solanaceous
fruits as do at least two species of Zonosemata. Cytologically the
chromosomes of Zonosemata also bear some resemblance to those of
R. striatella. Furthermore, unlike other Rhagoletis species, the egg
of striatella is stalked and somewhat like those of Z. electa (Fig. 26)
and Z. vittigera. These similarities, coupled with the fact that four
of the five species are from Mexico and Jamaica, suggest that Zono-
semata probably originated in Central or possibly South America from
some common ancestor with Rhagoletis. Further field work in these
regions may therefore eventually clarify the relationships between
these two genera.

BIOLOGY

The biology of Z. vittigera and Z. electa has been discussed in
some detail by several authors (Peterson 1923, Benjamin 1934, Bur-
dette 1935, Cazier 1962, and Foott 1963). Both species normally
infest certain native species of horsenettle ( Solanum spp., Solanacece) .

’Published with the aid of a grant from the Committee on Evolutionary
Biology, Department of Biology, Harvard University. Manuscript received
by the editor, August 30, 1965.

The presence of minute ocellar bristles in Carpomyia is the only character
that effectively separates this genus from Rhagoletis. This character is of
doubtful importance and Rhagoletis may eventually fall into synonymy
with Carpomyia (for detailed discussion see Bush 1966).

307

308

Psyche

[December

Since 1921, however, electa has become a serious pest of peppers and
eggplants in the eastern United States. Recently it was found attack-
ing these plants as far north as southwestern Ontario (Foott 1963).
The hosts of the three new species are not known, but they may also
be expected to infest the fruits of solanaceous plants.

There is evidence that the pepper and horsenettle populations, at
least in Ontario, have become somewhat ecologically isolated from
each other. Foott (1963) noted that larvae from horsenettle emerge
later and are considerably smaller than those emerging from peppers.
He offered three possible explanations for this observation : micro-
climatic conditions were different for the two hosts; nutritive quali-
ties of the fruits varied ; earlier availability of pepper fruits allowed
expansion of the early emerging fly population. As pointed out by
Bush (1966), the third possibility would establish allochronically
isolated populations on different hosts and permit divergence to prog-
ress rapidly in the absence of gene flow from the parent population.
This could result in the formation of two distinct host races. A
careful study of these populations would be extremely interesting as
little is known about the formation of host races and species in
phytophagous insects.

CYTOLOGY

Mitotic configurations in the larval brain and adult testes were
examined in Z. electa and Z. vittigera. Tissue was pretreated in a
saturated aqueous solution of coumarin for 5-8 minutes, fixed and
stained in proprionic orcein for 5 minutes, and squashed in a drop
of stain following the method of Bush (1962). Pretreatment of the
tissue with coumarin was necessary to produce well flattened meta-
phase plates and to locate the kinetichores.

The karyotypes of both species are indistinguishable. In the
male there are five pairs of very small metakinetic autosomes
and a heteromorphie pair of extremely long acrokinetic and hetero-
chromatic sex chromosomes (Fig. 1). Following coumarin pretreat-
ment the chromatids of the X chromosome at metaphase are usually
joined only at their extreme ends forming an oblong ring (Fig. 1,
X). The chromatids of the shorter Y chromosome are usually
closely approximated over much of their length and form a figure

Explanation of Plate 22

Figs. 1-3. Mitosis in neuroblast cells of Zonosemata electa: (1) meta-
phase; (2) prophase; (3) anaphase.

Figs. 4-8. Right wings of Zonosemata spp.: (4) electa, N. J., U. S. A.;
(5) vittigera, Texas, U. S. A.; (6) cocoyoc, paratype, Morelos, Mex. ;

(7) mlnuta, type, Jamaica, W. I.; (8) vidrapennis, paratype, Mexico, Mex.

11 fPl

Psyche, 1965

Vol. 72, Plate 22

Bush — Zonosemata

3io

Psyche

[December

eight (Fig. i, Y). At prophase (Fig. 2) the X chromosomes are
greatly elongated and are associated with a nucleolar organizer.
At late anaphase (Fig. 3) the chromatids of the sex chromosomes are
still in contact on the metaphase plate while the fully disjoined
autosomes have reached the poles. In both sexes a large heterochro-
matic irregular-shaped body is always visible in the nucleus during
interphase. However, it is not clear whether this represents only one
or both sex chromosomes. Both the autosomes and the sex chromo-
somes have a fuzzy appearance during all stages of division, suggesting
that some genetic activity may be in progress even during cell division.
The polytene chromosomes of the salivary gland, gut, and Malpighian
tubules were checked and found unsuitable for detailed analysis.

Zonosemata Benjamin

Spilographa Loew (in part), 1873, Smith. Misc. Coll. 256: 244-, 336.
Zonosema Coquillett, 1899, Jour. N.Y. Ent. Soc. 7: 261.

Phorellia Hendel (in part), 1914, Abh. u. Ber. Zool. Mus. Dresden (1912)
14: 28.

Zonosemata Benjamin, 1934, U.S. Dept. Agri. Tech. Bull. 401: 17-18. [Type
of genus, Trypeta electa Say, location unknown, apparently lost3; type
locality: Indiana]. — Aczel, 1951, Acta zool. Lilloana 9: 214. — Aczel,
1954, Dusenia 5 : 152-153.

Further details on the nomenclatorial history of this genus may
be found in Benjamin (1934) and Stone (1951).

generic diagnosis. Zonosemata may be distinguished from other
members of the tribe Trypetini by the following combination of
characters :

1. Predominantly yellow with black maculations, a cream colored
notopleural stripe and brown crossbands on the wing.

2. Dorsocentrals located closer to a line drawn between postalars
than to anterior supraalars.

3. One pair of well developed outer scapular bristles, inner pair
absent or if present (rarely) then minute.

4. Sex chromosomes diffuse, much elongated, heterochromatic and
heteropycnotic ; autosomes very small and diffuse.

5. Surstyli blunt, short and broad; not greatly elongated beyond
prensisetae.

6. Usually four pairs of lower fronto-orbital bristles.

7. Dorsum lacking pollinose microtrichia.

8. Larvae with minute poorly sclerotized stomal guard hooks.

3Although no type could be located for electa, Say’s original description
is clear enough to leave no doubt as to the identification of the type species.
A neotype was not designated pending a more thorough search for the type.

1965]

Bush — Zonosemata

31

GENERIC DESCRIPTION. Head (Fig. 30) : subquadrate in profile;
about 1. 1 times wider than high; all regions, including palps and
mentum, light yellow to yellowish-orange; ocellar triangle light
brown to brownish-black; vertex measured across anterior margin
of ocellar plate, narrower than maximum width of eye; eye about
1.5 times higher than wide; frons convex in profile, prominent at
antennae, slightly wider at vertex than at antennae ; antennae .5 - .6
length of face; third segment with sharp awl-shaped tip, more than
twice length of second ; artista black or dark brown, pubescent,
grading to yellow at base; face minutely pubescent. Genae moderately
narrow, .23-.26 height of head. Postcranial region and head slightly
concave; foveae and carina moderately developed; epistome slightly
upturned ; postgenae bulging. Usually four pairs convergent black
fronto-orbitals, occasionally only three pairs or with one or two extra
bristles on one side or both; two pairs black reclinate divergent
upper fronto-orbitals, upper approximately two-thirds length of
lower; ocellars strongly proclinate, divergent, approximately same
length as upper fronto-orbitals. Comb translucent yellow. Outer
verticals black, about two-thirds length black inner verticals; post-
orbitals yellow, short; genal bristles variable, yellow or black when
present, occasionally undifferentiated; yellow gular bristle weak or
undifferentiated. Thorax: entirely yellow or with brown and black
maculations; halteres yellow. Dorsum and scutellum covered with
short decumbent yellow or black setae, lacking any trace of pollinose
microtrichia. Normally one pair outer scutellars, rarely one or two
minute scapulars present; dorsocentrals in line or slightly before line
drawn between posterior supraalars; two mesopleurals ; all other
bristles normal for Trypetinae. Legs: all segments yellow except
for black shading along posterior surface of metathoracic tibia.
Prothoracic femur with one posterior-ventral and two dorsal rows
of stout bristles. Mesothoracic tibia with row of three to five short
semierect bristles on posterior surface. Metathoracic tibia with row
of short semierect bristles along posterior-lateral surface. Wing:
pattern consisting of transverse brown crossbands; Rx setulose over
entire length including node; R4+5 setulose; usually one to three
setae dorsally at junction of R4+5 and R2 + 3 ; anal cell drawn out
to point along Cu2 + 2nd A. Abdomen: tergites covered with fine
long decumbent setae ; long black bristles along posterior and lateral
margins of tergites III-V in male and II I- VI in female (Fig. 24)
with well developed internal apodemes. Sternite VI about .55 length
of ovipositor sheath. Sternites of male (Fig. 23) without well
developed internal apodemes. Genitalia: male epandrium globose,

Psyche, 1965

Vol. 72, Plate 23

L

)

l

J

j v /

Bush — Zonosemata

1965]

Bush — Zonosemata

313

usually black, covered with long bristles; surstyli short with prensi-
setae (or spurs) subterminal (Fig. 25) ; genital ring circular, without
apodemes (Fig. 27); ejaculatory apodeme fan-shaped (Fig. 19).
Vesica of aedeagus (Fig. 21) tubular, without convolutions or appen-
dages. Female — ovipositor sheath approximately same length as
tergite IV in dorsal view. Ovipositor (Fig. 17) short, margins
slightly convex with minute trifurcate tip and two pairs of sub-
terminal minute setae (Fig. 18) ; two elongate cylindrical spermathe-
cae covered with irregularly shaped minute scale-like papillae (Fig.
22) ; two elongate accessory glands. Ventral receptacle as in Fig. 20,
without central spine as in Rhagoletis. Egg : micropyle end elongated
into stalk (Fig. 26).

KEY TO THE GENUS ZONOSEMATA

The characters most readily used to distinguish the various species
in this genus are found in the wing pattern and distribution of
black maculations on the body. Other than size, little variation was
noted in such structural characters as the male and female genitalia
and the number, color, and position of major chaetae.

1. A pair of black presutural spots on the dorsum (Fig. 36) and a

black spot or stripe on the sternopleuron (Fig. 35). Host:

Solatium eleagnifolium vittigera Coquillett

(SW U.S.A., N Cent. Mex.)

No distinct black spots before the transverse suture or on the
sternopleuron 2

2. Last three segments of the abdomen each with large, irregularly

shaped dark brown to black spots (Figs. 15-cf, 16-$) ; dorsum
with broad U-shaped maculation with unforked arms (Fig. 33).
Host unknown minuta n. sp.

(Jamaica)

Pair of well developed spots on last one or two segments (i.e.,
Figs. 9-10) ; black U-shaped maculation on dorsum reduced or

Explanation of Plate 23

Figs. 9-16. Abdomen of Zonosemata spp., dorsal view: (9) $ electa,

N. J., U. S. A.; (10) $ electa, N. J., U. S. A. ; (11) 2 vittigera, Texas,
U. S. A.; (12) $ vittigera, Texas, U. S. A. ; (13) $ cocoyoc, Morelos,

Mex.; (14) $ vidrapennis, type, Mexico, Mex.; (15) $ minuta, type,

Jamaica, W. I.; (16) 2 minuta, paratype, Jamaica, W. I.

Figs. 17-27. Zonosemata electa, N. J., U. S. A.: (17) ovipositor; (18)
detail of ovipositor tip; (19) ejaculatory apodeme; (20) ventral receptacle;
(21) aedeagus; (22) spermatheca ; (23) $ sternites of abdomen; (24) 2
sternites of abdomen; (25) posterior view of $ genitalia; (26) egg;
(27) lateral view of $ genitalia showing details of fultella and genital
ring.

3H

Psyche

[December

absent; if present then usually with forked arms (i.e., Figs.

37-38) 3

3. Apical wing band recurved along R4 + 5 but not reaching sub-

apical band, forming the letter P (Fig. 6) ; last two segments
of abdomen each with pair of black spots (Fig. 13). Host

unknown cocoyoc n. sp.

(Mexico)

Apical crossband forming the letter V; not recurved along
R4+5; a single pair of black spots on last segment 4

4. Crossbands of wing narrow (Fig. 8) ; microtrichia lacking be-

tween medial and subapical crossbands in cells Ri, R3, R5 and
1st M2; spot on last abdominal segment large, triangular

shaped (Fig. 14). Host unknown vidrapenms n. sp.

( Mexico)

Crossbands of wing broad (Fig. 4) ; microtrichia present in all
cells between medial and subapical crossbands; spot on last
abdominal segment small, round (Figs. 9-10). Host: Solarium

spp electa (Say)

(E. U.S.A., SE Canada)

Zonosemata electa (Say)

Trypcta electa Say, 1829-30, Jour. Acad. Nat. Sci. Phil. 6: 185-186. [Type
not examined, apparently lost4; type locality: Indiana]. — Osten Sacken,
1858, Smith. Misc. Coll., Art. 1, 3: 79. — Loew, 1862, Smith. Misc.
Coll., Art. 1, 6: 58, 71. Stone, 1951, Proc. Ent. Soc. Wash. 53: 45-46.
T ephritis flavonotata Macquart, 1855, Dipt. Exot. Suppl. 5: 125. [Type not
examined0; type locality: De l’Amerique septintrionale. Baltimore (in
collection of J. E. Collin, Newmarket, England)].

Zonosema electa Coquillett, 1899, Jour. N. Y. Ent. Soc. 7: 261. — Phillips,

1923, Jour. N. Y. Ent. Soc. 31: 127-128, fig. 8.

Spilographa electa Loew, 1873, Smith. Misc. Coll. 256: 244, 336. — Snow,

1903, Kans. Univ. Quart. 11: 161. — Williston, 1905, Kans. Univ.

Quart. 13: 307. — Peterson, 1923, N. J. Expt. Sta. Bull. 373: 1-23,
pi. 1-3

Zonosemata electa Benjamin, 1934, U. S. Dept. Agri. Tech. Bull. 401: 19-20,
fig. 15. — Burdette, 1935, N. J. Expt. Sta. Bull. 585: 3-24, pi. 1-3.

— Phillips, 1946, Mem. Amer. Ent. Soc. 12: 96-98, 129, figs. 44, 89,
93, 191. — Anonymous, 1959, U. S. Dept. Agri., Agri. Res. Serv.,

Copp. Econ. Insect Rept. 9: 721-722. — Foote, 1960, Proc. Biol. Soc.
Wash. 73 : 114-116.

diagnosis. Excellent detailed descriptions of both larval and adult
stages have been presented by Peterson (1923), Benjamin (1934),

4See foot-note, p. 310.

’Macquart’s type has been discussed by Stone (1951) who has established
its identity with certainty with the help of Mr. J. E. Collin.

1965]

Bush — Zonosemata

315

and Phillips (1946) and will not be repeated here. Body and wing
measurements are presented in Table 1. This predominantly yellow
species can be distinguished by the following combination of charac-
ters: (1) large body size (Table 1) ; (2) absence of black markings
before the transverse suture on the dorsum and on the sternopleuron
(Figs. 29, 37) ; (3) yellow postscutellum ; (4) presence of only a
single pair of spots on the last segment of the preabdomen (Figs.
9-10); (5) crossbands of wing broad with medial and subapical
bands joined along the posterior margin (Fig. 4) ; (6) microtrichia
present between medial and subapical bands in cells Ra, R3, R5, and
1st M2. Other morphological features of electa are illustrated as
follows: genitalia (Figs. 17-22, 25, 27); sternites (Figs. 23-24);
head (Fig. 30) .

variation. There is a great deal of individual variation in this
species, particularly in the extent of black shading on the thorax and
hind femora. Figs. 29 and 37 therefore represent only the most
frequently encountered pattern in well-aged specimens whose color
has had time to develop fully. The U-shaped pattern of the dorsum,
for example, is often reduced to a small region along the sulcus
between the dorsum and scutellum, or is entirely absent. There is
also a tendency for individuals from Florida to be somewhat smaller
than those from either the North or Texas. A few of the Florida
specimens were as small as the average sized vittigera. Although
most specimens had four pairs of lower fronto-orbitals, numbers
ranging from three to as many as seven on one or both sides of the
frons were not uncommon.

hosts. Solarium carolinense Linn., S. aculeatissimum Jacq., S.
melongena Linn., Capsicum annum Linn., and infrequently Lyco-
persicum esculentum Mill. (Peterson 1932, Benjamin 1934).
parasite. O plus sanguineus (Ashmead) (Cazier 1962).
distribution ( Map i). This species ranges from central Florida
north to Massachusetts, southern Ontario, southern Illinois, eastern
Kansas, and northeastern Texas. The range of electa apparently
overlaps that of vittigera in the transition zone of eastern Texas as
far south as Brownsville (see also Foote i960).

Zonosemata vittigera (Coquillett)

Zonosema vittigera Coquillett, 1899, Jour. N. Y. Ent. Soc. 7: 261. [Lectotype
by present designation; locality: Eagle Pass, Texas; (J. Cram coll.)
(USNM, type no. 4398)].

Spilographa vittigera Aldrich, 1905, Smith. Misc. Coll. 46: 604.

Zonosemata vittiigera Benjamin, 1934, U. S. Dept. Agri. Tech. Bull. 401:
18, fig. 15. — Phillips, 1946, Mem. Amer. Ent. Soc. 12: 100-101, figs.

Psyche, 1965

Vol. 72, Plate 24

Bu:H — ZONOSEMATA

1965]

Bush — Zonosemata

317

43, 88, 98, 190. — Aczel, 1954, Dusenia 5: 152-157, figs. 19-27. — Foote,
I960, Proc. Biol. Soc. Wash. 73: 114-116.

Zonosemata variegata6 Aczel, 1954, Dusenia 5: 162, 164 figs. 19-27; estampa
II. C (nomen nudum).

diagnosis. Detailed descriptions of this species have been pre-
sented by Aczel (1954) (adult), and Phillips (1946) (larva).
Body and wing measurements are presented in Table 1. Z. vittigera
can be distinguished readily from all other members of this genus
by the presence of a pair of black dots before the transverse suture
(Fig. 36), and by the presence of a black stripe or distinct spot on
the sternopleuron (Fig. 35). The abdomen has a pair of baso-lateral
spots on the last segment in both sexes (Figs. 11, 12). Occasional
individuals may also have small brown spots on the penultimate
segments. The wing pattern of vittigera (Fig. 5) is very similar
to that of minuta (Fig. 7), but usually can be distinguished from
that of electa (Fig. 4) by the fact that the medial and subapical
bands are joined in the latter species.

variation. The intensity of the black pigmentation of the pre-
sutural spots varied a great deal in the specimens available for study,
particularly in those from Texas. Many of these specimens have the
presutural and sternopleural black spots generally diluted and the
postscutellum often predominantly yellow. Hybridization beween
electa and vittigera could account for the reduction in the degree
of black coloration in the latter species. However, this seems unlikely
as there is no indication of introgression, such as a tendency toward
reduction in size or an increase in the intensity or distribution of

‘'Although Aczel used the name vittigera throughout the text of his 1954
paper, he labelled the accompanying illustrations variegata. I have also
examined specimens bearing the name variegata on a type label in Aczel’s
handwriting in the USNM collection. Dr. Alan Stone of the USNM found
Aczel’s manuscript species variegata to be synonymous with Coquillett’s
vittigera after comparing the types at Aczel’s request (from correspondence
between Stone and Aczel, 1952, USNM). Aczel therefore apparently made
the necessary change in the text but failed to do so for the plate before the
paper was published.

Explanation of Plate 24

Figs. 28-29, 31-32, 35. Thorax, lateral view, Zonosemata spp.: (28) $

cocoyoc, paratype, Morelos, Mex. ; (29) $ electa , N. J., U. S. A.; (31)

$ minuta, type, Jamaica, W. I.; (32) $ vidrapennis, paratype, Mexico,

Mex.; (35) $ vittigera, Texas, U. S. A.

Figs. 33-34, 36-38. Thorax, dorsal view, Zonosemata spp.: (33) $ min-
uta, type, Jamaica W. I.; (34) $ vidrapennis, paratype, Mexico, Mex.;

(36) $ vittigera, Texas, U. S. A.; (37) $ electa, N. J., U. S. A.; (38) $
cocoyoc, paratype, Morelos, Mex.

Fig. 30. Head, electa $, N. J., U. S. A.

3 1 8

Psyche

[December

Species

n

HL

HW

HH

TL

WW

WL

OL

electa

cf

20

l-47±

•03

2.3 1 ± -04

2.lOrh .04

3-40± .04

2.562b .05

4-99~ .09

1. 16-

I.64

2.00—2.56

1.88 — 2.44

3.00-3.96

2.24—2.92

4.28 — 5.68

?

20

i-Si=±=

.02

2.37± .03

2.1 1± .03

3-47 ± -05

2.692b .05

5-i3± .07

1.67+: .03

1.28-

1.72

2.00—2.60

1.80—2.32

2.92-3.84

2.32-3.12

4.56-5.80

1.44—2.16

vittigera

c f

20

i.i9±

•03

i.84± .03

i.67± .03

2.492b .06

1.862b .03

3-83± .07

.96-

1.32

1.56—2.04

1.40— 1.92

2.04—2.84

1.60 — 2.08

3.20-4.36

?

20

I.20±

•03

i.90± .04

i-70± .05

2.53± .07

i.95± -05

4.00 zfc .08

1.122b .02

.92-

1.36

1.52— 2.10

1.24—2.00

1.84—2.96

1.48 — 2.24

3.00-4.52

1.00 — 1.28

vidrapennis
<$ (Holotype)

1

.1.12

1.88

1.60

2.80

2.32

4.68

minuta

cT (Holotype)

!

I.04

1.76

1.60

2.40

1.60

3-48

9

I

I.04

1.76

1.60

2.24

1.64

3.60

cocoyoc

9 (Holotype)

I

1.28

2.12

x.90

3-oo

2.52

5.08

1.64

9 (Paratype)

I

I.36

2.24

2.04

3.16

2.66

5.16

Table I. Body and wing measurements of Zonose/nata. Figures represent mean, standard error, and
range in mm. HL — head length; HW — head width; HH — head height; TL — -thorax length;
WL — wing length ; WW • — wing width ; OL — ovipositor length.

black maculation, in the sympatric specimens of electa examined
from Texas. Some specimens of vittigera were reared and appear
to be teneral. This could account for their lack of intense black
maculations.

Four specimens which appear to be vittigera have been examined,
each bearing the label ‘N.J., Sept. 7-8/ a locality well outside the
normal range of this species. Some of these specimens have spots on
both the penultimate and last segments. This has been noted in only
a few other representatives of this species from Mexico. In all
other respects, such as size, wing pattern, and body coloration, they
resemble vittigera. Without host data or additional information con-
cerning the collection of these specimens, it is impossible to decide
if they represent a distinct species. They are therefore tentatively
being treated as an adventive of vittigera to New Jersey until further
information becomes available.

host. Solanum eleagnifolium Cav. (Aczel 1954, Foote i960,
Cazier 1962).

parasite. Opius sanguineus (Ashmead) (Cazier 1962).

distribution (Map i). This species ranges from just north of
Mexico, D.F. in the central highlands of Mexico and the southwest
tip of Oklahoma to northeastern Texas. Its host plant now occurs
as on advent in Indiana, Illinois, Ohio, and Florida, but vittigera
has never been recorded from these areas. A single specimen from
San Francisco, California has been examined, but was collected in
a Santa Fe Pullman and therefore is not recorded on Map 1.

1965]

Bush • — Zonosemata

319

Zonosemata cocoyoc new species
types. Holotype Cocoyoc, MORELOS, Mexico, 8-VIII-56;
Trampa Cabo; (O. Hernandez) (USNM, type no. 68 1 1 1 ) . Para-
type 9> same data as holotype (USNM).

diagnosis. Zonosemata cocoyoc can be easily distinguished from
other members of the genus by the wing pattern in which the apical
and subapical bands form a letter P. The apical band extends along
vein R4 + 5, but does not reach the subapical band (Fig. 6). The
reduced black coloration of the thorax (Fig. 6). The reduced black
coloration of the thorax (Figs. 28, 38) and the presence of paired
spots on tergites V and VI of the female (Fig. 13) will also help
to identify this species. The male is unknown.

description. Body and wing measurements in Table 1. Head
indistinguishable in shape from electa ; all regions, including mentum
and palps, golden yellow to light yellow except black ocellar triangle.
Genal and gular bristles yellow, undifferentiated from other setae;
all other bristles normal and black. Thorax (Figs. 28, 38) : base
color golden yellow; notopleural and sternopleural stripes and light
region of dorsum and scutellum ivory. Black shading reduced on
dorsum and pleural regions as figured. Postscutellum mostly golden
yellow with some black shading along medial line and dorsal margin.
Legs: normal color and complement of bristles for genus. JVing
(Fig. 6) : medial and basal bands joined by a faint infuscated area
in cell Rj. Apical band joined to subapical band on costa and re-
curved along vein R4 + 5, but not reaching subapical band. Micro-
trichia present in all cells between medial and subapical crossbands;
absent in cells between medial and basal crossbands from costa to
M3 + Cu3. Abdomen (Fig. 13): base color yellow, covered with
long black decumbent setae. Tergite V with small pair and tergite
VI with large pair of black spots. Genitalia: ovipositor sheath

golden yellow, darker than rest of abdomen. Ovipositor tip similar
to electa.

host. Unknown; captured in a McPhail fermenting lure glass
trap.

distribution (Map i). Known only from the type locality.

Zonosemata minuta new species
types. Holotype ty, Montego Bay [Jamaica], 2-VII-57, (J. W.
Boyes) (USNM, type no. 68112). Paratype $, same data as type
(USNM).

diagnosis. This small species can be easily identified by the
distinctive U-shaped black pattern on the dorsum (Fig. 33), the

320

Psyche

[December

dark border around the humeral callus, and the presence of black
spotting on the last three segments of the abdomen in both male and
female (Figs. 15-16).

description cf- Body and wing measurements in Table 1 . Plead:
indistinguishable in shape from that of electa (Fig. 30) ; all regions
including mentum and palps golden yellow except black ocellar
triangle; post ocellars yellow or black; gular undifferentiated, genal
and all other bristles black. Thorax (Figs. 31, 33) : notopleural
and sternopleural stripes and light areas of dorsum and scutellum
yellow; dorsum and base of scutellum with broad dark brown to
black U-shaped pattern ; humeral callus in dorsal view emarginated
with black; pleural regions with black stripe extending from lower
half of mesopleuron to base of postscutellum ; dark brown to black
shading also present on lower half of sternopleuron. Postscutellum
brownish-black. Dorsum and scutellum covered with short light
yellow to brown decumbent setae. Legs: normal color and comple-
ment of bristles for genus. Wing (Fig. 7) : crossbands broad; basal

1965]

Bush — Zonosemata

321

band joined to medial band by faint infuscated area in cell (best
seen in transmitted light) ; microtrichia present in all cells of wing.
Abdomen (Fig. 15) : golden yellow to light yellowish-brown covered
with long black decumbent setae; tergite III with single pair of
irregular shaped black spots; tergite IV with pair of elongated
irregular bar-shaped markings; tergite V with large irregular tri-
angular-shaped spots. Genitalia: exposed portion on type indistin-
guishable from electa.

Description of 9: differs from male only in having more extensive
black shading on thorax, legs, and abdomen, particularly on dorsum
and metathoracic tibia. Genitalia: ovipositor tip exposed in para-
type, similar to electa.

host. Unknown.

distribution (Map 1). Jamaica, West Indies. Known only
from the type locality.

Zonosemata vidrapennis new species

types. Holotype cT , Oaxaca, OAXACA, Mexico, 16-IX-1933
(C.C. Plummer) (USNM, type no. 68113). Paratype <S , entre
Villa Guerrero y Tenanzingo, MEXICO, Mexico, 13-VIII-1961 ,
sweeping (F. Pacheco), slide no. no, G.L. Bush (USNM) (speci-
men badly damaged) .

diagnosis. Differs from all other Zonosemata in having narrow
wing bands and lacks the faint infuscated area joining the basal and
medial crossbands in cell along vein R (Fig. 8). The short
decumbent setae on dorsum are entirely black and the posterior
margin of the dorsum and anterior margin of the scutellum show
only a trace of the black horseshoe-shaped pattern (Fig. 34) normally
found in other members of this genus. The pleural regions are almost
entirely yellow except for black shading on the sternopleuron (Fig.
32). A pair of spots on the last abdominal segment are large and
triangular-shaped (Fig. 14). This species is known only from two
male specimens.

description cf • Body and wing measurements of type in Table 1.
Head: indistinguishable in shape from that of electa ; yellow to

brownish-yellow except black ocellar triangle. Genal and gular
bristles and setae on postgenae yellow; all other bristles black.
Thorax (Figs. 32, 34) : light brown to yellow; notopleural stripe,
sternopleural stripe and light regions of dorsum, scutellum, and
postscutellum golden yellow; small, ill-defined U-shaped black pat-
tern on anterior margin of scutellum and posterior margin of dorsum
in paratype, but almost absent in type. Dorsum and scutellum

322

Psyche

[December

covered with short black decumbent setae. Black shading on ptero-
pleuron becoming diffuse on mesopleuron. Legs: normal complement
of bristles; entirely yellow. Wing (Fig. 8): glassy in appearance;
crossbands narrow; basal band not joined to medial band in cell Rx,
and latter not joined to subapical band along posterior margin of
wing in cell Cui. Microtrichia present in all cells between subapical
crossband and apex of wing; absent from regions between basal and
subapical crossbands except along posterior margins of wing and in
cell R.! between short intercalary band and subapical band. Basal
two-thirds of R4 + 5 setulose. Abdomen (Fig. 24): base color

brownish-yellow, colored with short black decumbent setae. Tergite
V with pair of distinct triangular-shaped black spots on baso-lateral
surface. Genitalia: indistinguishable from that of electa.

host. Unknown.

distribution (Map i). Known only from the two localities
listed above.

Acknowledgments

I would like to thank Mr. J. P. Reed of the New Jersey Agri-
cultural Experiment Station, Rutgers University, and Mr. E. W.
Johnson, U.S. Department of Agriculture, Brownsville, Texas for
their assistance in locating suitable material for cytological studies.

Literature Cited

Aczel, M.

1954. Generos y especies da la tribus Trypetini 4.

El genero Rhagoletotrypeta y nuevas especies de Tomoplagia y
de Zonosemata (Diptera, Tephritidae) . Dusenia 5: 152-157.

Benjamin, F. H.

1934. Descriptions of some native trypetid flies with notes on their
habits. U. S. Dept. Agri. Tech. Bull. 401: 1-96, figs. 1-44.

Burdette, R. C.

1935. The biology and control of the pepper maggot, Zonosemata electa
Say. Trypetidae. N. J. Agri. Expt. Sta. Bull. 585: 3-31.

Bush, G. L.

1962. The cytotaxonomy of the larvae of some Mexican fruit flies in
the genus Anastrepha (Tephritidae, Diptera). Psyche 69: 87-101.

1966. The taxonomy, cytology, and evolution of the genus Rhagoletis in
North America (Diptera, Tephritidae). Bull. Mus. Comp. Zool.,
in press.

Cazier, M. A.

1962. Notes on the bionomics of Zonosemata vittigera (Coquillett) , a
fruit fly on Solanum (Diptera: Tephritidae). Pan-Pacific Ent. 38:
181-186.

1965]

Bush — Zonosemata

3 23

Foote, R. H.

1960. Notes on some North American Tephritidae, with descriptions of
two genera and two new species (Diptera). Proc. Biol. Soc.
Wash. 73 : 107-118.

Foott, W. H.

1963. The biology and control of the pepper maggot, Zonosemata electa
(Say) (Diptera: Trypetidae) in southwestern Ontario. Proc. Ent.
Soc. Ont. 93 : 75-81.

Peterson, A.

1923. The pepper maggot, a new pest of peppers and eggplants. Spilo-
grapha electa Say. (Trypetidae). N. J. Agri. Expt. Sta. Bull. 373:
4-23.

Phillips, V. T.

1946. The biology and identification of trypetid larvae (Diptera: Try-
petidae). Mem. Amer. Ent. Soc. 12: 161 pp.

Stone, A.

1951. The Rhagoletis of roses (Diptera, Tephritidae). Proc. Ent. Soc.
Wash. 53 : 45-48.

PSYCHE

INDEX TO VOL. 72, 1965

INDEX TO AUTHORS

Brown, JV . L., Jr. Contributions to a Reclassification of the Formicidae. IV.
Tribe Typhlomyrmecini (Hymenoptera) . 65

Bush, G. L. The Genus Zonosemata, with Notes on the Cytology of Twc
Species (Diptera-Tephritidae) . 307

Carpenter , F. M. Studies on North American Carboniferous Insects. 4.
The Genera M etropator , Eubleptus , Hapaloptera and Hadentomum. 175

Chapin, E. A. New Species of Chilocorini (Coleoptera: Coccinellidae). 148

Chickering, A. M. Five New Species of the Genus Tmarus (Araneae,
Thomisidae) from the West Indies. 229

Creighton, JJr. S. The Habits and Distribution of Macromischa subditiva
Wheeler (Hymenoptera: Formicidae). 282

Creighton, W. S., and IV. L. Nutting. The Habits and Distribution of
Cryptocerus rohweri Wheeler (Hymenoptera: Formicidae). 59

Ettershank, G. A New Species of Megalomyrmex from the Chilean Andes.
(Formicidae, Hymenoptera). 55

Evans, H. E. Further Studies on Epyrini (Hymenoptera, Bethylidae). 265

Evans, H. E. Simultaneous Care of More Than One Nest by Ammophila
azteca Cameron (Hymenoptera, Sphecidae). 8

Fairchild, G. B. Notes on Neotropical Tabanidae VI. A New Species of
Lepiselaga Macq. with Remarks on Related Genera. 210

Flint, 0. S., Jr. The Genus N eohermes (Megaloptera : Corydalidae) . 255

Gardner. B. T. Observations on Three Species of Phidippus Jumping
Spiders (Araneae: Salticidae). 133

Goulet, H. The Habitat of Platypatrobus Darlington (Coleoptera: Cara-
bidae). 304

Haskins, C. P. and R. M. Whelden. “Queenlessness,” Worker Sibship, and
Colony Versus Population Structure in the Formicid Genus Rhytido-
ponera. 87

Levi, H. IV. Techniques for the Study of Spider Genitalia. 152

Masner, L. The Types of Proctotrupoidea (Hymenoptera) in the Charles
T. Brues Collection at the Museum of Comparative Zoology. 295

325

Nutting , JV. L. Observations on the Nesting Site and Biology of the Arizona
Dampwood Termite, Zootcrmopsis laticeps (Banks) (Hodotermitidae).
113

Parsons, C. T. A Key to the North American Statira (Coleoptera: La-
griidae). 241

Porter, C. C. Picrocryptoides : A New Genus of the Tribe Mesostenini

from Southern South America (Hymenoptera, Ichneumonidae) . 167

Rcdner , J. H. and C. D. Dondale. A New Species of the Spider Genus
Xysticus (Araneae: Thomisidae) from Arizona. 291

Reichardt, H. Scolytoidea (Coleoptera) 5: Notes on Neotropical Platy-
podidae, Mainly from Central America. 159

Reis kind, J. A Revision of the Ant Tribe Cardiocondylini (Hymenoptera,
Formicidae). 79

Reiskind, J. Self-Burying Behavior in the Genus Sicarius (Araneae, Si-
cariidae). 218

Reiskind, J. The Taxonomic Problem of Sexual Dimorphism in Spiders and
a Synonymy in Myrmecotypus (Araneae, Clubionidae) . 279

Scherba, G. Observations on Microtus Nesting in Ant Mounds. 127

Taylor , R. JV. A Second African Species of the Dacetine Ant Genus
Codiomyrmex. 225

Taylor, R. JV. Notes on the Indo-Australian Ants of Genus Simpone Forel
(Hymenoptera, Formicidae). 287

Taylor, R. IV. The Australian Ants of the Genus Pristomyrmex , with a
Case of Apparent Character Displacement. 35

JVerner, F. G. A Redefinition of lschyropalpus , and Six New Species
(Coleoptera: Anthicidae). 191

Wheeler, G. C. and J. Wheeler. The Ant Larvae of the Subfamily Laptanil-
linae (Hymenoptera, Formicidae). 24

Wilson, E. O. Trail Sharing in Ants. 2

326

INDEX TO SUBJECTS

All new genera, new species and new names are printed in capital type.

A key to the North American Statira
(Coleoptera: Lagriidae), 241
A new species of Megalomyrmex
from the Chilean Andes (Formi-
cidae: Hymenoptera), 55
A new species of the spider genus
Xysticus (Araneae: Thomisidae)
from Arizona. 291
A redefinition of Ischyropalpus, and
six new species (Coleoptera: An-
thicidae), 191

A revision of the ant tribe Cardio-
condylini (Hymenoptera, Formi-
cidae), 79

A second African species of the
Dacetine ant genus Codiomyrmex,
225

Ammophila azteca, 8
Anthicidae, 191

Ants, 24, 35, 79, 127, 225, 282, 287,
Araneae, 133, 152, 218, 229, 279, 291
Azteca chartifex, 3
Bakeriella brasiliana, 270
Bakeriella depressa, 273
Bakeriella inca, 272
Bakeriella inconspicua, 270
Bakeriella rossi, 27 3
Bakeriella subcarinata, 271
Bethylidae, 265
Calyozina mexicana, 273
Camponotus beebei, 3
Carabidae, 305
Carboniferous insects, 175
Clubionidae, 279
Coccinellidae, 148
Codiomyrmex tetragnathus, 225
Coleoptera, 148, 159, 191, 241, 305
Contributions to a reclassification of
the Formicidae. IV. Tribe Typhlo-
myrmecini (Hymenoptera), 65
Corydalidae, 255
Cryptocerus rohiveri, 59
Curinus colombianus, 150
Cytology of Zonosemata, 307
Diptera, 307
Epyris crassipes, 276
Epyris, cubanus, 275

Epyris depressigaster, 275
Eubleptus danielsi, 180
Five new species of the genus
Tmarus (Araneae, Thomisidae)
from the West Indies, 229
Formicidae, 24, 59, 65, 79, 282, 287
Further studies on Epyrini (Hymen-
optera, Bethylidae). 265
Hadentomum americanum, 186
Hapaloptera gracilis, 183
Hodotermitidae, 113
Hymenoptera, 8, 24, 59, 65, 79, 265,
282, 287, 295

Ischyropalpus alvarengai, 205
Ischyropalpus decoratus, 204
Ischyropalpus eryngii, 200
Ischyropalpus freyi, 203
Ischyropalpus puteifer, 206
Ischyropalpus tibialis, 197
Jumping Spiders, 133
Lagriidae, 241
Lepiselaga colombiana, 215
Leptanilla revelierei sardoa, 27
Leptanilla scwanif 30
Leptome sites escheri, 31
Macromischa subditwa, 282
Megalomyrmex bicolor, 55
Megaloptera, 255
M etropator pusillus, 177
Microtus , 127
Myrmecotypus , 279
N eohermes angusticollis, 259
N eohermes calif ornicus, 257
N eohermes concolor, 262
N eohermes filicornis, 256
N eohermes matheri, 260
Neotropical Epyrini, 265
Neotropical Platypodidae, 159
Neotropical Tabanidae, 210
New species of Chilocorini (Cole-
optera: Coccinellidae), 148
Notes on Neotropical Tabanidae VI.
A new species of Lepiselaga
Macq. with remarks on related
genera, 210

Notes on the Indo-Australian ants
of genus Simpone Forel (Hymen-

327

optera, Formicidae), 287
Observations on Microtus nesting in
ant mounds, 127

Observations on the nesting site and
biology of the Arizona dampwood
termite, Zootermopsis laticeps
(Banks) (Hodotermitidae) , 113
Observations on three species of
Phidippus jumping spiders (Ar-
aneae: Salticidae), 133
Phaenochilus renipunctus, 151
Phidippus, 133

Picrocryptoides : A new genus of the
tribe Mesostenini from southern
South America (Hymenoptera,
Ichneumonidae) , 167
PICROCRYPTOIDES HESPERIOS, 173
PICROCRYPTOIDES WILLINKI, 171
Platypatrohus , 305
Platypus pint, 159
Platypus pulchellus , 160
Platypus DARLINGTONI, 162
Pristoviyrmex fovelatus, 38
Pristomyrmex quadridentatus, 42
Pristomyrmex thoracicus, 41
Pristomyrmcx wheeleri, 48
Proctotrupoidea, 295
Prosopidris papuna, 81
Prosopidris sima, 84
“Queenlessness,” worker sibship, and
colony versus population structure
in the Formicid genus Rhytido-
bonera , 87

Rhabdcpyris apache, 269
Rhabd epyris fortunatus , 269
Rhabdepyris gracilis, 268
Rhabdcpyris hirticulus, 269
Rhabdepyris muesebecki, 265
Rhabdcpyris multilineatus, 266
Rhabdcpyris nigripilosus , 268
Rhabdepyris nigriscapus , 266
Rhabdepyris virescens, 270
Rhytidoponera inornata, 97
Rhytidoponera metallica, 94
Rhytidoponera purpurea, 92
Rhytidoponera tasmaniensis, 96
Rhytidoponera victoriae, 98
Rhytidoponera violacea , 99
Salticidae, 133
Scolytoidea, 159

Scolytoidea (Coleoptera) 5: Notes

on neotropical Platypodidae, main-
ly from Central America, 159
Self-burying behavior in the genus
Sicarius (Araneae, Sicariidae),
218

Sicariidae, 218
Sicarius, 218
Simpone bakeri, 288
Simpone chapmani, 287
Simultaneous care of more than one
nest by Ammophila ezteca Cam-
eron (Hymenoptera, Sphecidae), 8
Sphecidae, 8
Spider Genitalia, 152
Statira basalis, 252
Statira colorata, 245
Statira croceicollis, 249
Statira defecta, 248
Statira dolera, 249
Statira gagatina gagatina, 252
Statira gagati?ia resplendens , 253
Statira hirsuta, 246
Statira huachucae , 247
Statira liebecki, 248
Statira nigromaculata, 246
Statira opacicollis, 247
Statira pluripunctata, 245
Statira pulchella, 246
Statira robusta, 248
Statira subnitida, 244
Studies on North American Carboni-
ferous insects. 4. The genera
Metropator, Eubleptus , Habalop-
tera and Hadentomum, 175
Techniques for the study of spider
genitalia, 152
Termites, 113
Tephritidae, 307
T esserocerus belti, 163
Tesserocerus deivalquei, 163
T esserocerus ericeus, 163
Tesserocranulus nevermanni, 165
The ant larvae of the subfamily
Laptanillinae (Hymenoptera, For-
micidae), 24

The Australian ants of the genus
Pristomyrmex, with a case of ap-
parent character displacement, 35
The genus N eohermcs (Megaloptera :
Corydalidae) , 255

The genus Zonosematz , with notes

328

on the cytology of two species
(Diptera-Tephritidae), 307
The habitat of Platypatrobus Dar-
lington (Coleoptera: Carabidae),
304

The habits and distribution of
Cryptocerus rohnveri Wheeler
(Hymenoptera : Formicidae), 59
The habits and distribution of Mac-
romischa subditlva Wheeler (Hy-
menoptera. Formicidae), 282
The taxonomic problem of sexual
dimorphism in spiders and a syn-
onymy in Myrmecotypus (Ar-
aneae, Clubionidae) , 279
The types of Proctotrupoidea (Hy-
menoptera) in the Charles T.
Brues Collection at the Museum
of Comparative Zoology, 295
Thomisidae, 291
Tmarus craneae, 230

T marus farri, 232
Tmarus insuetus, 234
Tmarus menotus, 236
Tmarus vertumus, 238
Trail sharing in ants, 2
Typhlomyrmex clavicornis, 69
Typhlomyrmex foreli, 76
Typhlomyrmex major, 71
Typhomyrmex prolatus, 72
Typhlomyrmex pusillus, 73
Typhlomyrmex rogenhoferi , 74
Xysticus humilis, 291
Zagreus costalimae, 148
Zagreus tetraspilus, 149
Zonosemata cocoyoc, 319
Zonosemata electa , 314
Zonosemata minuta, 320
Zootermopsis laticeps 113
Zonosemata vidrapennis, 321
Zonosemata vittigera , 315

329

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 B-455,
Biological Laboratories, Divinity Ave., Cambridge. Entomologists
visiting the vicinity are cordially invited to attend.

The illustration on the front cover of this issue of Psyche is a
reproduction of an unpublished drawing by W. M. Wheeler of
Cephalotes atratus (Linnaeus). [Courtesy of Miss A. E. Wheeler].

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Classification of Insects, by C. T. Brues, A. L. Melander and
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/I3